Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
Edited by
Richard E. Litz
Tropical Research and Education Center
and
The paper used for the text pages in this book is FSC certified. The FSC (Forest Stewardship
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Contents
Contributors vii
Preface ix
Acknowledgements xi
5. Reproductive Physiology 97
T.L. Davenport
6. Ecophysiology 170
B. Schaffer, L. Urban, P. Lu and A.W. Whiley
v
vi Contents
16. World Mango Trade and the Economics of Mango Production 606
E.A. Evans and O.J. Mendoza
Index 671
vii
viii Contributors
I. Kobiler, Department of Technology and Storage of Agricultural Products, Agricultural Research Organiza-
tion (ARO), The Volcani Center, PO Box 6, Bet Dagan 50250, Israel. E-mail: kobileri@volcani.agri.gov.il
U. Lavi, Department of Horticulture, Agricultural Research Organization (ARO), The Volcani Center,
PO Box 6, Bet Dagan 50250, Israel. E-mail: ulavi@agri.gov.il
T.-S. Lin, 111 Room, No. 4 Hall, Department of Horticulture, National Taiwan University, No. 1 Sec.
4 Roosevelt Road, 106 Taipei, Taiwan. E-mail: tslin@cc.ntu.edu.tw
R.E. Litz, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Home-
stead, FL 33031-3314, USA. E-mail: relitz@ufl.edu
P. Lu, EWL Sciences, PO Box 39443, Winnellie, NT 0821, Australia. E-mail: ping.lu@ewlsciences.
com.au
I. Maguire, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street,
Homestead, FL 33031-3314, USA. E-mail: imaguire@ufl.edu
O.J. Mendoza, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street,
Homestead, FL 33031-3314, USA.
I. Miyara, Department of Technology and Storage of Agricultural Products, Agricultural Research
Organization (ARO), The Volcani Center, PO Box 6, Bet Dagan 50250, Israel.
S.K. Mukherjee (deceased), Department of Agriculture, Calcutta University, 35 Ballygunge Circu-
lar Road, Calcutta 700 019, India.
M.T. Ombico, Fruit and Vegetable Laboratory, Food Science Cluster, College of Agriculture, Univer-
sity of the Philippines Los Baños, Laguna, 4031, Philippines.
J.E. Peña, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Home-
stead, FL 33031-3314, USA. E-mail: jepe@ifas.ufl.edu
R.C. Ploetz, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street,
Homestead, FL 33031-3314, USA. E-mail: rcp@ifas.ufl.edu
D. Prusky, Department of Technology and Storage of Agricultural Products, Agricultural Research Organiza-
tion (ARO), The Volcani Center, PO Box 6, Bet Dagan 50250, Israel. E-mail: dovprusk@volcani.agri.gov.il
A.C. de Queiroz Pinto, Private Consultant Tropical Fruits, SHCGN 706 Bloco P Casa 13, 70740-
716, Brasilia-DF, Brazil. E-mail: alberto.pinto@embrapa.br
S. Ram (deceased), Department of Horticulture, GB Pant University of Agriculture and Technology,
Pantnagar 263 145, India.
L.C. Raymundo, Fruit and Vegetable Laboratory, Food Science Cluster, College of Agriculture, Uni-
versity of the Philippines Los Baños, Laguna, 4031, Philippines. E-mail: lcr161940@yahoo.com
S. Salazar-García, Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Campo
Experimental Santiago Ixcuintla, Km. 6 Carret. Intnal. Tepic-Mazatlán, Apartado Postal 100, San-
tiago Ixcuintla, Nayarit 63300, Mexico. E-mail: samuelsalazar@prodigy.net.mx
B. Schaffer, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street,
Homestead, FL 33031-3314, USA. E-mail: bas@ifas.ufl.edu
R.J. Schnell, United States Department of Agriculture (USDA) Agriculture Research Service (ARS),
Subtropical Horticultural Research Unit/National Germplasm Repository, 13601 Old Cutler Road,
Miami, FL 33158, USA. E-mail: rschnell@ars-grin.gov
Z.-H. Shü, Department of Biological Science and Technology, Meiho Institute of Technology, 23 Ping
Kuang Road, Neipu, Pingtung, 91202, Taiwan. E-mail: zhshu2001@yahoo.com.tw
L. Urban, Directeur de l’UR Génétique et écophysiologie de la qualité des agrumes (GEQA), Centre
INRA de Corse, 20230 San Giuliano, France. E-mail: urban@corse.inra.fr
T.M. de Villa, Fruit and Vegetable Laboratory, Food Science Cluster, College of Agriculture, Univer-
sity of the Philippines Los Baños, Laguna, 4031, Philippines.
A.W. Whiley, Sunshine Horticultural Services Pty Ltd, 287 Dulong Road, Nambour, QLD 4560,
Australia. E-mail: whileys@bigpond.com
M. Wysoki, Department of Entomology, Institute of Plant Protection, The Volcani Center, Bet Dagan
50250, Israel. E-mail: manesw@volcani.agri.gov.il
E.M. Yahia, Facultad de Química, Universidad Autónoma de Querétaro, Querétaro, 76190 Qro, Mexico.
E-mail: yahia@.uaq.mx
Preface
The first edition of The Mango: Botany, Production and Uses appeared in 1997,
and went into an unprecedented second printing in the following year. Despite
the worldwide importance of the mango, this was the first book that was
devoted solely to this fruit crop species since the publication of The Mango by
Gangolly et al. in 1957 and The Mango: Botany, Cultivation and Utilization by L.B.
Singh in 1960. The appearance of The Mangoes: their Botany, Nomenclature, Hor-
ticulture and Utilization by Kostermans and Bompard in 1993 had provided a
much-needed taxonomic and systematic revision of mango and the related
Mangifera species; the Kostermans and Bompard book also stimulated interest
in the Mangifera spp. germplasm for breeding and rootstock development.
The Mango: Botany, Production and Uses (Litz, 1997) provided a fresh per-
spective of the mango. The authors represented several countries, including
India, Australia, Israel, the UK, France, USA, Mexico, Pakistan and South
Africa, and reflected the expansion of mango production outside its tradi-
tional areas of cultivation during the mid-20th century and the development
of new technologies in these new lands. The worldview of the first edition
was unique, and the authors were at the forefront of the advance of science
in support of mango production. I wish to particularly acknowledge L.A.
Milne (South Africa), R.V. Mosqueda-Vazquez (Mexico), S.K. Mukherjee (India)
and S. Ram (India), who contributed to the first edition, and who have passed
away since then.
Since 1997, other mango books have appeared: El Cultivo del Mango by V.
Galán Saúco in 1999 (Spain), Mango Cultivation edited by R.P. Srivastava in
1998 (India), A Cultura da Mangueira edited by P.J. de Carvalho Genu and
A.C. de Queiroz Pinto in 2002 (Brazil) and El Mango by E. Yahia Kazuz, J. de
J. Ornelas Paz and R. Ariza Flores in 2006 (Mexico). These books have gener-
ally targeted audiences in specific mango-producing countries. Drs Galán
Saúco, Pinto and Yahia are also contributors to the second edition of The
Mango: Botany, Production and Uses.
ix
x Preface
Much has happened in the decade following the appearance of the first
edition of The Mango: Botany, Production and Uses. China has emerged as the
second largest producer of mango fruit; India’s production is now less than
half of the world total. Fresh mangoes are now consumed worldwide and are
available year-round in the European Union (EU), North America and Japan.
The availability of fruit of a range of mango cultivars is increasing. Mango
products, including fruit nectars, leather, dried fruit slices, preserves, yogurt,
etc. have become widely popular outside the tropics.
The authorship of the second edition of The Mango: Botany, Production
and Uses represents the USA, Mexico, Brazil, Australia, the Philippines, Tai-
wan, India, Israel, France and Spain, and includes leading authorities in each
field. The subject matter of this book ranges from the most basic to the applied,
and is designed to be a compendium that will remain highly relevant for
researchers and growers for many years.
I would like to express my appreciation and thanks to all of the authors
for their persistence during the 3-year gestation period. I would like to
express my gratitude to Ian Maguire of the Tropical Research and Education
Center of the University of Florida for his photographic assistance. Financial
assistance provided by Dr Yungcong Li, also of the Tropical Research and
Education Center, for reproduction of colour plates is gratefully acknowl-
edged. Special thanks to Pamela A. Moon, Guillermo Padilla and Irene Perea
who tolerated me while I worked on this project.
Richard E. Litz
References
de Carvalho Genu, P.J. and de Queiroz Pinto, A.C. (eds) (2002) A Cultura da Mangueira.
EMBRAPA Informacao Tecnologica, Brasilia DF.
Galán Saúco, V. (1999) El Cultivo del Mango. Mundi-Prensa, Madrid.
Gangolly, S.R., Singh, R., Katyal, S.L. and Singh, D. (1957) The Mango. Indian Council
for Agricultural Research, New Delhi, India.
Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mangoes: their Botany, Nomencla-
ture, Horticulture and Utilization. Academic Press, London.
Litz, R.E. (ed.) (1997) The Mango: Botany, Production and Uses. CAB International,
Wallingford, UK.
Singh, L.B. (1960) The Mango: Botany, Cultivation and Utilization. Leonard Hill, London.
Srivastava, R.P. (1998) Mango Cultivation. International Book Distributing Co, Lucknow,
India.
Yahia Kazuz, E., de J. Ornelas Paz, J. and Ariza Flores, R. (2006) El Mango. Editorial Trillas,
S.A. de C.V., Mexico.
Acknowledgements
xi
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1 Introduction: Botany and Importance
1.1 Introduction 1
1.2 Description of Mango 2
The tree 2
Flowers 2
The fruit 3
The seeds and polyembryony 4
1.3 History of Cultivation 5
Origin of Mangifera indica 5
Domestication of mango 9
Distribution 10
1.4 Germplasm Conservation 11
Genetic erosion 11
Collection and documentation of Mangifera germplasm 12
Relevance of germplasm resources to mango improvement 12
1.5 Importance of Mango 12
Cultivars 12
1.6 Production and Uses 14
1.1 Introduction
Mango has become a major fruit crop of the tropics and subtropics, particu-
larly in Asia, where the mango has always been the most important fruit crop
and where it has been considered the ‘king of fruits’ (Purseglove, 1972). A
generation ago, the Green Revolution culminated, creating surpluses of sta-
ple and horticultural crops in many developing countries. The Green Revo-
lution was the result of nearly a century of effort of applying Mendelian
genetics to crop improvement (i.e. conventional breeding) together with the
optimization of agronomic and horticultural practices and the successful
management of insect pests and diseases. However, improvement of tree
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 1
2 S.K. Mukherjee and R.E. Litz
crops has lagged far behind field crops for several reasons: their heterogene-
ity, polyploidy, lengthy juvenile period, time required for evaluation of trees
in the field, and the relatively high cost of maintaining tree plantings. For the
most part, fruit cultivars continue to be ancient selections, many of which
have serious problems, including alternate bearing, lack of disease resistance,
low yields, etc. The rapid growth of mango production in recent years has
been due to its expansion into new growing regions of the New World, China
and parts of Africa; the planting of regular bearing selections; and the adop-
tion of modern field practices, which include irrigation management, control
of flowering, etc. Agricultural practices are currently undergoing another
revolution, as integrated pest and disease management replaces the earlier
reliance on agrichemicals, and emerging fields within biotechnology begin to
impact cultivar development.
The tree
Flowers
Mango flowers are borne on terminal pyramidal panicles, and are glabrous
or pubescent; the inflorescence is rigid and erect, up to 30 cm long, and is
widely branched, usually tertiary, although the final branch is always cymose.
The inflorescence is usually densely flowered with hundreds of small flow-
ers, which are 5–10 mm in diameter. The flowers are either monoecious or
polygamous, and both monoecious and polygamous flowers are borne
within a single inflorescence (Plate 1). The pistil aborts in male flowers. The
ratio of monoecious to polygamous flowers is strongly influenced by
Introduction: Botany and Importance 3
environmental and cultural factors. The flowers have four or five sepals and
petals that are ovate to ovoid to lanceolate and also thinly pubescent. The
floral disc also is four- or five-lobed, fleshy and large and located above the
base of the petals. There are five large, fleshy stamens, only one or two of
them being fertile; the remaining stamens are sterile staminodes that are sur-
mounted by a small gland. In addition, two or three smaller filaments arise
from the lobes of the nectaries. The stamens are central. The ovule is anatro-
pous and pendulous. It is believed that the flowers are cross-pollinated by
flies (see Davenport, Chapter 5, this volume).
Mukherjee (1951a, 1953) investigated the pollen morphology of mango
and 12 other Mangifera species. Their pollen grains were tricolpate of
almost the same size. Mondal et al. (1982, cited in Kostermans and Bom-
pard, 1993) attempted to correlate pollen morphology with taxonomic
relationships of 17 Mangifera species based upon different characteristics
of the exine and sporoderm. They demonstrated that all of the species of
section II (subgenus Limus) possess coarse exine; whereas there was no
clear correlation with pollen type in species within section I (subgenus
Mangifera).
The fruit
Description
The mango fruit is a large, fleshy drupe, containing an edible mesocarp of
varying thickness. The mesocarp is resinous and highly variable with respect
to shape, size, colour, presence of fibre and flavour. The flavour ranges from
turpentine to sweet. The exocarp is thick and glandular. There is a character-
istic beak that develops laterally on the proximal end of the fruit. A sinus is
always present above the beak. Fruit shape varies, including elongate,
oblong and ovate or intermediate forms involving two of these shapes. Fruit
length can range from 2.5 to > 30 cm, depending on the cultivar. The endo-
carp is woody, thick and fibrous; the fibres in the mesocarp arise from the
endocarp.
The mango fruit is climacteric (see Brecht and Yahia, Chapter 14, this
volume), and increased ethylene production occurs during ripening. Chloro-
phyll, carotenes, anthocyanins and xanthophylls are all present in the fruit.
The skin is generally a mixture of green, red and yellow pigments, although
fruit colour at maturity is genotype dependent. During ripening the chloro-
plasts in the peel become chromoplasts, which contain yellow and red pig-
ments (Krishnamurthy and Subramanyam, 1970; Akamine and Goo, 1973;
Salunkhe and Desai, 1984; Mitra and Baldwin, 1997). Peel colour obviously is
cultivar dependent (see Knight et al., Chapter 3, this volume). Fruit of ‘Bom-
bay Green’ is green; ‘Carabao’, ‘Manila’, ‘Mulgoa’ and ‘Arumanis’ are greenish-
yellow; ‘Dashehari’ and ‘Alphonso’ are yellow; and ‘Haden’, ‘Keitt’ and
‘Tommy Atkins’ have a red blush. The red blush is due to the presence of
anthocyanins (Lizada, 1991). The pulp carotenoids in ripe fruit also vary with
respect to cultivar (Mitra and Baldwin, 1997).
4 S.K. Mukherjee and R.E. Litz
Flavour
Flavour of the mango mesocarp is a function of carbohydrates, organic acids,
lactones, monoterpene hydrocarbons and fatty acids (Mitra and Baldwin,
1997). During fruit maturation, starch that accumulates in the chloroplasts is
hydrolysed to sucrose, glucose and fructose (Medlicott et al., 1986; Selvaraj
et al., 1989; S. Kumar et al., 1994); sucrose is present in slightly higher concen-
trations than either fructose or glucose. Organic acid content decreases dur-
ing ripening (Krishnamurthy and Subramanyam, 1970). The dominant
organic acid is citric acid, but glycolic acid, malic acid, tartaric acid and oxalic
acids are also present (Sarker and Muhsi, 1981; Medlicott and Thompson,
1985). The peach-like flavour of mangoes is attributed to the presence of lac-
tones (Lakshminarayana, 1980; Wilson et al., 1990).
Nutrition
Mango fruit contain amino acids, carbohydrates, fatty acids, minerals, organic
acids, proteins and vitamins. During the ripening process, the fruit are ini-
tially acidic, astringent and rich in ascorbic acid (vitamin C). Ripe mangoes
contain moderate levels of vitamin C, but are fairly rich in provitamin A and
vitamins B1 and B2. Perry and Zilva (1932) determined the vitamin A, C and
D content of the fruit of three Indian mango cultivars, and found that the
pulp of mangoes is a concentrated source of vitamin C. The pulp of mango
fruit contains as much vitamin A as butter, although vitamin D is not present
in a significant quantity. Fruit acidity is primarily due to the presence of malic
and citric acids. In addition, oxalic, malonic, succinic, pyruvic, adipic, galac-
turonic, glucuronic, tartaric, glycolic and mucic acids are also present (Jain et al.,
1959; Fang, 1965). Acidity is cultivar related; for example, immature Florida
cultivars have low acidity (0.5–1.0%) in comparison with ‘Alphonso’ (3%).
During ripening, acidity decreases to 0.1–0.2%. Following fruit set, starch
accumulates in the mesocarp. Free sugars, including glucose, fructose and
sucrose, generally increase during ripening; however, the sucrose content
increases three- to fourfold due to the hydrolysis of starch. Sucrose is the
principal sugar of ripe mangoes. The sucrose content of ripe fruit of three
Indian cultivars, ‘Alphonso’, ‘Pairie’ and ‘Totapuri’, ranges from 11 to 20%
representing 15 to 20% of the total soluble solids (Popenoe, 1932).
Mango seeds are solitary, large and flat, ovoid oblong and surrounded by the
fibrous endocarp at maturity. The testa and tegumen are thin and papery.
Embryos are dicotyledonous. Seeds of monoembryonic mango types contain
a single zygotic embryo, whose cotyledons can be unequal in size or lobed in
shape. The seeds of polyembryonic mango types contain one or more embryos
(Plate 2); usually one embryo is zygotic, whereas the remaining embryos are
derived directly from the nucellus, a maternal tissue. Nucellar embryos
apparently lack a suspensor. Polyembryony has also been reported in
Mangifera casturi, M. laurina and M. odorata (Bompard, 1993). Certain
Introduction: Botany and Importance 5
The largest number of Mangifera species occurs in the Malay Peninsula, the
Indonesian archipelago, Thailand, Indochina and the Philippines (Mukher-
jee, 1985; Bompard, 1989; see Bompard, Chapter 2, this volume). The most
recent classification of Mangifera species was based upon floral morphology
(Kostermans and Bompard, 1993) and included 69 species, most of which are
included in two subgenera Mangifera and Limus with another 11 species
occupying an uncertain position (Table 1.1). Eiadthong et al. (1999b) described
the phylogenetic relationships among Mangifera species using genomic
restriction fragment length polymorphisms (RFLPs) and amplification of
chloroplast DNA (cpDNA), and suggested that the Mangifera species should
be classified using molecular data. In the next few years, it is likely that
molecular biology will have a major impact on phylogenetic studies involving
mango and its relatives.
Mangifera species with a single fertile stamen are distributed in north-
eastern India, Myanmar, Thailand and the Malay Peninsula. Many of the
mango relatives have small fruits with thin, acidic flesh, large seeds, abun-
dant fibre and astringent resinous substances that are localized near the skin.
In addition to M. indica, edible fruit is produced by at least 26 other species
in the genus, primarily species found in South-east Asia (Gruezo, 1992).
Mangifera caesia, known as ‘binjai’ or ‘kemang’ in South-east Asia, is culti-
vated in Java, where it bears fruit in the mango off-season (Bompard, 1992a).
Mangifera foetida is less commonly cultivated due to its highly astringent
fruit; however, the fruit is widely used for pickling and as a substitute for
tamarind (Bompard, 1992b). Mangifera kemang and M. altissima are consumed
6
Table 1.1. Classification of Mangifera species according to Kostermans and Bompard (1993).
7
8 S.K. Mukherjee and R.E. Litz
Domestication of mango
Historical record
It is probable that mango cultivation originated in India, where De Candolle
(1884) estimated that mango cultivation appeared to have begun at least 4000
years ago. In the early period of domestication, mango trees probably yielded
small fruit with thin flesh. Such fruit can be found today in north-eastern
India and in the Andaman Islands (Anonymous, 1992). Folk selections of
superior seedlings over many hundreds of years would have resulted in
larger fruit with thicker flesh. Mukherjee (1950a, b) described many of these
primitive selections from Orissa in north-eastern India; they demonstrated
great variation in fruit shape and size.
The mango is a very important cultural and religious symbol of India.
Buddhist pilgrims Fa-Hien and Sung-Yun mentioned in their travel notes
that the Gautama Buddha was presented with a mango grove by Amradarika
(c.500 bc) as a place for meditation (Popenoe, 1932). According to Burns and
Prayag (1921), a mango tree is depicted in friezes on the stupa of Bharut,
which was constructed c.100 bc. Other travellers to India, including the Chi-
nese Hwen T’sung (ad 632–645), the Arabs Ibn Hankal (ad 902–968) and Ibn
Batuta (ad 1325–1349) and the Portuguese Lurdovei de Varthema (ad 1503–
1508), all described the mango. The Indian subcontinent was the birthplace
of some of the earliest highly developed civilizations, and over the centuries,
India exerted strong cultural, religious and commercial influence over South
and South-east Asia. In successive waves, Hinduism, Buddhism and Islam
were introduced into South-east Asia from India. To this day, many com-
monly used words in Indonesia are derived from both Sanskrit and Tamil.
One of the most widely used words for mango in Malaysia and Java (Indone-
sia) is ‘mangga’, which is derived from the Tamil ‘manga’. Traders and monks
from India possibly introduced superior selections of mango into South-east
Asia; however, vegetative propagation was unknown in India until after the
arrival of the Portuguese in Goa in the 15th century. Moreover, the most im-
portant mango selections of Thailand, Cambodia, Vietnam, Malaysia, Indo-
nesia and the Philippines historically have all been of the polyembryonic
type, and have traditionally been seed propagated. Until the establishment
of Portuguese enclaves on the coast of India beginning in the late 15th century,
mango cultivars did not exist in India, as there was no known method
for vegetatively propagating superior selections (see Iyer and Schnell,
10 S.K. Mukherjee and R.E. Litz
Chapter 4, this volume). However, under the Moghul emperor Akbar (1556–
1605), the best selections of seedling mangoes were propagated by approach
grafting and were planted in large orchards. The ‘Lakh Bagh’, a mango
orchard of 100,000 trees, was planted near Darbhanga in Bihar. Perhaps noth-
ing more eloquently attests to the importance of this fruit and the esteem in
which it was held than this vast mango orchard. The Ain-i-Akbari, an ency-
clopedic work that was written during the reign of Akbar, contains a lengthy
account of the mango, and includes information about the quality of the fruit
and varietal characteristics. There was evidently a strong body of informa-
tion about mango cultivation that had accumulated up to that time. Most of
the mango cultivars of India had their origin in those years, and have been
maintained under cultivation for over 400 years by vegetative propagation.
‘Alphonso’, ‘Dashehari’, ‘Langra’, ‘Rani Pasand’, ‘Safdar Pasand’ and other
mango cultivars were selected during that time. Relics of orchards from the
time of Akbar are found in different parts of India, and it has been suggested
that they could still provide valuable material for selection of superior mango
cultivars.
Distribution
Current distribution
The mango is cultivated commercially throughout the tropics and in many
subtropical areas. It is grown at the equator and at a latitude of 35–37q in
southern Spain. According to Knight and Schnell (1993), ‘The process that
began in Florida – introduction of superior germplasm from abroad followed
by selection of improved cultivars adapted to local conditions – is now
underway in many areas.’
The Mangifera species have their centre of diversity and origin in South-east
Asia, a region that has experienced great economic development in recent
years. Vast wooded areas have been completely or partially deforested either
for expanding agriculture or for removal of tropical hardwoods for export.
12 S.K. Mukherjee and R.E. Litz
This has caused great genetic erosion within many species and genera. The
Mangifera species, like many other tropical fruit trees, are canopy and emer-
gent trees of the tropical rainforest (Kaur et al., 1980). These trees are widely
scattered in the tropical rainforest, flower erratically and reproduce from
large seeds that deteriorate rapidly. As such, they are particularly vulnerable
and in danger of extinction.
Cultivars
A partial list of the principal mango cultivars has been provided in Table 1.2.
This list includes many cultivars that were identified in a survey of world
mango production compiled by Watson and Winston (1984). The distribution
of mango cultivars outside their centres of domestication can be attributed
primarily to three historical events: (i) the movement of Indian varieties
(monoembryonic) along the trade routes of the Portuguese to Africa and
South America; (ii) the spread of South-east Asian varieties (polyembryonic)
across the Pacific Ocean to Central and South America by the Spaniards; and
(iii) the identification of improved mango cultivars initially in Florida and
Introduction: Botany and Importance 13
The mango is the most important fruit of Asia, and currently ranks fifth in
total production (in metric tonnes) among major fruit crops worldwide, after
Musa (bananas and plantains) (105,815,354 t), Citrus (all types) (105,440,168
t), grapes (65,584,233 t) and apples (59,444,377 t) (FAOSTAT, 2006). According
to the Food and Agriculture Organization of the United Nations (FAO)
database (FAOSTAT, 2006), world mango production has increased from
16,903,407 t in 1990 to 28,221,510 t in 2005. Much of this new production has
occurred outside the traditional centres of mango culture of South and
South-east Asia. In 1990, India produced approximately 51% of the world’s
mangoes, but by 2005, India’s share had declined to approximately 38%,
despite the substantial increase in mango production since 1990 (from 8,645,405
to 10,800,000 t between 1990 and 2005). The current leading producing nations
after India include (in metric tonnes) China (3,450,000), Thailand (1,800,000),
Pakistan (1,673,900), Mexico (1,600,000), Indonesia (1,478,204), Brazil (1,000,000)
and the Philippines (950,000). Although world production has increased by
67% between 1990 and 2005, mango exports have increased almost sixfold
Introduction: Botany and Importance 15
References
Akamine, E.K. and Goo, T. (1973) Respiration and ethylene production during ontogeny
of fruit. Journal of the American Society for Horticultural Science 98, 381–383.
Angeles, D.E. (1992) Mangifera altissima Blanco. In: Verheij, E.W.M. and Coronel, R.E.
(eds) Plant Resources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO,
Wageningen, the Netherlands, pp. 206–207.
Anonymous (1992) Annual Report. Central Institute of Horticulture for Northern Plains,
Lucknow, India.
Aron, Y., Czosnek, H., Gazit, S. and Degani, C. (1998) Polyembryony in mango
(Mangifera indica L.) is controlled by a single dominant gene. HortScience 33,
1241–1242.
Blume, C.L. (1850) Museum Botanicum Lugduno – Batavum 1, 190–193.
Bompard, J.M. (1989) Wild Mangifera species in Kalimantan (Indonesia) and in Malay-
sia. Final Report. International Board for Plant Genetic Resources, Rome.
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2 Taxonomy and Systematics
J.M. Bompard
Les Mazes, Montaud, France
2.1 Introduction 19
2.2 The Genus Mangifera L. 20
Distribution 20
Ecology and habitat 20
2.3 Taxonomy and Systematics 22
Taxonomic history 22
2.4 Phytogeography 28
Species distribution 28
Subgenera and section distribution 29
2.5 Interspecific Molecular Characterization 30
2.6 Region of Origin of the Genus 31
2.7 Origin of the Common Mango 32
The common mango in South-east Asia 32
2.8 Conclusion 35
Potential contribution of wild species to mango cultivation 35
Source of rootstock 35
Hybridization 36
Potential of wild species 36
2.1 Introduction
The genus Mangifera is one of the 73 genera (c.850 species) belonging to the
family of Anacardiaceae, in the order of Sapindales. Anacardiaceae is a fam-
ily of mainly tropical species, with a few representatives in temperate regions.
Malesia, which is the phytogeographic region extending from the Malay
Peninsula south of the Kangar-Pattani line to the Bismarck Archipelago east
of New Guinea (Whitmore, 1975) contains more species in the Anacardiaceae
than any other area. Within Malesia occurrence is mainly in Western Malesia
(Ding Hou, 1978b).
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 19
20 J.M. Bompard
Apart from mango, several other cultivated fruit trees belong to the fam-
ily, for example the ambarella or Otaheite apple (Spondias dulcis Forst.) prob-
ably from Melanesia, and the yellow and purple mombins (Spondias mombin
L. and S. purpurea L., respectively) from tropical America, the Bouea species
from IndoMalesia, dragon plums (Dracontomelum spp.) from IndoMalesia
and the Pacific region, kaffir plum (Harpephyllum caffrum Bernh. ex K. Krause)
and the marula plum (Sclerocarya caffra Sond.) of southern Africa. The cashew
(Anacardium occidentale L.) is from tropical America and the pistachio (Pistacia
vera L.) from Iran and Central Asia. Anacardiaceous species also yield other
valuable products: wood (several genera), gums and resins (Pistacia spp.),
varnishes (Rhus spp. and Melanorrhoea spp., ‘lacquer trees’) and tanning
materials (Rhus spp. and Schinopsis spp.). It is also a family well known for
the dermal irritation produced by some of its members, such as the poison
ivies and oaks (Rhus spp.) in North America, rengas (Gluta spp.) in South-
east Asia and other species including some Mangifera species whose resinous
sap may induce a mild to strong allergic reaction.
Distribution
4 3
20° N 20° N
4
13
6
10° N
5 5 10° N
2
27
0° N 28 0° N
27 7 4
9 3
2
5
10° N 10° N
Fig. 2.1. Distribution of Mangifera species in the range of the genus. Numbers shown indicate
the number of wild species in each area: Sri Lanka, India and Sikkim, Andaman and Nicobar
Islands, Myanmar, Thailand, Indochina, China, peninsular Malaysia, Sumatra, Borneo, Java,
Lesser Sunda Islands, Sulawesi, Moluccas, the Philippines, New Guinea, and Solomon Islands
(the Caroline Islands not represented).
Taxonomic history
one group with four or five petals and the other group with four petals. He
considered the following sequence of morphological characters to be impor-
tant for classification: (i) texture of the leaves; (ii) number of fertile stamens;
(iii) prominence of veins; (iv) pilosity of inflorescences; and (v) leaf shape.
Pierre (1897) further divided the genus Mangifera into five sections based
on flower characters, i.e. number of stamens, the attachment of stamens to
the disk, and the style. Two of these five sections – namely section I Euan-
therae, with a short thick flower disc and 4–12 fertile stamens, and section V
Marchandora then consisting of M. camptosperma (currently considered a syn-
onym of M. gedebe) are still maintained as they form clear-cut sections.
In his monograph, Mukherjee (1949) recognized two unnamed sections,
conserving Hooker’s subdivision. Ding Hou (1978a) adopted the same
method in his revision of the Malesian Anacardiaceae recognizing only
Hooker’s two original sections and providing them with proper names and
synonyms: section Mangifera (section I Hooker, section Amba Marchand,
group A Engler, sections Euantherae and Marchandora Pierre) and section
Limus (section II Hooker, sections Limus and Manga Marchand, group B
Engler, and sections Eudiscus and Microdiscus Pierre).
SUBGENUS MANGIFERA. The subgenus Mangifera contains most of the species (47),
and is divided into four sections: (i) section Marchandora Pierre; (ii) section Euan-
therae Pierre; (iii) section Rawa Kosterm.; and (iv) section Mangifera Ding Hou.
Section Marchandora Pierre. This section has only one species, M. gedebe Miquel
(syn. M. camptosperma Pierre, M. inocarpoides Merr. and Perry, M. reba Pierre).
The labyrinthine seed is unique to this species, wherein the inner integu-
ments penetrate the cotyledons and form numerous irregular folds. The flat,
discus-like fruit has only a very thin mesocarp. Mangifera gedebe grows in
inundated places along rivers or lakes. The seed floats in water and is dis-
persed during periods of high water, and this may explain its wide distribu-
tion, from Myanmar through Malesia to New Guinea and the Bougainville
Island.
Section Euantherae Pierre. The three species in this section (M. caloneura Kurz
(syn. M. duperreana Pierre), M. cochinchinensis Engler and M. pentandra Hook.
f.) appear to be the most primitive among the species of the subgenus
Mangifera. The flowers are characterized by the presence of five fertile sta-
mens. The three species are mainly confined to Myanmar, Thailand, Indo-
china and the north of the Malay Peninsula. The region is in the transition
zone from the humid tropical rainforest to monsoon forest, and these species
show an adaptation to low rainfall. Mangifera cochinchinensis, which occurs in
south-eastern Thailand and in Vietnam, has small oblong fruits with a thin
seed; the fruits are much relished by local people in southern Vietnam,
although they are very acidic. Mangifera caloneura and M. pentandra are closely
26 J.M. Bompard
related, and can be mistaken for M. indica. However, their leaves are more
leathery, have a more conspicuously dense reticulation, and the panicles are
much more hirsute than the common mango. Mangifera caloneura occurs from
Myanmar through Thailand to Indochina, in lowland evergreen forests, as
well as in semi-deciduous forests. It is cultivated for its acidic-sweet fruit,
and has been planted along the streets of Vientiane and Ho Chi Minh City
(Saigon). Mangifera pentandra, apparently native to the northern Malay Pen-
insula close to the Kra isthmus transition zone, is found in old orchards, in
scattered locations, especially in Kedah and possibly also in peninsular Thai-
land. It is also grown in the Anambas Islands and in Sabah, where it might
have been introduced in early times. It is a prolific bearer, with small man-
goes, c.8 cm length, and ripening green or yellow. The pale orange, watery
pulp has a sweet taste and few fibres.
Section Rawa Kosterm. This group, consisting of nine species, is not well
delimitated. Most species have thick twigs and rather coriaceous leaves
seated on protruding pedestals. The small, hardly flattened ovoid or ellip-
soid fruits that are black or partly red at maturity in several species are also
characteristic. ‘Rawa’ is the Malay word for marsh, indicating that these spe-
cies usually are found in periodically or permanently inundated areas. The
five species that occur in west Malesia (M. gracilipes, M. griffithii, M. micro-
phylla, M. paludosa and M. parvifolia) grow primarily in the swamps of south
peninsular Malaysia, in central coastal areas of east Sumatra and western
Borneo, and occasionally in peripheral uplands. It has also been reported
from the Andaman Islands and from Thailand (Sreekumar et al., 1996; Eiad-
thong et al., 2000a).
Mangifera andamanica and M. nicobarica are endemics from the Andaman
and Nicobar Islands, respectively. Mangifera merrillii is a rare species endemic
to the Philippines and M. minutifolia is known solely from a single collection
from southern Vietnam. Mangifera griffithii and M. microphylla are the only
cultivated species within section Rawa. The former species is considered to
be representative of the section, and is cultivated along the eastern coast of
peninsular Malaysia and in western Borneo, and rarely in Sumatra. The fruits
are small (3–5 cm long) and oblong or ovoid; the skin is rose-red, turning
purplish black at maturity. The rind is thin and easily removed from the
orange-yellow pulp, which is juicy and pleasantly sweet. Different forms are
recognized by local people, according to the size and taste of fruits. Mangifera
microphylla is a related, but less well-known species, having thinner leaves
and a rather similar fruit.
Section Mangifera Ding Hou. With more than 30 species, section Mangifera is
by far the largest. The common mango and the related M. laurina belong
here. Species within the section have the same distribution range as the
genus. The section may be divided into three groups based on floral structure
and organ number variation: (i) those having pentamerous flowers; (ii) those
having tetramerous flowers; and (iii) an intermediate group of species hav-
ing both pentamerous and tetramerous flowers. Within these three groups, it
is possible to distinguish species with either puberulous or glabrous panicles.
Taxonomy and Systematics 27
2.4 Phytogeography
Species distribution
An examination of the present distribution of the genus shows that the larg-
est number of Mangifera species in either subgenera is found in western
Malesia on the Sunda shelf. A decreasing number of species occurs towards
the genus boundary east of Wallace’s line in east Malesia, and in its northern
and western range of distribution. While peninsular Malaysia and the islands
of Sumatra and Borneo have the highest diversity of species, the number of
species becomes gradually lower in east Malesia, especially in the Lesser
Sunda Islands, Moluccas and New Guinea. This is explained by the geologic
and paleogeographic features of the Malesian region which spans two large
partly submerged continental shelves, the Asiatic shelf (Sunda Shelf linking
the Malay Peninsula with the islands of Sumatra, Java, Borneo and Palawan)
and the Australasian shelf (Sahul Shelf linking the Aru islands and New
Guinea with Australia). During the last glaciation period (c.22,500–11,000 bp)
the shelves were regions of land uncovered by the lowering of sea level, and
present day peninsular Malaysia, Sumatra and Borneo were connected by
Taxonomy and Systematics 29
land bridges during the late period of maximal sea lowering. During the cool
periods of glacial maxima, the Malesian forest was reduced in extent, but
there is no evidence that it was reduced to isolated island forests. The Sunda-
land and Papuasian rainforest blocks are therefore comparable to refugia in
terms of species richness and the high degree of endemism (Whitmore, 1981).
Mangifera has undergone major species development in west Malesia, which
has remained relatively stable over a long period of time. The current vegeta-
tion of west Malesia probably differs very little from that at the end of the
Tertiary (van Steenis, 1950). A lower number of Mangifera species is found in
Java and the Philippines, regions less often connected with Asia during the
Pleistocene.
Only three species occur in New Guinea, which is largely covered with
rainforest. These include M. minor, M. mucronulata and the widely distributed
M. gedebe. Mangifera foetida also occurs, but may have been introduced. Mangifera
minor occurs from Celebes and the Philippines to the Solomon Islands; M.
mucronulata is found in the Moluccas, New Guinea and the Solomon Islands.
The distribution of these species suggests a late immigration of a Laurasian
genus from Sundaland via the Philippines, Sulawesi and the Moluccas into
New Guinea, which is supported by the geological history of the region. No
Mangifera species have ever been recorded from northern Australia.
Very few species are found in peninsular India and Sikkim. From present-
day distribution, there is little evidence of migration of species into the sub-
continent of India after its collision with Eurasia in the middle Eocene
(Audley-Charles et al., 1981). According to Mehrotra et al. (1998), fossil leaves
described as Eomangiferophyllum damalgiriensis Mehr. from the Upper Palaeo-
cene in north-eastern India are an analogue of the modern genus Mangifera.
Mangifera sylvatica occurs along the northern limit of the range of
Mangifera, with more or less discontinuity, from Sikkim to northern Thailand
and to the southern part of Yunnan, where it is reported in mountains up to
1900 m above sea level (Anonymous, 1980). The few species that grow in
southern China are very poorly known: M. austro-yunnanensis from western
Yunnan, M. persiciformis from south-eastern Yunnan and southern Guizhou
at latitudes up to 26°N and M. hiemalis, the ‘winter mango’ from Guangxi
near the northern border Vietnam. In the revised Flora of China (Min and Bar-
fod, 2008), M. austro-yunnanensis is considered to be conspecific with M.
indica, M. hiemalis is treated as a synonym of M. persiciformis, and M. laurina
is recorded from the lowland forests of south Yunnan.
The species distribution is especially meaningful when the ranges of the spe-
cies of each subgenus and section are considered separately.
Subgenus Limus
All species of the subgenus Limus are restricted to the Malesian area (M. foetida
and M. macrocarpa occurring in peninsular Thailand), whereas all the species
30 J.M. Bompard
with five fertile stamens, considered the most primitive condition, are con-
fined to west Malesia (M. decandra in Sumatra and Borneo; M. lagenifera in the
two latter areas and in peninsular Malaysia). Only M. caesia, M. foetida and
closely related M. leschenaultii occur in east Malesia.
Subgenus Mangifera
In the subgenus Mangifera, M. gedebe is the only species belonging to the sec-
tion Marchandora, and has the widest range within the genus, extending from
Myanmar through Malesia to New Guinea and Bougainville Island. Section
Euantherae is centred in the region from Myanmar to Vietnam. Mangifera
pentandra is only known from peninsular Malaysia, the Anambas Islands and
Borneo. Section Rawa is mainly in western Malaysia and shows notable
diversification in the swamps and peripheral uplands in the south of penin-
sular Malaysia, east central Sumatra (notably the Riau province) and west
Borneo. During the glacial period this area, termed the ‘Riouw pocket’ (Cor-
ner, 1978), formed a vast plain connecting the Malay Peninsula, Sumatra and
Borneo, and is believed to have been filled with swamps. Mangifera merrillii
is an endemic of the Philippines, M. minutifolia is an endemic of Vietnam, M.
andamanica and M. nicobarica are endemics of the Andamans and Nicobar
Islands. None of the species of section Mangifera occurring in mainland
South-east Asia, north of the isthmus of Kra, are found in eastern Malesia;
however, it would be interesting to assess the genetic relatedness of M. syl-
vatica and M. minor, and also M. laurina, which may prove to be phylogeneti-
cally very closely related.
He concluded that the genus had its origin somewhere in the Myanmar–
Thailand–Indochina area or in the Malayan area. Careful identification of the
greatest part of herbarium materials available has allowed a more accurate
delimitation of the distribution ranges of the Mangifera species, notably of the
subgenus Limus, and has revealed, among other things, that M. lagenifera
does not occur north of Kra isthmus contrary to Mukherjee’s assertions. Fur-
thermore, the ten-stamen species, M. decandra, which was described by Ding
Hou in 1972 and hence was unknown to Mukherjee, is confined to Borneo
and Sumatra, and to date has not been recorded from peninsular Malaysia.
32 J.M. Bompard
The Linnean binomial (Mangifera indica) indicates in this instance the place
where the common mango was selected and improved, and not necessarily its
place of its origin. It has been traditionally accepted that mango was domesti-
cated several millennia ago in India (see Mukherjee and Litz, Chapter 1, this
volume); however, it cannot be excluded that domestication occurred inde-
pendently in several areas, possibly in the south-western and south-eastern
regions of its centre of origin, or later differentiated in those two regions. This
hypothesis would account for the differences that exist between the local
polyembryonic cultivars of Myanmar, Thailand, Indochina and Indonesia,
and the monoembryonic Indian cultivars. Note that polyembryony occurs
Taxonomy and Systematics 33
also in the cultivated M. casturi, M. laurina and M. odorata. Aron et al. (1998)
have demonstrated that polyembryony in mango is under the control of a
single dominant gene.
According to Juliano (1937), Bijhouwer suggested that there were two
main centres of domestication of mango, ‘one in India with monoembryonic
mangoes, the other in the Saigon area, Indonesia and the Philippines with
polyembryonic mangoes’. The ‘Saigon’ area must in fact be extended to
southern Vietnam, other parts of Indochina, Thailand and Myanmar, which
were recognized by Valmayor (1962) as homes of polyembryonic mangoes.
Notwithstanding, the origin of polyembryonic mangoes is probably better
placed in Myanmar, and possibly the eastern part of Assam. According to
Brandis (1874), ‘in Burma, the mango is not generally grafted, and seeds of a
good kind, as a rule, produce fruit of a similar description’. There are only a
few polyembryonic mango cultivars in India. They are restricted to the south-
western coastal region, and geographically isolated from the polyembryonic
mangoes of Myanmar and South-east Asia. Analysis of genetic relatedness
using RAPD markers among polyembryonic and monoembryonic cultivars
grown in the west coast of southern India suggest that the polyembryonic
types are unlikely to have originated from India and might have been intro-
duced from South-east Asia (Ravishankar et al., 2004).
Indian Buddhist monks might have introduced the common polyembry-
onic mango to South-east Asia, first along land trade routes through Myan-
mar, where they might have found better races, and from there into insular
South-east Asia. It is well established that some local names of the common
mango currently used in parts of Indonesia are of Sanskrit origin (‘ampelam’
and its cognates), and are sometimes used to designate M. laurina, which is a
truly native species. Vernacular names do not always travel with a plant, and
even if they did so in the case of the common mango, it is very unlikely that
these introductions were the first ones and that they came obligatorily from
India. In the absence of a comprehensive classification of the innumerable
South-east Asian cultivated forms of the common and wild mangoes, includ-
ing the countless primitive races, we have to rely on linguistics and the rich
history and prehistory of this region.
Vernacular names
The different local names of the common mango in Indonesia (‘pauh’, ‘ampe-
lam’ and its variants, and ‘mangga’) bear evidence of a long history of con-
tacts with mainland Asia and India, and point to possible introduction at
different times from different places. In some parts of Indonesia, the vernacu-
lar names ‘paoh’ or ‘pauh’ refer either to primitive races of the common
mango, or to native species, as a rule the ones most closely resembling the
common mango, for example: ‘pauh asal’ (= native mango) for M. pentandra
in peninsular Malaysia; ‘pahohutan‘ or ‘pahutan’ (= forest mango) for M.
altissima in the Philippines; and ‘pao pong’ (= forest mango) for M. minor in
Flores, Lesser Sunda Islands. ‘Pau’ is a word belonging to Austronesian lan-
guages, nowadays spoken over a very wide area from Madagascar to the
Easter Islands by people who originate from mainland Asia. These languages
34 J.M. Bompard
2.8 Conclusion
Source of rootstock
Grafting experiments between M. indica and other species are reported in the
literature, for example budding of M. indica on M. foetida and M. odorata in
Java (Ochse and Bakhuizen, 1931), M. odorata on M. indica in the Philippines
(Wester, 1920), and M. indica on M. zeylanica in Sri Lanka (Gunaratman, 1946).
36 J.M. Bompard
Mangifera indica ‘Madu’ in Java, and M. laurina in Sabah have been used as
rootstocks for M. casturi. Trials of grafted M. caesia on M. indica (Wester, 1920)
and M. indica on M. kemanga or M. caesia (Ochse and Bakhuizen, 1931) were
unsuccessful, as these two species have distinct bark features and only remote
affinity with the common mango. Better compatibility can be expected using
species more closely related to the common mango within the subgenus
Mangifera. In West Kalimantan, M. laurina is occasionally used as a rootstock
for the common mango on periodically inundated riverbanks. It has been
tried as a rootstock by the Department of Agriculture in Sabah (Lamb, 1987).
Campbell (2004) reported that M. casturi, M. griffithii, M. laurina, M. odorata,
M. pentandra and M. zeylanica grafted on M. indica had a high percentage of
success.
Several species that can grow in permanently inundated areas (i.e. M.
gedebe, M. quadrifida, M. griffithii and other species of the section Rawa) repre-
sent a potential source of rootstock for the development of mango cultivation
on poorly drained soils or in areas liable to prolonged flood. Other species
may be a source of dwarfing rootstocks.
Hybridization
There is little doubt that wild mangoes are potentially valuable in breeding
programmes. Some species have important horticultural implications as they
demonstrate many desirable characteristics (Bompard, 1993). Fairchild (1948)
Taxonomy and Systematics 37
noted that crosses between the common mango and related five-stamen spe-
cies of the section Euantherae might produce hybrids with better pollinating
quality. Mangifera pentandra, which is grown in peninsular Malaysia and
Sabah, is a prolific bearer, due to its high proportion of hermaphrodite to
male flowers.
Stress resistance
In the Malesian rainforests, wild mangoes thrive well under an ever-humid
climate, without a prolonged dry season, i.e. is in areas with an annual rain-
fall > 4000 mm and no monthly mean < 100 mm and where the common
mango cannot be grown satisfactorily. Species, occurring in subtropical areas,
including primitive races of the common mango, or in high altitude tropical
forests, should be evaluated for cold tolerance, opening up the possibilities
for mango production in subtropical and Mediterranean areas. Mangifera lau-
rina and other species related to the common mango that grow in the rainfor-
est (e.g. M. minor in New Guinea) are apparently immune to anthracnose.
Sharma and Choudhury (1976) also observed that trees of an unknown wild
race found in the Tripura State (north-eastern India) were free from mango
malformation.
Acknowledgement
Thanks are due to Dr Dawn Frame who assisted in correcting the text.
Note
1Surveys were carried out in Kalimantan in cooperation with the Indonesian Institute of
Science (LIPI) and the Indonesian Commission on Germplasm, and in Malaysia with the
Forest Research Institute of Malaysia (FRIM).
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3 Important Mango Cultivars and
their Descriptors
3.1 Introduction 43
3.2 Criteria for Cultivar Description 44
3.3 Mango Cultivars 45
‘Alfa’ (Brazil) 45
‘Alphonso’ (India) 45
‘Amelie’ (West Africa) 46
‘Arumanis’ (Indonesia) 46
‘Ataulfo’ (Mexico) 46
‘B74’ (‘Calypso’™) (Australia) 47
‘Banganpalli’ (India) 47
‘Beta’ (Brazil) 47
‘Bombay Green’ (India) 47
‘Cambodiana’ (Vietnam) 48
‘Carabao’ (Philippines) 48
‘Chausa’ (India) 48
‘Cogshall’ (Florida, USA) 49
‘Coração de Boi’ (Brazil) 49
‘Dasheheri’ (India) 49
‘Espada’ (Brazil) 49
‘Ewais’ (Egypt) 50
‘Excellent Succari’ (Egypt) 50
‘Extrema’ (Brazil) 50
‘Fajri’ (India) 50
‘Fernandin’ (India) 51
‘Genovea’ (Egypt) 51
‘Glenn’ (Florida, USA) 51
‘Golek’ (Indonesia) 51
‘Haden’ (Florida, USA) 52
‘Himsagar’ (India) 52
‘Hindi Besennara’ (Egypt) 52
‘Hindi Khassa’ (Egypt) 52
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
42 (ed. R.E. Litz)
Mango Cultivars and Descriptors 43
3.1 Introduction
The mango (Mangifera indica L.) has traditionally been grown in an area that
extends southwards and eastwards from India through Myanmar and Vietnam
to Indonesia. It probably is not indigenous to the Philippines, where it has long
been cultivated (Valmayor, 1962), but Mangifera species are endemic there (Bon-
dad, 1982). This crop is best adapted to a warm tropical monsoon climate, with a
pronounced dry season followed by rains. Fruit of the best quality is usually
produced in such areas, but specific races are known to fruit in humid regions.
For example, some Mangifera species bear dependably on the island of Borneo,
where most standard cultivars do not mature normal crops of fruit. Numerous
Mangifera species closely related to the common mango are indigenous to
44 R.J. Knight et al.
Borneo and nearby parts of Malaysia and Indonesia, and this region is the prob-
able centre of diversity for this genus (see Bompard, Chapter 2 this volume).
Most crops long cultivated over an extended time and area show consider-
able diversity, reflecting different selection criteria in different regions of cul-
ture as well as genetic responses to varied environmental influences. Certainly
this is true of the mango. Indian cultivars differ markedly from those grown
in South-east Asian countries and in Egypt. An additional factor that has
promoted genetic diversity in the group recently has been the widespread
introduction of this crop into new areas of cultivation, many in the western
hemisphere, over the last 500 years. In this manner, genetically diverse germ-
plasm has been brought from widely dispersed areas of the original range of
the species and grown in mixed plantings where, through the cross-pollination
natural to the species, new genetic combinations have been made and selected
under many varying conditions of microclimate. In Florida, since the late 18th
century, enough such importations and genetic recombinations have occurred
to qualify the southern part of this state as a secondary centre of diversity for
the crop. A new group of mango clones designated the Florida cultivars has
been exported to Brazil, Israel, Australia and other places where the process of
increasing diversity under new and varying cultural and environmental con-
ditions continues (Knight and Schnell, 1994; Schnell et al., 2006).
Some Florida cultivars, most notably ‘Haden’, have been important in aid-
ing the establishment of a modern mango industry in other parts of the world
(Knight and Schnell, 1994), and the phenomenon first observed in Florida is now
occurring elsewhere; we are presented with the prospect of the importation of
cultivars of outstanding merit from their countries of origin to be grown, first
experimentally and then commercially, in new regions. For this reason it is
important to become familiar with the characteristics of a group of cultivars that
currently are known in the commerce and/or horticulture of one or more coun-
tries, and that may have potential for expanded culture or use in breeding.
that both leafy bracts and number of perfect flowers are influenced by local
conditions and vary in their expression with differing environments.)
Additional plant data used in initial evaluation include those for the flower,
diameter in millimetres, type (pentamerous, tetramerous or both), nature of
disc (swollen, broader or narrower than ovary, reduced or absent) and number
of stamens; fruit, length, width and thickness, weight, shape, skin colour (which
may be compared with reference cultivars), skin thickness, skin texture, ratio
of pulp to skin and stone, texture of pulp, adherence of skin to pulp, fibre in
pulp and its quantity and length, and stem insertion; and stone, length, weight,
veins and pattern of venation, presence or absence of fibres and their length.
Additional plant data for leaves, inflorescence and fruit have been col-
lected and some of these, notably season (maturity period), productivity, eat-
ing quality and attractiveness are quite important. Unfortunately, from the
viewpoint of those who expect to apply these criteria outside the Indian sub-
continent, reference cultivars are for the most part Indian and many are not
readily available outside India. Other important characters that have been
evaluated or proposed for evaluation include susceptibility to stress (drought,
wind, flooding), susceptibility to specific diseases and pests, molecular
markers, cytological characters and identified genes. Because of the extreme
comprehensiveness of this list and the limited availability of many of the
proposed descriptor evaluations at this time, we have tried to utilize such
information as is available to make the comparison, identification and evalu-
ation of specific well-known cultivars a practical possibility.
A list of mango cultivars that are of interest in areas other than their places of
origin, with descriptions intended to help differentiate them, follows (see
Plates 4–40). Spelling and name variants in some cases represent efforts to
transliterate from other orthographies to the Roman alphabet, and in others
reflect regional differences in usage.
‘Alfa’ (Brazil)
‘Alphonso’ (India)
Also known as ‘Gouverneur’ in the Caribbean. The tree is tall with a rounded,
dense canopy; the fruit is green to orange-yellow with the advance of the season,
rounded, 10–15 cm long by approximately 10 cm broad by approximately 7.8 cm
thick and weighing 300–600 g (average 360 g). The skin is thick and separated
with difficulty; the flesh is soft, juicy, melting, without fibre, a deep orange
colour, sweet and perfumed, free from turpentine, and provides the best of
mango tastes. Seed is monoembryonic in a medium-sized, elongate, narrow
stone that adheres to the flesh, having a few short, pliable fibres that are not
objectionable; the quality is excellent; the season early. The fruit closely resem-
bles that of ‘Julie’. ‘Amelie’ is exported to France, along with ‘Kent’, from
Burkina Faso, Ivory Coast and Mali. ‘Amelie’ is increasing in popularity on the
French market, chiefly in Paris and the surrounding area. It brings lower prices
than cultivars with blushed fruit because the consumer is not always aware
when it is ripe (Naville, 1985, 1986; R. LePrette, personal communication, 1996).
‘Arumanis’ (Indonesia)
Also referred to as ‘Harumanis’. The tree is vigorous and tall with a slightly
open canopy. The fruit (Plate 5) is greenish yellow with large, light-yellow
dots, elongate oblong with rounded base, 11–14 cm long by 6.6–7.5 cm broad
by 4.75–6.5 cm thick, weighing 200–350 g. The skin is thick, tough and easily
separated, the flesh firm and juicy with little fibre, lemon yellow, sweet,
slightly insipid with a strong aroma, of poor to fair quality. Seed is polyem-
bryonic in a thick, woody stone; this cultivar ripens midseason and bears
regularly. Relatively easy to propagate by graftage, scionwood survives well;
widely planted in humid parts of the world where many better-quality culti-
vars fail to fruit (R.J. Knight, Jr, personal communication, 1995).
‘Ataulfo’ (Mexico)
‘Banganpalli’ (India)
Also called ‘Beneshan’ and ‘Chappatai’. The tree is medium sized with a
rounded canopy; the fruit is primrose-yellow, ovate-oblique, large and the
skin smooth, thin and shiny, flesh firm to meaty with juice moderately abun-
dant, without fibre, maize-yellow, with pleasant aroma and sweet taste. Seed
is monoembryonic, in an oblong stone covered with sparse fibres; quality
good; ripens midseason and bears heavily (Singh, 1960).
‘Beta’ (Brazil)
Also called ‘Bhojpuri’, ‘Bombai’, ‘Hiralal Bombai’, ‘Kali Bombai’, ‘Laile Alipur’,
‘Malda’, ‘Sarauli’ and ‘Sheeri-Dhan’. The tree is tall and erect; the fruit (Plate
8) is apple green with ochre blush at the base and on some exposed parts,
dots abundant, with brown specks in the middle, ovate with beak almost
missing, medium sized, with tough, thick, non-adhering smooth skin; the
flesh is cadmium-orange, firm and juicy with scanty fibre just under the skin,
48 R.J. Knight et al.
very sweet with pleasant aroma, of very good quality; seed is monoembryonic
in a full, thick, medium-sized stone. This cultivar ripens early in the season
and is a medium bearer. ‘Bombay Yellow’ is said to be practically identical to
this cultivar but for a slight difference in fruit colour. The present ‘Bombay
Green’ is said to be a degenerate form of the original one (Singh, 1960). In
Jamaica it is sometimes called ‘Peter’, which suggests a confusion with ‘Pairi’,
but the Jamaican ‘Peter’ is without the bright red blush normal to ‘Pairi’.
‘Cambodiana’ (Vietnam)
Also known as ‘Xoai Voi’. The tree is moderately vigorous, with a dense,
rounded canopy; the fruit (Plate 9) is greenish yellow, unblushed with a few
small white dots, oblong to ovate, 9–11.5 cm long by 6.5–7.5 cm broad by
5–6 cm thick, weighing 220–340 g; the skin is thin, tender and adherent; the
flesh contains little fibre, is tender and melting, lemon yellow, sweet and
mildly subacid with a pleasant aroma; the seed is polyembryonic in a thick,
woody stone. Ripens early in the season. Brought to Florida in 1902, where it
gave rise to the ‘Saigon’ landrace (Campbell, 1992).
‘Carabao’ (Philippines)
The tree is vigorous, forming a large and dense canopy; the fruit (Plate 10) is
greenish to bright yellow, brushed with a few small green dots, long and
slender, with base rounded to slightly flattened, 11–13 cm long by 6.5–7 cm
broad by 6–6.5 cm thick, weighing 270–440 g; the skin is thick, medium tough
and easily separated; the flesh is without fibre, tender and melting, lemon
yellow, spicy and sweet with a mild aroma, of good to excellent quality; seed
is polyembryonic in a thin, papery stone. Ripens early in the season (Camp-
bell, 1992). This is a heavy bearer that may alternate; however, it can be
induced to fruit by potassium nitrate treatment in the tropics (Bondad and
Linsangan, 1979). It is highly resistant to bacterial black spot (Xanthomonas
campestris pv. mangiferaeindicae) in Queensland (Mayers et al., 1988). It was
introduced to Florida in 1909. ‘Carabao’ is important in commerce between
the Philippines and Japan and is increasingly imported into the USA.
‘Chausa’ (India)
Also called ‘Samar Bahisht Chausa’ and ’Khajari’. The tree is tall and spread-
ing; the fruit is canary yellow to raw sienna when fully ripe, with numerous
obscure medium-sized dots with minute specks inside them, oblong with
prominent beak, obtuse to rounded, medium sized; the skin is thin and some-
what adhering, pulp raw sienna, soft and juicy with scanty fine, long fibres
near the skin; the fruit is very sweet with a luscious, delightful aroma, of
excellent quality; seed is monoembryonic in a thick, medium-sized oblong
Mango Cultivars and Descriptors 49
stone with fine, short fibres all over the surface and a tuft of long fibres on
the ventral edge. Ripens late in the season and is a light bearer (Singh, 1960).
The tree is vigorous, precocious and productive; the fruit is greenish yellow and
intense red on the side exposed to the sun, cordiform, medium sized, pulp yel-
low and fibrous. The seed is polyembryonic. There are two seasons in São Paulo,
January–February and September–December. This is one of the best-known
commercial cultivars in São Paulo state (Sampaio, 1980; A.C. Pinto, personal
communication, 1996; L.C. Donadio, personal communication, 1996).
‘Dasheheri’ (India)
Also known as ‘Dasheri’ and ‘Aman Dusehri’. The tree is of medium height
and moderate vigour, spreading, with a rounded, medium-dense canopy; the
fruit is primrose to canary yellow with abundant light-yellow dots, oblong to
oblong-oblique with base rounded to obliquely rounded, medium sized, skin
smooth, medium thick, tough and non-adhering; the flesh is yellow, firm,
with almost no fibre, scanty juice and a delightful aroma, very sweet taste, of
excellent quality; seed is monoembryonic in a thick, medium-sized stone. Rip-
ens midseason and is heavy bearing; fruit keeps well (Singh, 1960).
‘Espada’ (Brazil)
The tree is tall and develops rapidly, with a dense canopy, very productive;
the fruit is intense green or greenish yellow, oblong-elongate with a concave
base, medium sized, with smooth, thick skin; the flesh has much fibre, is egg-
yellow, with a strong aroma of turpentine. The quality is considered good for
fresh consumption. The polyembryonic seed is in an oblong stone, covered
50 R.J. Knight et al.
with soft fibres and many nerves. There are two seasons per year in São
Paulo, January–February and November–December (Sampaio, 1980; A.C.
Pinto, personal communication, 1996).
‘Ewais’ (Egypt)
‘Extrema’ (Brazil)
The tree is upright, vigorous and productive. The fruit is yellow with green-
ish areas, ovate-reniform, weighing 350–400 g, with smooth and thin skin,
and yellow, watery flesh with almost no fibres with a moderately resinous,
agreeable taste. The quality is considered good for fresh consumption and
processing. The polyembryonic seed is in a large, fibrous stone. Ripens early
in the season (Sampaio, 1980; A.C. Pinto, personal communication, 1996).
‘Fajri’ (India)
Also spelled ‘Fazli’. The tree is of medium size and moderately vigorous,
with rounded, open canopy. The fruit is light chrome yellow with small,
dark-coloured fairly sparse dots, obliquely oval with base slightly rounded
and beak distinct to slightly prominent, large (averaging 14.3 cm long by
9.8 cm broad, weighing 500 g on average) with a medium-thick skin that is
Mango Cultivars and Descriptors 51
smooth with some inclination to be warty, and firm to soft, fibreless flesh of
a light cadmium yellow with a pleasant aroma and a sweet taste, having juice
that may be scanty to moderately abundant, of good to very good quality.
The seed is monoembryonic in a large, oblong stone that is covered with a
sparse, short and soft fibre. Ripens midseason to late (Gangolly et al., 1957; N.
Balasundaram, India, personal communication, 1990).
‘Fernandin’ (India)
The tree is moderately vigorous with a dense, rounded canopy; the fruit is
bright yellow with an attractive bright-red blush, ovate-oblique, averaging
12.2 cm long by 8.5 cm broad, weighing 450 g; the skin is rough and warty,
thick and adherent, flesh bright yellow, moderately to abundantly juicy,
thick, with no objectionable fibre, with delightful to piquant aroma and sweet
to very sweet, delicious taste, of superior quality; seed is monoembryonic;
season late (Gangolly et al., 1957; Singh, 1960).
‘Genovea’ (Egypt)
The tree is moderately vigorous, small to medium with dense, rounded can-
opy of compact growth; the fruit (Plate 11) is bright yellow with orange-red
blush, with numerous small yellow and white dots, oval to oblong with a
rounded base, 9.5–12.5 cm long by 7.5–8.5 cm broad by 7–8 cm thick, weigh-
ing 400–620 g; the skin is thin, tough and easily separated, flesh soft and juicy,
with little fibre, deep yellow, rich and spicy with a strong, pleasant aroma, of
excellent quality; seed is monoembryonic in a thick, woody stone. Ripens
early in the season and is a regular bearer. This is a seedling of ‘Haden’
(Campbell, 1992; Schnell et al., 2006).
‘Golek’ (Indonesia)
The tree is moderately vigorous with an upright, open canopy; the fruit
(Plate 12) is greenish yellow with an orange overlay and prominent white
dots, oblong with rounded base, 9.5–12.5 cm long by 6–8 cm broad by 5.5–
6.5 cm thick, weighing 200–365 g; the skin is thin, tough and easily separated;
52 R.J. Knight et al.
the flesh is soft and juicy with abundant fibre (not objectionable), deep yel-
low, sweet, insipid with a mild aroma, of poor to fair quality; the seed is
polyembryonic in large, woody stone with abundant fine fibre. Ripens mid-
season (R.J. Knight, Jr, personal communication, 1995).
The tree is vigorous, with a large, spreading canopy; the fruit (Plate 13) is
bright yellow with a deep crimson or red blush and numerous large yellow
dots, oval with a rounded base, 10.5–14 cm long by 9–10.5 cm broad by 8.5–
9.5 cm thick, weighing 510–680 g; the skin is thick, tough and adherent; the
flesh is firm and juicy with abundant fibre, deep yellow, rich and sweet with
a pleasant aroma, of good to excellent quality; the seed is monoembyonic in
a medium-thick woody stone. Ripens early to midseason and bearing is
sometimes irregular. This is a seedling of ‘Mulgoba’ × ‘Turpentine’ and is the
first of the Florida mango cultivars, introduced in 1910 and since grown in
many other countries. It is the seed parent of numerous other cultivars
(Campbell, 1992; Knight and Schnell, 1994; Schnell et al., 2006).
‘Himsagar’ (India)
The tree is vigorous, tall, with a dense, spreading canopy; the fruit (Plate 14)
is greenish yellow to bright yellow with no blush, with light-yellow dots,
ovate with a flattened base, 12–15 cm long by 8.5–9.5 cm broad by 7.5–8.5 cm
thick, weighing 465–585 g; the skin is thin, tough and easily separated; the
flesh is firm and juicy with no fibre, orange, rich and sweet with a mild aroma,
of good to excellent quality; the seed is monoembryonic in a thick, woody
stone. This is a late midseason cultivar that bears well (R.J. Knight, Jr, per-
sonal communication, 1995).
The tree is small to medium, moderately vigorous, with open canopy. The fruit
(Plate 16) is bright yellow with a crimson or bright red blush, numerous large
white dots, ovate with rounded base, 11.5–13 cm long by 8–9 cm broad by 6.5–
7.5 cm thick, weighing 340–450 g; the skin is medium-thick, tender and adherent;
the flesh is soft, tender, melting and juicy without fibre, lemon yellow, sweet and
mild with a pleasant aroma, of good quality; the seed is monoembryonic in a
thin, papery stone. The stone may be seedless following cool weather at flower-
ing time. This is an early, regular and heavy bearer. The fruit is usually soft with
a short postharvest life, but it is often exported from tropical America to Europe.
It is a seedling of ‘Lippens’ × ‘Haden’ (Campbell, 1992; Schnell et al., 2006).
Also called ‘St Julienne’. The tree is compact (dwarf), with a dense canopy;
the fruit (Plate 17) is greenish yellow with a light pink to maroon blush and
numerous small white dots, rounded with flattened apex, pronouncedly
compressed laterally, 7–9.5 cm long by 4–7.5 cm broad by 2–5.5 cm thick,
weighing 200–325 g with a thin, tender skin and soft, melting, juicy, orange
flesh with scanty fibre, of a rich, spicy flavour with a strong, pleasant aroma,
of good quality; seed is monoembryonic in a thin, papery stone. This cultivar
ripens midseason and is a regular producer of small crops. The fruit is often
severely infected with anthracnose disease, but its unique taste is preferred
by many West Indians, and it is exported to the London market (C.W. Campbell,
personal communication, 1996).
The tree is medium sized, moderately vigorous, upright with open canopy;
the fruit (Plate 18) is greenish yellow, with a pink or red blush, numerous
small white or yellow dots, oval, with rounded base, 13–15 cm long by
9–11 cm broad by 8.5–10 cm thick, weighing 510–2000 g; the skin is thick,
tough and adherent; the flesh is firm and juicy, with little fibre, lemon yellow,
sweet and mild with a pleasant aroma, of good to excellent quality; the seed is
monoembryonic in a thick and woody stone. This cultivar ripens late in the
season. It is a seedling of ‘Brooks’. After ‘Tommy Atkins’ it is the most com-
mercially important cultivar in the export mango industry of the western
54 R.J. Knight et al.
‘Kensington’ (Australia)
Also known as ‘Kensington Pride’ and ‘Bowen’. ‘Kensington’ has a large, vig-
orous tree with spreading canopy; the fruit (Plate 19) is yellow with an orange-
red blush on the shoulder, round ovate with a flattened base and a slight beak,
10.5–13 cm long by 8.5–9.6 cm broad by 7.5–8.5 cm thick, weighing 350–750 g;
the skin is thick, tender and adherent; the flesh is soft and juicy, with moderate
to little fibre, sweet with a characteristic flavour that makes it the most popu-
lar cultivar in Australian markets, of excellent quality; seed is polyembryonic
in a moderately thick, woody stone. This cultivar ripens midseason and it
bears well. It is unusually susceptible locally, in Florida, to damage by red-
banded thrips (Selenothrips rubricinctus (Giard.)), and may be killed by this pest
without adequate countermeasures (R.J. Campbell, personal communication,
1994; R.J. Knight, Jr, personal communication, 1995). It is moderately suscep-
tible to anthracnose and bacterial spot (Mayers et al., 1988).
The tree is large and vigorous with a dense, upright canopy; the fruit (Plate
20) is greenish yellow with a red or crimson blush, numerous small yellow
dots, oval, with rounded base, 11–13 cm long by 9.5–11 cm broad by 9.9.5 cm
thick, weighing 600–750 g; the skin is thick, tough and adherent; the flesh is
firm, tender, melting and juicy with little fibre, deep yellow to orange-yellow,
sweet with a rich flavour and pleasant aroma, of excellent quality; the seed
is monoembryonic in a thick, woody stone. Fruit ripens late midseason to
late and bearing may be alternate. It is a seedling of ‘Haden’ × ‘Brooks’, which
is a seedling of ‘Totapuri’ (‘Sandersha’) (Schnell et al., 2006). ‘Kent’ is not
commonly commercial in Florida because it is prone to storage disease, but
is a successful commercial cultivar in drier parts of Mexico, Central and
South America and West Africa (Campbell, 1992). It is highly susceptible to
bacterial black spot in Queensland (Mayers et al., 1988).
‘Khanefy’ (Egypt)
A cultivar of minor commercial importance. The fruit is large (475 g), green
with a yellow overlay and large, brown, smooth dots, ovate in shape (10.7 cm
long by 8.3 cm wide by 8.6 cm thick), with an adherent skin quite free of sur-
face disease; the flesh is yellow, often with jelly seed, juicy, with no objection-
able fibres and a bland flavour unacceptable to many Western palates. The
stone is moderately large (53 g) (Knight and Sanford, 1998).
Mango Cultivars and Descriptors 55
The tree is large, vigorous, with an open canopy made up of long branches;
the fruit is green when harvested (before the ripening process begins) turn-
ing to greenish yellow, oblong, 11.5–12.5 cm long by 5.5–6.5 cm broad by
5–6 cm thick, weighing 230–340 g; the skin is thin, tender and adherent; the
flesh is medium firm, tender and not very juicy with no fibre, pale yellow,
very sweet with an insipid taste and a mild, pleasant aroma, of fair to good
quality; the seed is highly polyembryonic in a medium-thin stone. This is a
regular producer (C.W. Campbell, personal communication, 1995). The fruit
is often consumed green.
‘Langra’ (India)
‘Mabrouka’ (Egypt)
The tree is moderately vigorous, medium sized, forming an open canopy; the
fruit (Plate 22) is greenish to bright yellow, with no blush and a few large rus-
set dots, oblong, sigmoid with rounded base, 15–17 cm long by 8.5–11 cm
broad by 5.5–7.5 cm thick, weighing 370–520 g; the skin is thin, tender and
adherent; the flesh is soft and juicy with medium fibre, orange, rich spicy and
sweet with a pleasant aroma, of fair to good quality; seed is polyembryonic
56 R.J. Knight et al.
in a thin, papery stone. This cultivar ripens early to midseason and bears
well. Shipped to North American markets from Haiti nearly 10 months of the
year (R.J. Campbell, personal communication, 2007).
‘Mallika’ (India)
The tree is a moderately vigorous dwarf with a dense canopy; the fruit (Plate
23) is bright yellow with no blush and numerous small, light-yellow dots,
oblong with rounded base, 10–12 cm long by 6.5–7.5 cm broad by 5–5.5 cm
thick, weighing 280–450 g; the skin is thick, tough and easily separating; the
flesh is soft, tender and juicy with little fibre, deep yellow to orange, rich,
strongly aromatic and sweet, of excellent quality; seed is monoembryonic in
a medium-thick and woody stone. This cultivar ripens midseason and is an
irregular producer. This cultivar came from crossing ‘Neelum’ and ‘Dashe-
hari’ (Singh et al., 1972; Campbell, 1992).
‘Manila’ (Mexico)
The tree is large, vigorous, with an upright, open canopy; the fruit (Plate 24)
is bright yellow, sometimes with a light-pink blush, a few small reddish dots,
long and slender with rounded base and bluntly pointed apex sometimes
with a small beak, 12.5–14 cm long by 5.5–6 cm broad by 5–5.5 cm thick,
weighing 180–260 g; the skin is thin, medium tough and easily separating;
the flesh is medium firm and juicy, with little to abundant fibre, deep yellow,
sweet, rich and spicy in taste with a pleasant aroma, of good to very good
quality; seed is polyembryonic in a medium-thick and woody stone. This
cultivar ripens early midseason and crops fairly dependably. For a long time
‘Manila’ has been the most popular mango in Mexico.
‘Manzanillo’ (Mexico)
The tree is large, of medium vigour with an upright canopy; the fruit is yel-
lowish orange with 75% of the surface blushed an intense dark red with
numerous dots, oval with moderately flattened base, averaging 12 cm long
by 10 cm broad by 7.5 cm thick, and 660 g in weight; the flesh is low in fibre,
slightly subacid and very palatable, quality high; seed is monoembryonic in
a relatively small stone. This cultivar ripens early in the season but spread
over a 60-day harvest period. It bears heavily without pronounced alterna-
tion and the fruit stores and ships well (Núñez-Elisea, 1984).
‘Mesk’ (Egypt)
A major commercial cultivar. The tree is vigorous, the fruit small to medium
sized (312.5 g), yellow with a red blush, with small, corky yellow dots;
Mango Cultivars and Descriptors 57
Also spelled ‘Mugoba’ and ‘Mulgova’. The tree is large, vigorous with open,
spreading canopy; the fruit (Plate 25) is bright yellow with a pink blush and
numerous large white dots, oval to ovate with flattened base, 8.5–10.5 cm
long by 6.5–7.5 cm broad by 5–6 cm thick, weighing 340–450 g; the skin
is thick, medium tough and adherent; the flesh is soft, tender, melting and
juicy, with little fibre, lemon yellow, rich spicy and sweet with strong, pleas-
ant aroma, of good to excellent quality; seed is monoembryonic in a thick,
woody stone. This cultivar ripens midseason to late and is a shy, irregular
bearer. Introduced to Florida in 1889 and called ‘Mulgoba’, this is the seed
parent of ‘Haden’, first of a series of cultivars known as the Florida group. A
question exists whether the cultivar known in Florida is identical with the
Indian cultivar or is a seedling rootstock that survived after the scion was
killed by cold. In either case its superior quality ensured its retention and
propagation (Campbell, 1992). Literature serves to compound the nomencla-
tural confusion, as illustrated by Gangolly et al. (1957) whose ‘Mulgoa’ fruit,
yellow overall and roundish oblique with a deeply depressed stem insertion,
does not resemble the cultivar introduced to Florida. Singh (1960), on the
other hand, portrays a rounded, lightly blushed greenish yellow fruit that
closely resembles the Florida mango. Furthermore, vegetative propagation
of selected chance seedlings has resulted in a variety of clonal types carried
under this name in India (Ratnam and Chellapa, no date, post-1954).
‘Nabeel’ (Egypt)
A minor commercial cultivar. The fruit is large (495 g), green with small yel-
low dots that are smooth; ovate-oblong (14 cm long by 9 cm wide by 7 cm
thick), with adherent skin relatively free of surface disease; the flesh is
orange, firm and juicy, without objectionable fibre, with a passable but not out-
standingly pleasing taste and acceptable quality. The seed is polyembryonic
in a large (56.6 g) stone (Knight and Sanford, 1998).
The tree is vigorous, medium sized with upright, dense canopy; the fruit
(Plate 26) is greenish to bright yellow with a slight pink blush and numerous
58 R.J. Knight et al.
small green dots, long and slender, sigmoid in shape with a rounded base,
17–19 cm long by 7.5–8.5 cm broad by 6.5–7.5 cm thick, weighing 340–580 g;
the skin is medium thick, tender and easily separated from the flesh which is
soft, tender and juicy with no fibre, lemon yellow, rich, spicy and very sweet
with pleasant aroma, of excellent quality; seed is polyembryonic in a thin,
papery stone. This cultivar ripens early midseason, fruits regularly and may
have multiple crops in one season (Campbell, 1992). It is highly resistant to
foliar infection, and resistant to fruit infection by bacterial black spot in
Queensland (Mayers et al., 1988).
‘Neelum’ (India)
The tree is moderately vigorous with a small, compact canopy; the fruit is
bright yellow with no blush and numerous small white dots, oval with flat-
tened or slightly rounded base, 9.5–11 cm long by 7.5–8.5 cm broad by
6–6.5 cm thick, weighing 230–300 g; the skin is thick, tender and easily sepa-
rating; the flesh is soft, melting and juicy with no fibre, deep yellow, mild and
sweet with a delightfully pleasant aroma, of good to excellent quality; seed is
monoembryonic in a medium-thick, woody stone. This cultivar is a late,
heavy bearer (Campbell, 1992).
The tree is moderately vigorous, small, upright with a dense canopy; the fruit
(Plate 27) is greenish yellow with a pink to red blush, numerous small green
dots, long and slender with a flattened base, 16–18 cm long by 7–8 cm broad
by 6–6.5 cm thick, weighing 340–500 g; the skin is thick, tough and easily
separating; the flesh is soft, melting, juicy with little fibre, pale yellow, mild
and sweet with a faint, pleasant aroma, of good eating quality; seed is poly-
embryonic in a thick, woody stone. This cultivar is an early, regular bearer.
Fruit is often eaten green (Campbell, 1992).
‘Okrung’ (Thailand)
The tree is moderately vigorous, medium sized and upright, forming a dense
canopy; the fruit (Plate 28) is green to greenish yellow with no blush and
numerous small white dots, oblong and sigmoid with a rounded base,
11–13 cm long by 5–6 cm broad by 4.5–5.5 cm thick, weighing 160–240 g; the
skin is thick, tough and medium adherent; the flesh is soft and juicy with
abundant fibre, yellow or greenish, mild, somewhat insipid and very sweet
with a pleasant aroma, of good quality; seed is polyembryonic in a thick,
woody stone. This cultivar ripens midseason, is a heavy producer and some-
times bears more than one crop/year (Campbell, 1992).
Mango Cultivars and Descriptors 59
The tree is vigorous, medium sized, forming a dense canopy; the fruit (Plate
29) is yellow-orange with a purple or lavender blush and numerous small
white dots, oblong with rounded base, 12–15.5 cm long by 9–10.5 cm broad by
8.6–9.5 cm thick, weighing 500–760 g; the skin is thick, tough and easily sepa-
rating; the flesh is firm and juicy, with little fibre, lemon yellow, mild and sweet
with a pleasant aroma, of good quality; seed is monoembryonic in a thick and
woody stone. This cultivar ripens late midseason to late and is a regular
producer. It is a ‘Haden’ seedling (Campbell, 1992; Schnell et al., 2006).
‘Pairi’ (India)
Also written ‘Pairie’, ‘Paheri’ and ‘Pirie’; synonyms are said to be ‘Peter’,
‘Peter Pasand’, ‘Grape’, ‘Gohabunder’, ‘Nadusalai’, ‘Rasjuri’ and ‘Yerra Goa’.
The tree is moderately vigorous, forming a dense, rounded canopy; the fruit
(Plate 30) is medium sized, green to yellow with a bright red blush, roundish,
skin smooth, thick, flesh golden yellow, slightly juicy, fibreless, with a deli-
cious subacid taste, of excellent quality; the thick stone covered with short,
bristly fibre encloses monoembryonic seed (Popenoe, 1927; Singh, 1960). This
cultivar has long been popular as a dooryard fruit tree in Hawaii.
The tree is moderately vigorous, forming a large, upright, tight canopy; the
fruit (Plate 31) is yellow-orange with a dark-red to crimson blush and a few
small white dots, oblong with rounded base, 12–15 cm long by 8.5–10 cm
broad by 6.5–7.5 cm thick, weighing 510–850 g; the skin is medium thick,
tough and adherent; the flesh is firm and melting with little fibre, orange-
yellow, mild and aromatic, of good quality; seed is monoembryonic in a
medium-thick woody stone. This is a late midseason cultivar and is a regular
bearer. It is a seedling of ‘Haden’ (Schnell et al., 2006). In Florida it is of minor
commercial importance (Campbell, 1992). It is grown in Israel and is the seed
parent of ‘Naomi’. It is attracting increased attention in the western hemi-
sphere export market as a result of its superior eating quality.
‘Rosa’ (Brazil)
The tree is medium sized, of slow growth with a rounded canopy; the fruit
(Plate 32) is yellow to rose-red on the side exposed to sun, oblong-cordiform
and medium sized; the skin is thick and smooth; the flesh is firm and mod-
erately juicy, fibrous, golden yellow, moderately sweet with a turpentine
aroma, of ordinary quality, susceptible to anthracnose disease; the seed is
polyembryonic in a small, oblong stone. This cultivar ripens midseason to
60 R.J. Knight et al.
late. It is one of the most important commercial cultivars in the Federal Dis-
trict of Brazil, used for juice as well as fresh consumption, and is one of the
most well-known cultivars in Brazil (Sampaio, 1980; L.C. Donadio, personal
communication, 1996; A. Pinto, personal communication, 1996).
The tree is vigorous, with a moderately open, symmetrical canopy; the fruit
(Plate 33) is dark yellow with a prominent dark-red to purple blush that cov-
ers most of its surface, oval with rounded base and rounded apex, 9–11.5 cm
long by 7–8 cm broad by 6.5–7 cm thick, weighing 280–340 g; the skin is
medium thick, tough and easily separating; the flesh is firm and medium
juicy, fibreless, deep yellow, mild and sweet with a weak, pleasant aroma,
of fair to good quality; seed is monoembryonic in a thick, woody stone.
This cultivar ripens midseason to late (Campbell, 1992). It is a seedling of
‘Haden’ × ‘Brooks’ (Schnell et al., 2006), and the seed parent of ‘B74’. It alter-
nates severely, and in ‘on’ years the fruit may be clustered so heavily that it
becomes diseased before maturity, thus ‘Sensation’ is not of commercial
importance. It is highly resistant to bacterial black spot in Queensland (May-
ers et al., 1988), but often has severe internal breakdown (browning, water
soaking) (A.W. Whiley, personal communication, 1996).
‘Suvarnarekha’ (India)
Also called ‘Swarnarekha’ and ‘Sundri’. The tree is moderately vigorous and
tall, with a rounded, dense, spreading canopy; the fruit is light cadmium yel-
low with a blush of jasper red and abundant small, light-coloured dots, ovate
oblong with a base slightly flattened, of medium size, 11 cm long by 8.2 cm
broad, weighing 400 g; the skin is medium thick, easily separated, flesh soft,
fibreless, primrose yellow with a pleasant aroma, sweet taste and abundant
juice, of medium to good quality; seed is monoembryonic in an oblong-oval
stone covered with soft, short fibre. Ripens early in the season early and is
heavy bearing (Gangolly et al., 1957).
‘Tahar’ (Israel)
The tree is vigorous, medium sized, with an upright, dense canopy; the fruit
is bright yellow with a dark-red blush and numerous small white dots, ovate
with flattened base, 11.5–13 cm long by 8.9–9.5 cm broad by 7.5–8 cm thick,
weighing 360–520 g; the skin is thick, tough and easily separating; the flesh is
soft and juicy with little fibre, deep yellow, mild, aromatic and slightly insipid
with a strong odour not appreciated by many, of fair to good quality; seed is
monoembryonic in a medium-thick woody stone. This cultivar ripens in late
midseason and bears well in Israel (Campbell, 1992).
Mango Cultivars and Descriptors 61
‘Taimour’ (Egypt)
A major commercial cultivar. The tree is vigorous; the fruit (Plate 34) is large
(500 g), ripening late in the season, dark green with large light-brown dots,
smooth in texture, ovate-oblong (12.8 cm long by 8.4 cm wide by 8 cm thick),
with non-adherent skin of intermediate thickness, quite free from surface
disease; flesh is orange, firm (free of jelly seed) and juicy with no objection-
able fibre, of a delightfully rich, sweet taste and excellent quality. The seed is
polyembryonic in a medium-sized (50.8 g) stone (Knight and Sanford,
1998).
The tree is vigorous, with a dense, rounded canopy; the fruit (Plate 35) is
orange-yellow, with a crimson or dark-red blush and numerous small, white
dots, oval to oblong, with broadly rounded base, 12–14.5 cm long by 10–13 cm
broad by 8.5–10 cm thick, weighing 450–700 g; the skin is thick, tough and
adherent; the flesh is firm and medium juicy; with a medium amount of fibre,
lemon to deep yellow, mild and sweet with a strong pleasant aroma, of fair to
good quality; seed is monoembryonic in a thick, woody stone. This cultivar
ripens early to midseason. It is a ‘Haden’ seedling (Schnell et al., 2006).
‘Tommy Atkins’ is the most important commercial cultivar in the western
hemisphere export mango market; it is highly resistant to anthracnose dis-
ease and handling and shipping stress, and a consistent, heavy producer
(Campbell, 1992). ‘Jelly seed’ (internal breakdown) is a serious problem in
the moist subtropics and tropics outside Florida, where the mango is grown
on calcareous, well-drained soil (A.W. Whiley, personal communication,
1996).
‘Totapuri’ (India)
The tree is vigorous, with a large, spreading, rounded canopy; the fruit (Plate
37) is bright yellow with a few large white dots, occasionally with a pink
blush, oval with a flattened base, 7.5–8.5 cm long by 6.5–7.5 cm broad by
6–6.5 cm thick, weighing 140–200 g; the skin is thick, tough and easily sepa-
rating; the flesh is firm and juicy, with abundant coarse fibre, lemon yellow,
rich, aromatic, spicy, resinous and sweet with a strong, pleasant aroma, of
poor to fair quality; seed is polyembryonic in a thick, woody stone. This cul-
tivar ripens early midseason to late midseason and is a heavy producer but
may alternate. It is commonly used as a grafting stock (Campbell, 1992).
‘Vallenato’ (Colombia)
The tree is vigorous, with an upright, dense canopy; the fruit (Plate 38) is
bright yellow, with a crimson blush, oblong with flattened base, 8–9 cm long
by 7–8 cm broad by 6–7 cm thick, weighing 195–340 g; the skin is thin, tough
and adherent; the flesh is firm, juicy with abundant fine fibre (not objection-
able), pale yellow, mild and sweet with a strong, pleasant aroma, of good to
excellent quality; seed is monoembryonic. This cultivar ripens in early mid-
season (R.J. Campbell, personal communication, 1995).
The tree is moderately vigorous, with a large, open canopy; the fruit (Plate
39) is bright yellow with a bright red or crimson blush, oval with rounded
base, 9–11.5 cm long by 7.5–9.5 cm broad by 7–8 cm thick, weighing 250–
520 g; the skin is thick, tough and easily separating; the flesh is quite firm,
melting and juicy with little fibre, orange-yellow, rich, spicy and sweet with
a strong, pleasant aroma, of good to excellent quality, but susceptible to inter-
nal breakdown; seed is monoembryonic in a medium-thick, woody stone.
This cultivar ripens in late midseason and is a regular, heavy producer. It is a
seedling of ‘Haden’ (Campbell, 1992; Schnell et al., 2006).
‘Zebda’ (Egypt)
3.4 Conclusion
The mango fruit’s nutritional value, aesthetic and gustatory appeal have
assured its growing importance in non-traditional markets since the late
1950s, as it has been introduced to consumers previously unacquainted with
it. Furthermore, the migration of large populations from South-east Asia and
other regions where this fruit is a traditional crop to metropolitan centres
where it has not been well known has created a permanent demand for it in
these new markets. An additional factor permitting market expansion has
been the growing mango production in areas previously unimportant in
world commerce such as Mexico, Brazil, Australia, West Africa, Israel, Flor-
ida and the Canary Islands. The fact that most new markets are remote from
areas of production has necessitated selection of cultivars for fresh market
sale that are dependably productive and resistant to harvest, handling and
shipping stress, with relatively long shelf life, for example ‘Tommy Atkins’,
‘Keitt’ and ‘Madame Francis’. The fruit quality of mango cultivars well suited
to packing and shipping has been a secondary consideration, and is gener-
ally not so high as that of cultivars acknowledged to be superior for eating.
Economic factors obviously must dictate what is grown for the fresh market.
The commercial market for processed mango products permits other
cultivars to be utilized, and these may vary with the product that is mar-
keted. Cultivars chosen for purée or juice preparation are likely to be quite
different from those used for manufacture of chutney or other products
requiring pulp that maintains its integrity after it is cooked. ‘Totapuri’ (‘Sand-
ersha’) or ‘Turpentine’, for example, considered mediocre for fresh consump-
tion, can be used to prepare excellent chutneys, as can many ‘criollo’ types in
the West Indies. ‘Tommy Atkins’ makes outstandingly good dried fruit sec-
tions, sweet and aromatic, even though its fresh-fruit quality is generally
conceded not to be high. Mango butter and mango leather are other products
that are appreciated by many who know them (see Raymundo et al., Chapter
17, this volume; Campbell and Campbell, 1983; Campbell and Smith, 1987).
As more fruit that is wholesome, but not of export quality, becomes available
in areas of increasing production, it is likely that processed mango products
will become more common.
64 R.J. Knight et al.
Table 3.1. Ratings of selected mango cultivars grown in Florida (Source: Knight, 1993).
Cultivar Shapea Sizeb Firmnessc Colourd Anthracnosee Fibref Tasteg Yieldh Scorei
‘Alphonso’ 3 5 7 2 3 7 9 1h x
‘Boribo’ 3 8 8 4 7 9 5 6 x
‘Carabao’ 5 6 7 3 5 9 8 6h x
‘Haden’ 3 9 8 8 5 7 7 3h x
‘Keitt’ 4 10 9 6 8 9 8 8 ///
‘Kensington’ 3 8 7 7 7 8 7 6 /
‘Langra’ 2 6 8 3 5 8 8 3h x
‘Pope’ 3 9 5 7 2 8 8 1 x
‘Tommy 3 9 9 9 9 6 6 7 ///
Atkins’
‘Van Dyke’ 3 7 10 9 7 8 7 6 ///
aRatings of 1 (round) to 5 (long) indicate fruit shape, not its desirability.
bRatings below 6 justify discard; those of 7 and above show size only, not merit.
cRatings of 1–10 where 1 = least and 10 = most.
dRatings of 1–10 where 1 = least and 10 = most.
eRatings of 1–10 where 1 = most and 10 = least susceptible.
fRatings of 1–10 where 1 = most and 10 = least.
gRatings of 1–10 where 1 = worst and 10 = best.
hTrends markedly towards alternate bearing.
iOne or more checks (/) show overall value; (x) indicates no commercial acceptability.
Acknowledgements
For their help in reviewing portions of this chapter and/or contributing vast
quantities of information on mango cultivar descriptors and attributes, the
authors are profoundly grateful to the following: Dr N. Balasundaram, Head,
Sugarcane Breeding Institute, Regional Centre, Karnal, India 132001; the late
Dr Carl W. Campbell, Tropical Research and Education Center, 18905 SW 280
Mango Cultivars and Descriptors 65
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4 Breeding and Genetics
4.1 Introduction 68
4.2 Origin of Cultivars and Distribution 68
Mangifera species and mango 68
History of cultivation 69
Impact of Florida mangoes 70
4.3 Reproductive Mechanisms 70
Polyembryony 70
Floral biology and pollination 71
Incompatibility 72
Cytology 72
4.4 Inheritance of Characters 73
Dwarfness, regular bearing and precocity 73
Flesh colour 74
Skin colour 74
Flowering time 74
Beak 74
Disease resistance 74
Other horticultural traits 75
4.5 Breeding Objectives 75
General objectives 75
Specific objectives 76
4.6 Methods of Breeding 78
Selection from open-pollinated seedlings 78
Controlled pollination 79
4.7 Handling of Hybrid Populations and Selection 80
Criteria for initial selection 80
Pre-selection 81
Potential for marker assisted selection (MAS) 81
Molecular markers 81
4.8 Minimizing Problems in Breeding 83
Heavy fruit drop 83
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 67
68 C.P.A. Iyer and R.J. Schnell
4.1 Introduction
History of cultivation
Mango has been cultivated in India for at least 4000 years and over 1000 vari-
eties are recognized there (Mukherjee, 1953). Almost all of them are selec-
tions made from naturally occurring, open-pollinated seedlings. However,
based on random amplification of polymorphic DNA (RAPD) analysis, Rav-
ishankar et al. (2004) felt that the mono- and polyembryonic types of Indian
mango cultivars have a different genetic base, and that the polyembryonic
types might have been introduced from South-east Asia and are unlikely to
have originated in India. Mango culture gradually spread to tropical and
subtropical countries throughout the world, where selections were made
that were adapted to particular growing conditions. Thus, selection by man
has played the most significant role in the development of new mango culti-
vars. The explorers who tasted the mango in the regions of its origin were
enchanted with its aromatic qualities, ambrosial flavour and creamy, smooth
and silky texture, and introduced the fruit to other tropical regions. The
spread of Hinduism and Buddhism assisted in the distribution of mangoes in
South-east Asia. The Chinese traveller Hwen T’sang who visited India in the
first half of the 7th century ad returned to China with the mango. The mango
was known in Baghdad in the 7th century.
The Persians or Omanis may have taken mangoes to East Africa around
the 10th century ad. The fruit was introduced to East and West Africa in the
early 16th century by the Portuguese and thence into Brazil. After being
established in Brazil, the mango was carried to the West Indies, being first
planted in Barbados by about 1742 and later in the Dominican Republic and
Jamaica (about 1782). The mango was introduced into Mexico from the Phil-
ippines by the Spanish and also from the West Indies (Morton, 1987). Duval
et al. (2006) developed microsatellite markers for studying the genetic vari-
ability of Caribbean mangoes and concluded that there were two routes of
mango to the French West Indies, namely, cultivars grown in Central Amer-
ica (Mexico) and South America (Colombia) introduced from South-east Asia
and also from former French colonies in the Indian Ocean. As the mango
70 C.P.A. Iyer and R.J. Schnell
adapted to new locales, new cultivars were selected based on local adapta-
tion and fruit preferences.
The first recorded successful introduction of mango into Florida (USA) was
made in 1861 (Knight, 1980). The earliest introductions were from the West
Indies and India, followed by the introduction of several hundred accessions
in the 20th century from South-east Asia, India and from other mango-growing
areas of the world (Florida Mango Forum, 1951). The introduction of mangoes
into Florida and subsequent development of a Florida group of mangoes has
been reviewed by Knight and Schnell (1994). The Florida mango cultivars are
unique in that they are hybrids between Indian cultivars (primarily mono-
embryonic) and the South-east Asian cultivars (primarily polyembryonic)
selected under south Florida conditions. The mango breeding system favours
out-crossing. Therefore, the proximity of numerous genotypes of disparate
geographical origins led to the production of many new seedlings by inter-
pollination in Florida (Knight and Schnell, 1993). Florida selections are
therefore not the result of a formal breeding programme. Early Florida selec-
tions were made by growers and enthusiasts and historical information is
often anecdotal. The Florida Mango Forum, established in 1938 for the
advancement of mango production, documented historical information on
the parentage of Florida cultivars in their proceedings. In addition to the
United States Department of Agriculture (USDA) Germplasm Resources In-
formation Network (GRIN) database, several sources compile information
on Florida mango selections and introduction of accessions to Florida (Ruehle
and Ledin, 1956; Singh, 1960; Campbell and Campbell, 1993; Schnell et al.,
2006). With the exception of South-east Asia, Australia and some African
countries, which cultivate mostly locally selected varieties, the majority of
countries produce cultivars developed in Florida, i.e. ‘Haden’, ‘Tommy
Atkins’ and ‘Keitt’ (Galan Sauco, 1997). These Florida selections are now
widely grown commercial cultivars affording production stability across
many environments (see Mukherjee and Litz, Chapter 1, this volume).
Nucellar embryos
Mangoes can be classified into two groups, monoembryonic and polyembry-
onic, based on their mode of reproduction from seeds. In general, monoem-
bryonic seeds are found in the sub-tropical group (Indian type) and the
polyembryonic seeds in the tropical group (South-east Asian). Monoembry-
onic mango seeds each contain a single zygotic embryo, and hence only one
seedling per seed that is of probable hybrid origin. Polyembryonic mango
Breeding and Genetics 71
seeds can contain one or more embryos, one of which is usually, but not
always zygotic. Adventitious embryos develop from the nucellus, a maternal
tissue surrounding the embryo sac, and consequently the seedlings of poly-
embryonic mangoes are genetically identical to the maternal parent. Adven-
titious embryos can also originate by direct budding from the cotyledons and
hypocotyls of other nucellar embryos (Juliano, 1934). According to Mahesh-
wari and Rangaswamy (1958), the nucellar cells destined to form adventi-
tious embryos are recognizable by their dense cytoplasm and starchy
contents. They gradually push into the embryo sac cavity where they divide
and differentiate into embryos.
Inheritance of polyembryony
Polyembryony is genetically determined. Leroy (1947) considered that adven-
tive embryony probably reflects the effect of one or more recessive genes.
This view was supported by Sturrock (1968), whose study of the progenies of
monoembryonic mango hybridized with polyembryonic cultivars indicated
that monoembryony was possibly a dominant trait. In contrast, Aron et al.
(1998) and Brettell et al. (2004) observed that polyembryony in mango is con-
trolled by a single dominant gene. Schnell et al. (2006) reported that 58 of the
Florida cultivars had been classified with 50 being monoembryonic and eight
polyembryonic. Information from the Florida cultivars parentage analysis
using 25 microsatellite markers supported the findings of Aron et al. (1998)
where polyembryony was found to be dominant. ‘Haden’ is a cross of the
monoembryonic ‘Mulgoba’ and the polyembryonic ‘Turpentine’. If we assume
that a single dominant gene controls this trait, all of the Indian cultivars in
Florida must be homozygous recessive and the ‘Turpentine’ parent of ‘Haden’
must have been heterozygous. The evidence suggests that ‘Haden’ inherited
the recessive allele from ‘Turpentine’, as all identified progeny of ‘Haden’ are
monoembryonic with the exception of ‘Winters’. The most probable pollen
parent of ‘Winters’ is ‘Ono’, a polyembryonic cultivar from Hawaii. The fre-
quency of this dominant allele is low in the Florida population and absent
from the Indian cultivars in Florida. In view of these interesting findings, and
since a thorough knowledge of inheritance of polyembryony is essential for
speculating the origin of M. indica, more work on these lines is warranted.
Flowers begin to open early in the morning and anthesis has generally
been completed by noon. The greatest number of flowers opens between 9
and 10 a.m. Although the receptivity of the stigma continues for 72 h after
anthesis, it is most receptive during the first 6 h; however, there are reports
that the stigma can become receptive even before anthesis has occurred
(Singh, 1960). The minimum time required for pollen grains to germinate is
1.5 h (Sen et al., 1946; Singh, 1954; Spencer and Kennard, 1955). Singh and
Singh (1952) observed 98% pollen viability after 11 months in storage at 7°C
and 25% relative humidity (RH), and 65.7% viability after 24 months of stor-
age at 0°C and 25% RH.
Mango is cross-pollinated, which is carried out by insects such as the
common housefly, honeybees and thrips, and possibly by other insects
al-though to a lesser extent. Pollination by wind and gravity has been sug-
gested to occur in mango (Popenoe, 1917; Maheshwari, 1934; Malik, 1951). In
nature, > 50% of flowers do not receive any pollen. Some workers had sug-
gested that self-pollination in certain cultivars can also occur quite frequently
(Dijkman and Soule, 1951). Studies by Issarakraisila and Considine (1994)
have shown that for polyembryonic ‘Kensington’, a night temperature of
< 10°C results in pollen grains with low viability (< 50%). The optimum
temperature for normal meiosis is between 15 and 33°C with 70–85%
viability.
Incompatibility
Cytology
Chromosome number
Information on the cytology of mango is quite limited. Only a few Mangifera
species (i.e. M. indica, M. caloneura, M. sylvatica, M. foetida, M. caesia, M. odorata
and M. zeylancia) have been studied, and were found to have chromosome
numbers of 2n = 2x = 40 and n = x = 20 (Mukherjee, 1950, 1957; Roy and
Visweswariya, 1951). Chromosome numbers and ploidy status of other species
are yet to be studied. The only exception to this chromosome number that has
been reported to date (Roy and Visweswariya, 1951) involves ‘Vallikolamban’,
which was reported to be tetraploid (2n = 4x = 80), although subsequent stud-
ies have indicated that it is only a diploid (Majumder and Sharma, 1990).
Polyploidy
Mango has been referred to as an allopolyploid. Due to the presence of
secondary associations at metaphase of meiosis, Mukherjee (1950) suggested
that the basic chromosome number of Mangifera is n = 8. In addition, the high
number of somatic chromosomes and the correspondingly high number of
nucleolar chromosomes led him to conclude that mango is an allopolyploid.
However, the evidence used to arrive at this conclusion is not unequivocal.
In fact, the molecular marker evidence is antithetical to this conclusion.
Results from Duval et al. (2005), Viruel et al. (2005) and Schnell et al. (2005,
2006) using microsatellite markers all indicate that M. indica is diploid.
Although many wild Mangifera species are potentially valuable for crop
improvement, they are yet to be exploited. Mukherjee (1963) felt that the
different Mangifera species could intercross easily, based on the success ob-
tained with interspecific crosses between M. zeylanica and M. odorata.
High heterozygosity in the cultivars that are used in hybridization and the
inadequate number of hybrid progenies realized has made accurate genetic
analysis in mango very difficult. However, based on limited data, some indi-
cations are available which would be useful in selecting parents in breeding
programmes designed with specific objectives. In studies of the distribution
of different traits in seedlings derived from open-pollination (where the pol-
len parent is unknown), Lavi et al. (1989) observed: (i) there is no maternal
effect on juvenile period and fertility; (ii) there is a slight effect of the female
parent on fruit taste and size; (iii) there is a maternal parent effect on harvest
season and fruit colour.
Flesh colour
Sharma (1987) considered that additive gene action may be involved in the
inheritance of flesh colour; however, studies involving monoembryonic
‘Alphonso’ and ‘Neelum’ have indicated that light yellow is dominant over
orange-yellow (Iyer, 1991).
Skin colour
With regard to skin colour of fruit, Sharma (1987) observed that when red
cultivars were crossed with green cultivars, the F1 seedlings exhibited vari-
ous gradations of red. Iyer and Subramanyam (1987) also found a wide array
of colours in the hybrids when the coloured monoembryonic ‘Janardhan
Pasand’ was crossed with green-fruited cultivars, indicating that colour is
mediated by a number of loci.
Flowering time
Beak
Disease resistance
General objectives
Specific objectives
Dwarfness
Because of the obvious benefits of comparatively dwarf trees for orchard man-
agement and fruit quality, attempts have been focused on obtaining hybrids
with a dwarf tree framework. Breeding for dwarfness is important in mango,
since a consistent dwarfing effect of any rootstock has not been established to
date. The Indian cultivars that could be useful as a source for dwarfness
include ‘Kerla Dwarf’, ‘Janardan Pasand’, ‘Manjeera’, ‘Amrapali’, ‘Creeping’
(Iyer and Subramanyam, 1986) and ‘Nileswar Dwarf’ (Singh, 1990).
Regular bearing
The causes of irregular bearing vary from region to region. In general, the
main reason for alternate bearing, particularly in subtropical India, is the
lack of initiation of vegetative growth soon after fruiting. However, two cul-
tivars, ‘Neelum’ and ‘Bangalora’ (‘Totapuri’), which are regular bearers, have
been extensively used as either of the parents in a hybridization programme
to transfer the regular bearing habit to hybrids. ‘Neelum’ has been observed
to be a good combiner and has contributed to the evolution of many regular-
bearing Indian hybrid cultivars. However, ‘Bangalora’ is not a suitable par-
ent since the hybrids possess very prominent beaks and their fruit quality is
invariably poor. The regular bearing Florida cultivars (i.e. ‘Tommy Atkins’,
‘Keitt’, etc.) also have potential as parents.
Fruit colour
Most of the commercial cultivars in South-east Asia possess green skin.
Efforts are underway to produce new hybrid cultivars that retain the good
qualities of these fruits together with attractive skin colour, so that they will
occupy a better position in international trade. Since good skin colour has
been shown to be transmissible to hybrids from suitably coloured parental
cultivars, a number of cultivars with coloured skin are being used for hybrid-
ization. In general, the attractively coloured Florida cultivars have been
found to be suitable parents. In addition, there are several Indian cultivars
(e.g. ‘Janardan Pasand’, ‘Suvarnarekha’, etc.) that would be suitable for use
as parents for this purpose.
In Florida, the skin colour of the mango is an important factor and red
skin is considered essential for mangoes shipped to northern markets. In the
past, the evaluation of mango colour has been subjective and based on visual
Breeding and Genetics 77
ratings. Large errors are associated with these types of ratings, which makes
evaluation based on fruit colour difficult. Ayala-Silva et al. (2005) used a colo-
rimeter to quantify fruit colour, quality and differentiation among cultivars.
Mango colour was measured with a Minolta Chroma Meter CR-400 (Osaka,
Japan) portable tristimulus colorimeter and fruit chromaticity was recorded
in Commission Internationale de l’Eclairage (CIE) L*, a* and b* colour space
coordinates. In this system of colour representation the values L*, a* and b*
describe a uniform three-dimensional CIE colour space. If the L*, a* and b*
coordinates are known, then the colour is not only described, but also located
in quantifiable space. Maternal half-sib families (MHS) of the mango culti-
vars, ‘Keitt’, ‘Tommy Atkins’, ‘Tyler Premier’, ‘Mamita’, ‘White Alfonso’ and
‘Sandersha’ were evaluated along with two parental check clones, ‘Tommy
Atkins’ and ‘Keitt’. Significant differences were found for each of the L*, a* and
b* colour space coordinates. Further work is underway to estimate the herita-
bility of these traits to estimate their usefulness for breeding and selection.
Disease resistance
MANGO MALFORMATION. Although no breeding work has been reported that spe-
cifically addresses disease or pest resistance/tolerance, cultivars are known to
show varying degrees of susceptibility to biotic stress (see Ploetz and Freeman,
Chapter 8, this volume ). Mango malformation, caused by Fusarium subglutin-
ans, is a very serious disease that has threatened the very survival of the
mango industry in many subtropical mango-growing regions. As there are
no reliable cultural and chemical control measures available, breeding for
resistance/tolerance has been attempted using ‘Bhadauran’ as the resistant
parent; however, all of the F1 hybrids were susceptible to the disease (Sharma
and Majumder, 1988a). In this respect, the observations of Ram et al. (1987)
are very encouraging. Out of 102 cultivars screened, three of them, namely,
‘Bhydayam Dula’, ‘Samar Bahist Rampur’ and ‘Mian Sahib’, were free of
malformation and could be tried as one of the parents in hybridization.
Gupta (1976) has listed those cultivars that are most tolerant of this disease –
‘Neelum’, ‘Zardalu’, ‘Bangalora’, ‘Totapuri-Khurd’ and ‘Janardan Pasand’ –
and hence could be valuable in breeding programmes.
In India, almost all cultivars are selections that were made from naturally occur-
ring open-pollinated seedlings. All of the Florida cultivars were selected from
open-pollinated seedling progenies; none has come from a controlled breeding
programme. Among the 64 Florida cultivars evaluated in the parentage analysis
by Schnell et al. (2006), the genetic background was found to be based on as few
as four Indian cultivars and the polyembryonic cultivar ‘Turpentine’. Two Indian
cultivars, ‘Mulgoba’ and ‘Sandersha’, are in the background of most Florida
types with ‘Amini’, ‘Bombay’, ‘Cambodiana’, ‘Long’, ‘Julie’ and ‘Nam Doc Mai’
making lesser contributions. In the parentage analysis ‘Turpentine 10’ was iden-
tified as a most probable paternal parent for ‘Haden’. The polyembryonic seed-
ling races of Cuba and Florida were considered the same by Popenoe (1920) who
called them the West Indian race (commonly known as ‘Turpentine’ in Florida).
‘Haden’ was reported as the maternal parent for ten cultivars included in the
analysis, but based on the parentage analysis, 31 cultivars were found to have
‘Haden’ as one of the most likely parents. Likewise, the other important early
Florida selection ‘Brooks’ is the parent of seven cultivars. ‘Haden, ‘Brooks’ and
seedlings of ‘Haden’ and ‘Brooks’ have contributed disproportionately to the
Florida group. In Florida, modern selection and breeding programmes for
mango have focused on cultivars with exceptional production, red skin, disease
resistance and extended shelf life. Methodology for crop improvement consists
of collecting seeds from selected maternal parents with desired characteristics
and growing them in close proximity to desirable male parents. Seedlings are
screened by leaf aroma and horticultural traits, leading to a field population of
thousands of candidate seedlings (Campbell and Zill, 2006).
Breeding and Genetics 79
Controlled pollination
Hand pollination
The traditional, cumbersome method involving the continued pollination of
flowers on a panicle over several days when the flowers are open has now
been replaced in India with more efficient methods. The current method
involves the pollination of a limited number of flowers per panicle (maxi-
mum of ten), utilizing a larger number of panicles since it is very rare that a
panicle bears more than one fruit to maturity. Using this method, fruit set as
high as 3.85% can be achieved compared to the 0.23–1.57% efficiency involv-
ing other methods (Mukherjee et al., 1968; Singh et al., 1980).
Caging
The enclosure of two desirable parents of synchronous flowering in a screen
house with pollinating insects provides a more practical method of hybridiza-
tion. This method has been used in Israel (Degani et al., 1993), Brazil (Pinto and
Byrne, 1993) and South Africa (Cilliers et al., 1996). A standardized caging
technique for mango breeding was previously used in India following the
discovery of self-incompatibility in some of the most popular commercial culti-
vars (Sharma and Singh, 1970). This procedure involves the planting of self-in-
compatible (female) and male parents in specially prepared breeding plots, and
are enclosed in an insect-proof cage into which freshly reared houseflies, or any
other suitable pollinator, are introduced to effect cross-pollination (Sharma
et al., 1972).
Polycross mating
A polycross is simply the use of a number of advanced selections or current
commercial clones planted in a design that maximizes the chance for
cross-pollination. The polycross design has been extensively used in
sugarcane breeding where small flower size and low numbers of seedlings
per cross make controlled pollinations difficult. At USDA Subtropical Horti-
culture Research Station (SHRS) in Miami the clones ‘Haden’, ‘Tommy
Atkins’, ‘Kent’, ‘Keitt’ and ‘Nam Doc Mai’ have been used to produce new
seedlings for selection. Five clones of each genotype were planted, with at
least one plant of each genotype next to all other genotypes. Over 1000 seed-
lings from known mother trees are planted as maternal half-sib families. The
pollen parent of superior selections is determined using microsatellite
markers.
Primary selection from the hybrid progeny is based on: (i) precocity; (ii) fruit
size and shape; (iii) skin colour; (iv) fruit characteristics (high pulp to stone
ratio and freedom from fibre and physiological disorders); and (v) fruit qual-
ity. Following this preliminary evaluation, selected hybrids are retained for
Breeding and Genetics 81
Pre-selection
Trees have a long juvenile phase, and the development of pre-selection meth-
ods is important for discarding inferior seedlings at a very early stage, obvi-
ating the need for maintaining a large number of seedlings for long periods.
This can save time, land and labour. Leaf flavour has been reported to be
directly correlated with fruit flavour (Majumder et al., 1972; Whiley et al.,
1993). Emergence of new growth flushes, simultaneously with fruiting or
immediately after harvest, is indicative of regular bearing (Sharma et al.,
1972). A higher phloem to xylem ratio, associated with dwarfing, has been
used effectively as a pre-selection criterion. Genotypes in which the ratio
exceeds 1.0 are least vigorous, those with a ratio between 0.6 and 1.0 are of
medium vigour and those with a ratio of less than 0.6 are most vigorous
(Kurian and Iyer, 1992). In addition, higher levels of phenolics in the apical
bud is associated with reduced vigour and dwarfing (Iyer, 1991). Although
Majumder et al. (1981) indicated that low stomatal density is an indicator of
dwarfness this has not been confirmed by other workers (Iyer, 1991). Regular
bearing mango cultivars have low polyphenol oxidase (PPO) activity (cate-
cholase and cresolase) compared to alternate bearers (Sharma, 2003). Sharma
et al. (2000) observed that a strong positive correlation existed between the
incidence of floral malformation and both enzyme activity (catecholase and
cresolase) and phenolic content and speculated that PPO activity can be used
as a biochemical index for screening mango germplasm against malforma-
tion disease.
More than 65 microsatellite markers have been developed for mango and
these are easily used to verify parentage using a software package such as
cervus (Marshall et al., 1998). When caging trees or using the polycross mat-
ing design it is possible to identify the male parent from a set of potential
male parents. This has been useful in cacao breeding where mistakes in pol-
lination have lead to the estimation of unreliable breeding values for parental
clones. The development of linkage maps and identification of quantitative
trait loci (QTL) for productivity and quality traits has led to a very successful
MAS in cacao (Schnell et al., 2007). This could serve as a model for future
mango breeding and selection efforts.
82 C.P.A. Iyer and R.J. Schnell
Molecular markers
propagation error in the collection (i.e. plants that had been incorrectly
labelled or grafted) was estimated to be 6.13%. When compared by origin,
the Florida cultivars were more closely related to Indian than to South-east
Asian cultivars. Unbiased gene diversity (Hnb) of 0.600 and 0.582 was found
for Indian and South-east Asian cultivars, respectively, and both were higher
than Hnb among Florida cultivars (0.538). When compared by horticultural
type, Hnb was higher among the polyembryonic types (0.596) than in the
monoembryonic types (0.571).
To date 63 microsatellite markers have been developed for mango
(Duval et al., 2005; Honsho et al., 2005; Schnell et al., 2005; Viruel et al., 2005).
This number is more than adequate for genetic diversity studies and for par-
entage analysis as has been demonstrated by Schnell et al. (2006); however, it
is not enough to develop a saturated linkage map for the 20 linkage groups
of mango. Developing an additional 200 microsatellite or single nucleotide
polymorphic markers is a major objective of the USDA Agriculture Research
Service (ARS) programme in Miami over the next 2 years. Three experimen-
tal populations have been developed and planted in the field as mapping
populations. The first population is an F2 population derived from self-polli-
nation of ‘Tommy Atkins’ consisting of 168 seedlings that was planted in the
field in 1995. The second population is an F2 population derived from self-
pollination of ‘Haden’. A total of 224 seedlings from a single isolated ‘Haden’
tree have been in the field for 3 years. Phenotypic data collection is in prog-
ress for both of these populations. The development of a saturated linkage
map and the identification of QTL for important traits are objectives for the
USDA-ARS programme in Miami for the next 5 years.
Heavy fruit drop ultimately results in few hybrid fruits, despite the large
number of flowers used for cross-pollination. While many recommendations
are available to minimize mango fruit drop with growth regulators, these
have not been very useful in breeding programmes where the number of
flowers remaining in a panicle is very low. Iyer and Subramanyam (1972)
suggested that embryo culture could be used to rescue hybrid embryos, and
Sahijram et al. (2005) developed in-vitro techniques to rescue immature mango
embryos from controlled crosses and recovered hybrid plants.
Polyembryony
1992, respectively) and 36 and 64% with ‘Madu’ and ‘Golek’, respectively
(Schnell and Knight, 1992).
Despite the many problems associated with mango breeding for cultivar devel-
opment, many useful hybrids have been released. The earliest attempts were
probably made in the West Indies to combine the good qualities of the Indian
mango with the indigenous types by controlled pollination (Brooks, 1912).
India
‘Alphonso’ but free of ‘spongy tissue’, has a good shelf life and is not suscep-
tible to fruit fly attack. ‘Arka Anmol’ is a heavy bearer with good keeping
quality (Iyer and Subramanyam, 1993). ‘Arka Neelkiran’ is free of spongy
tissue and has excellent skin colour.
‘Ratna’ is a cross between ‘Alphonso’ and ‘Neelum’ that was carried out
at the Fruit Research Station, Vengurla, Maharashtra; it has a larger fruit size,
fruit quality similar to ‘Alphonso’ and is free of ‘spongy tissue’ (Salvi and
Gunjate, 1988). A parthenocarpic mango cultivar, ‘Sindhu’, has been devel-
oped at this station as a result of back-crossing ‘Ratna’ with ‘Alphonso’ (Gun-
jate and Burondkar, 1993).
Two hybrid cultivars were released from the Fruit Research Station in
Sangareddy, Andhra Pradesh. ‘Au-Rumani’ (‘Rumani’ × ‘Mulgoa’) is a regular
and prolific bearer with fibreless flesh. ‘Manjira’ (‘Rumani’ × ‘Neelum’) is a
dwarf, regular and prolific bearer with good quality fruits.
The Paria Research Station in Gujarat developed three mango hybrids,
‘Neelphonso’ (‘Neelum’ × ‘Alphonso’), ‘Neeleshan Gujarat’ (‘Neelum’ × ‘Bane-
shan’) and ‘Neeleshwar’ (‘Neelum’ × ‘Dashehari’). These hybrids are supe-
rior in TSS, total sugars and vitamin C, in addition to their dwarfing habit,
with respect to their parents (Sachan et al., 1988).
Other countries
USA
Mango hybridization was reported from Hawaii in the 1920s, but no out-
standing problem appears to have been addressed or solved (Pope, 1929). A
number of crosses have been reported in Florida (Young and Ledin, 1954;
Sturrock, 1969), but all of the Florida cultivars are chance seedlings and none
came from controlled pollinations.
Israel
There is an extensive breeding programme in Israel aimed at producing
higher yielding cultivars with good quality, attractive fruit and with longer
harvest periods. Several hundred seedlings from open and controlled polli-
nations have been evaluated, and 14 of them have been identified as being of
interest (Lavi et al., 1993). The rootstock breeding programme is aimed at
developing rootstocks resistant to or tolerant of soil stresses, i.e. calcareous
soils, saline irrigation water and heavy non-aerated soils that predominate in
the mango-growing regions of Israel. Several interesting monoembryonic and
polyembryonic rootstocks have been selected (Lavi et al., 1993), but none has
performed better than ‘13-1’, the currently preferred rootstock in Israel (Gazit
and Kadman, 1980).
Australia
A breeding programme to develop a new cultivar which retains the charac-
teristic flavour of ‘Kensington’, but with improved productivity, greater dis-
ease resistance, enhanced skin colour and better postharvest performance,
Breeding and Genetics 87
Brazil
Breeding has been initiated in the tropical savannah of Brazil to develop cul-
tivars that are dwarf and with good quality fruit. Hybridizations have
involved local, Indian and Florida cultivars. ‘Amrapali’ and ‘Imperial’ were
good male parents to confer dwarfing in the progeny (Pinto and Byrne, 1993).
Out of 2088 seedlings in the field, 209 seedlings were selected in the first
year and 42 of these were later identified as promising, from which four have
been released as new cultivars (Pinto et al., 2004). These four are: ‘Alfa’ (‘Mal-
lika’ × ‘Van Dyke’), which is semi-dwarf, high yielding and regular bearing;
‘Beta’ (‘Amrapali’ × ‘Winter’), high yielding and moderately resistant to
anthracnose and Oidium; ‘Roxa’ (‘Amrapali’ × ‘Tommy Atkins’), with excel-
lent fruit quality; and ‘Lita’ (‘Amrapali’ × ‘Tommy Atkins’), high yielding
with excellent fruit quality.
South Africa
The South African breeding programme at the Citrus and Subtropical Fruit
Research Institute (CSFRI) is based on introductions, open-pollination and
mass selection. Four new cultivars have been released: ‘Heidi’, ‘Neldawn’,
‘Neldica’ and ‘Ceriese’. In addition, 12 promising selections have been iden-
tified for further evaluation (Marais, 1992).
4.10 Mutations
Somatic mutations
a month earlier (Young and Ledin, 1954). ‘Rosica’ from Peru, is a bud mutant
of ‘Rosado de lca’. Unlike its parent, ‘Rosica’ is high yielding and regular
bearing, and does not produce seedless fruits (Medina, 1977).
Oppenheimer (1956), after a survey of many orchards in India, reported
wide variability in the performance of trees of the same clone within a single
orchard. Mukherjee et al. (1983) conducted a survey of mangoes in eastern
India and identified some superior clones. Singh and Chadha (1981), in a
study of orchards of ‘Dashehari’, located four clones which were superior in
performance. Singh et al. (1985) isolated two high-yielding clones from
orchards of ‘Langra’. Within ‘Kensington’, strains have also been identified
that show improved resistance to bacterial black spot (Whiley et al., 1993).
Roy (1950) observed a mutant of ‘Alphonso’ with respect to fruit shape,
and suspected it to be a mericlinal chimera. Pandey (1998) has described
seven clones of ‘Alphonso’: ‘Alphonso Behat’ and ‘Alphonso Bihar’ from Bihar,
‘Alphonso Batli’, ‘Alphonso Black’ and ‘Alphonso Bombay’ from Maharashtra,
‘Alphonso Punjab’ from Punjab and ‘Alphonso White’ or ‘Bili Ishada’ from
the North Canara district of Karnataka. Rajput et al. (1996) assembled several
‘Dashehari’ variants and after 14 years of observation, reported that the clone
‘Dashehari 51’ was superior with respect to yield and regular bearing. Other
somatic mutants include: ‘Cardozo Mankurad’ with large fruits of attractive
colour and high yields from ‘Mankurad’ of Goa; dwarf selections from the
‘Rumani’ and ‘Bangalora’ (Ramaswamy, 1989); development of ‘Paiyur’, a
dwarf selection from ‘Neelum’ (Vijaya Kumar et al., 1991); ‘Rati Bangana-
palli’ and ‘Nuzuvid’ from ‘Banganapalli’ (Anonymous, 1999); and ‘MA-1’,
regular bearing and high yielding with resistance to ‘spongy tissue’ from
‘Alphonso’ (Mukunda, 2003).
In Thailand, Chaikiattiyos et al. (2000) selected clone ‘SKoo7’ (now known
as ‘Kaew Sisaket’) from 320 ‘Kaew’ plants; ‘SKoo7’ has higher yield and
superior quality. Jintanawongse et al. (1999) also made superior selections for
yield and fruit quality from ‘Nam Dok Mai’, ‘Khiew Sawoey’, ‘Rad’ and
‘Nang Klang Wau’ and DNA fingerprints of all these clones were made for
comparison with the parental clone.
For these studies, it is important to conduct a replicated cultivar evalua-
tion trial against standard commercial cultivars to establish that these varia-
tions are stable and not due to environmental responses. The use of genetic
markers should be explored to confirm that the new clones are genetically
distinct from the original cultivar.
Induced mutations
seedlings, and found that dosages above 5 kR are lethal for mango and that
the lethal dose required for 50% mortality (LD50) lies between 2 and 4 kR.
Effective dosages of the chemical mutagens, ethane methyl sulfonate (EMS)
and N-nitroso methyl urea (NMU), were 1.5 and 0.05%, respectively. The
spectrum of mutations induced by physical and chemical mutagens was
observed to be more or less the same, indicating the high sensitivity of certain
loci. The mutants included dwarfness, changes in shape and serration of
leaves and in TSS content in ‘Dashehari’. As in other perennial crops, muta-
genesis techniques that can allow useful traits to be targeted, as well as
isolating mutated sectors from a chimera, are essential.
Bompard (1993; see Bompard, Chapter 2, this volume) has made a compre-
hensive study of the wild Mangifera species and enumerated their potential
use in breeding. Mangifera laurina, which has subglabrous and laxly flowered
panicles and is well adapted to areas with perpetual wet climates, is resistant
to anthracnose. Mangifera orophila from Malaysia and M. dongnaiensis from
Vietnam are both restricted to mountain forests 1000–1700 m above sea level
and their hybrids with mango could extend cultivation into temperate zones.
Mangifera magnifica is completely free of fibres; M. rufocostata and M.
swintonioides have an off-season bearing habit; M. pajang (endemic to Borneo)
and some strains of M. foetida have good quality fruits. Similarly ‘Wani’ from
Bali and Borneo, the best variety of M. caesia, has a distinctive taste. Mangifera
casturi from South Kalimantan is a prolific bearer with small, black, sweet
fruits having good potential. Mangifera altissima is reportedly resistant to
mango pests, such as hoppers, tip borers and seed borers (Angeles, 1991).
Sharma and Choudhury (1976) observed that wild Mangifera trees identified
in Tripura State (north-eastern India) were free of mango malformation. The
wild species could also contribute to higher productivity. Fairchild (1948)
observed that crosses between five-stamened mango and the Indian mango
(only one fertile stamen) could produce hybrids having better pollinating
quality. The interspecific compatibility of these species with M. indica must
be verified before they can be utilized in hybridization programmes (as sug-
gested by Bompard, 1993; Kostermans and Bompard, 1993; see Bompard,
Chapter 2, this volume).
4.12 Conclusions
and more rapid screening of hybrid populations have enabled the release of
many hybrid mango cultivars of commercial value. Because of the world
market’s demand for mangoes with specific qualities, the synthesis of new
cultivars has become imperative. Rapid strides in molecular biology and in
other aspects of biotechnology have opened up new approaches in plant
breeding. The development of polymerase chain reaction (PCR)-based
genetic markers, specifically microsatellites, and their application to classical
breeding offer tremendous potential for mango improvement. The develop-
ment of a saturated linkage map and the identification of QTL for important
traits will allow the implementation of a MAS programme. The introduction
of specific genes for disease resistance from cultivars and wild species into
popular cultivars should soon be a reality. Without resorting to these new
technologies, mango breeding will continue to be a slow process.
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5 Reproductive Physiology
T.L. Davenport
University of Florida, Florida, USA
5.1 Introduction 98
5.2 Phenology 99
5.3 Shoot Development 100
Vegetative shoots 102
Reproductive shoots 104
5.4 Flowering Mechanisms 105
Shoot initiation 105
Induction 106
Florigenic promoter (FP) or stimulus 108
Vegetative promoter (VP) 110
5.5 Environmental Influence on Vegetative and Reproductive Development 111
Temperature 111
Water relations 113
Effect of N on flowering 114
Photoperiod 116
5.6 Hormonal Influence on Flowering 116
Ethylene 116
Auxin 117
Cytokinins 118
Gibberellins 119
Plant growth retardants 121
5.7 Photoassimilate Influence on Flowering 123
5.8 Horticultural Manipulation of Flowering 123
5.9 Conceptual Flowering Models 124
Carbohydrate-regulated flowering models 124
Hormone-regulated flowering models 127
5.10 Floral Management 133
5.11 Floral Biology 134
Sex ratio 134
Environmental determinants of sex ratio 134
Physiological determinants of sex ratio 135
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 97
98 T.L. Davenport
5.1 Introduction
Flowering and fruit set are the most critical of all events occurring after estab-
lishment of a tree crop. Given favourable growth conditions, the timing and
intensity of flowering greatly determine when and how much fruit are pro-
duced. Many important details about flowering are becoming clearer, espe-
cially in herbaceous plants, at the physiological, biochemical and molecular
levels (see reviews by Searle, 1965; Zeevaart, 1976, 2006; Bernier et al., 1981,
1993; Halevy, 1985–1986; Bernier, 1988; Kinet, 1993; Boss et al., 2004; Komeda,
2004; Putterill et al., 2004; Corbesier and Coupland, 2005).
Cool temperatures in the subtropics stimulate mango flowering and age
of the last vegetative flush has an important bearing on its ability to flower in
marginally cool or warm temperatures of the tropics (van der Meulen et al.,
1971; Davenport, 2000, 2003). Consequently, mango flowering can be en-
hanced during its normal season or manipulated to occur at other times of
the year in the tropics. For example, potassium nitrate (KNO3) can stimulate
out-of-season flowering in mangoes in tropical latitudes (Barba, 1974; Núñez-
Elisea, 1985; Davenport, 1993; Protacio, 2000), although this treatment has
not always been dependable. Various aspects of mango flowering and/or
fruit set have been reviewed (Singh, 1958a, 1979; L.B. Singh, 1960, 1977;
Chacko, 1986, 1991; Chadha and Pal, 1986; Davenport, 1993, 2000, 2003; Dav-
enport and Núñez-Elisea, 1997; Singh et al., 2005), and M.J. Soule (1950) pub-
lished an extensive annotated bibliography of the older literature related to
mango reproduction.
Understanding mango flowering is essential to efficiently utilize man-
agement systems that extend the flowering and crop production seasons.
Recent studies of mango flowering have resulted in conceptual models that
help explain the physiological basis of flowering (Chacko, 1991; Cull, 1991;
Kulkarni, 1991, 2004; Whiley et al., 1991; Davenport and Núñez-Elisea, 1997;
Davenport, 2000, 2003). Control of flowering allows growers to harvest their
crops at the most profitable times. Increasing the season of availability
improves competitiveness in the international marketplace, and promotes
the most efficient use of resources as costs of inputs continue to rise.
This chapter addresses the physiology of mango flowering, early fruit set
and retention. Cultivar names and type of embryony (i.e. monoembryonic or
Reproductive Physiology 99
5.2 Phenology
Growth of mango and other tropical trees is not continuous (Nakasone et al.,
1955; Halle et al., 1978; Verheij, 1986; Davenport, 1993, 2000, 2003). Apical
buds spend most of the time in rest. Growth occurs as intermittent, ephem-
eral flushes of shoots from apical or lateral buds (Naik and Mohan Rao, 1942;
Singh, 1958a, b). Stems are quiescent or resting terminal vegetative structures
on branches from which shoot growth occurs. Shoots are elongating vegeta-
tive or reproductive structures that emerge from apical or lateral buds of
stems. Vegetative shoots develop a prescribed number of nodes during
growth before entering a resting state as a stem. Depending on environment,
periods of stem rest are generally short in young plants but usually last sev-
eral months between episodes of growth in mature trees. Vegetative growth
generally occurs up to three or four times a year on individual branches,
depending upon cultivar and growth conditions.
Development of the vegetative shoot from initiation of growth to full
elongation requires 3–6 weeks, depending on the cultivar and climatic condi-
tions (Whiley et al., 1991). During this period, 10–20 new leaves are generally
produced before returning to a resting state. These rhythmic episodes of
extension growth are recorded on each branch as segments consisting of
compressed internodes interspersed with long internodes, that is articulate
growth (Tomlinson and Gill, 1973). Davenport (1992, 2003, 2006) referred to
regions of compressed internodes as intercalations and the entire segment of
long internodes terminating in an intercalation as an intercalary unit. The
number of intercalations between each branching point indicates the number
of vegetative growth episodes or flushes that have occurred between each
flowering flush.
Flushes of vegetative growth occur on groups of stems borne on scaffold-
ing branches in isolated sections of tree canopy. Flushing stems are usually
connected at some common branch point within the tree limbs. Asynchro-
nous flushes of growth at various times in random portions of a tree canopy
may appear to be continuous growth but are simply flushes occurring in
various parts of the total canopy over time. Flowering flushes generally occur
after extended periods of stem rest in the low-latitude tropics or during cool
winter months in the high-latitude tropics and subtropics. Like vegetative
flushes, reproductive flushes are usually asynchronous in tropical climates
100 T.L. Davenport
Lateral meristematic
primordia Apical dome
(includes meristem)
Fig. 5.2. Stylized cross-section of apical bud showing positions of apical meristem,
lateral meristems and leaf primordia.
Stem
Fig. 5.3. Axillary bud of resting mango stem. Leaf petioles (arrows).
begin to elongate and branch at each node forming secondary, tertiary and
quaternary lateral meristems. Each branch point in the lateral inflorescence
from the panicle axis to the floral pedicels bears a floral bract (i.e. partially
developed vestigial leaf primordium) (Fig. 5.4). The distal half of the panicle
structure is derived from newly formed nodes laid down by cell divisions in
the apical meristem prior to returning to a resting state. Mixed shoots, bear-
ing both leaves and inflorescences at each node, result from development of
both the primary leaf primordia and the lateral meristems, which form the
inflorescences in the same nodes as leaves.
Vegetative shoot induction, thus, involves stimulating development of
leaf primordia from resting buds while repressing development of lateral
meristems. Leaf primordia then follow a predetermined cascade of genetic
signals resulting in leaf development at each node. Because all shoots emerge
from resting buds, a vegetatively induced event does not involve simply
inhibition of flowering. The putative inductive signal directing differentia-
tion of leaf primordia onto leaves upon initiation is termed a vegetative pro-
moter (VP) rather than a floral inhibitor.
Shoots bearing only inflorescences (generative shoots) result from induc-
tive development of lateral meristems and suppression of leaf primordial
development. A predetermined cascade of flowering gene signals is activated
in lateral meristems resulting in lateral cymose inflorescences terminating
with flowers. A distinct florigenic promoter (FP) may be responsible for spe-
cific activation of the lateral meristems of mango. Mixed shoot induction
results in combined development of leaf primordia and lateral meristems.
Vegetative shoots
Vegetative shoots bear only leaves (Fig. 5.5). The anatomy of mango vege-
tative shoot development has been described (Singh, 1958b; Chaikiattiyos
Reproductive Physiology 103
3°
2°
º
4° 1°
5°
1 Bract
1°
Axis
1°
Pedicel
1° Bract
1° 3° Bract
Pedicel
2° Bract
2°
Pedicel 1°
1° Bract 1° 2° Bract Pedicel
Axis Pedicel
Fig. 5.4. Diagram and photos of mango inflorescence depicting the panicle axis and
primary (1°), secondary (2°) and succeeding levels of pedicel and cymose floral archi-
tecture. Vestigial leaf promorida (floral bracts) are depicted at the base of each level of
pedicel architecture.
et al., 1994). Vegetative shoots may arise either from axillary buds, if no apical
bud exists due to flowering in the previous flush, or from the apical bud
when present. The latter is considered extension growth or addition of an
intercalary unit on the existing stem, but the developmental events during
shoot formation from either apical or lateral buds are basically the same.
Cells in the leaf primordia of initiating buds begin to form individual leaves
in the proximal portion of the vegetative shoot. Soon thereafter, the apical
meristem activates to form more nodes bearing leaf primordia and lateral
meristems. These newly formed leaf primordia develop as the distal portion
of the vegetative shoot if environmental conditions remain vegetatively
inductive (Núñez-Elisea et al., 1996). Newly elongating vegetative shoots are
green in most cultivars but may be bronze or red in others. Fully expanded
104 T.L. Davenport
Fig. 5.5. Stylized diagrams and photomontage of shoot types found in mango. Transition
shoots shift from vegetative to floral (V/F) or floral to vegetative (F/V). Arrow ( )
represents individual leaves; floral diagram ( ) represents lateral inflorescences.
leaves are a shade of red, depending upon cultivar and cultural conditions
and are thin and limp from lack of lignification. The apical buds of vegetative
shoots generally become quiescent before completion of the limp, red-leaf
stage (Núñez-Elisea and Davenport, 1995). Internodes are compressed at the
apex, and leaf development is arrested thereby forming a bud with protec-
tive outer scales, inner leaf primordia, lateral meristems and the apical mer-
istem. Fully expanded leaves become light green and stiff as they become
lignified and suberized. Vegetative shoots are mature when leaves become
dark green, which occurs when they are c.2 or 3 months old.
Reproductive shoots
1958b; L.B. Singh, 1960; Sturrock, 1966; Ravishankar et al., 1979; Scholefield,
1982; Scholefield et al., 1986). The complexes of primary to quaternary branch-
ing lateral structures of the inflorescence each terminate with three cymose
flowers. The terminal flower opens first, followed by two subtending lateral
flowers. These complexes form the lateral inflorescence structures emerging
from the central axis of the panicle. The central axis extension also terminates
in a similar fashion. Morphological stages of floral buds and panicle develop-
ment were described by Shu (1981) and Oosthuyse (1991a). Reece et al. (1949)
described the development of inflorescences initiated in lateral buds when
the terminal bud is missing. There are more nodes in dormant apical buds
and their bracts are more developed than in axillary buds; however, floral
evocation is indistinguishable.
Generative shoot development in apical buds initially involves swelling
of the lateral meristems and their bud scales. Each axillary meristem devel-
ops as an inflorescence on a primary peduncle. The apical meristem then
forms new lateral meristems and leaf primordia for the distal portion of pan-
icle development if floral inductive conditions persist (Núñez-Elisea et al.,
1996). Panicles may be open or compact, depending upon internode elonga-
tion, which is cultivar dependent (L.B. Singh, 1960), but the architecture gen-
erally conforms to that in Fig. 5.5. Mixed shoots develop under weak floral
inductive conditions (i.e. in the low-latitude tropics). Both leaves and pri-
mary pedunculate inflorescences develop from the same nodes (Fig. 5.5).
Leaf primordia and lateral meristems develop as leaf and floral structures,
respectively.
Shoot initiation
also stimulates shoot initiation. Reece et al. (1946, 1949), Mustard and Lynch
(1946), Núñez-Elisea and Davenport (1992b), Núñez-Elisea et al. (1996) and
Davenport et al. (2006a) observed that the vegetative or reproductive fate of
mango buds remains undetermined until after shoot growth is initiated.
Reece et al. (1949) proposed that a putative signal that triggers initiation of
shoot development is separate and different from the inductive signal, which
determines the fate of the shoot. Removal of apical buds by pruning stimu-
lates initiation of axillary shoots (Singh and Singh, 1956; Núñez-Elisea and
Davenport, 1992b; Núñez-Elisea et al., 1996; Davenport et al., 2006a). Defolia-
tion of the apical whorl of five to ten leaves also stimulates shoot initiation
in dormant apical buds (Núñez-Elisea et al., 1991; Núñez-Elisea and Daven-
port, 1995). The fate of shoots that emerge in response to these initiation stim-
uli, however, is determined by other factors that are prevalent at the time of
initiation. Tip pruning, for example, during warm summer months results in
initiation of vegetative shoots from axillary buds, whereas pruning during
cool winter months usually results in initiation of axillary inflorescences.
Induction
Target cells within leaf primordia and lateral meristems are competent to
respond to inductive signals; for example when initiated to grow under veg-
etatively inductive conditions, individual leaf primordia develop as leaves
and subtending lateral meristems associated with each developing leaf
develop as dormant axillary buds with protective bracts. These axillary buds
may develop in subsequent flushes as vegetative shoots when initiated in
vegetatively inductive conditions or as axillary inflorescences under floral
inductive conditions. Under strongly floral-inductive conditions, leaf pri-
mordia fail to develop beyond the bract stage, become dormant, and lateral
meristems develop. Each lateral meristem forms nodes consisting of leaf pri-
mordia and meristems that are influenced by the putative floral-inductive
stimulus, which suppresses development of newly formed leaf primordia.
Subsequently formed meristems form pedunculate structures that terminate
in cymose inflorescences borne on each tertiary peduncle (Fig. 5.4). Forma-
tion of the primary, secondary, tertiary and quaternary peduncles, as well as
pedicels of inflorescences are always accompanied by a subtending, aborted
bract or vestigial leaf at each node (Fig. 5.4). Such development is attributed
to a sequence of gene expression (Coen et al., 1990; Coen and Meyerowitz,
1991; Weigel et al., 1992; Coen and Carpenter, 1993; Lumsden, 1993; Yanofsky,
1995). Shoot initiation during weakly floral-inductive conditions activates
growth of leaf primordia to develop leaves and the lateral meristems to pro-
duce peduncles bearing lateral inflorescences in each node of mixed shoots.
The bases of each pedicel branch within each lateral inflorescence also bear a
vestigial leaf.
Upon termination of cell divisions in the apical meristem at the end of a
flushing period, no more nodes are formed. The apical bud of vegetative
shoots becomes quiescent, and the resting leaf primordia, bracts and lateral
meristems are poised to resume growth at a later date. When reproductive or
mixed shoots become quiescent, the lateral meristems ultimately develop
determinant cymose inflorescences. The most distally located meristem is
possibly the determinant extension of the central axis forming the terminal
cymose floral group.
Chimeric shoots (Fig. 5.5) can occur in mango trees when shoot initiation
occurs during floral inductive conditions. They display inflorescences on one
side of the longitudinally bisected shoot and leaves on the other. The shoot
axis is red on the floral side of red fruiting cultivars (typical of panicles) and
green on the vegetative side (typical of vegetative shoots). This difference
in the two sides extends to the apical bud, which bears an undeveloped
inflorescence on the floral side and leaf bracts on the vegetative side. The
explanation for this spatial differentiation is that target nodes on each side of
the apical bud respond to the different inductive signals at the same time.
The apical meristem is not implicated except to form more nodes for the lat-
eral inductive responses on each side in the second portion of growth. Differ-
ences in inductive signals on each side of an existing shoot probably cause
the differential response. This phenomenon indicates that the fate of nodes
on each side of the shoot cannot be attributed to a single mother cell in the
apical meristem. The inductive response must involve cells formed in later
108 T.L. Davenport
Early flowering work provided evidence for the presence of a graft transmis-
sible floral stimulus (i.e. florigen) that was induced in leaves and was trans-
located to buds to stimulate floral development (Chailakhyan, 1936; Zeevaart
and Boyer, 1987). Florigen was functionally conserved across plant species
(Lang, 1965, 1984; Zeevaart, 1976; Lang et al., 1977). Floral induction in most
plants involves sensing of some environmental cue (i.e. daylength, water
stress or vernalizing temperature) in some organ (e.g. leaves). A putative flo-
ral stimulus or alteration in the ratio of florigenic to anti-florigenic compo-
nents may be translocated to target cells in meristems (Bernier et al., 1981).
Photoassimilate movement from leaves in phloem facilitates its transport to
buds where it can interact to initiate flowering (King and Zeevaart, 1973).
Until recently, a floral stimulus could not be identified. Alternative hypoth-
eses were proposed that nutrient diversion to the meristems could be
involved (Sachs and Hackett, 1983) or that floral induction might be con-
trolled by multiple factors, including the putative floral stimulus, photoas-
similates and phytohormones (Bernier et al., 1993).
Molecular biology of flowering in the facultative, long-day, model plant,
Arabidopsis thaliana (reviewed in Zeevaart, 2006 and Aksenova et al., 2006),
has provided insight into the nature of the floral stimulus (FP). A network of
four interacting genetic signalling pathways may result in flowering in
response to photoperiodic, vernalization, gibberellin and autonomous envi-
ronmental cues (Perilleux et al., 1994; Mouradov et al., 2002; Perilleux and
Bernier, 2002; Boss et al., 2004; Komeda, 2004; Putterill et al., 2004; Corbesier
and Coupland, 2005). The photoperiodic pathway involves activation of
the CONSTANS (CO) gene that encodes a zinc-finger protein, which in
turn induces expression of the FLOWERING LOCUS T (FT) gene in the
phloem tissue of leaves. FT is the terminal, integrating gene of the four path-
ways regulating flowering in Arabidopsis. Its transcribed mRNA was initially
thought to be the FP that is transported in phloem to buds (Huang et al.,
2005); however, evidence indicates that the translated protein product of FT
is translocated to Arabidopsis buds (Corbesier et al., 2007). Analogous proteins
encoded by Hd3a, an ortholog of FT in rice (Tamaki et al., 2007), and the aspen
ortholog, PtFT1, which along with CO regulates the timing of flowering and
growth cessation of Populus trichocarpa (Bohlenius et al., 2006), appear to be
the FP. In the buds, the protein product of FT is thought to combine with the
bZIP transcription factor (FD) protein to activate transcription of floral iden-
tity genes (i.e. APETALA1) to begin floral expression (Abe et al., 2005; Wigge
et al., 2005). Similar mechanisms are likely to exist in mango.
Zhang et al. (2005) and Davenport et al. (2006b) isolated a CONSTANS-
like gene (MiCOL) from mango leaf DNA. CO is a circadian expression gene
interacting with the photoperiodic pathway in Arabidopsis (Putterill et al.,
Reproductive Physiology 109
(PBZ) reduces the time in rest necessary to allow floral induction during
warm temperature conditions by c.1 month (Davenport, 2003), thus increas-
ing the potential to produce reproductive shoots in younger stems when ini-
tiated to grow. PBZ and uniconazole, triazole compounds that inhibit kaurene
oxidase in the gibberellin-synthesis pathway (Dalziel and Lawrence, 1984;
Rademacher, 1991), stimulate production of flowering shoots during weakly
inductive conditions (Burondkar and Gunjate, 1991, 1993; Tongumpai et al.,
1991a; Voon et al., 1991; Nartvaranant et al., 2000; Yeshitela et al., 2004a).
Application of PBZ to mango trees bearing 1-month-old stems produced
inflorescences when bud break was initiated 3 months later by foliar applica-
tion of KNO3 (Davenport, 2003).
Vegetative or reproductive induction at the time of shoot initiation is
governed by the ratio of the putative floral promotive to inhibitory compo-
nents (Lang et al., 1977; Lang, 1984; Kulkarni, 1988a; see Bernier et al., 1981 for
additional references). The mango floral inhibitor should be viewed as an
age-dependent VP. The presence of an age-regulated VP in mango leaves,
which moves with the temperature-regulated FP and photoassimilates in
phloem, may explain the induction of specific receptors by this promoter in
targeted leaf primordia to cause development of leaves in vegetative or
mixed shoots. A gradual decrease in the level or influence of the VP may
cause vegetative shoots to develop when initiation occurs on 2-month-old
stems, and generative or mixed shoots when initiation occurs in stems from
4- to 7-month-old stems, given the constantly warm daily temperatures
maintaining a low level of FP in both situations.
Temperature
Water relations
In the absence of cool temperatures, mango trees in the tropics may flower in
response to irrigation or rain following periods of water stress lasting 6–12
weeks or more (Pongsomboon, 1991). Plant water stress has been presumed
to provide the stimulus for flowering (reviewed in Whiley, 1993; Chaikiatti-
yos et al., 1994; Schaffer et al., 1994; Davenport and Núñez-Elisea, 1997); how-
ever, most of these studies have failed to substantiate prolonged tree water
deficit as a successful agent for floral induction.
Experiments with container-grown trees fail to produce inflorescences
after 8 weeks of water deficit (Wolstenholme and Hofmeyr, 1985). Under
glasshouse conditions (27°C day/22°C night; relative humidity (RH) ≥ 90%),
container-grown, monoembryonic cultivars were water stressed through
deficit irrigation for 14 days, resulting in an average leaf xylem water poten-
tial of −3.9 MPa (Davenport, 1992; Núñez-Elisea and Davenport, 1992a,
1994b). Following resumption of irrigation, all trees grew vegetatively. Sim-
ilarly, only vegetative growth was obtained when container-grown trees
were deprived of irrigation for 36 days during summer, although leaf xylem
water potentials of −3.78 MPa were attained (Núñez-Elisea and Davenport,
1994b). Water stress imposed on plants during the cool autumn months
114 T.L. Davenport
(night temperatures < 15°C) do not increase the proportion of apical buds
forming inflorescences, but expedited shoot initiation after rewatering
(Núñez-Elisea and Davenport, 1994b). These results demonstrated that cool
temperatures provide inductive conditions, whereas relief of water stress
accelerated shoot initiation under cool, inductive temperatures. Flowering
was delayed when container-grown monoembryonic mangoes were water-
stressed at 18°C day/15°C night (Chaikiattiyos et al., 1994). Water-stressed
trees held at 29°C day/25°C night did not flower.
Mango trees growing in the low-latitude tropics may flower after an
extended period of mild water stress (Harris, 1901; Collins, 1903; Kinman,
1918; Gangolly et al., 1957; Gangolly, 1960; L.B. Singh, 1960). Pongsomboon
et al. (1991) observed flowering in field-grown trees in the tropics following
6 weeks of withholding water. The primary impact of water stress appears to
be prevention of shoot initiation during stress. The accumulating age of stems
is greater in water-stressed trees than in trees maintained under well-watered
conditions that promote frequent vegetative flushes (Davenport, 1992, 1993;
Schaffer et al., 1994). This delay in flushing may provide more time for accu-
mulation of a putative FP (Schaffer et al., 1994) or reduction in the level of a
putative VP (Davenport and Núñez-Elisea, 1997; Davenport, 2000). Some
cultivars appear to be better adapted to such delays in growth and perform
better in dry environments in the tropics.
Effect of N on flowering
Chemical bud forcing is most effective in the tropics where distinct wet
and dry seasons prevail. The response to chemical bud forcing by NO3− and
ethephon diminishes at latitudes > 22° N or S (Mosqueda-Vázquez and de
los Santos de la Rosa, 1981). Their effect may involve the decline of night
temperatures from ≥ 20°C around the equator to ≤ 10°C between 22° and 27°
N or S latitude during winter months or by late summer vegetative flushes.
Trees in the wet or dry subtropics at 25° N or S have not responded to treat-
ments (Davenport, 1993).
Stems must be sufficiently mature, dark green with a minimum age of 4
months since the previous limp, red-leaf stage in easily induced cultivars
and 5 months for more recalcitrant cultivars to obtain a reproductive shoot
response in the low-latitude tropics (Davenport, 2003). Bueno and Valmayor
(1974) indicated that leaves must be brittle when hand-crushed. Núñez-
Elisea (1986, 1988) reported that stems must be at least 6 months old. Trees
that experience autumn dry periods become responsive to treatments as
early as October (northern hemisphere). Groups of stems within tree cano-
pies are produced through asynchronous flushes of growth, and vary in age;
only a few are responsive to the first inductive spray. Subsequent biweekly
applications cause flowering in canopy sectors as they reach the age-depen-
dent requirement for initiation. Early and out-of-season flowering and fruit-
ing can thereby be achieved.
KNO3 may be floral inductive in mango (Barba, 1974); however, trees in
the upper latitude tropics typically flush vegetatively rather than produce
bloom when either KNO3 or NH4NO3 is sprayed between June and Septem-
ber (N. Golez, personal communication, the Philippines, 1989). The warm,
rainy season producing frequent flushes of growth during this period is con-
ducive to a vegetative response to the sprays. These results indicate that
KNO3 and NH4NO3 stimulate shoot initiation but do not determine bud
morphogenesis. In buds released after KNO3 or NH4NO3 treatments, the
ratio of leaf-generated FP to VP and not NO3– causes initiating buds to become
reproductive. Kulkarni (1988b, 2004) suggested that the floral stimulus is
present in stems when buds are forced in response to KNO3 and suggested
that KNO3 may also sensitize buds to the floral stimulus. Davenport (2003),
T.L. Davenport and J. Oleo (2006, unpublished data) and F. Ramirez and T.L.
Davenport (submitted for publication) observed 100% vegetative shoots
when 4% KNO3 was foliar applied to 2-month-old stems; whereas, applica-
tion of the same spray treatment to 4.5-month-old stems on trees in the same
orchards resulted in 100% reproductive shoots.
Trees with high leaf N levels rarely flower in the tropics. Lack of flower-
ing is always due to frequent vegetative flushes of growth, especially during
the rainy season. Mango trees must have leaf N levels of 1.4% or less in order
to suppress frequent flushes of vegetative growth (Davenport, 2003). Leaf N
levels of < 1.1% suppress frequent flushes but also provide insufficient nutri-
tion to support good cropping. Thus, 1.1–1.4% N levels in leaves appear to be
optimum for good commercial production and control of flowering time in a
managed orchard. The application of KNO3 to the foliage of the resting stems
4–5 months after the limp, red-leaf stage will cause a flowering response.
116 T.L. Davenport
Photoperiod
Ethylene
Auxin
Although auxin may have a critical role in floral induction of mango (Chadha
and Pal, 1986; Hegele et al., 2006), there is little supporting evidence. The
application (L.B. Singh, 1961; Singh and Singh, 1963; Bakr et al., 1981; Pandey
and Narwadkar, 1984) and analysis of auxin in leaves (Paulas and Shanmu-
gavelu, 1989; Sivagami et al., 1989), stems (Chen, 1987) and shoots (Chacko et al.,
1972b) have been reported in relation to mango flowering. These studies are
inconclusive due to inconsistencies in purification and analytical methodolo-
gies (Davenport and Núñez-Elisea, 1997).
Auxin may indirectly stimulate root-produced cytokinins through initia-
tion of new root growth. Auxin is transported basipetally from growing
118 T.L. Davenport
shoots and leaves to roots (Goldsmith, 1968; Cane and Wilkins, 1970; Wilkins
and Cane, 1970; Goldsmith and Ray, 1973; Lomax et al., 1995) and stimulates
root initiation (Hassig, 1974; Wightman et al., 1980). The efficacy of various
auxins for stimulating adventitious rooting of mango marcots and cuttings
was reviewed by Davenport and Núñez-Elisea (1997).
Auxin inhibits shoot initiation (Davies, 1995) and confers apical domi-
nance by preventing axillary bud break. Leaf-produced auxin and petiolar
auxin transport capacity declines as leaves age (Veen, 1969; Veen and Jacobs,
1969; Davenport et al., 1980). The interaction of decreasing auxin and accu-
mulating cytokinins in resting buds may explain the cyclic nature of shoot
initiation. The ratio of cytokinin to auxin levels in buds regulates shoot
initiation (Skoog and Miller, 1957; Bangerth, 1994; Cline et al., 1997; Beveridge
et al., 2003).
Cytokinins
Gibberellins
should utilize plants grown under defined conditions with specific environ-
mental controls for evaluation of cause and effect. Finally, extraction and
purification protocols should include quantifiable internal standards and
use of sensitive unambiguous analytical techniques.
Triazoles
PBZ is being used (except in the USA where it has not been cleared for use)
to stimulate enhanced or early flowering. It is best applied to the soil due to
its low solubility, long residual activity and lack of efficient foliar uptake
(Rademacher, 2000b). PBZ applied as a soil drench (1–20 g active ingredient
(ai)/tree) reduces internode lengths and causes earlier and enhanced flower-
ing in mango trees (Hasdiseve and Tongumpai, 1986; Haw, 1986; Hongsb-
hanich, 1986). Depending on climate, residual activity lasts for c.2 years
(Kulkarni, 1988a). These results have been confirmed in different locations in
the tropics (Davenport and Núñez-Elisea, 1997; Yeshitela et al., 2004a, b).
Nartvaranant et al. (2000) recommended soil application of PBZ at 1–1.5 g
ai/m of canopy diameter to achieve flowering in 90–120 days if the trees are
stimulated to flush. Davenport (2003) observed that such treatments allowed
a reduction of c.1 month in the time required for stem rest before stimulating
them to initiate reproductive shoots using KNO3. PBZ also reduces alternate
122 T.L. Davenport
bearing of some cultivars (Hillier and Rudge, 1991; Burondkar and Gunjate,
1993; Rao, 1997; Rao et al., 1997; Rao and Srihari, 1998; Vijayalakshmi and
Srinivasan, 1999). Cultivars that tend to flower with minimal inductive impe-
tus are more responsive and can be induced to flower out-of-season using
PBZ (Tongumpai et al., 1989). Núñez-Elisea et al. (1993) demonstrated that
application of PBZ and uniconazole advanced bud break of containerized
trees in controlled environment chambers, but cool temperatures were neces-
sary to induce flowering. Initiated shoots were induced to be vegetative in
warm temperatures. The greater proportion of purely reproductive panicles
in treated plants (compared with controls) suggests that triazoles impact the
level of a putative VP, probably a gibberellin. Whiley (1993) suggested a sec-
ondary mechanism for the floral promotive action of PBZ on mangoes, not-
ing inconsistent responses in the literature between cultivars, environments
and application times.
Application of PBZ reduces the number of panicles, despite increased
fruit set (Goguey, 1990). Davenport (1987, 1994) observed neither growth
inhibition nor enhanced or early flowering in response to root drenches or
bark banding with uniconazole (1–5 g ai/tree) in trees growing in alkaline,
calcareous soil. He reported that new shoot growth was stunted with
extremely short internodes when trees were severely pruned soon after or as
long as 3 years after treatment. Yield was severely reduced due to the lack of
normal growth flushes. The growth stunting effect continued for 7 years after
pruning. Davenport (1994) warned that use of triazole plant growth retar-
dants for control of tree growth, flowering or yield must be done with con-
siderable caution, especially if severe pruning of the trees is anticipated.
Residual uniconazole or PBZ applied as a soil drench or bark band is appar-
ently retained in high concentrations in main scaffolding branches. In Cen-
tral and South America, growers utilize PBZ annually to stimulate early
flowering. A test tree should be severely pruned to determine if the trees are
affected by PBZ to anticipate the orchard response to later severe pruning.
Certain gibberellins (i.e. GA1) are necessary for shoot elongation. Inhibi-
tion of bud break and shoot elongation in response to application of the
growth retardants cycocel (Maiti et al., 1972) and triazoles (Kulkarni, 1988a;
Burondkar and Gunjate, 1991, 1993; Tongumpai et al., 1991a; Kurian et al.,
1992; Winston, 1992; Kurian and Iyer, 1993a, b; Núñez-Elisea et al., 1993; Wer-
ner, 1993) have been reported. Elongation of panicles is inhibited, especially
by high levels of triazoles (Kulkarni, 1988b; Winston, 1992; Davenport, 1994;
Salomon and Reuveni, 1994). Inflorescences in treated trees may become
compact, improving opportunities for disease and insect attack (Winston,
1992). Kurian et al. (1992) associated reduced cytokinin levels in leaves with
inhibition of shoot initiation in plants treated with soil drenches of PBZ. Ele-
vated, concentration-dependent levels of phenolic compounds were also
found in resting apical buds of PBZ-treated trees (Kurian et al., 1994). They
suggested that low cytokinin activity and high phenolic levels in buds con-
tributed to inhibition of shoot initiation.
The combined impact of the gibberellin synthesis-inhibiting triazoles on
shoot initiation, induction, and elongation implies that several different
Reproductive Physiology 123
Flowering may be regulated by C:N ratios with high levels being conducive
to flowering (Kraus and Kraybill, 1918). Photoassimilates reaching the apical
bud from leaves was central to several theories of floral induction (Sachs,
1977; Bernier and Sachs, 1979; Bernier et al., 1981, 1993; Bernier, 1988) includ-
ing mango (Mallik, 1951; L.B. Singh, 1960; Chacko and Ananthanarayanan,
1982; Rameshwar, 1989) and other species (Allsopp, 1965; Sachs, 1977; Mishra
and Dhillon, 1978; Ramina et al., 1979; Bernier et al., 1981; Sachs and Hackett,
1983). The theory of photoassimilate diversion to the apical bud (Sachs et al.,
1979) is the basis for the carbohydrate-regulated flowering models (see
below). Sugars are utilized during panicle development (Ravishankar and
Mohan Rao, 1982). Starch reserves and C:N ratios have been correlated with
flowering (Mishra and Dhillon, 1978; Suryanarayana, 1978a, b, c; Chacko and
Ananthanarayanan, 1982; Whiley et al., 1988, 1989, 1991; Robert and Wolsten-
holme, 1992; Shivashankara and Mathai, 1995), and the subject has been
reviewed (L.B. Singh, 1960, 1972; Singh, 1979; Chacko, 1986, 1991; Chadha
and Pal, 1986; Pandey, 1989). Starch accumulation during extended periods
of canopy rest prior to flowering provides supportive evidence, but there is
little consensus regarding the role of carbohydrates and N in flowering.
Photoassimilates may be necessary for floral induction. If a florigenic
promoting gene product is synthesized in leaves in small amounts, it must be
able to move to those buds via phloem. Due to the requirement for high sol-
ute concentrations to motivate phloem flow, the low concentration of the FP
could not cause fluid movement through sieve tubes of the phloem on its
own. The much higher concentrations of photoassimilated sugars carried by
water loading into the phloem in leaves passively transports the FP towards
the various sinks, including respiring buds, where they are utilized for floral
induction.
Several conceptual models have been proposed that attempt to explain the
physiological basis of mango flowering. Each model should be viewed as a
collection of integrated ideas, which require rigorous testing for validity
within the context of the models. A useful model should explain how flower-
ing and vegetative growth is regulated in all cultivars and races in both
humid and dry climates in the tropics and subtropics. It should also be sup-
ported by the preponderance of research evidence. The flowering models are
either carbohydrate-regulated or hormone-regulated.
Cull (1987, 1991) presented a holistic approach for tree crop research and
management to maximize sustainable fruit production. This concept is based
Reproductive Physiology 125
High
· Stem girdling MISSING LINK
starch
· Root pruning
GROWTH
INCREASED ASSIMILATE
GROWTH STIMULATION · High temperature
ASSIMILATE supply DIVERSION
CHECK and high · High humidity
to SHOOT APEX from SHOOT APEX
gibberellin · High soil moisture
T.L. Davenport
· Mild nitrogen
Sugar
stress
High
nitrogen
· High reserves
· Growth · Low reserves
· Efficient
retardants · More wood
assimilate
· Inhibitors formation
partitioning
Frequent High
Dwarf/precocious Over vigorous
flushing of gibberellin
cultivars cultivars
roots and shoots levels
e.g. ‘Irwin’ e.g. ‘Kensington’
HEREDITY JUVENILITY
Fig. 5.6. Chacko’s Assimilate Supply and Diversion Flowering Model, a carbohydrate-regulated flowering model (Source: Chacko, 1991).
Reproductive Physiology 127
SHOOT FORMATION. Two distinct events must occur for vegetative or repro-
ductive growth to occur in resting apical or lateral buds of mango: (i) the
Reproductive Physiology 129
bud(s) must be initiated to grow (shoot initiation); and (ii) at the time of ini-
tiation, shoot development (i.e. vegetative, mixed, or generative) is deter-
mined (induction). Although conditions for floral induction may be present
prior to shoot initiation, determination of that inductive condition in buds is
not made until initiation occurs. Initiation and induction events are regulated
by different signals and each may be manipulated by different stimuli.
Removing the apical whorl of leaves or tip pruning physiologically mature
stems stimulates shoot initiation in apical or lateral buds, respectively. If
containerized plants are maintained in warm temperatures (30°C day/25°C
night) following initiation, vegetative shoot growth is induced. If they are
kept under cool conditions (18°C day/10°C night), initiating shoots are
induced to be generative. In either of the two temperature regimes without
pruning, they do not initiate shoots until the natural flushing event occurs
much later. They become vegetative or reproductive according to the tem-
perature at the time of shoot initiation. If transferred from cool to warm tem-
peratures before shoot initiation, new shoot growth is induced to be vegetative.
Induction is therefore determined at the time of shoot initiation, and plants
rapidly lose their floral inductive potential when removed from the cool envi-
ronment. Determination of shoot type can be reversed during morphogenesis
by transferring containerized trees from warm-to-cool or cool-to-warm con-
ditions (Batten and McConchie, 1995; Núñez-Elisea et al., 1996).
Henny, 1995). Auxin inhibits shoot initiation (Davies, 1995) and confers api-
cal dominance by preventing axillary bud break. Leaf-produced auxin and
petiolar auxin transport capacity declines as leaves age (Davenport et al.,
1980). Auxin and cytokinins may therefore be involved in the periodic cycle
of bud break.
A critical balance of these two phytohormones, possibly modulated by
GA3, may regulate shoot initiation. During a rest period, the inhibitory action
of auxin transported to buds decreases with time; whereas, cytokinin lev-
els in buds increase (Chen, 1987). When a critical cytokinin/auxin ratio is
achieved, the buds are stimulated to grow, thereby resetting the cycle with
initiation of new shoots. The interaction of auxin and cytokinin to control
bud break in plants is a concept that is supported by molecular studies (see
review by Nordstrom et al., 2004). Moreover, initiation of shoot growth occurs
following pruning, defoliation or the application of thidiazuron (Núñez-
Elisea et al., 1990). Vigorous cultivars (Whiley et al., 1989) and young, small
trees under vegetatively promotive conditions flush frequently with only
short periods of rest; however, this cycle slows with age. Old centennial trees
flush infrequently (N. Golez, personal communication, the Philippines, 1989).
Foliar or soil-applied NO3− stimulates initiation of reproductive shoots
only if applied after resting stems have attained an age to overcome any veg-
etatively inductive influence. In contrast, high N in soils leads to high N lev-
els in leaves resulting in frequent vegetative flushes. The mechanism whereby
NO3− stimulates shoot initiation is unknown.
Seeds are rich sources of auxin and gibberellins, which contribute to the
strong inhibition of bud break commonly observed on fruit-bearing mango
stems. The longer that fruit are attached to stems, the longer the postharvest
inhibition may last in the stem (Kulkarni and Rameshwar, 1989; Kulkarni,
1991).
Water stress inhibits shoot initiation by its direct impact on cell division
and elongation possibly by interfering with translocation of cytokinins from
roots. There is little evidence that water stress is directly involved in induc-
tive processes. During water stress, roots continue to grow and produce cyto-
kinins (Itai and Vaadia, 1965; Itai et al., 1968; Wu et al., 1994). Reduced xylem
flux due to limited soil hydration, and transpiration due to increased sto-
matal resistance during water stress may reduce the amount of cytokinins
reaching stems. After rewatering, the increased levels of cytokinins in roots
may translocate to and accumulate in buds. Auxin synthesis and transport
from leaves are reduced during water stress (Davenport et al., 1980) and may
require several days for correction after rewatering. This rapid shift in the
cytokinin/auxin ratio of buds may explain the shooting response that occurs
soon after relief of water stress. GA3 may act with auxin to inhibit shoot ini-
tiation (Davenport et al., 2001b). Early flowering in plants treated with PBZ
may be a response to lowered gibberellin levels, thus lowering the level of
initiation inhibitor.
This model could explain why sectors of tree canopies flush in the trop-
ics. Mango trees flush often and synchronously throughout the canopy when
they are young. With advancing age, the frequency of flushing is reduced
Reproductive Physiology 131
branches may occur at any time of the year in trees growing in low-latitude
tropics. High proportions of mixed shoots are commonly found in these con-
ditions, indicating the marginally floral-inductive ratios present under these
conditions. In contrast, flowering in younger stems having higher levels of
VP is observed only when initiation occurs in cool, floral-inductive tempera-
tures. More flowering occurs throughout the canopy when stems are exposed
to cool temperatures, attributable to the higher ratio of up-regulated FP to
resident VP.
Genetic differences in base levels of the putative FP and/or VP or the
receptors of these components could explain the range in flowering responses
in tropical and subtropical cultivars and why a cultivar grown in an environ-
ment different from that in which it was selected is less productive. Cultivars
selected in the subtropics usually flower as well in the low-latitude tropics as
those selected in the tropics. Cool temperatures in the subtropics sometimes
cause earlier flowering in tropical cultivars than those selected in the sub-
tropics. Kulkarni (1991) demonstrated that several multi-flowering cultivars
can induce flowering in receptor graft plants and cause a range of the flower-
ing response of the receivers to donors. Some cultivars may produce higher
base levels of putative FP than others. These are the same cultivars that read-
ily flower under warm temperatures and flower early during cool winter
months. The Comprehensive Conceptual Flowering Model suggests that
flowering can occur at any time in any cultivar regardless of origin so long as
stems are sufficiently old to reduce the VP level to below the critical FP/VP
ratio when initiation occurs.
Although the putative FP, perhaps a product of an ortholog of the Arabi-
dopsis FT gene, has not been identified, the VP may be a gibberellin. Triazoles
and other plant growth retardants that inhibit gibberellin synthesis, promote
strong and out-of-season flowering under conditions that would normally
be marginally or non-floral inductive.
flushes until close to the normal flowering period impact the flowering abil-
ity of young shoots. This may explain the occurrence of chronic alternate
bearing in some cultivars.
Sex ratio
Inflorescences that emerge during the middle and end of the flowering sea-
son produce two and seven times more perfect flowers, respectively, than the
early breaking inflorescences (Majumder and Mukherjee, 1961; Singh et al.,
1966). This response correlates with higher temperatures later in the flower-
ing season. In controlled-environment studies, low temperatures (15°C
day/10°C night) reduced the proportion of perfect flowers, particularly in
tropical, polyembryonic cultivars relative to subtropical, monoembryonic
cultivars (Sukhvibul et al., 1999).
et al., 1994). If only one or two fruits were set on each terminal, the tree would
carry an unusually heavy crop. It is unlikely that reduced perfect flower
numbers due to cool temperatures during inflorescence development is
directly responsible for poor fruit set and yields. Pollen viability, growth and
ovule fertilization are probably the main factors contributing to low fruit set
under these conditions.
Pollen
Pollen grains are 20–45 Pm long and are oblong when dry and more
spherical when hydrated (Popenoe, 1917; Jivanna Rao, 1923; Bijhouwer, 1937;
Reproductive Physiology 137
Pollination
Wind
Early investigators concluded that the species is wind pollinated (Hartless,
1914). Initially wet pollen dries to a powdery consistency on anthers soon
after anthesis in dry conditions (Pimentel et al., 1984), whence it is likely to be
liberated in moving air or via gravity to adjacent stigmas on the same and
138 T.L. Davenport
nearby flowers (Naik and Mohan Rao, 1943; Mallik, 1957). Singh (1954a) and
S.N. Singh (1961) suggested, however, that the amount of pollen moving in
air streams was too low for wind to be a pollination vector. They did not
report the location of pollen-collecting slides or take into account the close
proximity of flowers within inflorescences or numbers of open flowers in the
canopy. Panicles bagged to exclude pollinating insects were reported to set
fruit (Free and Williams, 1976), which were retained to maturity, thereby con-
firming that mango pollen can be transferred by air movement or gravity
(Bijhouwer, 1937; Mallik 1957). The tacit assumption that open-pollinated
flowers are exclusively crossed is likely to be incorrect, although mango may
favour cross-pollination.
Insect
Popenoe (1917) reasoned that pollen transfer occurs primarily within flowers
by insects. Panicles bagged to exclude insect visitation generally result in less
fruit set than on panicles in the open (Popenoe, 1917; Musahib-ud-din and
Dinsa, 1946; Mallik, 1957; Free and Williams, 1976; Jiron and Hedstrom, 1985).
Insects working mango flowers include Diptera, Hymenoptera, Lepidoptera
and Coleoptera (Popenoe, 1917; Simao and Maranhao, 1959; Randhawa and
Damodaran, 1961b; McGregor, 1974; Anderson et al., 1982; Jiron and Hed-
strom, 1985). Flies of various genera are common on mango flowers (Pope-
noe, 1917; Burns and Prayag, 1921; Bijhouwer, 1937; Singh, 1954a; Spencer
and Kennard, 1955; Eardley and Mansell, 1993). Polistes wasps are observed
on mango flowers but are considered to be ineffectual for pollen transfer
(Spencer and Kennard, 1955; Free and Williams, 1976; Wolfenbarger, 1977).
Honeybees (Hymenoptera) are occasional visitors (Young, 1942; Simao and
Maranhao, 1959; Smith, 1960; Morton, 1964; Jiron and Hedstrom, 1985; Mac-
Millan, 1991; Du Toit and Swart, 1993, 1994; Eardley and Mansell, 1993, 1994),
but only if other more inviting flowers are not present (Spencer and Kennard,
1955; Free and Williams, 1976; McGregor, 1976). They are assumed to be the
most effective pollinators of mango and may be more effective if hives are
placed in orchards during flowering (Du Toit and Swart, 1993, 1994). Ander-
son et al. (1982) recorded actual pollen transfer on mango flowers by insects
and found, in order of importance, the most efficient pollinators to be wasps,
bees, large ants and large flies.
With few exceptions (Mallik, 1957), pollen deposition rates are generally
low (Naik and Mohan Rao, 1943; Mukherjee, 1951). Differences in pollination
rates can be attributed to environmental conditions during flowering, differ-
ing attraction of insects to specific cultivars, proximity of more attractive
flowering species or a combination of the above. Young (1942) observed that
insects visit only 10–12% of available flowers. Depending on weather condi-
tions, insect activity on mango flowers is usually continuous from early
morning to late afternoon, but nocturnal activity of some species has also
been reported (Jiron and Hedstrom, 1985).
The role of insects in cross-pollination is not understood. Anderson et al.
(1982) observed various insects carrying pollen to and from flowers and
noted pollination subsequent to those visits; however, they made no distinction
Reproductive Physiology 139
5.13 Stenospermocarpy
Abscission of non-fertilized and fertilized flowers is normal. Fruitlet abscis-
sion from pea size on is often associated with embryo abortion (Chandler,
1958; U.R. Singh, 1961; Singh, 1964; Lakshminarayana and Aguilar, 1975;
Ram et al., 1976) and is referred to as stenospermocarpy (Soule, 1985). Steno-
spermocarpy in mango is unusual (Chacko and Singh, 1969a) but occurs
regularly in some cultivars (Núñez-Elisea and Davenport, 1983; Whiley et al.,
1988). Stenospermocarpic fruitlets have slower growth rates than seeded
fruit, generally become misshapen and fail to reach full size.
140 T.L. Davenport
Fruit set and retention of mango was recently reviewed by Singh et al. (2005).
Abscission of flowers and fruitlets is accomplished by rapid formation of a
separation layer in the abscission zone in the pedicel-peduncle junction (Bar-
nell, 1939). U.R. Singh (1961) described formation of the abscission zone dur-
ing floral ontogeny and of the separation layer during abscission of flowers
and fruitlets. The majority of panicles lose all fruitlets (Núñez-Elisea and
Davenport, 1983). The pattern of fruitlet abscission is asymptotic with the
greatest losses occurring during the first weeks after anthesis (Núñez-Elisea
and Davenport, 1983; Prakash and Ram, 1984; Searle et al., 1995). Except for
the tendency to retain fruit in the distal portion of panicles, abscission of
flowers and fruitlets is random. It can involve fruitlets regardless of size or
location.
Of the 8–13% of perfect flowers setting fruit, < 1% reach maturity (Bijhou-
wer, 1937; Sen, 1939; Naik and Mohan Rao, 1943; Mukherjee, 1949b; U.R.
Singh, 1960; Randhawa and Damodaran, 1961a; Singh, 1978; Gunjate et al.,
1983; Prakash and Ram, 1984). Generally, most fruit are set on the most distal
spike portion of panicles (Chadha and Singh, 1963; Núñez-Elisea and Daven-
port, 1983). Fruit loss has been associated with embryo abortion, resulting in
blackened or shrivelled embryos (Singh, 1954a, 1964; Chandler, 1958; U.R.
Singh, 1961; Sharma and Singh, 1972; Ram et al., 1976) after the fruit is sepa-
rated from the tree (Núñez-Elisea and Davenport, 1983).
Sex ratio
The perfect/staminate floral ratio in panicles may influence fruit set and pro-
ductivity (Naik and Mohan Rao, 1943; Singh, 1954b; Singh and Singh, 1959;
U.R. Singh, 1960). Mallik (1957) noted that more perfect flowers are formed
in ‘on’ than ‘off’ years of alternate-bearing cultivars. Other studies, however,
Reproductive Physiology 141
have demonstrated that the number of perfect flowers does not correlate
with subsequent yield (Randhawa and Damodaran, 1961a) so long as the
proportion of perfect flowers is not < 4% (Singh, 1964, 1971). Most fruit are
borne in the distal portion of panicles (Shawky et al., 1977), which may be
correlated with the high ratio of perfect to staminate flowers there. Schole-
field and Oag (1984) estimated that one mature fruit is harvested for each 169
perfect flowers in the distal half of the panicle; whereas 592 perfect flowers
are required to produce one fruit in the proximal half. Therefore, intrinsic
factors other than sex ratio regulate fruit set.
Mineral nutrients
Boron is one of seven micronutrients required for normal plant growth. The
physiological function of B is unknown (Hu and Brown, 1994), although it is
essential for floral development, pollen germination, pollen tube growth,
embryo development and growth of organs (i.e. fruit) (Vasil, 1963; Agarwala
et al., 1981; Dell and Huang, 1997; Shorrocks, 1997). Deficient soils are com-
monly found in mango-producing areas of Australia, Thailand, Central and
South America and Africa where symptoms are common (Aitken et al., 1987;
Singh et al., 2005). Boron applications to deficient mango trees increase normal
fruit set (Robbertse et al., 1990; Raja et al., 2005). Fruitlet abscission in mangoes
has also been attributed to zinc (Zn) deficiency (Jiron and Hedstrom, 1985).
Hormonal control
Auxin
Research demonstrating improved fruit set and retention following applica-
tion of several auxin analogues to pre-anthesis panicles or to panicles bearing
fruitlets of various sizes has been reviewed (Davenport and Núñez-Elisea,
1997; Singh et al., 2005). NAA is the most effective auxin analogue for improv-
ing fruit retention (Prakash and Ram, 1986; Khan et al., 1993). Initial fruit set
was substantially increased when sprays of 200 mg/l indole acetic acid (IAA)
were applied to developing panicles (Singh et al., 1965). A 300–400% increase
in fruit set resulted when NAA (40 or 50 mg/l) was sprayed at the pre-anthesis
stage (Ram, 1983; Singh and Ram, 1983; Prakash and Ram, 1986). Chen (1981)
reported no effect on fruit retention when 5 mg/l of either naphthaleneacet-
amide or β-naphthoxoyacetic acid were applied three times at 2-week inter-
vals to panicles in which fruit had reached 4 mm in diameter.
Despite increased fruit retention of mango using exogenous applications
of auxins, few studies have examined endogenous auxins in fruit as related
to retention (Chacko et al., 1970a, b; Ram et al., 1983; Prakash and Ram, 1984).
Singh and Singh (1974) were unable to detect significant differences in endog-
enous auxins or inhibitors when comparing alternate and regular bearing
cultivars. Chen (1981) observed lower levels of auxin-like substances in
142 T.L. Davenport
mesocarp and calyx tissues of abscised fruits than those of intact fruits. Simi-
lar decreases in auxin and gibberellins with an increase in abscisic acid as
fruitlets abscised were reported by Bains et al. (1999). The interaction of auxin
in fruit and abscission zones to maintain mango fruit retention is not clear.
Continuous auxin synthesis and basipetal transport to the abscission
zone is critical for maintenance of plant organs, including fruit (Crane, 1964;
Nitsch, 1965; Morgan et al., 1977; Davenport et al., 1980; Roberts and Osborne,
1981). Increased mango fruit set and retention in response to exogenously
applied auxins confirms this requirement; however, other hormonal factors
also appear to be involved. Developing seeds are rich sources of all the known
classes of phytohormones, including auxins (Crane, 1964; Nitsch, 1965; Chacko
et al., 1970a, b, c; Chen, 1981). Hence, exogenous enrichment of auxin in the
presence of other seed-produced phytohormones facilitates increased fruit
retention. In contrast, NAA (10 and 20 mg/l) spray-applied to bagged, self-
pollinated flowers, does not result in development of stenospermocarpic
fruits beyond the marble size (Venkataratnam, 1949; Chacko and Singh,
1969a, b). Similarly, applications of 250 or 500 mg/l GA3 or 250 mg/l BA
alone to panicles does not promote production of stenospermocarpic fruits
(Chacko and Singh, 1969a, b). Supplying exogenous β-naphthoxyacetic acid
(10 mg/l), BA (250 mg/l) and GA3 (250 and 500 mg/l) together in multiple
sprays until half grown, however, resulted in retention of several seedless
fruit to maturity. Chen (1983) and Oosthuyse (1995b) observed that gibberel-
lin, cytokinin and auxin reduce fruit drop of open-pollinated fruitlets of some
cultivars. Thus, although auxin is important for maintaining the abscission
zone, the presence of other phytohormones appears to be important for fruit-
let development (Chacko et al., 1970a, b; Ram, 1983; Ram et al., 1983).
Cytokinins
Although cytokinins are not generally thought to be associated directly with
abscission, Ram (1983) and Ram et al. (1983) concluded that low cytokinin
levels during fruit development might contribute to fruit loss. Chen (1983)
observed a correlation of low cytokinin levels in stenospermocarpic fruits
with abscission at the marble stage of growth. Application of 250 mg/l BA to
bagged panicles does not promote production of seedless fruits (Chacko and
Singh, 1969a, b). The synthetic cytokinin, N-(2-chloro-4-pyridyl)-N’-phenylurea
(CPPU) also does not improve fruit set when applied alone at a rate of 10
mg/l to post-anthesis panicles (Oosthuyse, 1995b). The role of cytokinins in
separation events remains inconclusive.
Gibberellins
Gibberellins do not appear to be directly linked to the onset of abscission
(Chacko et al., 1970c, 1972a; Ram and Pal, 1979; Chen, 1981; Ram, 1983). Spray
applications of GA3 to pre- and post-anthesis panicles to increase fruit set
and retention have been inconsistent. Increased yield (Teaotia et al., 1967;
Singh and Ram, 1983; Rajput and Singh, 1989) and production of seedless fruit
(Kulkarni and Rameshwar, 1978) have been reported from these treatments,
but Chacko and Singh (1969a, b) observed no such effects. Chen (1983) and
Reproductive Physiology 143
Inhibitors
Abscisic acid (ABA) is possibly involved in fruitlet abscission. Although cor-
relations exist between certain inhibitors and abscission of mango fruitlets,
no clear cause and effect relationships have been established. Fruit drop was
correlated with levels of an acidic inhibitor, possibly ABA (Chacko et al.,
1970b, 1972a; Singh and Singh, 1974; Ram, 1983; Prakash and Ram, 1984).
Chen (1981) reported similar changes in putative ABA with maximum levels
occurring during early fruit drop and with advancing age of fruits. Putative
ABA levels in abscised and retained fruits were compared and were highest
in the calyx and mesocarp of abscised fruitlets.
Ethylene
Ethylene has the greatest immediate impact on flower and fruitlet abscission.
Van Lelyveld and Nel (1982) reported higher levels of ethylene in abscised
fruitlets compared with those retained on trees. Núñez-Elisea and Davenport
(1983, 1984, 1986) examined the dynamics of ethylene production in intact
and excised fruitlets from onset to separation. Increased production began in
explants about 26 h postharvest and increased logarithmically until fruit sep-
aration. Abscission of the fruitlets began 48 h after the onset of enhanced
ethylene production. Similar results with avocado fruitlet abscission experi-
ments (Davenport and Manners, 1982) indicate that the onset of ethylene
production in intact fruitlets is spontaneous in individual fruitlets followed
by abscission 48 h later. The pericarp provided the bulk of ethylene for induc-
tion of abscission processes; the pedicel produced no ethylene. There was
reduced fruit drop in response to inhibitors of ethylene production and action
(Singh and Ram, 1983; Naqvi et al., 1990, 1992). Whereas increased peroxi-
dase (Van Lelyveld, 1978) and polyphenol oxidase activities have been
reported in abscissed mango fruitlets (Van Lelyveld and Nel, 1982), Núñez-
Elisea and Davenport (1984) observed no changes in peroxidase activity or
protein levels prior to separation of fruitlets.
144 T.L. Davenport
Photoassimilates
Wolstenholme and Whiley (1995) discussed the ecophysiology of the mango
as a basis for preharvest management. They proposed that the adaptive sur-
vival strategies of the mango explain its notoriously poor cropping perfor-
mance. Mechanisms that impart tolerance to heat, drought and flood stresses,
which the tree has developed for survival in harsh environments, have come
at considerable carbon cost with the resultant diversion of photoassimilate
resources away from fruiting.
There is abundant evidence that heavy cropping in tree crops exhausts
stored reserves (Jones et al., 1975; Kaiser and Wolstenholme, 1994; Whiley et al.,
1996) and that current photosynthate is often unable to satisfy the demands
of fruit set and fruit growth after heavy and prolonged flowering (Chacko
et al., 1982). There are significant genotypic differences in photoassimilation
rates between low- and high-yielding cultivars growing in both the tropics
and the subtropics of Australia (Chacko et al., 1995; Searle et al., 1995). At each
location, photoassimilation rates were considerably greater on the higher-
yielding cultivar, and this difference was maintained from flowering through
to fruit maturation. 14C studies during the fruit set and abscission period also
demonstrated strong discrimination in the movement of assimilates, which
was dominated by randomly located fruit on panicles of the low-yielding
cultivar (Chacko et al., 1995). In contrast, assimilate discrimination to fruitlets
was less severe in the high-yielding cultivar with a more even distribution of
photoassimilates. It was concluded that the availability and distribution of
photoassimilates during the fruit set and establishment stages was largely
responsible for the yield differences between the cultivars.
Supporting evidence for the role of photoassimilates in fruit set and
retention also comes from enrichment studies (Schaffer et al., 1999). Container-
grown plants that flowered in the open were transferred to controlled-
environment glasshouse rooms immediately after the completion of anthesis.
Temperatures were 28°C day/20°C night while the atmospheric carbon diox-
ide (CO2) concentrations were 350 or 600 Pmol/mol. Photoassimilation of
trees in the CO2-enriched rooms was approximately 60% greater than those
held at partial pressures of 350 Pmol/mol CO2. Fruit retention and final yield
were significantly higher on those trees grown at the partial pressure of
Reproductive Physiology 145
600 μmol/mol CO2. Higher levels of available assimilates during the fruiting
cycle appear to benefit fruit retention and yield.
5.16 Conclusions
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6 Ecophysiology
6.1 Introduction
6.2 Photosynthesis
Introduction
The net CO2 assimilation rate (Anet) in C3 plants is a function of the carboxyla-
tion rate (Vc), the oxygenation rate (Vo) and the rate of CO2 evolution in light
that results from processes other than photorespiration, sometimes called
‘day respiration’ (Rd):
Anet = Vc – 0.5Vo – Rd (6.1)
Rd is usually inferred from measurements of leaf CO2 exchanges after 5 min
in the dark (i.e. ‘night respiration’ Rn). However, it has been repeatedly
shown that Rd is lower than Rn (see Atkin et al. (2000) for review), so that light
is known to inhibit respiration, with a Rd/Rn value ranging from 30 to 100%
(see Peisker and Apel (2001) for review). Urban et al. (2008) established the
following linear regression for Rd of mango leaves: Rd = 0.35Rn – 0.21, which
may be used to infer Rd from Rn for photosynthetic photon flux (Q) values
above 170 Pmol photons/m2/s.
Currently, modelling of Anet often uses the Harley et al. (1992) version
of the Farquhar et al. (1980) model. According to this model, Anet can be
expressed as:
Anet = (1 – 0.5O/(WCi))min(Wc, Wj, Wp) – Rd (6.2)
where O represents the partial pressure of oxygen (O2) in the intercellular air
spaces (Pa), W the specificity factor of ribulose-1,5-bisphosphate carboxylase/
oxygenase (Rubisco). Ci is the partial pressure of CO2 in the intercellular air
spaces (Pa), Wc the carboxylation rate limited by the amount, activation state
or kinetic properties of Rubisco (Pmol CO2/m2/s), Wj the carboxylation rate
limited by the rate of ribulose bisphosphate regeneration (Pmol CO2/m2/s),
and Wp the carboxylation rate limited by triose phosphate utilization in
sucrose and starch synthesis (Pmol CO2/m2/s).
Usually O is set as 21 kPa (21%). The variable W, which characterizes the
ratio of the affinities of CO2 and O2 for ribulose-1,5-bisphosphate in the active
site of Rubisco, can be calculated from the CO2 compensation point ** (the
CO2 concentration at which photosynthesis equilibrates with respiration):
W = 0.5O/* * (6.3)
where W = 2220 mol CO2/mol O2 at 25°C for ‘Cogshall’ mango leaves (Urban
et al., 2008), which is lower than those given by Epron et al. (1995): 2100–2900
Ecophysiology 173
mol CO2/mol O2. Rubisco’s large subunit is encoded by a single gene in the
chloroplast genome, and no post-transcriptional modifications have been
discovered so far. It is thus very unlikely that W can change in the short term
(Spreitzer and Salvucci, 2002).
The internal partial pressure of CO2 (Ci) is one of the two major variables
of photosynthesis (with the photosynthetically active photon flux density).
It may be calculated from the supply function:
Ci = Ca – Anet/gb – Anet/gs (6.4)
where Ca is the partial pressure of CO2 (Pa) in ambient air, gb represents the
leaf boundary layer conductance (mol H2O/m2/s), and gs is the stomatal
conductance of water (H2O) (mol H2O/m2/s).
Stomatal conductance is the major factor controlling Anet. It ranges from
c.0.02 to c.0.4 mol H2O/m2/s in ‘Cogshall’ mango leaves and may be linearly
related to Anet (Urban et al., 2002, 2003, 2006). The slope of the relationship
between gs and Anet however is affected by drought (Fig. 6.1). Variations in
the slope of this relationship reflect changes in photosynthetic water use effi-
ciency and are not well understood.
It must be stressed that using Ci as the driving variable of photosynthesis
is much debated. It has been advocated that Cc, the partial pressure of CO2 at
the site of carboxylation, should be utilized instead. Using Ci implies that the
following assumptions have been made: Cc = Ci and gm = 0, where gm repre-
sents mesophyll conductance, also called liquid phase resistance, which
0.50
0.45
0.40
0.35
Wet
gs (mol H2O/m2/s)
0.20
0.15
Dry
0.10 y = 0.009x + 0.024
R2 = 0.695
0.05
0
0 5 10 15 20
Anet (μmol CO2/m2/s)
Fig. 6.1. The relationship between stomatal conductance (gs) and net photosynthesis
(Anet) in mango leaves from well-irrigated (■) and drought-stressed (▲) 12-year-old
‘Cogshall’ trees (Source: redrawn from Urban et al., 2006).
174 B. Schaffer et al.
1. The photosynthetically active photon flux density (Q), which is the major
driving variable of photosynthesis. Gross photosynthesis is determined by
Q while Ci determines the proportion of photorespiration, and thus net
photosynthesis. One of the major environmental factors affecting Ci is water
availability in the root zone through its effect on gs.
2. Leaf nitrogen concentration (Na), which is not a rate-determining factor
of photosynthesis, unlike Q, but may be considered as a rate-limiting factor.
In other words, Na sets the photosynthetic potential of a leaf (i.e. photosyn-
thetic capacity). We shall see below which factors influence Na in mango
leaves.
3. Leaf temperature influences leaf photosynthesis. Net photosynthesis is
positively correlated with leaf temperature in a normal range. Leaf tempera-
ture (Tl) is not a driving variable of photosynthesis but it is the single most
important rate-determining factor after Q. In addition, extreme temperatures
may influence photosynthesis through their damaging effects. Kinetics of
enzymes involved with photosynthetic reactions collectively comprise an
additional set of factors that influence leaf net photosynthesis.
176 B. Schaffer et al.
140
120
Vcmax (μmol CO2/m2/s)
100
80
60 y = 41.52x – 15.52
R2 = 0.87
40 y = –201.64x–1 + 173.41
R2 = 0.88
20
0
1.0 1.5 2.0 2.5 3.0 3.5 4.0
(a) Na (g N/m2)
250
200
Jmax (μmol/m2/s)
150
100
y = 66.94x – 15.40
R2 = 0.83
50 y = –330.44x–1 + 291.55
R2 = 0.86
0
1.0 1.5 2.0 2.5 3.0 3.5 4.0
(b) Na (g N/m2)
Fig. 6.2. Relationship between (a) the maximum rate of carboxylation (Vcmax) and (b)
the light-saturated rate of electron transport (Jmax), and nitrogen concentration per unit
leaf area (Na). Measurements were performed on mango leaves of 3-year-old ‘Cogshall’
trees (●), standard leaves ({) and leaves close to developing fruits (
) of 11-year-old
‘Cogshall’ trees. Best fit lines for pooled data correspond to the linear (_) and the ax–1 +
b (…) models (Source: redrawn from Urban et al., 2003).
Light
Light exposure
Plants allocate nitrogen resources within the canopy to enhance photosyn-
thetic capacity at locations exposed to high incident light levels, thus maxi-
mizing whole plant carbon gain (Field and Mooney, 1983; Hollinger, 1996;
Carswell et al., 2000). For leaves of a given age and for a given nitrogen sup-
ply, leaf N per unit leaf area appears to be strongly related with light expo-
sure (DeJong and Doyle, 1985; Le Roux et al., 1999, 2001; Rosati et al., 1999,
2000). Photosynthetic light acclimation of leaves may result from changes in
either leaf nitrogen concentration (Nm) or mass-to-area ratio (Ma) because
Na = MaNm. Lynch and González (1993) observed a negative correlation
between Nm and light exposure in the tropical fruit tree Borojoa patinoi, but
such a behaviour is rare; positive correlations between Nm and light exposure
are more commonly observed. In addition, photosynthetic light acclimation
of leaves may result from changes in partitioning of total leaf N among the
different pools of the photosynthetic machinery (Evans, 1989). In mango,
light acclimation of photosynthesis results mainly from changes in Ma, and
to a lesser extent from changes in allocation of total leaf N at low irradiance;
whereas changes in Nm play only a minor role (Fig. 6.3). Light acclimation of
mango leaves thus follows a pattern similar to peach leaves (Le Roux et al.,
1999; Walcroft et al., 2002).
Light intensity
Photosynthesis of ‘Cogshall’ mango trees increases with increasing levels of
light intensity to reach a maximum at Q = 1200 Pmol photons/m2/s (L.
Urban, unpublished data). Whiley et al. (1999) measured Q at 1284 Pmol pho-
tons/m2/s for field-grown ‘Kensington Pride’ trees growing in subtropical
Queensland, Australia, which is well below full sunlight (full sunlight ≥ 2000
Pmol photons/m2/s). Such a high threshold is a typical feature of sun plants.
Individual leaves are rarely able to utilize full sunlight; whole trees consist of
many leaves that shade each other, so that only a small fraction of a tree’s
leaves are exposed to full sun at any given time of the day, while the rest of
the leaves receive subsaturating photon fluxes in the form of small patches of
light that penetrate through gaps of the leaf canopy. Because the photosyn-
thetic response of whole trees is the sum of the photosynthetic activity of all
the leaves, only rarely is photosynthesis saturated with light at the whole-
tree level.
While most leaves experience subsaturating light intensities, well-exposed
leaves of the upper-crown may receive excessive quantities of light. Those
leaves must dissipate the absorbed light energy in excess to prevent damage to
the photosynthetic apparatus. Moderate decreases in maximal quantum
efficiency (i.e. quantum efficiency of dark-adapted leaves Fv/FmPredawn) are
178 B. Schaffer et al.
3.0
1.0
0.0
0.0 0.2 0.4 0.6 0.8 1.0
(a) Gap fraction
3.0
Tree # 1
y = 1.42x + 1.43
2.5
R2 = 0.90
2.0
Tree # 2
Na (g/m2)
y = 1.33x + 1.44
1.5 R2 = 0.79
1.0
0.5
0.0
0.0 0.2 0.4 0.6 0.8 1.0
(b) Gap fraction
Fig. 6.3. Relationship between (a) leaf nitrogen concentration per unit mass (Nm) and
(b) leaf nitrogen concentration per unit leaf area (Na) and the gap fraction for mango
leaves measured in the crown of two 3-year-old ‘Cogshall’ trees. Gap fractions were
measured as an indicator of light exposure. Measurements were performed on leaves
< 2 months old (●), 8 months old (■), 12–14 months old (▲) and 17–20 months old
(♦) (Source: Urban et al., 2003).
Q, on ‘Cogshall’ trees. Whiley et al. (1999) measured Amax of 15.2 Pmol CO2/
m2/s for ‘Kensington Pride’ trees growing in a subtropical climate in Queen-
sland, Australia. However, values > 16 Pmol CO2/m2/s were observed on
field-grown trees of ‘Tommy Atkins’, ‘Haden’ and ‘Irwin’ on sunny days
during the wet season in tropical regions of Australia (P. Lu, unpublished
data). This is much higher than citrus (< 10 Pmol CO2/m2/s), but substan-
tially lower than plum (approx. 26 Pmol CO2/m2/s). Whiley et al. (1999) esti-
mated the light compensation point to be 29 Pmol photons/m2/s for leaves
of non-stressed, field-grown mango trees, which is much higher than that
attributed to shade-tolerant species (< 10 Pmol photons/m2/s) (Harvey,
1979). The data show that mango trees are basically sun-adapted plants.
Leaf temperature
Chilling temperatures
Fv/FmPredawn decreases in mango leaves with decreasing temperature, while
chilling reduces quantum efficiency (Whiley et al., 1999; Sukhvibul et al.,
2000; Weng et al., 2006a, b). The decrease in Fv/FmPredawn may be interpreted
to reflect sustained engagement of zeaxanthin in photoprotective energy dis-
sipation. Decreases in quantum efficiency correspond to a decrease in the
rate of electron flow. It may be argued that sustained zeaxanthin-dependent
180 B. Schaffer et al.
4.0
3.5
3.0
Vcmax /Vcmax at 25°C
2.5
2.0
1.5
1.0
0.5
0.0
15 20 25 30 35 40 45 50
(a) Tl (°C)
2.0
1.8
1.6
1.4
Jmax /Jmax at 25°C
1.2
1.0
0.8
0.6
0.4
0.2
0.0
15 20 25 30 35 40 45 50
(b) Tl (°C)
Fig. 6.4. Temperature response functions adjusted to the (a) maximal rate of car-
boxylation (Vcmax) and (b) the light-saturated rate of photosynthetic electron flux (Jmax),
normalized to the mean value at 25°C in leaves from ‘Cogshall’ mango seedlings.
The data scatter represents the real scatter at each temperature. Reference values at
25°C were computed for each of eight leaves, taken from young trees from two
origins (● and {), and a unique temperature response was adjusted over the range of
normalized data. Tl, the leaf temperature; the dotted lines correspond to Equation 9;
the solid lines correspond to Equation 10 (no deactivation energy component).
energy dissipation reduces the risk of formation of singlet oxygen 1O2 in the
antennae, while decreases in J lower the risk of electrons reducing O2 to anion
superoxide O2− in the photosynthetic electron transport chain (Adams et al.,
2005). In other words, decreases in Fv/FmPredawn and quantum efficiency cor-
respond to adaptative mechanisms against the effect of cold, when photosyn-
thesis is low and there is an imbalance between the quantity of light energy
absorbed and the quantity of energy used in the photochemical reactions of
Ecophysiology 181
The CO2 concentration in the earth’s atmosphere has been increasing rapidly
since the early 20th century and is continuing to rise, primarily due to burn-
ing of fossil fuels (Houghton, 2005). Earth’s atmospheric CO2 concentration
is currently about 370 Pmol CO2/mol (Houghton, 2005) and is projected to
reach 600 Pmol CO2/mol by 2050. Elevated ambient CO2 levels will undoubt-
edly affect cropping systems since atmospheric CO2 concentrations can sig-
nificantly affect plant growth and productivity (Idso and Kimball, 1991;
Houghton, 2005). There is little published information concerning the effects
of elevated ambient CO2 levels on physiology, growth and production of
tropical fruit trees, including mango.
Schaffer et al. (1997) exposed leaves of field- and container-grown ‘Kens-
ington’ (syn. ‘Kensington Pride’) trees to short durations (several minutes) of
varying ambient CO2 concentrations. They found that under saturating light
levels for photosynthesis, net photosynthesis increased as ambient CO2 con-
centration increased up to 1200 Pmol CO2/mol. At ambient CO2 concentra-
tions > 1200 Pmol CO2/mol, net photosynthesis stabilized, probably due to
leaves reaching their maximum biochemical capacity to fix carbon. Studies
with ‘Cogshall’ mango trees indicated that when Ca increases, stomata close
swiftly and Ci may become very unpredictable (L. Urban, unpublished data).
Therefore, using Ci may be preferable to Ca for quantifying short-term effects
of elevated CO2 concentations on Anet of mango. Saturating CO2 levels may
often be reached at Ci = 800 Pmol CO2/mol air.
Long-term (6–12 months) exposure of ‘Kensington’ mango trees to an
atmospheric CO2 concentration of 700 Pmol/mol resulted in higher net CO2
assimilation rates than in leaves of plants grown at atmospheric CO2 concen-
trations of 350 Pmol/mol when net CO2 assimilation was measured at the
same CO2 concentration as the growth environment. However, carboxylation
efficiency (the amount of CO2 fixed per mole of ambient CO2) was lower for
plants in the CO2-enriched environment compared to plants in the ambient
(350 Pmol CO2/mol) environment (Schaffer et al., 1997). Although further
studies are needed to determine the effects of long-term exposure to elevated
CO2 concentrations on mango growth and productivity, it appears that
182 B. Schaffer et al.
Humidity
Flooding
0.6
0.5
0.3
0.2
0.0
1 2 3 4 5 6 7 8
LAVPD (kPa)
Fig. 6.5. Correlation between leaf stomatal conductance and leaf-to-air vapour
pressure deficit (LAVPD) during the dry and wet season for ‘Irwin’ (closed circles)
and ‘Kensington’ mango trees (open circles). Trees were well irrigated during the dry
season and all measurements were taken when the Q > 300 Pmol photons/m2/s
(n = 12) (Source: P. Lu, unpublished data).
a result of these elements becoming more soluble when calcareous soils are
flooded (Larson et al., 1991b, 1992).
Internal factors
Leaf age
Leaf characteristics (i.e. photosynthetic capacity and the amount of N per
unit area) are generally strongly influenced by leaf age, with maximum val-
ues being observed when leaves have just completed full expansion (Con-
stable and Rawson, 1980; Marshall and Biscoe, 1980; Dwyer and Stewart,
1986; Field, 1987; Wilson et al., 2000; Frak et al., 2001). Chlorophyll content is
three to four times lower in young than in mature mango leaves (Zude and
Ludders, 1997). Similarly, the concentration of Rubisco is lower in young
than in mature, green leaves (Nii et al., 1995). In contrast to many other plant
species, once mango leaves are mature the relationship between Na and irra-
diance does not seem to be affected by leaf age (Urban et al., 2003). The Na
values may remain high in old leaves experiencing high irradiance. This
indicates that changes in Na in mango leaves are influenced by irradiance
and not age, at least during the first year.
250
A Q = 400 μmol photons/m2/s
y = 84.5e–0.0313x
200 R2 = 0.69
J (μmol electrons/m2/s)
0
0 10 20 30 40
Starch (g/m2)
Fig. 6.6. The relationship between the total photosynthetic electron flux (J) measured
at photosynthetic photon flux (Q) = 400 ('), 1200 (●) and 2000 ({) Pmol photons/
m2/s, and the amount of starch per unit leaf area. Best fit lines at each Q were assessed
from measurements performed on both girdled and non-girdled mango leaves before
flowering. Data were used to establish the following relationship: J = (0.0434Q +
72.8)*e–0.0412[starch]a (Source: Urban and Alphonsout, 2007).
186 B. Schaffer et al.
decrease in leaf carbohydrate content during the first month following the
onset of flowering, suggesting that the effect of carbohydrate accumulation
on photosynthesis is mediated by sink activity. Apart from its negative effect
on the carbon budget of mango trees, girdling appeared to be rather harm-
less. However, leaf N concentration decreased, which indicates that there
may indeed exist long-term negative effects of girdling on photosynthetic
capacity. The width of bark (phloem) removed may be critical with respect to
the intensity of the effect of girdling on the tree. Whiley et al. (2006) girdled
the trunks of ‘B74’ (‘Calypso’™) mango trees in the Northern Territory of
Australia in autumn (as soon as they had come out of the wet season). The
girdles were no more than the thickness of a pruning saw (about 1 mm) and
healed within 6 weeks. In the first and third years after girdling, the trees had
significantly higher yields than non-girdled trees on which, coincidentally,
fruit matured early, thus giving market advantage. There was no significant
difference in yield in the second year of treatment between girdled and non-
girdled trees. The first and third years had strong natural induction while the
second year gave poor flowering across all varieties in the district. Thus, dur-
ing years of strong induction, this type of girdling most likely provided extra
carbohydrate reserves to drive flowering and support fruit set and retention
while in the off-flowering year there was sufficient carbohydrate reserves to
support reproductive activity. In contrast to observations with ‘Cogshall’
mango trees (Urban and Alphonsout, 2007), there was no evidence of long-term
effects of narrow girdles on leaf N of ‘B74’ mango trees (Whiley et al., 2006).
However, when wider girdles are made, tree recovery may take much longer
leading to sustained physiological disruption.
Proximity of inflorescences
While the effects of water stress and high light, temperature and atmospheric
CO2 concentration on photosynthesis are increasingly well described, very
little is known about the effect of phenology, and especially of flowering on
photosynthesis of mango. There is some evidence that flowering may have
an effect on photosynthesis. Flowering-associated decreases in Anet and gs
were observed in sweet cherry (Roper et al., 1988) and mango (Shivashankara
and Mahai, 2000; Urban et al., 2004a). Lack of precise knowledge about the
effect of flowering on photosynthesis may impair our ability to adequately
simulate photosynthesis, especially for tropical fruit trees for which flower-
ing often extends over a long period of time. Mango flowering can last for >
2 months. Therefore, its effect on photosynthesis should not be overlooked.
Urban et al. (2004a) showed that the decrease in Anet in mango leaves close to
inflorescences is not attributable to a gs-associated decrease in Ci or to an
increase in Rd. Rd was lower in leaves close to inflorescences than in standard
leaves. If any, the effect of Rd on Anet was a positive one. This study suggested
strongly that the decrease in Anet was due to a decrease in the electron flow in
photosystem II, but failed to provide direct evidence for it as well as the ele-
ments for interpretation. Using a modelling approach, Urban et al. (2008)
confirmed that there is a decrease in the total light-driven photosynthetic
electron flux in leaves close to inflorescences and showed that the decrease
Ecophysiology 187
In this section, theoretical concepts of plant water relations are briefly out-
lined to help interpret the effects of environmental factors on mango water
relations (see also Nobel (1983) and Baker (1984)).
An important concept in plant water relations is water potential (Ψ),
which is a measure of the free energy of water. For pure water, Ψ = 0. As sol-
utes are added to water, its free energy decreases and < becomes more nega-
tive. Water moves along a gradient from higher to lower (more negative) <.
< can be expressed as:
< = <S + <p + <m (6.11)
where <S is osmotic (or solute) potential which refers to the effect of solutes
on the change in free energy of water; <p is the hydrostatic or pressure poten-
tial also referred to as the turgor pressure; and <m is the matric potential,
which is generally negligible in plant cells.
In plant cells, <p is generally positive or equal to 0. However, in xylem
tissue of transpiring plants, <p is negative (under tension). The driving force
for transpiration is the vapour pressure difference between the leaf (consid-
ered to be water saturated) and the surrounding air. Thus, water moves from
a greater to a lower (or more negative) <p and hence along a decreasing <
gradient. The cohesive forces of the H2O molecules allows the xylem water to
remain in a continuous column even though there is a negative <p.
Plant water stress can be determined from Eqn 6.11 and from changes in
<. The components of <, such as <S and <p, can often be used to define the
sources of water stress. The drought tolerance of mango highlights some
unique aspects of physiology of this tree with respect to its water manage-
ment. Typical mango environments in the tropics impose extreme water
stress and high evaporative demand for prolonged periods. Adaptive strate-
gies of mango trees include a deep root system (Sukonthasing et al., 1991),
desiccation-tolerant surface feeder roots and drought avoidance mechanisms
thought to be mediated by a comprehensive system of resin canals distrib-
uted throughout the tree (Venning, 1948; Joel, 1980; Joel and Fahn, 1980a, b;
Pongsomboon, 1991) and rapid stomatal closure. Plants with laticfers or resin
ducts/canals have been reported to be drought tolerant due to extended
maintenance of turgor following the withdrawal of water (Downton, 1981;
Kramer, 1983; Kallarackal et al., 1990). While the mechanism of turgor main-
tenance remains unresolved, it is believed that the latex or resin is probably
involved in the modulation of plant water status (Kallarackal et al., 1990).
The differentiation, structure and distribution of resin canals in mango has
been described by Venning (1948), Joel (1980) and Joel and Fahn (1980a, b, c).
Resin canals are present in trunks, shoots, leaves and fruit (exocarp) of mango
in close association with the vascular tissues. The resin contains mainly ter-
penes, but phenols and protein-carbohydrate mucilage are also present (Joel
and Fahn, 1980c). In well-watered trees, the resin is under positive pressure
and freely exudes from damaged or cut surfaces (Pongsomboon, 1991). In
studies of the development of water deficit in container-grown ‘Kensington’
Ecophysiology 189
mango trees, loss of turgor occurred in expanding leaves when leaf water
potential (<l) reached –1.2 Mpa. In mature leaves turgor was not lost until <l
reached –1.75 MPa (Pongsomboon, 1991). Necrotic leaf areas appeared when
<l reached approximately –3.2 MPa with permanent wilting developing at
−3.45 MPa. This is high (less negative) compared with a <l of –6.6, –5.0 MPa
for orange (Citrus sinensis) and macadamia (Macadamia integrifolia), respec-
tively (Fereres et al., 1979; Stephenson et al., 1989). Thus, mango leaves toler-
ate less internal water stress than woody perennial fruit trees from more
mesic environments. With mango, however, the permanent wilting point
was reached 36 days after withholding water compared to 10 days for simi-
larly sized macadamia trees (Stephenson et al., 1989). The higher critical
threshold of <l and the longer period of survival for ‘Kensington’ mango
indicates that drought tolerance is based on more effective water regulation
to prevent desiccation and on maintenance of leaf turgor rather than resis-
tance by tissues to damage.
Reich and Borchert (1988) observed that stomatal regulation in mango
significantly reduced the rate of development of internal water deficit when
compared with four other tropical tree species. In water-withholding studies,
the radial expansion of mango trunks continued when most of the other
species were shrinking, indicating that mango trees could better tolerate
drought conditions and maintain photoassimilation rates. This is consistent
with the decrease in gs/Anet observed by Urban et al. (2006) as a consequence
of drought (Fig. 6.1). A decrease in gs/Anet indicates that there is an increase
in photosynthetic water use efficiency.
In containerized ‘Kensington’ mango trees, there was a linear correlation
between stomatal conductance and <l during the development of water
stress (Pongsomboon, 1991). In contrast, with avocado and macadamia,
stomatal response was much more rapid with a curvilinear relationship
between <l and stomatal conductance, and stomatal closure reached at –1.2
and 3.0 MPa, respectively. The slower response of stomatal conductance to
<l in mango trees appears to be related to a more effective mechanism for
the mediation of water deficit development compared with avocado and
macadamia.
Pongsomboon (1991) monitored leaf water potential, osmotic potential of
resin (<r) and osmotic potential of the whole leaf tissue (<S) in container-
grown ‘Kensington’ mango trees when water was withheld for a 45-day
period. When tissues were fully hydrated, <l and <r were higher than <S. For
40 days of the drying cycle, <l and <r declined at a similar rate; however, <S
declined to about –1.2 MPa within 4 days where it remained stable until 18
days into the drying cycle. There was a subsequent decline in <S for 28 days
after withholding water when it stabilized at –2.0 MPa, remaining constant
for another 12 days. Pongsomboon (1991) suggested that osmotic adjustment
occurred, probably mediated through the resin as water deficit developed. It
appears, therefore, that the energy investment by the tree in a resin canal sys-
tem is justified by the vital drought-avoidance benefits conferred by main-
taining turgor and preventing wilting under prolonged periods of water
stress. Further investigations are required to substantiate these conclusions.
190 B. Schaffer et al.
Light
In an orchard, light distribution within and between tree canopies can have
a profound effect on growth and development of the fruit. We have previ-
ously discussed the effect of light on photosynthesis and defined the opti-
mum light levels required for mango leaves. When light levels fall below the
Ecophysiology 191
100
2006 light
2006 shade
80 2005
60
PLC
40
20
0
–4 –3 –2 –1 0
Xylem pressure (MPa)
and northern sides of the tree. This information establishes an important con-
cept with respect to the light regime but does not quantify the absolute light
levels required for anthocyanin development. Further research is necessary
to establish physiological parameters from which pruning and orchard man-
agement strategies can be developed.
Temperature
at which shoot growth ceases is approximately 15°C (mean value for ten cul-
tivars) (Whiley et al., 1989). Subsequent studies (Issarakraisila et al., 1991)
have confirmed that 15°C is the critical minimum growth temperature for
shoots of ‘Kensington’.
Stress-inducing temperatures which prevent shoot growth have been
shown to promote floral induction in mangoes, but this is outside the scope
of this discussion. For further information of the effects of temperature on
pollination, floral initiation and fruit development, see Davenport, Chapter
5, this volume and Schaffer et al. (1994). We again emphasize that although
mango is a ‘heat-loving’ crop well adapted to the hot, semi-arid subtropics
and monsoonal tropics, in these environments it experiences extremes of heat,
drought and evaporative demand that may cumulatively reduce potential
production capacity.
Drought
trees. Irrigation at 7-day intervals resulted in the greatest yield with the larg-
est fruit, especially during the early harvest period (Larson et al., 1989).
Irrigation, therefore, particularly during the first 4–6 weeks following
fruit set, increases individual fruit size and yield. This is a critical period of
fruit development since it is when cell division is most rapid and cell walls
are developed. Even slight reductions in plant water status during this period
can have adverse effects on fruit growth and retention (Pongsomboon, 1991).
Although drought tolerance of the mango tree is well known, this comes at
considerable cost to tree performance, particularly in areas with prolonged
dry seasons that extend through flowering and fruiting. Irrigation is there-
fore one of the most powerful tools to alleviate non-lethal yet potentially
yield-reducing drought stress.
Flooding
(Schaffer et al., 1994). It is likely that these roots facilitate the absorption and
translocation of O2 to submerged roots and their development is a common
morphological response of many woody plants to root anoxia. The develop-
ment of adventitious roots has not been reported for flooded, field-grown
trees and they may only form on young trunks after extended flooding peri-
ods, which usually do not occur under normal production conditions.
Vegetative growth of mango trees generally declines when trees become
flooded for > 2–3 days. When trees in a limestone soil in containers were
flooded for > 110 days, there was a 94% reduction in shoot extension growth,
while flooding for approximately 10 days resulted in a 57% reduction in
shoot extension growth (Larson et al., 1991c). In a subsequent study, the stem
radial growth (a more sensitive indicator of tree growth than shoot extension
growth) of mango trees decreased 2 weeks after roots were submerged.
Flooding for > 14 days also significantly reduced root dry weight, resulting
in an increased shoot to root ratio (Larson et al., 1991c). These adverse effects
of flooding on the growth of mango trees are expected as reduced net photo-
synthesis and presumably higher root respiration limit the availability of
carbon-based assimilates required for growth.
Wind
Most fruit trees benefit from wind protection, particularly during the estab-
lishment years when the disruption of physiological processes results in a
significant depression of growth in young trees. In addition, wind also causes
abrasions to the skin of fruits, particularly when they are small, which
develop into unsightly blemishes by the time they are fully grown thereby
reducing quality and market value. However, the cost of windbreaks may
not be offset by higher returns. In some mango-producing regions, winds are
not sufficiently strong to justify the cost of wind protection. Until recently,
wind protection in South Africa was not recommended for mangoes due to
the loss of potential cropping space by ‘living’ windbreaks, their potential to
create frost pockets, and the likelihood of promoting the incidence of flower
and fruit diseases through increased humidity (Van der Meulen et al., 1971);
however, the value of windbreaks is well appreciated today in South Africa
(B.N Wolstenholme, personal communication, Pietermaritzburg, South
Africa, 1995).
In studies with ‘Kensington’ mangoes in Australia, where artificial wind-
breaks were constructed to shelter trees from the prevailing summer south-
easterly winds, a 600% increase in yield was recorded in the first year
following wind protection (Mayers et al., 1984). This significant improvement
in tree performance was a result of better growth of trees which set and held
more fruit per panicle, suffered less damage to leaves (cuticle fracturing) and
had reduced fruit loss from bacterial black spot (caused by Xanthomonas camp-
estris pv. mangiferaeindicae) compared to the wind-exposed trees. These results
indicate that wind can have a significant effect on mango productivity from
the reduction of both growth and yield through undisclosed physiological
196 B. Schaffer et al.
mechanisms, and a decreased level of bacterial black spot infection. The pro-
vision of windbreaks in orchards is expensive with decisions to be made on
the use of either ‘living’ or artificial shelters. Requirements for wind protec-
tion will vary depending on site circumstances, and all factors pertaining to
crop performance will require careful consideration.
Salinity
Salt stress in mango trees produces symptoms similar to those described for
other species (Harding et al., 1958; Ehlig, 1960; Kadman, 1964; Bingham et al.,
1968). Mild symptoms of chloride toxicity are scorched leaf tips and margins
and leaf curling, while in more severe cases growth ceases, leaves abscise and
trees die. Necrotic areas develop on leaves of trees exposed to high sodium
levels (Jindal et al., 1976; Kadman et al., 1976; Gazit and Kadman, 1980). Irri-
gation water concentrations of 20–60 mM sodium chloride (NaCl) or sodium
sulfate (Na2SO4) reduced leaf area and changed the branching structure of
container-grown mango trees, suggesting that salinity resulted in reduced
leaf cell elongation, and affected the activity of the terminal meristem
(Schmutz and Lüdders, 1993). As the duration of the exposure to saline con-
ditions increased, transpiration decreased exponentially (Schmutz and Lüd-
ders, 1993). In a later study, Schmutz (2000) found that following a gradual
increase in salinity of the nutrient solution applied to potted polyembryonic
‘13-1’ rootstocks (from 0 to 120 mM NaCl over 15 days), Amax significantly
declined despite there being no visible leaf symptoms of salt toxicity. The
decline in Amax occurred within 6 days of beginning the salinity treatment,
which was 15 mM NaCl for 3 days followed by 30 mM NaCl for 3 days. This
indicates that photoassimilation in mango is quite sensitive to exposure of
trees to salinity.
There is considerable variation in salinity stress of mango, both within
and between populations of mono- or polyembryonic mango ecotypes. Based
on the results of limited studies, there appears to be greater salt tolerance in
polyembryonic than in monoembryonic populations (Jindal et al., 1975; Kad-
man et al., 1976). In seedling populations from mono- and polyembryonic
cultivars irrigated for 2 years with water containing approximately 10 mM
chloride, most plants developed leaf scorching after 6 months which gradu-
ally became more severe, culminating in degeneration and death. However,
some seedlings which had no damage or only slight toxicity symptoms were
mostly of the polyembryonic ‘13-1’ rootstock cutivar or related types (Kad-
man et al., 1976). Leaf analyses revealed that the chloride concentration in
tolerant seedlings (0.68–0.77%) was greater than in susceptible seedlings
(0.43–0.55%). In addition, tolerant plants had lower leaf concentrations of
potassium, calcium and magnesium than saline-sensitive seedlings, possibly
a result of comparative nutrient dilution since vegetative growth was greater
for saline-tolerant than for saline-sensitive seedlings. Kadman et al. (1976)
also suggested that the mechanism of chloride tolerance in ‘13-1’ was based
on greater physiological tolerance of chloride concentrations in leaf tissues,
Ecophysiology 197
500
350 μmol CO2/mol
600 μmol CO2/mol
400
Dry weight (g)
300
200
100
0
Old Old New New Trunk Roots
leaves branches leaves branches
Plant part
Fig. 6.8. Partitioning of dry matter in ‘Kensington’ mango trees grown for 6 months in
atmospheric CO2 concentrations of 350 or 600 Pmol/mol. Bars represent means (n = 6
trees) ± standard error (Source: redrawn from Schaffer et al., 1999).
198 B. Schaffer et al.
50
350 μmol CO2/mol
40 600 μmol CO2/mol
20
10
0
Skin Flesh Testa Seed Total fruit
Fruit tissues
Fig. 6.9. Partitioning of dry matter in ‘Kensington’ fruit of trees grown for 6 months
in atmospheric CO2 concentrations of 350 or 500 Pmol/mol. Bars represent means
(n = 33–49 fruit) ± standard error (Source: redrawn from Schaffer et al., 1999).
pruning to increase the percentage of leaves exposed to > 60% of full sunlight
will increase the photosynthetic efficiency of the canopy with a potential
improvement in yield. Schaffer and Gaye (1989) increased light interception
of mango by removing 25% of the canopy. This resulted in higher chlorophyll
concentrations in leaves of pruned canopies later in the year. Although net
photosynthesis was not measured in that study, it is likely that the higher leaf
chlorophyll concentrations in the pruned canopies resulted in higher photo-
synthetic rates.
Light utilization of mango can be enhanced by pruning, but the timing of
such treatments is critical. For example, in the subtropics, shoots produced
following harvest generally flower 3–5 months after being exposed to induc-
tive (cool) temperatures. Therefore, trees should be pruned immediately after
harvest to improve light penetration and contain tree size. However, in the
tropics there is a shorter period between the cessation of summer growth and
flowering and summer-grown shoots of many cultivars fail to induct that
year (Scholefield et al., 1986; see Davenport, Chapter 5, this volume). There-
fore, due to the removal of potential flowering sites, summer pruning of
mango trees in the tropics generally reduces yield in the following season.
Growth control of mango trees through the development of dwarfing
rootstocks, low-vigour cultivars and pruning strategies to increase light pen-
etration within tree canopies and by entire orchards, may improve yield per-
formance of this crop. Improved information on light interception, critical
leaf to fruit ratios and relationships between shoot maturity and floral induc-
tion with respect to genotype/environmental interactions will enhance the
development of improved cultural practices for mango production. It is per-
tinent to emphasize that few evergreen fruit trees are as precocious as mango,
or as large at maturity. Orchardists should take advantage of the precocity
and of light interception principles by initial high-density planting, with
hedgerows perhaps the best option. As trees become crowded, however, effi-
ciency of light interception is compromised and remedial action before this
occurs is required to maintain productivity.
6.6 Conclusions
Although much literature in the past stressed problems associated with low
temperature on mango growth and production, sustained high temperatures
with associated soil moisture stress during fruit ontogeny and high evapora-
tive demand are perhaps the major reasons for relatively low yields in mango
orchards worldwide, especially in the tropics. Although mango trees have a
number of survival mechanisms that allow them to cope with stressful envi-
ronments, these come at a considerable energy cost thereby potentially reduc-
ing the availability of carbon-based inputs for fruiting. It is likely that annual
assimilation gains and resource availability during critical developmental
periods are inadequate for sustained high yields of quality fruit. These prob-
lems can be alleviated by development of improved germplasm with adapta-
tion to specific environments (Whiley et al., 2006). In the future, greater effort
Ecophysiology 201
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7 Fruit Diseases
7.1 Introduction
Postharvest diseases can reduce fruit quality and cause severe economic
losses, due to decay resulting in completely unmarketable and blemished
fruits that are often sold in less demanding local markets, where the prices
are considerably lower than export prices. It is clear to the producer that
quality at the time of harvest cannot be improved but merely maintained for
a limited period of time. Harvesting fruits at the optimal stage, with respect
to size and maturity, can, therefore, ensure peak quality and maximum shelf-
life potential. Thus, managing total tree health can contribute to reducing
postharvest losses. It is known that older and neglected orchards may become
a profuse inoculum source for postharvest diseases. Furthermore, preharvest
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
210 (ed. R.E. Litz)
Fruit Diseases 211
7.2 Anthracnose
Mango anthracnose is caused by Glomerella cingulata (Stoneman) Spauld. &
H. Schrenk (anamorph: Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. In
Penz.), C. gloeosporioides Penz. var. minor J.H. Simmonds (Fitzell and Peak,
1984) and Colletotrichum acutatum J.H. Simmonds (Freeman et al., 1998). The
pathogen also causes blossom blight, leaf blight and, in severe cases, tree
dieback (Ploetz, 1994; Ploetz and Prakash, 1997). The form-genus Colletotri-
chum Corda (form-order Melanconiales; form-class Coelomycetes; subdivi-
sion Deuteromycotina) comprises imperfect fungal species which exist as
Glomerella (subdivision Ascomycotina) in their sexual, teleomorphic or perfect
212 D. Prusky et al.
state. These fungal pathogens occur worldwide, and the genus is synony-
mous with anthracnose. Leaf anthracnose appears as irregular black necrotic
spots on both sides of the mango leaves. Lesions often coalesce and form
large necrotic areas, frequently along the leaf margins. Lesions develop pri-
marily on young tissue, and conidia are formed in lesions of all ages. Under
favourable conditions, the fungus can invade the twigs and cause dieback
(Ploetz et al., 1996). Panicle anthracnose or blossom blight can affect both the
inflorescence stalk and the individual flowers; in the stalk, elongated dark
grey to black lesions appear; the blighted flowers are dry, and their colour
ranges from brown to black. Fruits smaller than pea-size can be infected and
abort; whereas, larger fruits that are aborted because of normal self-thinning
or other physiological causes are usually mummified. The resulting mummi-
fied fruit are invaded saprophytically by C. gloeosporioides, and the fungus
sporulates abundantly on them.
Although field anthracnose causes considerable damage, the vast losses
inflicted by postharvest anthracnose are of far greater economic importance.
Postharvest anthracnose appears on the fruit surface, as rounded brown to
black lesions with an indefinite border. Lesions > 2 cm in diameter are fairly
common. Lesions of various sizes can coalesce and cover extensive areas of
the fruit, typically in a tear-drop pattern that develops from the basal towards
the distal end of the fruit. Lesions are usually restricted to the peel, but in
severe cases the fungus can invade the pulp. In advanced stages of the dis-
ease, the fungus produces acervuli, and abundant orange to salmon-pink
masses of conidia appear on the lesions.
contain osmolytes (i.e. glycerol) that are needed for generating the osmoti-
cum from which high turgor pressure will drive the invasive forces of the
penetrating hyphae through the small appressorial pores (0.5 Pm in Col-
letotrichum sublineolum). The osmolyte is generated by the metabolism of
stored glycogen, trehalose and lipids and, at an in vivo concentration of at
least 3.3 M in mature-melanized appressoria (de Jong et al., 1997), this osmo-
lyte is capable of generating turgor pressures as high as 8 MPa (Howard et al.,
1991; Money and Howard, 1996). Melanized appressoria appear to be quite
capable of a forcible, non-enzymatic penetration of an intact host surface.
However, Dickman et al. (1982) suggested that attack on papaya by C. gloeo-
sporioides depended on cutinase production by the pathogen. The germi-
nated appressoria in Colletotrichum develop single infection hyphae that
grow and extend into the waxy cuticle, reach the first layers of pericarp
cells, and then become quiescent for long periods of time (Coates et al.,
1993; Prusky, 1996).
In recent years, the taxonomy of Colletotrichum has been clarified by the
adoption of molecular biological techniques that involve the use of poly-
merase chain reaction amplification, alignment of nucleotide sequences, and
the construction of dendograms, phylogenetic trees and similarity matrices
from the data generated. For example, Sherriff et al. (1994) and Sreenivasap-
rasad et al. (1996) used homologies between the nucleotide sequences from
amplified non-transcribed regions (internally transcribed spacers 1 and 2
(ITS1, ITS2)) and the large subunits (domains 1 and 2) of ribosomal DNA
extracted from a wide range of isolates to both justify and resolve the taxo-
nomic status of Colletotrichum species. Bailey (1997) proposed the adoption of
species aggregates based on these and other data.
Conidia are formed abundantly in the mango canopy (Fitzell and Peak, 1984)
which, therefore, is considered to be the primary source of inoculum. In the
field, C. gloeosporioides produces conidia on lesions on leaves, twigs, panicles
and mummified fruits (Arauz, 2000). Conidia can be rain-splashed onto other
leaves or flowers, where they can cause secondary infections. Developing
fruit can be infected, and some isolates can cause preharvest fruit loss (Gan-
totti and Davis, 1993). In the case of postharvest anthracnose, developing
fruits are infected in the field, but the infections remain quiescent until the
onset of ripening, which occurs after harvest. Once the climacteric period of
the fruit starts, lesions begin to develop but there is no fruit-to-fruit infection.
In this context Prusky and co-workers (Guetsky et al., 2005) suggested that
the capability of C. gloeosporioides to cause early disease symptoms in unripe
fruits depends on the activation of laccases by specific strains of the fungus.
Details of these systems are discussed in the following section.
Colletotrichum gloeosporioides requires free water or RH > 95% to enable
conidial germination and appressorium formation (Fitzell et al., 1984; Dodd
et al., 1991). However, conidia can survive for 1–2 weeks under low RH and
214 D. Prusky et al.
Resorcinol-5-(12-heptadecenyl) Resorcinol-5-(pentadecyl)
HO HO HO HO
(CH2)11 (CH2)14
CH CH2
CH CH3
(CH2)3
CH3
Fig. 7.1. Chemical structure of the 5-substituted resorcinols isolated from mango
fruits.
Fruit Diseases 215
values at which the pathogen is quiescent (Eshel et al., 2002). Ambient alkal-
ization by Colletotrichum is achieved by active secretion of ammonia, which is
produced as a result of protease activity and deamination of amino acids.
The pathogenicity of C. gloeosporioides and expression of the virulence factor
PL-B both depend on raising the ambient pH (Drori et al., 2003). This modu-
lation of environmental pH has been used as the basis for a new approach to
disease control in mango fruits, and is discussed below.
Management
Cultural control
Since the development of mango anthracnose is dependent on moisture or
high RH, orchards ideally should be established in areas with a well-defined
dry season, to allow for fruit development under conditions unfavourable
for disease development. In the tropics, mango flowering usually occurs dur-
ing dry seasons, but anthracnose incidence of > 90% is common in fruits that
develop during the rainy season (Arauz, 1999). In contrast, the incidence and
severity of mango anthracnose can be close to zero in fruits that develop
completely in the dry season, without the application of any other control
measure (Arauz, 2000).
Considerable effort has been invested in understanding and managing
mango flowering. Flowering can be advanced by several weeks by applying
potassium nitrate sprays to mature foliage (Núñez-Elisea, 1985). The growth
retardant paclobutrazol, alone or followed by potassium nitrate sprays, can
also be used to advance flowering (Núñez-Elisea et al., 1993). Both treatments
could contribute to the manipulation of the flowering season to a less sensi-
tive period. Field sanitation of the tree itself is difficult to practise. Elimina-
tion of dry panicles and mummified fruits is time consuming. Bagging results
in reduced anthracnose severity, but it also reduces the red colour of the fruit,
which could reduce consumer appeal (Hofman et al., 1997).
Resistant cultivars
Although all commercial mango cultivars are susceptible to anthracnose,
some are less susceptible than others; ‘Tommy Atkins’ and ‘Keitt’ are less
susceptible than ‘Irwin’, ‘Kent’, ‘Haden’ and ‘Edward’ (Campbell, 1992).
Chemical control
In extreme situations, in which fruit develops entirely under disease-favouring
conditions, seasonal applications of up to 25 sprays of protective and sys-
temic fungicides have been reported (Dodd et al., 1997). However, few fungi-
cides are approved in importing countries for use on mango. Therefore, the
Fruit Diseases 217
Biological control
A number of microorganisms with in vitro or in vivo activity against C. gloeo-
sporioides have been isolated (Jeffries and Koomen, 1992), but few examples
had been used commercially in the field until Korsten (2004) isolated Bacillus
spp. from leaf and fruit surfaces, and effectively controlled anthracnose of
mango. Postharvest control was achieved with semi-commercial preharvest
sprays or postharvest packing house dips, sprays or ultra-low-volume appli-
cations. Integrated treatments involving antagonists combined with quarter-
strength or recommended dosages of fungicides such as prochloraz or sodium
hypochlorite also effectively suppressed postharvest anthracnose of mango.
Commercializing the antagonists in South Africa (Korsten, 2004) and in the
Philippines proved to be difficult because of the limitations set by the local
registration guidelines, and the effect of product formulation on antagonist
performance in commercial applications.
Postharvest control
Traditionally, postharvest control of mango anthracnose has aimed at eradi-
cation of quiescent infections on the fruit. Such eradication is achieved com-
mercially by thermal and chemical treatments, or a combination of both
(McMillan, 1987). Dipping fruit in hot water alone is moderately efficient;
temperatures of 50–55°C for 3–15 min have been recommended, with the
higher temperatures corresponding to the shorter exposures. Fruit from Latin
America entering the USA market must undergo a quarantine hot water
treatment to eliminate fruit fly (Ceratitis capitata and Anastrepha spp.) larvae;
the fruit is immersed in water at 46°C for 90–120 min, depending on variety
and fruit size. The efficacy of the fruit fly quarantine treatment varies from 60
to 85% for elimination of anthracnose infections (McGuire, 1991). Hot water
treatments leave no chemical residue on the fruit and could be a good anthra-
cnose control option for organically-produced mangoes or for mangoes tar-
geted for markets in the USA, where no fungicides are currently approved
for postharvest use. Temperature and time controls are critical, because fruits
can easily be damaged by overexposure to heat. Several fungicides have been
applied after harvest to control mango anthracnose. Benomyl, at rates vary-
ing from 500 to 1000 Pg/ml, was used as a postharvest dip in the past, but its
use is no longer permitted. Thiabendazole at 1000–2000 Pg/ml is also effec-
tive, and there is interest in its registration for postharvest use with mango,
218 D. Prusky et al.
to inoculation with the fungus, but not when fruit were inoculated with the
pathogen first (Korsten et al., 1992), which indicated that the quiescent phase
of the fungus was not affected by the antagonist. Other approaches to disease
control using biological methods included the use of a non-pathogenic strain
of Colletotrichum magna that colonizes the fruit endophytically and prevents
infection by C. gloeosporioides (Prusky et al., 1993); and the expression of an
antifungal peptide in the yeast Saccharomyces, which controlled postharvest
diseases caused by C. coccodes (Jones and Prusky, 2001).
Pathogenesis
Symptoms
Alternaria rot of mango has been increasingly reported as an important
pathogen that causes blossom disease and postharvest fruit rot in ripening
fruits in Australia, Egypt, India, Israel and South Africa. The symptoms are
small, black circular spots that develop around the lenticels. Initially, the
spots are concentrated around the stem end of the fruits, where there are
large numbers of lenticels. The spots can grow and coalesce to become a sin-
gle spot that covers a significant part of the fruit surface. At first, the decay is
firm and does not penetrate the pulp more than 1–2 mm, but later the disease
progresses into the flesh, which darkens and becomes soft (Prusky et al.,
1983). Symptoms of alternaria rot are more limited, darker and firmer than
those of anthracnose. The former pathogen also attacks mango leaves, and
symptoms can be observed throughout the year. The pathogen may also
attack mango inflorescences, resulting in a significant decrease in fruit set.
Epidemiology
The main sources of inoculum are conidia released from infected leaves,
twigs and inflorescences; however, Alternaria spores easily can be found in all
the dry tissues of mango trees in the orchard. Conidia are transferred to the
fruit by air currents and in dew runoff (Ploetz et al., 1994). Germination of
conidia depends on the RH in the orchard during fruit growth. The area of
quiescent Alternaria infection on mango fruit at harvest increased as the num-
ber of hours of exposure to RH ≥ 80% increased over 320 h (Prusky et al.,
220 D. Prusky et al.
1983). Regions with the highest potential for disease incidence are located
close to the 85–90% RH isolines during the fruit growth period. The interme-
diate regions lie between 75 and 85% RH, and the lowest potential risk is in
the dry regions, where the prevailing RH is < 75% (Prusky et al., 1992).
Management
Stem-end rots of mango fruit present one of the most serious postharvest
problems that affect this industry worldwide. They become more prominent
as orchards become older. Losses increase when fruits are stored for pro-
longed periods at low temperatures or when fruits ripen at temperatures
> 28°C. The stem-end rot diseases are caused by a variety of fungal patho-
gens including: D. dominicana (anamorph of Botryosphaeria dothidea), Dothi-
orella mangiferae, L. theobromae (Botryodiplodia theobromae), Phomopsis mangiferae
and Pestalotiopsis mangiferae, among which Botryosphaeria is the dominant
pathogen (Darvas 1991; Johnson et al., 1992; Sangchote, 1998). Stem-end rot
diseases cause heavy losses in mangoes during storage (Prusky et al., 1992;
Mitra, 1997; Kobiler et al., 2001).
Pathogenesis
Johnson and co-workers (1993) have suggested that spores of the pathogen
may germinate and penetrate into the host tissue through wounds, and
remain as endophytes in the branches of mango trees.
Symptoms
Depending on the fungus involved, a variety of symptoms may develop at
the stem and as the fruit ripens. Infections by L. theobromae (B. theobromae), D.
dominicana synonym Fusicoccum aesculi (anamorph of B. dothidea) and D.
mangiferae, cause the formation of diffuse, water-soaked tissue that spreads
222 D. Prusky et al.
from the stem end as fingerlike projections, which darken and coalesce into
circumpedicular lesions or lobed margins (Johnson et al., 1992; Slippers et al.,
2004, 2005). Necrosis remains beneath the cuticle and may penetrate through-
out the fruit flesh. Superficial mycelia may appear around the pedicel and
ruptures of the skin or, in some cases, may penetrate through the epidermis.
The ascomata of D. mangiferae are initially embedded, either separately or
grouped in complex stromata, and they finally erupt through the epidermis
and open. The spore wall is dark and thick, and becomes thinner towards the
end. Conidiophores are hyaline, cylindrical, smooth and branched at the
base. A watery fluid may drain from the stem or from surface ruptures (Kor-
sten, 2004). Diseased fruit could infect a whole box of fruits by direct contact,
and thereby spread the pathogen in the box. In the case of injured fruit,
lesions could appear at some distance from the stem. Stem-end rot diseases
also have a major effect on the flavour of the fruit.
The disease may also cause tip and branch dieback and cankers on mango
trees (Ramos et al., 1991). Anamorph morphology is commonly used to iden-
tify Botryosphaeria spp. (Jacobs and Rehner, 1998; Slippers et al., 2004), but the
morphological distinctions among the anamorphs of some of the closely
related species are not clear. Recent studies based on DNA sequence data have
highlighted taxonomic groups and relationships in Botryosphaeria (Jacobs and
Rehner, 1998; Slippers et al., 2004). These data, combined with morphological
characteristics, could clarify the current taxonomic confusion.
Phomopsis mangiferae is a weak parasite of less economic importance than
the species above; it produces a dark, circumpeduncular lesion with defined
edges that spreads relatively slowly but penetrates deeply into the flesh.
Fruiting bodies may develop on the affected surface. Phomopsis mangiferae
can also be distinguished by a dark pinhead-size pycnidial fruiting body
(Johnson et al., 1992). The lesion caused by Phomopsis may be similar to the
stem-end symptoms cause by C. gloeosporioides and A. alternata. However, the
lesions of the latter two diseases penetrate only to a depth of 10–20 mm.
Lesions of L. theobromae can be distinguished by their wrinkled black
edges which have a velvety appearance. In the dark zone, pycnidial masses
are formed (Johnson et al., 1992). In affected plants, twigs die back from the
tips and into old wood, giving a scorched appearance to the limb with abun-
dant gum secretions from branches, stems and the main trunk. Pestalotiopsis
mangiferae appears as silvery grey spots that vary in size, and are usually
sharply outlined by a dark border. The fungus may appear as a member of
the complex of stem-rot pathogens.
Epidemiology
Management
Cultural control
Johnson et al. (1992) demonstrated that infection of mango fruit before har-
vest occurred through endophytic colonization of pedicel tissue by Botry-
osphaeria spp. present from previous growth flushes, and the possibility of
pruning to promote growth flushing was tested as a means to reduce inocu-
lum in the stem tissue from which new-season inflorescences emerged. Cooke
et al. (1998) reported that the levels of endophytic organisms such as Botry-
osphaeria spp. were reduced significantly when a pruning programme was
implemented in mango orchards as a preharvest control measure. Korsten
(2006) found that prevention of water stress during fruit development and
maturation, and avoidance of placing fruits on the ground suppressed dis-
ease development. He also suggested that harvesting of immature fruit
should be avoided; fruit should be cooled to 13°C immediately after harvest
and chemical treatment, and stored in a well-ventilated place.
224 D. Prusky et al.
Chemical control
Postharvest control of Botryosphaeria spp. was achieved by postharvest dip-
ping, spraying or ultra-low-volume application of benomyl (where possible).
Prochloraz or sodium hypochlorite also effectively suppressed postharvest
stem-end rot of mango (Plan et al., 2002; Korsten, 2006). A combined treat-
ment of wax and hot water (55qC) provide very effective control of most
postharvest pathogens (Sangchote, 1998), but in some cases partial-vacuum
infiltration improved disease control, which suggests that control efficiency
may have been reduced because the fungicide did not reach the pathogen
(Plan et al., 2002).
A treatment consisting of HWB and prochloraz followed by 2,4-dichloro-
phenoxyacetic acid (2,4-D) diluted in wax, reduced side and stem-end decay
by 50–70%, and improved fruit quality during prolonged storage (Kobiler et al.,
2001). The best control was obtained by concentrations of 2,4-D ranging from
75 to 175 Pg/ml; this efficiently controlled side rots caused by A. alternata and
the stem-end rots caused by A. alternata, Phomopsis spp. and Lasiodiplodia
spp. The combination of HWB, prochloraz application and 2,4-D treatment
reduced the incidence of stem-end rot after 4 weeks of storage at 14°C and 7
days of shelf life at 20°C from 86 to 10% in ‘Tommy Atkins’ and from 63 to
12% in ‘Keith’ (Kobiler et al., 2001).
Biological control
Bacillus licheniformis, on its own or alternated with copper oxychloride, has
been evaluated as a preharvest spray treatment to control mango fruit dis-
eases. Preharvest applications of B. licheniformis at 3-week intervals from
flowering until harvest controlled moderate levels of anthracnose, and of soft
rot caused by Botryosphaeria, which suggests a potential treatment for com-
mercial preharvest applications (Silimela and Korsten, 2006).
Other pathogens may attack mango fruits after harvest through occasional
wounds, and cause severe diseases, such as black mould caused by Aspergillus
spp. and transit rot caused by Rhizopus spp. Disease control begins in the field,
and is followed by postharvest sanitation, and avoidance of latex burn (stain)
and mechanical injury. A hot water treatment consisting of 46°C for 60–120
min and fungicides can be used, depending on the cultivar (Spalding and
Reeder, 1986). HWB at 55°C for 20 s shows good control (Prusky et al., 1999).
7.6 Conclusions
Long periods in transit, new marketing approaches for mangoes (e.g. ‘Ready
to Eat’) and stringent international standards and requirements have raised
the need for improved approaches to disease control, in order to preserve
fruit quality. Integrated postharvest treatment has provided a more durable,
Fruit Diseases 225
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Fruit Diseases 229
8.1 Introduction
Diseases affect all tissues and developmental phases of mango. Mango dis-
eases have been reviewed in the first edition of this book (Litz, 1997) and by
Singh (1968), Cook (1975), Snowdon (1990), Ploetz et al. (1994) and Ploetz
(2003). Lim and Khoo (1985), Prakash and Srivastava (1987) and Ridgeway
(1989) have also reviewed this subject. This chapter focuses on foliar, floral
and soilborne diseases of mango. Each is discussed with respect to signifi-
cance, geographical distribution, history, symptoms, causal agent(s), epide-
miology and management. These diseases are caused mainly by eukaryotes
(Domain Eukaryota) among which the true fungi, Eumycota (Ascomycota
and Basidiomycota), predominate. Other less important eukaryotes include
the fungus-like oomycetes (Oomycota), nematodes (Metazoa), and parasitic
plants and green algae (Plantae). With one exception, relatively minor pathogens
of mango are prokaryotes in the Domain Eubacteria; all are Gram-negative
J-proteobacteria. None of these diseases is caused by other plant-pathogenic
eukaryotes (protozoa), prokaryotes (D- and E-proteobacteria, Mollicutes and
Firmicutes), or the nucleic acid-based pathogens, the viruses and viroids.
Diseases above ground are the most important and conspicuous problems on
mango. Since many of the pathogens that incite foliar diseases also affect
panicles, diseases on each are combined in this section, as are a few diseases
that affect the branches and trunks of mature trees.
Algal leaf spot, also known as red rust, is caused by a parasitic green alga,
Cephaleuros virescens Kunze (synonyms: Cephaleuros parasiticus Karst, Ce-
phaleuros mycoidea Karst) (family Trentepohliaceae, division Chlorophyta)
(Lim and Khoo, 1985). It is a common problem on mango and many other
tropical and subtropical plants (Joubert and Rijkenberg, 1971). Although
algal leaf spot can cause serious problems on tea Camellia sinensis (L.) O.
Kuntz, black pepper, Piper nigrum L., and other important crops, it is usually
debilitating on mango only in poorly managed orchards (Lim and Khoo,
1985). In the latter cases, abiotic or biotic stresses, such as mites, insects and
other foliar diseases, can increase the severity of this disease.
Conspicuous symptoms of algal leafspot are orange to rust-coloured,
velutinous patches on both leaf surfaces (Plate 43) (Lim and Khoo, 1985).
They are initially 5–8 mm in diameter, but can merge to involve large, irregu-
lar sections of the leaf. They later assume a dull, greyish green colour, and
eventually become bleached patches. The alga can also affect twigs and
branches, causing the bark to crack as the pathogen’s filaments extend into
the host cortical tissues. The orange tufts produced by C. virescens are the
algal thallus located beneath the host cuticle. They produce erect cells, some
Foliar, Floral and Soilborne Diseases 233
Fig. 8.1. Branch of the thallus of Cephaleuros virescens that has terminated in several
oval sporangia (Photograph courtesy of T.-K. Lim).
Alternaria leafspot
Fig. 8.2. Conidia and conidiophores of Alternaria alternata (Source: Ellis, 1971).
Foliar, Floral and Soilborne Diseases 235
Anthracnose
Anthracnose is the most important disease of mango (Cook, 1975; Lim and
Khoo, 1985; Ploetz, 2003), particularly on fruit in humid, high rainfall envi-
ronments. It is usually replaced by, and is less important than, other diseases
in dry production areas. Anthracnose can also be a serious problem on foli-
age and flowers. Crowded and moist conditions in nurseries can result in
significant damage to young leaves and, in extreme cases, new orchards have
been devastated (Bose et al., 1973). Blossom blight, which has been attributed
to one of the anthracnose agents but is also caused by other fungi, is covered
separately below.
Symptoms
On panicles, necrotic flowers abscise leaving persistent peduncles. Small, cir-
cular dark spots also develop on pedicles and peduncles. Lesions may enlarge
and coalesce to form large patches of necrotic, dark brown tissue. With suf-
ficient rainfall, salmon- to orange-coloured fructifications of the pathogen
develop on the affected tissues. On leaves, lesions are dark brown and sur-
rounded by chlorotic haloes, have irregular, rounded margins, and are not
delimited by veins (Fig. 8.3). Lesions are 0.5–1.0 cm in diameter on mature
leaves, but can expand on young leaves. Eventually, large, necrotic patches
can develop that deteriorate and fall from the leaf giving it a tattered appear-
ance. Although different mango cultivars are known to vary considerably in
their resistance to anthracnose on fruit, reports on the foliar and floral resis-
tance of different cultivars and whether resistances of the various organs are
related have not been published.
236 R.C. Ploetz and S. Freeman
Aetiology
In most production regions, anthracnose is caused by the ascomycete fungus,
Colletotrichum gloeosporioides Penz. (teleomorph: Glomerella cingulata (Stonem.)
Spauld. and Schrenk.) (Cook, 1975; Snowdon, 1990; Ploetz, 2003). In Austra-
lia, Florida USA, India, Japan and Taiwan, Colletotrichum acutatum Simmonds
(teleomorph: Glomerella acutata) plays a minor role (Fitzell, 1979; Prakash,
1990; Weng and Chuang, 1995; Taba et al., 2004; Tarnowski, unpublished). In
Colombia, Colletotrichum boninense J. Moriwaki, Toy. Sato and T. Tsukiboshi
has been reported (Afanador-Kafuri et al., 2003).
Colletotrichum spp. cause diseases on several subtropical and tropical
hosts (Jeffries et al., 1990; Freeman et al., 1998). Cultures of the fungus on
potato dextrose agar (PDA) are greyish white to dark grey and usually pro-
duce an aerial mycelium ranging from a thick mat to sparse tufts (Holliday,
1980). Conidia are hyaline, unicellular and either cylindrical with obtuse ends
or ellipsoidal with a rounded apex and a narrow, truncate base. They are
7–20 × 2.5–5 Pm, and are formed on hyaline to faintly brown conidiophores in
Foliar, Floral and Soilborne Diseases 237
(d)
50μ
(a)
(e)
(b)
10μ
(c) (f )
(g)
Fig. 8.4. (a) Acervulus and emergent setae, (b) conidiophores, (c) conidia, (d) perithecium, (e)
asci, (f) ascospores and (g) appressoria at hyphal termini of Glomerella cingulata (anamorph:
Colletotrichum gloeosporioides) (Source: from Commonwealth Mycological Institute (CMI)
description no. 315).
acervuli that are irregular in shape and about 500 Pm in diameter. Setae are
4–8 × 200 Pm, one- to four-septate, brown and slightly swollen at the base and
tapered at the apex. Hyphopodia have been used to distinguish isolates of C.
gloeosporioides and C. acutatum (Du et al., 2005), but provided ambiguous
results in Florida (Palmateer et al., 2007).
Characterization and taxonomic identification of Colletotrichum spp. has
relied on morphology and host range (Freeman et al., 1998; Du et al., 2005). In
general, C. gloeosporioides (Fig. 8.4) produces longer and narrower conidia
than C. acutatum (Fig. 8.5), as well as circular vs lobed hyphopodia. However,
variability in culture and host range has made these criteria unreliable for
species identification (Adaskaveg and Hartin, 1997; Freeman et al., 1998). Col-
letotrichum gloeosporioides and C. acutatum are species complexes with a large
number of hosts that are so broadly defined that the names are of ‘limited use
to the taxonomist or plant health practitioner’ (Du et al., 2005). Several molec-
ular tools have been implemented to differentiate within and among Col-
letotrichum spp., including: species-specific polymerase chain reaction (PCR)
primers, random amplified polymorphic DNA (RAPD) and arbitrarily
primed PCR (Freeman and Rodriguez, 1995; Afanador-Kafuri et al., 2003);
A+T-rich DNA analyses (Freeman et al., 1993); sequence analyses of the inter-
nal transcribed spacer (ITS) regions of ribosomal DNA (rDNA) (Sreenivasap-
rasad et al., 1996; Freeman et al., 2001; Afanador-Kafuri et al., 2003) and
MAT1-2 mating type sequences (Du et al., 2005).
238 R.C. Ploetz and S. Freeman
(c)
10 μ
(d)
(a) (b)
25 μ 10 μ
Fig. 8.5. (a) Acervulus, (b) conidiogenous cells and setae, (c) conidia and (d) appres-
soria of Colletotrichum acutatum, anamorph of Glomerella acutata (Source: from CMI
description no. 630).
that the mango isolates comprise a C. gloeosporioides population that was dis-
seminated worldwide from a single source, perhaps as an endophyte.
A recent study identified Colletotrichum spp. that infect mango, passion-
fruit (Passiflora spp.) and tamarillo (Solanum betacea cav. Sendt.) in Colombia
and assessed whether cross-infection between the host species occurred
(Afanador-Kafuri et al., 2003). With species-specific PCR primers, most of the
mango isolates were identified as C. gloeosporioides. However, DNA of the
passionfruit isolates and single isolates from tamarillo and mango were not
amplified by either C. acutatum- or C. gloeosporioides-specific primers; they
were identified later as C. boninense (Freeman, unpublished data). Further
molecular analyses determined that the isolates of C. gloeosporioides from
mango were heterogeneous, but that the population of C. boninense from pas-
sionfruit, tamarillo and mango was uniform; it may not be host specific.
The origins and diversity of C. gloeosporioides on mango require more
study. Furthermore, whether distinct populations of this diverse species
occur on different mango cultivars and organs, and whether disease reac-
tions on one mango organ could be used to predict those on another should
be determined. These results would be relevant to host resistance and
improvement of the crop, as well as the chemical and cultural control of this
important disease.
Apical necrosis
Apical necrosis was first reported in Spain in 1991, and now occurs in Israel,
Italy, Portugal and possibly Egypt (Cazorla et al., 1998, 2006; F.M. Cazorla,
Málaga, 2007, personal communication). The disease can be quite damaging,
and limits production when panicles are affected. Apical buds, leaves and
panicles are susceptible (Fig. 8.6a–c), but fruit are not (Cazorla et al., 1998).
Vegetative and floral apices are affected by a dark-brown to blackish necrosis
(Fig. 8.6a, c). Necrosis on leaves begins as blackened, water-soaked lesions,
1–3 mm in diameter, that can coalesce and expand to cover large areas (Fig.
8.6c). Necrosis also extends from affected buds to petioles, through the leaf
midrib, and can cover large areas (Fig. 8.6a). A milky bacterial exudate often
develops on affected apical buds, but infrequently on petioles (Fig. 8.6b).
Large portions of the canopy and high numbers of flowers can be killed.
Fig. 8.6. Symptoms caused by the apical necrosis pathogen, Pseudomonas syringae
pv. syringae. (a) Extensive necrosis on young stem, apical bud, petioles and leaves;
(b) bacterial exudate on necrotic stem; and (c) death of a developing floral panicle and
associated leaf necrosis (Photographs courtesy of F.M. Cazorla).
Foliar, Floral and Soilborne Diseases 241
The causal bacterium, Pseudomonas syringae pv. syringae van Hall, affects
many perennial fruit crops (Hirano and Upper, 1983; Kennelly et al., 2007). It
is an epiphyte and is generally not an aggressive pathogen. Disease usually
develops after high populations of the bacterium develop in host tissues as a
result of host predisposition. Strains from mango produce an antimetabolite
toxin, mangotoxin, which plays a role in pathogen virulence and symptom
development (Arrebola et al., 2007).
Cold, wet weather and host genotype are primary factors in the develop-
ment of apical necrosis (Cazorla et al., 1998, 2006). ‘Tommy Atkins’, ‘Lippens’
and ‘Manzanillo’ are very susceptible whereas ‘Keitt’ and ‘Sensation’ are less
so. Apical necrosis is managed in commercial orchards with Cu-containing
pesticides, although there has been a recent increase in control failures and
Cu resistance in Spain and Portugal (Cazorla et al., 2002, 2006). These out-
breaks have been associated with several different Cu-resistance plasmids in
the causal bacterium. Carzorla et al. (2006) determined that the plant resis-
tance activator acibenzolar-S-methyl and the phosphonate derivative fosetyl-Al
provided control comparable to Bordeaux mixture, and that the latter treat-
ment might protect plants due to the protective film it provides against
wound entry for the pathogen.
Bacterial black spot is a destructive leaf, stem and fruit disease in many pro-
duction areas (Gagnevin and Pruvost, 2001). In India, the disease is called
bacterial canker or black canker due to the cankers it causes on the stems of
some cultivars (Prakash et al., 1994). It can be the most important disease
where fungal-induced diseases are well controlled (Gagnevin and Pruvost,
2001). Bacterial black spot has been identified in Australia, Myanmar, the
Comoros, India, Japan, Kenya, Malaysia, Mauritius, New Caledonia, Paki-
stan, the Philippines, Réunion, Rodrigues, South Africa, Taiwan, Thailand
and the United Arab Emirates (UAE) (Fukuda et al., 1990; Pruvost et al., 1992;
Prakash et al., 1994; Gagnevin and Pruvost, 1995, 2001; Kishun, 1995; Ah-You
et al., 2007b). Given the ease with which the pathogen is disseminated in
propagation materials and its wide, confirmed distribution, bacterial black
spot may be more widely spread than is currently recognized.
Symptoms
Mango leaves, stems and fruit are affected (Manicom and Pruvost, 1994;
Gagnevin and Pruvost, 2001). On leaves, water-soaked spots are initially 1–3
mm in diameter. As they enlarge, they become raised, black and angular, are
limited by veins and surrounded by chlorotic haloes (Fig. 8.7). These lesions
are larger and more conspicuously raised than those caused by other xan-
thomonads that have been recovered from other species in the Anacardiaceae
(Ah-You et al., 2007a). Lesions can merge to form large necrotic patches, and
bacteria may ooze from lesions during wet conditions. Old lesions dry out,
turn white or grey, and crack. Defoliation occurs in severe cases. Anthracnose
242 R.C. Ploetz and S. Freeman
(a)
(b)
Fig. 8.7. (a) Symptoms of bacterial black spot on the undersurface of a mango leaf,
caused by Xanthomonas axonopodis pv. mangiferaeindicae (Photograph courtesy
of O. Pruvost). (b) Symptoms of bacterial spot on the undersurface of a mango leaf,
caused by a yellow-pigmented xanthomonad (Photograph courtesy of R.C. Ploetz).
Bacterial black spot lesions are larger and more raised than those of bacterial spot.
lesions are not raised or as black and angular as those caused by bacterial
black spot. On branches, bacterial black spot lesions are dark and cracked
along the long axis (Fig. 8.8). They develop only on highly susceptible culti-
vars and are often associated with wounds. Conspicuous, star-shaped lesions
are produced on fruit.
Aetiology
Diverse xanthomonads have been recovered from mango and other hosts in
the Anacardiaceae (Gagnevin and Pruvost, 2001; Ah-You et al., 2007a). Only
some of these cause symptoms of bacterial black spot. Early reports that this
disease is caused by Pseudomonas mangiferae-indicae (Patel et al., 1948) and
Foliar, Floral and Soilborne Diseases 243
Erwinia mangiferae (Steyn et al., 1974) are erroneous. The pathogen’s place-
ment in Pseudomonas was probably due to its production of non-pigmented
colonies in culture (P. syringae pv. syringae causes a different disease of mango,
apical necrosis – see above). Cook (1975) indicated that E. mangiferae is a
saprophyte that reached high populations in old lesions.
Pathological, cultural, biochemical, physiological, serological and genetic
data indicate that strains of the pathogen from different production areas are
diverse (Sanders et al., 1994; Gagnevin and Pruvost, 1995, 2001; Kishun, 1995;
Pruvost et al., 2005). Genetic diversity is greatest among strains from South-
east Asia, suggesting that this region of host diversity is also a centre of
pathogen diversification (Gagnevin and Pruvost, 1995).
The pathogen has a single flagellum, is Gram-negative, rod shaped and
0.4–0.5 × 1.0–1.5 Pm (Manicom and Wallis, 1984). It is oxidase negative and
does not reduce nitrates to nitrites. It cannot use asparagine as a sole carbon
and nitrogen source, but is able to hydrolyse starch, esculin, gelatin and
casein. On artificial media, colonies are cream coloured. (The latter trait is
atypical for xanthomonads, which are usually yellow in culture.) Yellow-
pigmented xanthomonads have been recovered from mango in Brazil,
Réunion, South Africa and Florida USA. These strains cause non-raised leaf
lesions (Fig. 8.7b), do not cause fruit or stem lesions, and should not be
244 R.C. Ploetz and S. Freeman
under high disease pressure (Pruvost et al., 1989). During rainy weather,
applications of Cu-based bactericides are recommended. The application
schedules for these compounds focus on protecting fruit, and vary according
to the length of time fruit are exposed to wet conditions (Manicom and Pru-
vost, 1994). Although agricultural antibiotics (e.g. various formulations of
streptomycin sulfate or nitrate) have been reported to be effective (Misra and
Prakash, 1992; Viljoen and Kotzé, 1972), resistance that develops to these
products after continuous use limits their long-term effectiveness against
this disease. Biological control measures have not been widely researched.
Pruvost and Luisetti (1991a) reported little success. In India, Kishun (1994)
indicated that a strain of Bacillus coagulans from the phylloplane of mango
was effective against strains of the pathogen, although control of bacterial
black spot in the field was not reported.
Black-banded disease
and sooty blotch are not dependent on honeydew and grow directly on host
surfaces.
Aetiology
Black mildew and sooty mould are similar in appearance, but their respec-
tive causal agents are distinct (Lim and Khoo, 1985). The black or dark mil-
dews are a group of mostly tropical obligate plant pathogens that produce
two types of hyphopodia (Alexopoulos et al., 1996). Capitate hyphopodia are
lobed appressoria from which infection haustoria are formed, whereas
mucronate hyphopodia function as conidiogenous cells. Black mildew of
mango is caused by Meliola mangiferae Earle (Sordariomycetes, Ascomycota)
(Fig. 8.9).
In contrast, the fungi that cause sooty moulds are diverse saprophytes
that require honeydew to colonize plant surfaces. In Malaysia, Lim and Khoo
(1985) listed coelomycetes (Polychaeton), hyphomycetes (Tripospermum) and
loculoascomycetes (Antennulariella, Chaetothyrium, Limacinula and Scorias),
whereas the reported agents in India were hyphomycetes (Leptoxyphium,
10 mm (b)
(a)
1 mm
10 μm
(c) 250 μm
(d)
Fig. 8.9. (a–c) Signs of black mildew, and (d) microscopic features of the causal
agent, Meliola mangiferae (Source: from CMI description no. 1355).
Foliar, Floral and Soilborne Diseases 247
Blossom blight
Blossom blight can reduce fruit set and production considerably since flow-
ers and large areas of the panicle can be killed. When this disease was con-
trolled with fungicides in the Philippines, a 55–80% increase in fruit set
occurred (Pordesimo, 1982).
Symptoms
Blossom blight starts as a wilt of the affected part of the inflorescence that is
often curved, the ‘shepherd’s crook symptom’ (Fig. 8.10). The peduncle
blackens and dies back from the tip. Internally, discoloration advances
beyond the surface lesion. Large black lesions can develop lower on the
peduncle, and once it is girdled the apex dies.
248 R.C. Ploetz and S. Freeman
Fig. 8.10. Symptoms of blossom blight on panicles of mango. Note the blighted,
curved terminals and almost complete lack of fruit set (Photograph courtesy of
D. Benscher).
Aetiology
The cause of blossom blight is confused. Colletotrichum gloeosporioides has
been reported most frequently as the responsible fungus (Fitzell et al., 1984;
Jeffries et al., 1990), and A. alternata has also been reported to attack panicles
and reduce fruit set (Cronje et al., 1990). Powdery mildew (see section below)
also damages panicles, but its symptoms are distinct from those of blossom
blight. In South Africa, symptoms caused by A. alternata and C. gloeosporioides
are small, mainly superficial black spots, 1–2 × 2–5 mm, on the peduncle
(Darvas, 1993; Lonsdale and Kotzé, 1993). Rather than blossom blight, Lonsdale
and Kotzé (1993) indicated that these pathogens caused blossom spot. In con-
trast, Lonsdale and Kotzé (1993) reported that Dothiorella mangiferae caused
extensive, systemic damage, and Darvas (1993) indicated that Dothiorella domini-
cana is the only fungus that caused typical symptoms of blossom blight.
Crous et al. (2006) placed these fungi in a new genus, Neofusicoccum (see
Decline disorders section below). Work is needed to determine the distribu-
tion of Neofusicoccum-incited panicle disease, and the identity of the most
important blossom blight pathogens worldwide.
Decline disorders
Several diseases of mango have been variously termed blight, canker, decline,
gummosis, twig blight, tip dieback and stem bleeding. They have similar
symptoms and aetiologies.
Symptoms
These widespread problems are not well understood. Symptoms include: (i)
marginal scorching of leaf lamina; (ii) foliar symptoms of nutritional defi-
ciencies, particularly of iron (Fe) and manganese (Mn); (iii) vascular discolor-
ation (Fig. 8.11a); (iv) dieback of small branches basipetally from the terminal
that may or may not progress to defoliation (Fig. 8.11b and c); (v) gummosis,
an oozing of a clear or cloudy exudate either from terminal buds or from
branches, scaffold limbs or trunks (Plate 45); and (vi) root degeneration (Lim
and Khoo, 1985; Ploetz et al., 1996a; Ploetz and Prakash, 1997).
Aetiology
Diverse biotic and abiotic factors may be primary causes of decline symp-
toms or predisposing agents (McSorley et al., 1980; Kadman and Gazit, 1984;
Schaffer et al., 1988; Ploetz and Prakash, 1997). Fungi are the most common
agents. They are endophytes that also cause stem-end rots on mango fruit,
and are usually secondary pathogens that cause disease on weakened, pre-
disposed hosts (Johnson et al., 1992; Ploetz and Prakash, 1997; Slippers et al.,
2005; Slippers and Wingfield, 2007). Several species cause all or some of the
above symptoms when used to individually inoculate plants (Ploetz et al.,
1996a). Their frequent association with one another in affected tissues may
indicate that these symptoms usually develop, or develop more severely,
after multiple infections.
Several of these pathogens are in the Botryosphaeriaceae (Dothidiomy-
cetes, Ascomycota). The taxonomy and nomenclature of these fungi has been
confused, and ‘phylogenetic understanding of major groups within Botry-
osphaeria remains poor’ (Crous et al., 2006). With 28S rDNA sequence data,
Crous et al. (2006) examined natural relationships among available members
of the family. Ten lineages were distinguished, most of which contained ana-
morphs with distinct morphological features. New relationships were revealed
250 R.C. Ploetz and S. Freeman
Fig. 8.11. Among the symptoms that are associated with mango decline are:
(a) internal/vascular discoloration and branch terminal death (tip dieback) that may not
(b), or may be associated with defoliation (c) (Photographs courtesy of D. Benscher).
in some of the lineages that necessitated the renaming of several taxa (Crous
et al., 2006). These new names and holomorphs are used below when discuss-
ing the taxa that occur on mango.
Lasiodiplodia theobromae (Pat.) Griffon and Maubl.) (synonyms: Botryodi-
plodia theobromae Pat., Diplodia natalensis Pole-Evans, and Diplodia theobromae
(Pat.) W. Nowell) is the most common and widespread cause of decline dis-
eases of mango (Ploetz and Prakash, 1997), and affects many other host plants
in the tropics (Punithalingam, 1976). Crous and Palm (1999) declared B. theo-
bromae, a nomen dubium. Denman et al. (2000) reduced D. natalensis and L.
theobromae to synonymy with D. theobromae. However, adopting this change
was questioned by Burgess et al. (2006), who noted that five species of Lasio-
diplodia fell in a phylogenetic clade that had 100% bootstrap support; it was
distinct from a clade that included species of Diplodia and Dothiorella. The
teleomorph of L. theobromae, formerly Botryosphaeria rhodina (Cooke) Arx
(synonym: Physalospora rhodina Cooke), is usually not found in nature. In the
study of Crous et al. (2006), the genus Botryosphaeria was reserved for the type
species Botryosphaeria dothidea (Moug.:Fr.) (anamorph: Fusicoccum aesculi
Corda), which was in a different clade than L. theobromae. However, Crous et
al. (2006) refrained from renaming B. rhodina until the poorly resolved clade
in which it resided could be clarified with work with additional isolates and
analyses.
Lasiodiplodia theobromae attacks weakened trees that are predisposed by:
high and low temperatures; drought; high RH; hardpan soils; sun scorch;
and tar and tanglefoot (Muller, 1940; Das Gupta and Zacchariah, 1945;
Alvarez-García and López-Gracía, 1971; Acuña and Waite, 1977; Ploetz et al.,
1996a). It is often an endophyte, infects wounded plants, and is found in soil,
on dead twigs, mummified fruit and on organic debris beneath trees (Johnson
et al., 1992).
Foliar, Floral and Soilborne Diseases 251
(b)
(d)
(c)
(a)
500 μ
10 μ
Fig. 8.12. Pycnidia (a and b), conidiogenous cells and immature conidia (c) and
mature and immature conidia (d) of Lasiodiplodia theobromae (Source: from CMI
description no. 519).
(d)
(e)
(c)
10 μ
(b)
50 μ
(a)
Fig. 8.13. (a) Vertical section of stroma, (b) part of pycnidial wall and conidiophores,
(c) conidiophores, (d) conidia, and (e) the Scytalidium-like synanamorph of Neoscyta-
lidium dimidiatum (Source: from CMI description no. 274).
Conidia are 18.8–30.4 × 4.5–7 Pm, hyaline, and single celled (Fig. 8.14). The
teleomorph is occasionally produced on OA and has been found in litter
beneath avocado and mango trees (Johnson, 1994b; Michailides et al., 1999).
On twigs, pseudothecia are subglobose to pyriform, 210 × 120 Pm, and im-
mersed beneath the epidermis. On OA, ascostromata are hemi-lenticular and
up to 10 mm wide. Asci are eight spored, bitunicate and irregularly biseriate.
Ascospores are hyaline, single celled, fusiform and 16–25 × 4.5–9.5 Pm.
Mango decline is an important disorder in Florida USA, Israel and
other areas that have calcareous soils (Schaffer, 1994; Ploetz et al., 1996a).
Symptoms include interveinal chlorosis and marginal necrosis of leaves,
(b)
(a)
10 μ
(c)
500 μ
Fig. 8.14. (a) Conidia, (b) conidiophores and (c) a vertical section of a conidioma of
Fusicoccum aesculi, anamorph of Botryosphaeria dothidea (Source: Sutton, 1980).
Foliar, Floral and Soilborne Diseases 255
Management
Controlling decline disorders of mango is difficult. Techniques that could
detect these pathogens in plants would be useful to identify pathogen-free
propagation materials. The internal location and the diversity of fungi that
are involved decrease the opportunities for controlling these disorders with
fungicides (Peterson et al., 1991). Because significant movement of some of
these pathogens may occur via wind and rainsplash, strategic applications of
broad-spectrum protectant fungicides may be effective at certain times of the
year (Lonsdale and Kotzé, 1993), but have not been tested. In India, dieback
was managed by pruning affected portions of the canopy and treating the
wounded areas with a 5:5:50 Bordeaux mix (Prakash and Raoof, 1989). Man-
agement of the controllable predisposing factors, such as drought stress, may
also be beneficial. A better understanding of the epidemiology of these dis-
eases would assist these efforts. Pruning to force synchronous flushes of
foliar growth might enable the avoidance of windows of infection for certain
pathogens (Johnson, 1994b).
256 R.C. Ploetz and S. Freeman
(a)
(a) (b)
(b) (c)
(c)
Fig. 8.15. Decline symptoms induced on ‘Tommy Atkins’ plants artificially inoculated
with isolates of: (a) C. gloeosporioides; (b) Neofusicoccum parvum; and (c) L. theobro-
mae (Photographs courtesy of D. Benscher).
Gall and scaly bark disorders of mango are known in several producing
regions. These diseases are usually minor problems.
Symptoms
In India, bark scaling develops as deep cracks along the entire rootstock por-
tion of the plant, and cracks may penetrate the phloem and become necrotic
(Prakash and Srivastava, 1987). These symptoms resemble those of a scaly
bark disorder, ‘cuarteado’, in Colombia (Cook, 1975). In Hawaii, similar
symptoms developed on mango seedlings (Cook et al., 1971). The bark from
the soil line to the first branches was rough and scaly, and xylem pegs, 5 mm
long, were evident when the bark was removed around leaf scars and
secondary branches.
In Mexico, a disorder known as ‘nanahuate’, ‘bolas’ or ‘buba of mango’,
causes galls, 5–10 cm in diameter, which resemble a cauliflower, are initially
light green, but become dark brown as they die (Fig. 8.16) (Angulo and
Villapudua, 1982). The galls remain attached to trees for many years, and
severely affected branches die. Similar symptoms are found in Florida USA,
and are associated with pruning injuries. Larger galls have also been noted
in Puerto Rico, as well as the US Department of Agriculture (USDA)
Agriculture Research Service (ARS) in Miami and University of Florida
in Homestead (Fig. 8.17) (Ploetz et al., 1996b; R. Rodriguez, personal
communication). Some of the latter galls are large, have rough, scaly exteri-
ors, and are usually found on the main trunk or scaffold limbs of affected
trees. Cracks may penetrate the phloem and become necrotic, but the branch
death that is associated with galls in Mexico and India has not been
observed.
Foliar, Floral and Soilborne Diseases 257
Fig. 8.16. Galls of the ‘buba’ type in Haiti (Photograph courtesy of Carolyn Cohen,
USDA, Animal and Plant Health Inspection Service (APHIS)).
Fig. 8.17. Large, 30-year-old gall on ‘Langra’ in the USDA-ARS mango germplasm
repository in Miami (Photograph courtesy of R.C. Ploetz).
Aetiology
Fusarium decemcellulare C. Brick (synonym: Fusarium rigidiuscula (Brick) Snyd.
and Hans.) causes these diseases in Florida USA, Mexico and Venezuela
(Malaguti and de Reyes, 1964; Angulo and Villapudua, 1982; Ploetz et al.,
1996b). Colonies on PDA are dark carmine-red on the underside. The fungus
produces microconidia in false heads or chains on branched and non-
branched monophialides (Fig. 8.18). Large macroconidia, 92–55 × 7–5.5 Pm,
are produced in slimy yellow sporodochia, c.1 mm in diameter. The fungus’s
258 R.C. Ploetz and S. Freeman
(a)
(b)
(c)
20 μ
Fig. 8.18. (a) Ascus and ascospores of Albonectria rigidiuscula, and (b) micro-
conidia and conidiophores, and (c) macroconidia and conidiophores of its anamorph,
Fusarium decemcellulare (Source: from CMI description no. 21).
Epidemiology
Isolates of F. decemcellulare from mango are only mildly aggressive (Ploetz
et al., 1996b), and require wounding in order to infect. A recent outbreak of
scaly bark in a commercial mango orchard in Florida USA was associated
with pruning wounds. In the cushion gall disease on cacao, F. decemcellulare
interacts with several different insect pests and pathogenic agents (Holliday,
Foliar, Floral and Soilborne Diseases 259
1980; Ploetz, 2007). These insects may facilitate infection and disseminate the
pathogen. Insect-F. decemcellulare interactions have not been investigated on
mango.
Management
No pesticides have been identified to control this problem. Measures that
should be helpful include the removal and destruction of affected branches
and trees in the orchard, disinfestation of pruning equipment to ensure that
the pathogen is not spread during pruning operations, and the use of healthy
planting material in new orchards.
Grey leafspot
Leaf blight
This disease has been reported in India and Nigeria (Hingorani et al., 1960;
Cook, 1975; Okigbo, 2001; Okigbo and Osuinde, 2003), and the causal fun-
gus, Macrophoma mangiferae Hingorani and Sharma (Ascomycota), has also
been intercepted in shipments to the USA from Mexico (Systematic Mycol-
ogy and Microbiology Laboratory, USDA-ARS, Beltsville). This is not a
serious problem.
260 R.C. Ploetz and S. Freeman
50 μm (a)
(b)
(c)
10 μm
Fig. 8.19. (a) Vertical section of an acervulus, (b) mature conidia, and (c) conidiog-
enous cells and developing conidia of Pestalotiopsis mangiferae (Source: from CMI
description no. 676).
Malformation
Symptoms
Malformation affects vegetative and floral meristematic tissues (Fig. 8.20)
(Ploetz, 2001). Vegetative malformation is most serious on seedlings and
small plants in nurseries, especially where seedlings are grown beneath
affected trees, a common practice in the Middle East (Ploetz et al., 2002;
Youssef et al., 2007). Vegetative malformation also occurs on mature trees.
Apical and axillary buds produce misshapen shoots with shortened inter-
nodes and dwarfed leaves that are brittle and recurve towards the sup-
porting stem (Fig. 8.20). Shoots may not expand fully, resulting in a bunched
appearance (i.e. the ‘bunchy-top’ symptom of the disease).
Floral malformation is most important. Affected inflorescences usually
do not set fruit or fruit are aborted. Primary or secondary axes on affected
panicles are shortened, thickened and highly branched (Fig. 8.20). Malformed
panicles produce up to three times the normal number of flowers, range from
one-half to two times the normal size, and have an increased proportion of
male to perfect flowers. Malformed panicles may also produce dwarfed and
distorted leaves (exhibit phyllody).
Aetiology
The aetiology of malformation has been controversial for almost as long as
the disease has been recognized (Ploetz, 2001). Suggested causes include
mites (Narasimhan, 1954), nutritional problems (Prasad et al., 1965), physio-
logical or hormonal imbalances (Dang and Daulta, 1982; Singh and Dhillon,
1989), viruses (Kauser, 1959) and unknown causes (Kumar and Beniwal,
1991). Summanwar et al. (1966) demonstrated that a fungus, identified then
as Fusarium moniliforme Sheld., was responsible for the floral phase of this
disease. Varma et al. (1974) later showed that F. moniliforme also caused
262 R.C. Ploetz and S. Freeman
(a)
(b)
Fig. 8.20. Among the symptoms caused by malformation are: (a) in panicles, an in-
crease in the size and number of flowers and interspersed floral and vegetative organs
(phylody); and (b) in vegetative shoots, compact or retarded growth of buds and brittle,
dwarfed and recurved leaves. Symptoms in (a) are on ‘Haden’ in Michoacan, Mexico
and are associated with an undescribed species in the Gibberella fujikuroi species
complex, whereas (b) is on a ‘Van Dyke’ plant artificially inoculated with an isolate of
Fusarium mangiferae (Photographs courtesy of R.C. Ploetz).
Foliar, Floral and Soilborne Diseases 263
(b) (d) (f )
Fig. 8.21. Microscopic features of Fusarium mangiferae: (a) and (b), macroconidia;
(c) and (d), microconidia, and (e) and (f), microconidia in situ on carnation leaf agar.
Scale bars for (a)–(d) = 25 Pm, and (e)–(f) = 50 Pm (Photographs courtesy of Suzanne
Bullock).
Fig. 8.22. Microscopic features of Fusarium sterilihyphosum: (a) and (b), macroconidia; (c) and
(d), microconidia; (e) and (f), coiled hyphae; and (g) and (h), microconidia in situ on carnation
leaf agar. Scale bars for (a)–(d) = 25 Pm and (e)–(f) = 50 Pm (Photographs courtesy of Suzanne
Bullock).
Three other taxa have been associated with mango malformation. Fusar-
ium oxysporum Schlecht emend. Snyder and Hansen (Fig. 8.23) was reported
in Egypt and Mexico (Bhatnagar and Beniwal, 1977; Kumar and Beniwal,
1991), but these reports appear to be erroneous since bona fide, vouchered
specimens have neither been described nor shown to cause the disease (Plo-
etz, 2001; Rodríguez-Alvarado et al., 2008). Fusarium sp. nov. (Britz et al., 2002)
and F. proliferatum (Gibberella intermedia (Kuhlman) Samuels, Nirenberg and
Seifert) were recovered from malformed mango trees in Malaysia (Leslie,
1995), but their pathogenicity has not been determined.
Epidemiology
Although malformation has been reproduced with F. sterilihyphosum and the
unnamed taxa from Brazil and Mexico, no work has been conducted on the
epidemiology of disease that is caused by these pathogens. Thus, results
below are from work on F. mangiferae or what is presumed to be this species.
Fusarium mangiferae is spread by grafting and in infected nursery stock
(Prakash and Srivastava, 1987). Since seed do not appear to harbour the
fungus (Saeed and Schlosser, 1972; Youssef et al., 2007), seedlings should be
disease free. Microconidia of F. mangiferae are probable infective propagules
since they are the primary spores that are produced by the fungus and form
profusely on dead malformed tissues. The disease spreads slowly in orchards,
perhaps because conidia of the pathogen die quickly when exposed to sun-
light (Manicom, 1989). Experimentally, populations of F. mangiferae in infected
panicles in Egypt and Israel declined rapidly during the summer (Youssef
et al., 2007). Wounding enhances infection and subsequent disease develop-
ment (Ploetz, 2001).
266 R.C. Ploetz and S. Freeman
(a) (b)
(c)
(e) (d)
Fig. 8.23. Microscopic features of Fusarium oxysporum: (a) sporodochia, (b) macro-
conidia, (c) microconidia in false head on monophialide, (d) terminal and intercalary
chlamydospores, and (e) macroconidia and microconidia (Photographs courtesy of
K. O’Donnell).
GFP-labelled microconida of
Fusarium mangiferae
Aceria mangiferae
20.0 μm
Fig. 8.24. Body of the mango bud mite, Aceria mangiferae, to which green
fluorescent protein (gfp)-labelled microconidia of Fusarium mangiferae have adhered
(Photograph courtesy of E. Gamiel-Atinsky).
F. mangiferae, these infections are not systemic and do not appear to result in
symptom development (Youssef et al., 2007).
The localized and variable levels of infection by F. mangiferae that have
been noted in diseased and non-symptomatic tissue (Ploetz, 1994; Youssef
et al., 2007), suggest that there are thresholds of infection, whereby malfor-
mation develops only after a sufficient proportion of a host meristem is colo-
nized by the pathogen. This hypothesis is supported by the long latent period
that exists before symptoms develop in artificially inoculated plants and the
hormonal perturbations that probably occur when meristematic tissues are
infected with this pathogen (van Staden et al., 1989; van Staden and Nichol-
son, 1989; Ploetz, 2001).
Management
Management of malformation can be difficult. New plantings should be
established with pathogen-free nursery stock. Scion material should never be
taken from an affected orchard, and any affected plants that are observed in
the nursery should be removed and burned immediately. Nurseries should
not be established in orchards that are affected by malformation. Once the
disease is found in an orchard, control is possible, but time consuming. In
these cases, cultural management has been most effective (Narisimhan, 1959;
Singh et al., 1974; Manicom, 1989). All affected terminals and the subtending
three nodes are cut from trees, removed from the field and burned. Unfortu-
nately, producers may be unwilling to devote the effort that is required to
ensure that this approach succeeds. In addition, it may be difficult to impose
this treatment on large trees.
A diverse array of pesticides, hormones and growth regulators has been
tested against malformation, but these measures have been marginally effec-
tive. Singh et al. (1994) obtained moderate control with sulfates of cobalt (Co),
cadmium (Cd) and nickel (Ni) in India, but it is unlikely that these toxic com-
pounds could be used safely. Darvas (1987) reduced the percentage of mal-
formed inflorescences from 96% to 48% by injecting ‘Keitt’ trees with the
fungicide fosetyl-Al. This reduction was significant (P < 0.05), but the increase
in fruit yield, 46–95 kg of fruit/tree, was not. Other fungicidal compounds
have been generally less effective (Diekman et al., 1982; Chakrabarti and
Ghosal, 1989). In general, the protected, internal location of the pathogen in
affected trees makes it difficult to control this disease. When applied as foliar
sprays or via continuous drip irrigation, prochloraz reduced the severity of
malformation significantly in Israel, but this was dependent on the timing of
application (Freeman et al., unpublished data). Although disease was not
completely controlled, this and other systemic fungicides might be useful in
future integrated management programmes that would incorporate other
measures such as removal of symptomatic terminals and use of tolerant
cultivars.
Prakash and Srivistava (1987) indicated that ‘There is great variation in
the susceptibility of existing varieties.’ Unfortunately, controlled inoculations
have not been used to determine cultivar resistance, and these reports have
come from non-replicated tests; cultivars listed as ‘resistant’ may have come
Foliar, Floral and Soilborne Diseases 269
from healthy nursery stock or may have escaped infection once planted in
the field (Ploetz, 2001). For example, Bastawros (1996) reported that two
newly introduced cultivars in Egypt, ‘Kent’ and ‘Keitt’, were immune (0%
disease); however, these cultivars are susceptible in Florida USA (Ploetz,
unpublished data). Controlled inoculations with grafted plants of different
genotypes and quantified levels of virulent isolates of F. mangiferae have not
been reported.
Recently, Ploetz (unpublished data) utilized previously described proto-
cols (Freeman et al., 1999) to assess malformation development on grafted
plants of diverse genotypes. Disease development was affected by: the length
of time after inoculation and inoculated apical buds remained dormant after
inoculation; the isolate of F. mangiferae that was used for inoculation; and
mango genotype. Virulent isolates, patience (latent period ranged from 40 to
210 days), and sufficient replication were needed to successfully conduct
screenings for response to malformation. Future work is warranted to investi-
gate attributes that are related, and might predict resistance, to this disease.
The symptoms of malformation suggest that a hormone imbalance
occurs in affected tissues. Singh and Dhillon (1989) assayed levels of indole
acetic acid (IAA), gibberellic acid (GA3) and zeatin in malformed and healthy
mango seedlings. Whereas IAA and GA3 levels were, respectively, about ten
and five times lower in malformed plants, levels of zeatin were about five
times higher. Van Staden and colleagues (Nicholson and van Staden, 1988;
van Staden and Nicholson, 1989; van Staden et al., 1989) examined specific
cytokinins produced by mango and ‘F. moniliforme’ (presumably F. mangiferae).
They determined that the cytokinin complements in healthy and malformed
panicles differed qualitatively and quantitatively, and that the pathogen pro-
duced some of the hormones and metabolites that were implicated in dis-
ease development. However, it was impossible to assign unequivocal roles
for production of hormones by the host and pathogen and the subsequent
development of symptoms. For example, whether production of hormones
by the pathogen directly caused the noted changes or whether hormone pro-
duction by the host was somehow altered in the presence of the pathogen
was not clear. Additional work would be needed to clarify these interactions,
and whether F. sterilihyphosum and the unnamed pathogens in Brazil and
Mexico also produce cytokinins or other hormones in affected plants.
Parasitic plants
The family Loranthaceae contains several parasitic plant species that affect
mango. In Malaysia, Dendrophthoe (fomerly Loranthus) pentandra Linn. is
the most important species (Lim and Khoo, 1985). Other Dendrophthoe spp.,
Elytranthe spp. and Viscum spp. are also known in Malaysia, but are less
important. In India, Dendrophthoe falcata (formerly Loranthus longiflorus) is
common, and other, less frequently encountered, species include Macrosolen
cochinchinensis, Helicanthes elasticus and Elytranthe capitellata (Majumder
and Sharma, 1990). These parasites are usually only important in neglected
270 R.C. Ploetz and S. Freeman
Phoma blight
Phoma blight is widespread in India (Prakash and Singh, 1977). It occurs only on
old leaves. Initially, lesions are minute and yellow-brown (Prakash and Singh,
1977). As they expand they darken to brown or cinnamon, become irregular, and
may ultimately develop dark margins and dull-grey centres. In severe cases,
necrotic patches as large as 13 cm in diameter may form that cause defoliation.
The disease is caused by Phoma glomerata (Corda) Wollenw. and Hochapf
(Prakash and Singh, 1977). It forms globose to obpyriform, light-coloured to car-
bonaceous pycnidia that average 30–400 Pm in diameter. Pycnidia have one to
three ostioles, form singly or in clusters, and have short necks. On PDA, pyc-
nidia and conidia are abundant. Conidia are hyaline to dark coloured, ovoid to
ellipsoid, unicellular or occasionally bicellular, and average 8.3 × 3.2 Pm.
Phoma leafspot
Another Phoma sp. causes a leafspot in India (Prakash and Singh, 1976b), and
is referred to as phoma leafspot. On young leaves, Phoma sorghina (Sacc.)
Boerema. Doren. and Vankest causes irregular to roughly circular, water-
soaked spots, up to 2.5 mm in diameter. Lesions are brown with a yellow to
brown margin. Lesions on midribs are elongated and more conspicuous, and
may coalesce to up to 14 cm in diameter. They can be confused with those
caused by anthracnose.
Pink disease
(a)
100 μ
(b) (f )
(e)
(c)
(d) 20 μ
Fig. 8.25. (a) Conidium-bearing pustule, and (f) conidiogenous cells and conidia
of Necator decretus, and (b) hymenium, (c) basal hyphae, (d) immature and mature
basidia, and (e) basidiospores of its teleomorph, Erythricium salmonicolor (Source:
from CMI description no. 511).
272 R.C. Ploetz and S. Freeman
formed during wet conditions, and conidia and basidiospores are dispersed
by rainsplash and wind.
Pink disease management relies on early detection and removal of
affected tissues from orchards. When removal of syptomatic tissues is imprac-
tical, control depends upon treatment with fungicides. Several protectant
and systemic fungicides are effective (Lim, 1994). They should be used as
soon as symptoms are evident, and as long as the disease is present. All cul-
tivars of mango that have been tested in Malaysia are susceptible (Lim and
Khoo, 1985).
Powdery mildew
Symptoms
Mango cultivars vary in their response to powdery mildew (Palti et al., 1974).
On the most susceptible cultivars, virtually all foliar, floral and fruit parts of
the plant are affected (Plate 46). Powdery growth can cover all tissues on
panicles, resulting in a brown, shrivelled necrosis. Since fruit set and reten-
tion can be affected, the disease can have a profound impact on yield. Foliage
can also be damaged significantly, and young leaves are most susceptible.
White, powdery coatings of conidia develop on either side or both sides of
leaves, depending on the cultivar. When damage occurs on the undersides of
leaves it is often restricted to the mid-rib. In all cases, leaves become dis-
torted, and affected areas turn purple and ultimately necrotic.
Aetiology
Powdery mildew is caused by Oidium mangiferae Berthet, a host-specific fun-
gus (Prakash and Srivistava, 1987; Ploetz and Prakash, 1997). It was first
described in Brazil (Berthet, 1914), and is now recognized in most mango-
producing regions (Palti et al., 1974). Conidium and haustorium traits indi-
cate that O. mangiferae belongs to the Erysiphe polygoni group (Johnson, 1994a).
Although the pathogen was originally classified as Erysiphe cichoracearum by
Wagle (1928), Uppal et al. (1941) noted that it produced saccate and lobed
appressoria, which are not characteristic of E. cichoracearum. The pathogen
has no known teleomorph, a common trait for powdery mildew fungi in the
tropics (Holliday, 1980). Conidia of O. mangiferae are unicellular, hyaline,
elliptical to barrel-shaped and measure 33–43 × 18–28 Pm (Uppal et al., 1941;
Palti et al., 1974). They are produced in large numbers on host surfaces, and
impart a powdery appearance to affected tissues (Plate 46). The lengths of
germ tubes vary depending upon RH, and they terminate in appressoria. Glob-
ular haustoria form in host epidermal cells. Conidiophores are of the pseudoid-
ium type, with two to four septa and a straight basal cell (Boesewinkel,
1980).
Foliar, Floral and Soilborne Diseases 273
Epidemiology
Powdery mildew is most severe during cool, dry weather. Conidia are dis-
seminated by wind and are released on a diurnal basis (Schoeman et al.,
1995). Peak spore release, between 11:00 to 16:00 h, was positively correlated
with hourly temperature and negatively correlated with RH, vapour pres-
sure deficit and leaf wetness (all P < 0.01). Conidia germinate at temperatures
ranging from 9 to 32°C (23°C is optimal), and at RH as low as 20% (Palti et al.,
1974). Since germination occurs in such a wide range of RH, disease develop-
ment is usually independent of this parameter. Infection can occur within
5–7 h, and conidia are produced within 5 days of infection. Disease develop-
ment occurs within a very broad range of temperatures, 10–31°C. Gupta
(1989) reported that dry weather encouraged disease development.
Management
Mango cultivars vary in their resistance to powdery mildew (Palti et al.,
1974). ‘Zill’, ‘Kent’, ‘Alphonso’, ‘Seddek’ and ‘Nam Doc Mai’ are very sus-
ceptible; ‘Haden’, ‘Glenn’, ‘Carrie’, ‘Zebda’, ‘Hindi be Sennara’, ‘Ewaise’ and
‘Keitt’ are moderately susceptible; and ‘Sensation’, ‘Tommy Atkins’ and
‘Kensington’ are slightly susceptible (Ploetz et al., 1994; Nofal and Haggag,
2006). In India, Tiwari et al. (2006) reported that ‘Baigan Phalli’, ‘Barbalia’,
‘Dabari’, ‘Dilpasand’, ‘Khirama’, ‘Nagarideeh’, ‘Oloor’ and ‘Totapari’ were
highly resistant and ‘Amrapali’ was most susceptible.
Schoeman et al. (1995) recommended that the first fungicide application
to control this disease should occur when panicles begin to change colour.
Assuming an effective period of 3 weeks for a given application, they con-
cluded that applications should continue every third week until panicle sus-
ceptibility decreased at the end of fruit set. Powdery mildew is easily
controlled with S (Johnson, 1994a). Other fungicides are effective, but many
have negative environmental impacts (Ray, 2003; Tavares et al., 2004). Foliar
sprays of di-potassium hydrogen orthophosphate (K2HPO4) and potassium
di-hydrogen orthophosphate (KH2PO4), systemic fungicides, and an alterna-
tion of fertilizer and systemic fungicides inhibited powdery mildew on pan-
icles (Reuveni et al., 1998; Nofal and Haggag, 2006). Treatments of the
fertilizers with half or a quarter of the recommended rate of sterol-inhibitor
fungicides and Kresoxym-methyl provided protection comparable with or
superior to that of standard fungicides alone (Oosthuyse, 1998; Reuveni et al.,
1998). Sulfur can burn flowers and young fruit during warm, sunny condi-
tions (Johnson, 1994a), and three fungicides used during bloom, dinocap,
fenbuconazole and hexaconazole, can reduce pollen germination (Dag et al.,
2001).
Scab
This is a disease that is known by several different names in Brazil and the
Middle East and is the only one that routinely kills mango trees. ‘Seca’ (dry-
ing), ‘murcha’ (withering), branch blight and Recife sickness in Brazil, was
first recognized in Pernambuco in 1938, and is now also found in Bahia,
Goias, the Federal District, Rio de Janiero and São Paulo (Ribeiro, 1997; Colo-
simo et al., 2000; Silveira et al., 2006). It threatens neighbouring states due to
its efficient movement in infected propagation materials, on pruning equip-
ment, and via a mobile beetle vector.
In 1998, a disease termed ‘sudden decline’ began to kill trees in Oman
(Al Adawi et al., 2003, 2006), about the same time a similar problem (i.e. quick
decline or sudden death) was observed in Pakistan (Malik et al., 2005). In
many ways these diseases resembled seca. Circumstantial evidence sug-
gested that the disease was introduced from Brazil by a producer with
orchards in Oman and Pakistan (M. Deadman, Muscat, 2005, personal com-
munication). By 2007, many mango-producing areas in Oman and Pakistan
were affected and uncontrolled dissemination of infected germplasm may
have spread the disease elsewhere in the region. Its spread into India should
be investigated (A.W. Cooke, Indooropilly, 2007, personal communication).
Symptoms
Symptoms include: discoloration of the vascular cambium; exudation of an
amber-coloured gum from the trunk and branches, particularly from galler-
ies of the putative beetle vector of the pathogen; wilting; rapid death of
branches and entire trees without defoliation; and a scorched appearance of
dead trees (Plate 47) (Junqueira et al., 2002; Al Adawi et al., 2006). On grafted
trees, scions, rootstocks or both may be susceptible and exhibit vascular
Foliar, Floral and Soilborne Diseases 275
Aetiology
Ceratocystis fimbriata Ellis and Halst. sensu lato (s.l.) (anamorph: Thielaviopsis
sp.) was reported in Brazil in the 1930s (Viegas, 1960; Ribiero, 1980; Silveira
et al., 2006), and is recognized as the primary cause of seca. Diplodia recifiensis
Batista (= Lasiodiplodia theobromae?) was indicted as the cause of Recife sick-
ness in Brazil (Batista, 1947), but it probably plays no role or a secondary role
in the development of this disease (see below). In Oman, C. fimbriata s.l.
causes sudden decline, but L. theobromae and Ceratocystis omanensis Al Subhi,
M.J. Wingf., M. van Wyk and Deadman are also associated with the disease
(Al Adawi et al., 2006; van Wyk et al., 2007). The ease with which L. theobromae
and the difficulty with which C. fimbriata s.l. are recovered from affected trees
may have been responsible for previous assumptions that ‘D. recifiensis’
caused Recife sickness in Brazil and L. theobromae caused sudden decline in
Oman (Batista, 1947; Ploetz and Prakash, 1997; Al Adawi et al., 2003, 2006).
Ceratocystis contains many pathogens, particularly of trees (Kile, 1993).
The wide host range of C. fimbriata s.l. led Webster and Butler (1967) to
hypothesize that it was a species complex, and DNA sequences have begun
to delineate some of the host-specific, often morphologically indistinct, taxa
in the species (van Wyk et al., 2007). A contemporary view is that C. fimbriata
sensu stricto (s.s.) specifically refers to the cause of black rot of sweet potato
(Ipomoea batatas L.) on which it was first described (Halsted and Fairchild,
1891). Other cryptic, monophyletic lineages of C. fimbriata s.l. have been
described as distinct species (Engelbrecht and Harrington, 2005; Johnson
et al., 2005; van Wyk et al., 2005, 2007), and more will likely follow.
Two new Ceratocystis spp. have been described on mango in the Oman
Gulf region. Ceratocystis omanensis belongs to the Ceratocystis moniliformis
Hedgc. s.l. species complex (Al Subhi et al., 2006), which are typically not
primary pathogens. Ceratocystis omanensis is a minor pathogen on mango.
The primary sudden decline agent in Oman and Pakistan, C. fimbriata s.l.,
represents a monophyletic lineage based on ITS, E-tubulin and translation
elongation factor (TEF) 1-D DNA sequence comparisons, and it has unique
morphological characteristics; it was renamed Ceratocystis manginecans M.
van Wyk, A Al Adawi and M.J. Wingf. sp. nov. (van Wyk et al., 2007).
On 2% malt extract agar (MEA), colonies of C. manginecans are greyish
olive and have a banana odour (van Wyk et al., 2007). Hyphae are smooth
and segmented (Fig. 8.26). Ascomatal bases are globose, black and (153–)192–
254(–281) Pm in diameter; ascomatal necks are dark brown, lighter towards
the apices (514–)557–635(–673) Pm long, (25–)32–42(–48) Pm, wide at the
276 R.C. Ploetz and S. Freeman
(c)
base, and (14–)16–22(–26) Pm wide at the tip; and ostiolar hyphae are hya-
line, divergent and (42–)45–59(–69) Pm long. Asci are evanescent, and
ascospores are hyaline, hat-shaped, 3–4 Pm long, and 4–5 Pm wide without,
and 7–8 Pm wide within the sheath. Primary conidiophores are phialidic,
lageniform, hyaline, (72–)81–109(–144) Pm long, 5–7(–9) Pm wide at the base,
6–8(–9) Pm wide at the broadest point, and 3–6 Pm wide at the tip. Secondary
conidiophores are tube like, flared at the mouth, short, hyaline, (59–)65–
77(–84) Pm long, 5–8 Pm wide at the base and (5–)6–8 Pm wide at the tip.
Primary conidia are hyaline, cylindrical, (15–)23–29(–33) Pm long, and 3–6
Pm wide; and secondary conidia are hyaline, barrel shaped, (8–)9–11(–12) Pm
long, and 5–7(–8) Pm wide. Chlamydospores are brown, thick-walled, glo-
bose to subglobose, (11–)12–14 Pm long and 9–11(–12) Pm wide.
Two isolates of C. fimbriata s.l. from mango in Brazil (CBS 114721 and CBS
600.70) have been compared to isolates of C. manginecans (van Wyk et al.,
2005, 2007). They are similar to, but differ significantly from, C. manginecans.
They reside with C. manginecans in a clade that contains other New World
Foliar, Floral and Soilborne Diseases 277
Epidemiology
Genotype has a profound impact on disease development, and severe epi-
demics occur wherever susceptible rootstocks and/or scions are used. Greater
disease develops when trees are stressed, although it is not clear whether this
results from an increased attraction of the vector to stressed trees or reduced
disease resistance in the host. The associated pathogens are moved easily in
infected germplasm, the route by which the diseases have spread in Brazil
and probably to Oman. Pruning implements also move the pathogen, and soil,
once infested with chlamydospores of the pathogen, can be a long-term reser-
voir of inoculum. Insect dissemination plays a particularly insidious role.
Beetles (Coleoptera: Scolytidae) are closely associated with seca in Brazil
(Batista, 1947; Viegas, 1960; Piza, 1966; Ribiero, 1980). Batista (1947) indicated
that Xyleborus affinis was the sole vector of D. recifiensis. In contrast, Ribiero
(1980) reported that Hypocryphalus mangiferae Stebbing was the primary vec-
tor of C. fimbriata s.l. (Fig. 8.27). It produced galleries in the cambium of
affected trees (Plate 47a), and was the only scolytid found on healthy, as well
as diseased, trees. Hypocryphalus mangiferae is also associated with the dis-
eases in Oman and Pakistan, where C. manginecans is recovered from the
insect and trees are commonly found with insect probing damage before dis-
ease develops (Al Adawi et al., 2006; van Wyk et al., 2007).
The interactions between H. mangiferae and the Ceratocystis pathogens
of mango are incompletely understood. In olfactometer tests in Brazil,
H. mangiferae was attracted to cultures of C. fimbriata s.l., and larvae of the
insect were raised to adulthood on the fungus (Ribiero, 1980). Several other
species, many of which are in the genus Xyleborus, were also associated with
seca, but because they were found only in diseased trees they appeared to be
opportunistic feeders on C. fimbriata s.l. rather than primary vectors. Although
the sequence of events in Brazil and the Oman Gulf has not been studied, it
is probable that H. mangiferae contaminates its body with these pathogens
while feeding in diseased trees and subsequently disseminates the pathogen
to healthy trees.
Hypocryphalus mangiferae is thought to be native to some of the same
areas in southern Asia where mango evolved (Wood, 1982; Butani, 1993;
278 R.C. Ploetz and S. Freeman
Fig. 8.27. Hypocryphalus mangiferae, vector of the seca and sudden decline
pathogens (Photograph courtesy of R.C. Ploetz).
Atkinson and Peck, 1994; Mukherjee, 1997). Thus, the insect would have
been introduced into Brazil and would have been a new encounter, rather
than coevolved, relationship with C. fimbriata s.l. (van Wyk et al., 2007). In
contrast, if C. manginecans were introduced into Oman and Pakistan from
Brazil, it may have then established a relationship with a native insect.
Although the available information suggests that the H. mangiferae–
Ceratocystis interactions on mango were recent, opportunistic encounters in
the New World, additional work is needed.
Management
Given the ease with which these pathogens are moved and their destructive
impact, preventing their dissemination to new areas must be a high priority.
Pathogen-free propagation material should be used whenever new plantings
are established and germplasm is moved. Clean pruning implements should
be used in affected areas, and should be frequently disinfested with bleach,
formalin or other disinfestants (Junqueira et al., 2002). Trees that have been
killed by the disease must be removed and destroyed as they are significant
reservoirs of inoculum and infested vectors. Where partially resistant culti-
vars are grown, the removal and burning of affected branches and treatment
of the exposed branch stubs with Cu fungicides is recommended (Ribeiro
et al., 1995; Ribeiro, 1997).
Managing these diseases with fungicides on susceptible cultivars would
be a challenge. External applications of protectant or systemic fungicides
would probably be ineffective, given the internal, protected location of the
pathogen. Injecting fungicides, as is done to control Dutch elm disease, might
be effective. However, this would probably not be allowed where concerns
Foliar, Floral and Soilborne Diseases 279
Stigmina leafspot
Fig. 8.28. Conidia of Stigmina mangiferae, cause of stigmina leaf spot (Source: from
CMI description no. 097).
Nematode damage
Phytophthora diseases
(a) (b)
Fig. 8.30. (a) Sporangia, (b) oogonia with amphygynous antheridia and oospores,
and (c) chlamydospore of Phytophthora palmivora (Source: from CMI description
no. 831).
284 R.C. Ploetz and S. Freeman
Fig. 8.31. (a) Semipapillate sporangia and (b) oogonia of Phytophthora citricola
(Source: from CMI description no. 114).
the Spanish Type Culture Collection, CECT 20567, caused root rot on
‘Florida’ and lesions on leaves and stems of seedlings of ‘Gomera 3’.
The oomycete, Pythium vexans de Bary, can cause root rot and wilt of seed-
lings (Lim and Khoo, 1985). In Malaysia, this pathogen caused seedling losses
of up to 30% in nurseries. Symptoms included wilting of foliage, which
initially becomes pale green, but later develops necrotic patches. Roots
develop a wet, blackened necrosis that begins in fine roots and progresses to
larger roots and the root collar. Death of seedlings often occurs. Lim and
Khoo (1985) indicated that overcrowding, excessive moisture and the use of
polybags favoured this disease.
Prakash and Singh (1980) reported that the basidiomycete Rhizoctonia
solani Kuhn (teleomorph: Thanatephorus cucumeris (Frank) Donk) caused root
and damping off of seedlings in India (Fig. 8.32). Affected tissues become
soft, dark brown or black, and seedlings may ultimately become completely
girdled and collapse. Mycelia and sclerotia of the pathogen form conspicu-
ously on affected tissues.
Sclerotium rot
This disease has been reported in Brazil (Almeida et al., 1979), India (Prakash
and Singh, 1976a) and the Philippines (Palo, 1933). The causal fungus, Sclero-
tium rolfsii Sacc. (teleomorph: Athelia rolfsii (Curzi) Tu and Kimbrough;
Foliar, Floral and Soilborne Diseases 285
(d)
(a)
(b)
20 μ
(c)
Fig. 8.32. (a) Sclerotial cells, (b) mycelium and (c) monilioid cells of Rhizoctonia
solani, and (d) basidia and basidiospores of its teleomorph, Thanatephorus cucumeris
(Source: from CMI description no. 406).
Verticillium wilt
Verticillium wilt of mango was first reported in Florida USA (Marlatt et al.,
1970). The disease was originally attributed to Verticillium albo-atrum Reinke
and Berth., but this was before Verticillium dahliae Kleb. was recognized as a
distinct species. Since the causal fungus described by Marlatt et al. (1970)
formed microsclerotia, it is clear that V. dahliae was involved (Fig. 8.33).
Symptoms of the disease include a ‘firing’ and necrosis of leaves, usually
in a portion of the canopy. Sectoral development of the disease often does not
progress to other portions of the trees, which may recover. Killed leaves usu-
ally remain attached to the tree, and the xylem of affected branches is dis-
coloured brown (Fig. 8.34). Verticillium wilt is relatively uncommon, and is
286 R.C. Ploetz and S. Freeman
(b)
10 μ
Fig. 8.33. (a) Verticilliate conidiophore, (b) phialospores and (c) immature and (d) mature
microsclerotia of Verticillium dahliae (Source: from CMI description no. 256).
found on land where susceptible vegetable crops (i.e. potato, tomato and
aubergine) were recently grown (Pohronezny and Marlatt, 1982). New mango
orchards should not be planted on such sites.
(a) (b)
10 μ (c)
20 μ (d)
Fig. 8.35. (a) Upper and (b) lower surface of sporophore, (c) basidiospores, and (d) hyphae of
Rigidoporus lignosus (Source: from CMI description no. 198).
8.4 Conclusions
As mango production continues to increase in different regions, so will the
scope and types of disease problems. Although new fungicides and bacteri-
cides will be developed in the future, it is probable that reliance on these
tools will diminish. In recent years, the regulation of pesticides has increased
while the number of new compounds that have been registered for use has
decreased. Since it appears certain that this trend will continue, the problems
posed by diseases must be solved increasingly with alternative disease
control strategies.
Foliar, Floral and Soilborne Diseases 289
Acknowledgements
The authors thank Francisco Cazorla for comments on apical necrosis; Ber-
nard Slippers and Mike Wingfield for comments on the decline section; Syl-
via Fernandez, John Leslie, Christiano Lima, Kerry O’Donnell and Gerardo
Rodriquez for information on malformation; Ali Obaid Al-Adawi and Mike
Wingfield for comments on seca and sudden decline; and Robert Knight for
translating Portuguese articles on seca. The following individuals are thanked
for pictures: Francisco Cazorla, T.-K. Lim, Oliver Pruvost, Carolyn Cohen,
Suzanne Bullock, Efrat Gamliel-Atinsky, Eric Palevsky and Tony Cooke.
Kevin Hyde, editor of Fungal Diversity, is thanked for permission to use
micrographs of Ceratocystis manginecans.
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9 Physiological Disorders
V. Galán Saúco
Instituto Canario de Investigaciones Agrarias, Tenerife (Canary Islands), Spain
9.1 Introduction
Physiological disorders can be defined as ailments that have not been caused
by infecting organisms. Unlike mango malformation, for example, which has
been considered as a physiological disorder in the past (IBPGR, 1989) but has
a phytopathological origin, true physiological disorders cannot be transmit-
ted from plant to plant, mechanically or by insect bites. For the most part,
they are a result of some form of physical damage or of an altered physiology
of the tree or fruit. In the particular case of fruit disorders – the most preva-
lent and important physiological disorders of mango – according to Subra-
manyan et al. (1971), they are essentially the result of imbalances in metabolism
induced by some factor or factors in the pre- or postharvest environment
that leads to cell collapse and, typically, the appearance of waterlogged or
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 303
304 V. Galán Saúco
brown areas on some part of the fruit. The complexity of events leading to
the occurrence of physiological disorders makes it more difficult to pinpoint
the causal factors than is the case of symptoms caused by pathogens or pests.
Physiological disorders occur relatively frequently in many fruit and
vegetable species. Some well-known examples include bitter pit, water core
and internal breakdown in apple (Atkinson et al., 1980) and in watermelon
(Singh et al., 1975; Cirulli and Ciccarese, 1981), blossom end rot in tomato and
pepper (Winsor and Adams, 1987), mesocarp discoloration and pulp spot in
avocado (Van Lelyveld et al., 1984; Bower and Cutting, 1987) and yellow pulp
in banana (Melin and Aubert, 1973).
Although physiological disorders of annual crops and temperate fruits
have been studied extensively, tropical fruits have only recently been stud-
ied in order to understand and control physiological disorders. This is largely
due to the rapid expansion of these crops during the past few decades.
Preharvest factors that predispose mango fruit to physiological disorders
include growing location, orchard condition, tree nutrition and condition at
the time of harvest. Postharvest storage conditions, such as temperature,
oxygen and carbon dioxide levels, packaging and surface coating treatments,
are also potential contributing factors to the occurrence of these disorders
(Subramanyan et al., 1971).
Evidence of specific metabolic changes that occur in mango fruits that
express this type of disorder is scarce. Probably due to this lack of knowl-
edge, disorders are usually named for the associated environmental factor,
for example chilling injury, or by the altered appearance that the physiologi-
cal disorder confers to the fruit. Clear examples of the latter are internal fruit
breakdown, which is by far the most prevalent physiological disorder in
mango, lenticel spot, fruit cracking and black tip disorder.
calcium (Lim and Khoo, 1985) and also what Velasco Cárdenas (1974) calls
‘ablandamiento del pico’ (literally peak softening).
Cheema and Dani (1934) first reported an IFB disorder, but perhaps the
most comprehensive work published on physiological disorders of the mango
fruit was that of Malo and Campbell (1978), who described the different
degrees of tissue decomposition in fruit of ‘Tommy Atkins’. The comprehen-
sive work of Wainwright and Burbage (1989) reviews the symptomatol-
ogy, chemical changes and causes of IFB, illustrating its occurrence in many
cultivars in virtually all the producing areas of the world. Losses from IFB
vary geographically and also among cultivars, and can affect 100% of the
harvested fruit (Subramanyan et al., 1971; Malo and Campbell, 1978; Bro-
drick and Thord-Gray, 1982; Galán Saúco et al., 1984; Santos Filho et al., 2002;
Cracknell Torres et al., 2003a).
Symptoms
Histology
Altered physiology
A reduction in firmness, total soluble solids, total pectin and pectinase activ-
ity was observed by these researchers in the affected mesocarp tissue, con-
firming the results of Van Lelyveld and Smith (1979) and Roe and Bruemmer
(1981), both of whom also observed greater activity of both pectinase and
malic enzyme in affected pulp of ‘Alphonso’. On the other hand, the higher
level of nitrogen, the lower levels of calcium, and the higher nitrogen to cal-
cium ratio observed by Cracknell Torres and Galán Saúco (2003) in the
affected tissue of ‘Tommy Atkins’ and ‘Lippens’ is in agreement with the
studies of Young (1960) and Young and Miner (1961) but in conflict with
results of Krishnamurthy (1981), who found no differences among calcium
levels between affected and unaffected mesocarp tissues. Other differences
in chemical composition have been detected in affected and unaffected tis-
sues (Subramanyan et al., 1971; Patkar et al., 1983; Nuevo et al., 1984; Gupta
et al., 1985) and further research should be devoted to ascertain the true
nature of this disorder.
Causes
Cultivar susceptibility
in Brazil (Ferreira, 1989), and ‘Gomera 1’ in the Canary Islands (Cracknell Tor-
res et al., 2003a). There is at least one documented case of IFB in Australia
involving fruit of the polyembryonic ‘Kamerunga White’ (Winston, 1984).
Cultivar-related differences with respect to IFB incidence have been
described by Cracknell Torres et al. (2003a). They studied 28 cultivars and
observed that ‘Edward’, ‘Gomera 1’, ‘Irwin’, ‘Valencia Pride’, ‘Mabroka’, ‘Ah
Ping’ and ‘Heidi’ were almost free of this disorder. Modern Israeli cultivars,
such as ‘Shelly’ and ‘Tango’ have also exhibited low frequency of IFB (Lavi et
al., 1997a, b). Observations of the extent of spongy tissue in various hybrids
and selfed progenies of ‘Alphonso’ clearly indicate that this disorder is
genetically determined, and ‘Alphonso’ is apparently homozygous recessive
for this character (Iyer and Subramanyan, 1992).
Control
Despite the close relationship that exists between nutrition and IFB, very few
practical recommendations have been proposed for controlling this problem.
There is general agreement with the observations of Young (1960) and Young
and Miner (1961) that maintaining a low leaf content of nitrogen (<1.2%) and
a calcium level ≥ 2.5% minimizes the amount of affected fruits. High nitrogen
levels enhance vegetative growth and rapid fruit development. Calcium
moves only in the xylem. Calcium concentration in organs such as fruits,
which have a low rate of transpiration and which are preferably supplied via
the phloem, can result in calcium levels falling below the critical level
required for membrane integrity and cell wall stability (Marschner, 1995).
The antagonistic effect of nitrogen fertilization, especially when ammonium
salts are applied, and calcium uptake and accumulation on leaves and fruits,
is well documented in many fruit species (Shear, 1975; Lewis et al., 1977; Lud-
ders, 1979) and has been clearly demonstrated for mangoes (Young et al.,
1962, 1965).
Lime application has also been recommended to increase the cation
exchange percentage to values ≥7.0 (Ferreira, 1989). According to Schaffer
(1994), IFB has been corrected in ‘Keitt’ by calcium application either to the soil
as calcium carbonate (CaCO3) or as a foliar calcium nitrate (Ca (NO3)2) spray.
Cultural practices such as mulching have been cited as being beneficial
for reducing IFB. Mulching lowers the soil temperature and thereby miti-
gates the reflected or rising heat that eventually affects the fruit (Katrodia,
1988; Lad et al., 1992). The accompanying reduction in the transpiration
stream of the tree and the consequent minor mobilization of calcium to the
fruit (Bangerth, 1979) may also be important.
Certain rootstocks can improve calcium uptake and accumulation and
provides an important new procedure for controlling IFB. Fieldwork was
initiated in Malaysia at the end of the last century (Tengku Ab.Malik, 1996),
but further studies remain unreported.
Early harvesting at the green-ripe stage has been recommended as a
measure to reduce IFB (Young, 1957; Young and Miner, 1961; Galán Saúco
Physiological Disorders 309
et al., 1984; Winston, 1984), but for some cultivars this practice results in
lower quality fruits. The elimination at harvesting of fruits that do not exude
latex (Mead and Winston, 1991) and avoiding the exposure of fruit to direct
sunlight during harvest (Gunjate et al., 1982) have been recommended as prac-
tical methods to reduce the incidence of IFB in mangoes in the market. Accord-
ing to Santos Filho et al. (2002), postharvest hydrothermal treatments should
be avoided in fruits coming from orchards with a history of IFB incidence.
Preharvest dips of 0.5–2.0% of calcium chloride (CaCl2), applied from the
second month after fruit set until harvest, can increase calcium content in the
fruit, and has been reported to be an effective control measure for spongy
tissue in fruit of ‘Alphonso’ (Gunjate et al., 1979). In contrast, foliar sprays
with calcium have been reported to be ineffective for increasing leaf cal-
cium content (McKenzie, 1995), and IFB has even been increased by applica-
tion of calcium, which indicates that a nutrition imbalance within the fruit
was induced (Oosthuyse, 1997).
An integrated approach to control insidious fruit rot incidence in ‘Haru-
manis’ has been developed in Malaysia. This includes maintaining soil pH
close to 6.0, late pruning, application of calcium sprays (2% CaCl2) to the
fruits 4–6 weeks after flowering, supplemental irrigation, monitoring nitro-
gen and potassium fertilization and anticipated harvesting (Tengku Ab.
Malik, 1992a, b). Galán Saúco (1999) suggested the following general inte-
grated management measures: avoid planting sensitive cultivars (i.e. ‘Sensa-
tion’, ‘Tommy Atkins’, etc.); use appropriate rootstocks (i.e. ‘Tangkai Panjang’
or others with high calcium absorption capacity); harvest at the green-ripe
stage; use an appropriate fertilization programme that is rich in calcium and
poor in nitrogen; avoid excessive irrigation close to fruit maturity; and
mulch soil in regions with hot summers.
There are notable differences in IFB resistance among cultivars. Because
of the widespread incidence of this problem, breeding for rootstocks and sci-
ons for IFB resistance is critical for resolving this problem (Iyer and Degani,
1997). Iyer and Subramanyan (1992) have described the progress of breeding
studies to overcome IFB in India, obtaining hybrids of ‘Alphonso’ by other
Indian cultivars free from spongy tissue.
Biotechnological approaches also merit attention. Litz and Lavi (1997),
based on work done in tomatoes to control jelly seed (Smith et al., 1990;
Oeller et al., 1991), have suggested that control of IFB could be achieved by
manipulating ripening using the antisense strategy, either by blocking ethyl-
ene biosynthesis or with polygalacturonase, but no work has been reported
in mango so far to test this suggestion.
Outside India, the only known occurrence of black tip is from the Guang-
dong province of China (Zhang et al., 1995). In both India and China, the
disorder occurs in areas close to brick-making kilns, particularly where trees
are more exposed to fume-laden winds coming from the brick works. Although
not fully assessed, it seems that fluorine is the causal agent of this disorder
(Zhang et al., 1995), although a dry hot climate may enhance the effect of
gases. Differences in susceptibility among Indian cultivars have been reported
by Ram (1988), who also suggested the possibility of varietal selection for
orchards in the vicinity of brick kilns. Majumder and Sharma (1985) recom-
mend a minimum distance between kiln and trees of 1.6 km in the path of
prevailing winds and 0.8 km on the other sides, as well as recommending
prevention by spraying three times with borax (0.6%) and caustic soda (0.8%),
for example prior to flowering, during flowering and at fruit set.
Spots and marks on green fruit can have several different causes: cold dam-
age or chilling injury, usually seen as small raised brown or discoloured spots
on the skin, or pitting surrounded by sunken areas affecting the epidermis
and the endocarp (Lizada et al., 1984). Storage temperatures below 13°C have
been identified as major causes of chilling injury, and must be avoided (Meurant
et al., 1997). High temperatures or sudden exposure to sunlight can cause
sunburn, which is characterized by bleached or yellow patches on the skin,
which may become leathery, yellow-brown to black, and lightly sunken.
Trees should be well irrigated during fruit filling and harvested fruit must
always be kept (even briefly) in full shade.
Wind damage, resulting from prolonged rubbing of the fruit against
leaves or dead twigs, requires adequate windbreaks and timely pruning of
dead wood. If hail damage is likely to be a frequent problem, the whole
orchard will need to be protected by nets. Damage caused during harvest,
storage and transport, can only be reduced by careful handling. Sapburn,
312 V. Galán Saúco
caused by prolonged contact with latex leaking onto the skin from a cut stem,
can be minimized by suitable harvesting and packing operations. High levels
of carbon dioxide (CO2) during storage provoke the development of off-
flavours and small internal lesions, as well as exudating brown tissues
(Chaplin, 1986) and inhibition of ripening (Thompson, 1971).
A disorder named pulpa negra, literally ‘black flesh’, which speaks quite
clearly as to the symptoms, has been reported in Mexico, especially in ‘Haden’
(Mora et al., 1998). Although the problem was detected in fruits that had been
stored at 13°C for more than 20 days, it is not entirely clear that low tempera-
ture is the only cause as it also occurred in fruits reportedly stored only at
room temperature.
While some slight cracking of the bark around the trunk may be normal,
some cultivars, like ‘Manzanillo’ or ‘Gomera 4’, are prone to fissuring, with
an unusually high degree of suberification. This disorder is common in the
Canary Islands, Spain, and it is more pronounced on the sides of the trunk
that are wetted by sprinklers than on the less exposed sides (Fernández
Galván and Galán Saúco, unpublished). This phenomenon of stem cracking
has been also associated with bad orchard management, particularly with
bad irrigation practices (Mora et al., 1998). Some rootstocks, including
‘Gomera 1’ in the Canary Islands, frequently exhibit gall-like growths along
the branches, thought to be linked to climatic factors which facilitate
swelling of lateral buds without being enough to provoke flushing and the
corresponding shoot elongation.
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10 Pests
10.1 Introduction
Mango, like most fruit trees, is usually attacked by two or three key pests,
several secondary pests and by a large number of occasional pests in local-
ized areas where it is grown. Worldwide lists of pests of mango have been
published by Laroussilhe (1980), Tandon and Verghese (1985) and Veeresh
(1989). The pests of mango in India (Srivastava, 1998; Anonymous, 2006),
Australia (Anonymous, 1989), Pakistan (Mohyuddin, 1981), Israel (Wysoki et al.,
1993; Swirski et al., 2002), the USA (Peña, 1993), West Africa (Vannière et al.,
2004), Brazil (Assis and Rabelo, 2005), Central America (Coto et al., 1995) and
Puerto Rico (Martorell, 1975) have also been described. Some publications
contain check lists of mango pests and most contain details of life histories and
control of mango pests (Morin, 1967; Golez, 1991; Murray, 1991).
Of c.322 species of insects and mites that have been recorded as minor
and major pests of mango, 127 (39%) are foliage feeders, 87 (27%) are fruit
feeders, 36 (12%) feed on the inflorescence, 33 (10%) inhabit buds and 39
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 317
318 J.E. Peña et al.
(12%) feed on branches, the trunk and roots. The four key pests (fruit flies,
seed weevils, tree borers and mango hoppers) require annual control mea-
sures. Secondary pests generally occur at sub-economic levels, but can
become serious pests as a result of changes in cultural practices and cultivar
or because of indiscriminate use of pesticides against a key pest. For exam-
ple, Mohyuddin and Mahmood (1993) reported that scale insects became
serious pests following non-judicious use of insecticides against fruit flies.
Similarly, mites, Oligonychus spp., are secondary pests of mango, which can
become serious because of human intervention. Occasional or incidental
pests also can cause economic damage only in localized areas at certain times.
The majority of pests reported here are of this category.
With current world emphasis on quality fruit for local consumption and
export, insects that blemish fruit by feeding, scratching or ovipositing in the
pulp or seed can cause high losses. Only fruit flies, seed weevils and lepi-
dopterous larvae actually penetrate the fruit pulp and seed. The feeding of
other pests (e.g. Othreis materna (L.), Gonodonta pyrgo (Cram.), Gonodonta clo-
tilda (Stoll) and Leptoglossus stigmai (Herbest)) often extends only into the
pulp of ripening mangoes (Angeles and Requena, 1966).
Fruit flies
Anastrepha
Anastrepha spp. are endemic to the western hemisphere and their range
extends from the southern USA to northern Argentina and includes the
Caribbean islands (Aluja, 1994) (Plate 54). Twelve Anastrepha species have
been purportedly associated with mango (Norrbom, 2004). Of these, A. obli-
qua, A. ludens (Loew) and A. suspensa (Loew) stand out as economically
important pests of mangoes (Aluja et al., 1996; Norrbom, 2004). Anastrepha
obliqua is reportedly the most common fruit fly pest in the Americas (Jirón
and Hedström, 1988; Nascimento et al., 1992). This species is the most com-
mon fruit fly pest of mangoes in Mexico, Costa Rica, Honduras and Guate-
mala (Soto-Manitiú et al., 1987; Jirón and Hedström, 1991; Aluja et al., 1996;
Camargo et al., 1996; Sponagel et al., 1996), but it also attacks mangoes in
Cuba, Puerto Rico, Jamaica, El Salvador and Venezuela (Norrbom, 2004). In
Mexico, A. ludens commonly attacks mangoes at higher elevations, while A.
obliqua dominates at lower altitudes (Aluja et al., 1996). In Brazil and Ecuador,
mangoes are mainly attacked by A. fraterculus (Wiedemann), but A. turpiniae
Stone, A. serpentina Wiedemann, A. pseudoparallela (Loew) and A. zuelaniae
Stone have been reported (Zucchi, 1988; Rebouças et al., 1996; Arias and Jines,
2004; Norrbom, 2004; Barbosa et al., 2005; A. Malavasi, January 2008, São
Paulo, personal communication).
Birke et al., 2006). Life expectancy varies greatly depending on the host on
which the larvae developed and environmental conditions (for example see
Toledo and Lara 1996, and Malavasi and Zucchi, 2000), but some adults can
live for >150 days (Aluja et al., 2000).
Abundance of A. obliqua populations has been positively correlated with
temperature and negatively correlated with relative humidity (RH) (Herrera
and Viñas, 1977). However, studies by Celedonio-Hurtado et al. (1995) and
Aluja et al. (1996) demonstrated the lack of a clear relationship between rain-
fall and Anastrepha fly captures in mango orchards in Mexico. They indicated
that overall population fluctuation patterns can vary greatly among orchards
within a fairly small geographic region.
Bactrocera
Bactrocera spp. are pests of mango in Africa and Australasia (Drew, 1989;
Leblanc and Allwood, 1997; Leblanc et al., 1997; Tenakanai, 1997; Hancock
et al., 2000; Hollingsworth et al., 2003; Clarke et al., 2005) (Plate 55). The
common species reported on mango include B. tryoni (Frogatt), B. zonata
(Saunders), B. dorsalis (Hendel), B. neohumeralis (Hardy), B. jarvisi (Tryon), B.
papayae Drew and B. frauenfeldi (Schiner) (Umeya and Hirao, 1975; Drew and
Hancock, 1994; Hollingsworth et al., 2003). Two species, B. phillippiensis Drew
& Hancock and B. occipitalis Bezzi, have been recorded for Palau, Pacific
Islands (Secretariat of the Pacific Community, 2005), and recently, a new spe-
cies, B. invadens Drew, Tsuruta and White, was reported for West Africa
(Kenya, Benin) (Lux et al., 2003; Vayssières et al., 2005). Bactrocera correcta
(Bezzi), B. caryeae (Kapoor), B. curcubitae (Coquillett), B. diversa (Coquillett)
and B. tau (Walker) have been reported in India (Australian Government,
2004).
BIOLOGY OF BACTROCERA FRUIT FLIES. The biology of dacine fruit flies was most
recently reviewed by Fletcher (1987); additional details can be found in
Christenson and Foote (1960), Bateman (1972), Robinson and Hooper (1989),
White and Elson-Harris (1992) and Aluja and Norrbom (2000). As with most
pestiferous flies, females within Bactrocera insert their eggs beneath the fruit
skin, especially in ripening fruit; white banana-shaped eggs are usually
deposited in clusters, hatching after 1.5–20 days (White and Elson-Harris,
1992; Messing, 1999). A single female can lay >1000 eggs over her lifetime
(White and Elson-Harris, 1992). One generation requires c.37 days with a
period of 19 days for egg to adult transformation; eggs hatch 38 h after ovi-
position; larvae develop in 7–8 days and adults emerge in 10–11 days (Mess-
ing, 1999). There are usually three larval instars. The larvae tunnel into the
fruit, contaminating it with frass and providing entry for fungi and bacteria.
Depending on factors such as temperature conditions and type of host, larval
development can be completed in 7–8 days (Messing, 1999) but can take up
to 2 weeks (White and Elson-Harris, 1992). When the infested fruit is imma-
ture, the fruit ripens prematurely and is unfit for marketing. Fully-grown
larvae c.7 mm long drop to the ground and enter the soil where they pupate.
Pests 321
After emergence, the females require a protein source for egg maturation
(White and Elson-Harris, 1992). Studies with B. dorsalis in India (Singh, 1991)
indicated that pupal period was longest (18 days) at 15°C and shortest (6
days) at 35°C. Warm, humid weather is favourable for Bactrocera fruit flies
and pest populations build up as mango ripening occurs. Bactrocera popula-
tions decrease during dry periods.
Syed et al. (1970) reported that up to 30% of mango fruit were attacked by
B. dorsalis in July and August. Mohyuddin and Mahmood (1993) reported
that mango fruit are heavily attacked in Central Punjab during July and
August, with up to 35% of the fruit being damaged by B. dorsalis and B. zonata.
Vayssières et al. (2005) reported the presence of B. invadens after the first sig-
nificant rains in mid-April, reaching >900 males captured/trap/week. Trap
captures peaked at 1800 flies/trap/week in mid-June when the presence of
B. invadens was related to ripening of different mango cultivars. Ekesi et al.
(2006) observed that B. invadens shared mango fruit with Ceratitis cosyra in
Africa and suggested that B. invadens is predominantly a lowland pest.
Ceratitis
Eight Ceratitis spp. have been reported to attack mango fruit. The Mediter-
ranean fruit fly C. capitata (Wiedemann) is a common polyphagous pest in
mango-growing areas of Hawaii USA, Israel, Australia, Spain, Mexico,
Réunion and Brazil and elsewhere in South America (Etienne, 1966; Morin,
1967; Galán-Saúco, 1990; Harris et al., 1993; Barbosa et al., 2005; Woods et al.,
2005) (Plate 56). In Africa the most common species are C. cosyra (Walker), C.
fasciventris (Bezzi), C. rosa (Karrsch), C. anonae (Graham) and less frequently
C. capitata (Wiedemann) (Lux et al., 2003); whereas, C. catoirii Guer. occurs in
Réunion (Etienne, 1968). Ceratitis quinaria and C. silvestrii are considered of
economic importance in Benin (Vayssières et al., 2005). Ceratitis cosyra is
broadly distributed across Africa and causes enormous damage, which can
result in total loss of the crop. On average about 20–30% of mango produc-
tion is lost due to this fly species in various African countries (Lux et al.,
2003).
Lopez (1969) reported that high concentrations of McPhail traps reduced the
build-up of fly populations and protected mangoes from severe injury dur-
ing certain periods of the year. However, the McPhail trap has several draw-
backs. It is expensive, breaks easily, is cumbersome to service and, most
importantly, is quite inefficient. Aluja et al. (1989), working in a mixed mango
orchard in Chiapas, Mexico, found that only 31.1% of Anastrepha spp. flies
landing on the McPhail trap were caught with many flies entering the trap
but then escaping. Due to the low efficiency of the McPhail trap it is being
replaced with Multi-Lure® traps, which provide new trap designs. Dry
synthetic-food-based lures have also been developed, i.e. BioLure® (Suterra
LLC, Inc., Bend, Oregon) (Heath et al., 1995, 1997; Epsky et al., 1999) and
Nu-Lure® (Advanced Pheromone Technologies) (Robacker and Warfield,
1993; Robacker et al., 1997; Robacker, 2001).
Fruit fly presence has also been monitored in Australia using Dakpot®
fruit fly traps hung beneath the tree canopy (Anonymous, 1989). Methyl
eugenol is considered the most powerful male lure for oriental fruit flies.
Methyl eugenol was used for successful monitoring, control and erradication
of B. dorsalis in Oahu Hawaii USA (Steiner and Lee, 1955), Rota Island (Steiner
et al., 1965) and Okinawa, Kume, Miyako and Uaekama Islands (Japan) (Iwa-
hashi, 1984). It has been used for monitoring B. umbrosa (F.) in the Philippines
(Umeya and Hirao, 1975), and is used to lure B. invadens in Africa, which is
unlike other African Dacini species that are attracted to Cue-lures (Lux et al.,
2003; Anonymous, 2005). In Palau, Pacific Islands, two lures are used to
attract mango flies: Bactrocera fraeunfeldi (Schiner) is attracted to Cue-lure,
and B. occipitalis and B. philippinensis Drew and Hancock to methyl eugenol
(Secretariat of the Pacific Community, 2005). Bactrocera dorsalis and B. umbrosa
were monitored and controlled by mass trapping of males with methyl
eugenol and infestations were brought to sub-economic levels in Pakistan
(Mohyuddin and Mahmood, 1993). However, concern over the carcinogenic-
ity of methyl eugenol (Waddell et al., 2004) calls for the development of other
para-pheromones to attract Bactrocera fruit flies. Trimedlure is still consid-
ered an important para-pheromone for the Mediterranean fruit fly, with the
exception of C. cosyra adults, which are attracted to terpinyl acetate and not
to trimedlure (Steck, 2003). The attractiveness of mango compounds is
currently being investigated. For example some of the volatiles emitted by
‘Tommy Atkins’ mangoes, i.e. terpenes (p-cymene and limonene), are
attractive to C. capitata adults (Hernández-Sánchez et al., 2001).
Many questions linger with respect to the optimal trap number and time
for trap placement in mango groves. In Naru, to produce mango free of B.
frauenfeldi, 300–400 traps baited with methyl eugenol plus a toxicant were
placed every square kilometre and trapping density was increased around
mango plantings (Anonymous, 2002). Even though large numbers of traps
can be utilized to increase detection sensitivity, the cumulative costs and
logistical considerations do not make this option practical. Traps with spe-
cific and effective lures that can detect the F1 generation at low trap densities
(5–10 traps/km2) would fit the description of a good detection and monitor-
ing device (Tween, 1993).
324 J.E. Peña et al.
Sampling for earlier fruit fly stages can be used to demonstrate that the
fruit is not susceptible to fruit fly attack. For instance, the Caribbean fruit fly,
A. suspensa, may not attack green mango fruit (Peña et al., 2006b). Peña et al.
(2006b) initiated research to determine if the Caribbean fruit fly will attack
green ‘Tommy Atkins’ mangoes and infest it under field and forced labora-
tory conditions. Through a sequential collection of fruit from fruit-fly infested
mango orchards, fruit were dissected for eggs and larvae. At the same time,
fruit were stored and held for puparia emergence. In addition fruit were
exposed in cages to wild fruit flies and traps were placed to verify the pres-
ence of fruit flies. Estimating the time that ‘Tommy Atkins’ fruit remain
immature and therefore non-hosts for fruit flies, may provide a window for
mango exports in some fruit fly-infested areas. Lux et al. (2003) also mention
that small growers tend to harvest fruit before maturation as a strategy to
evade fruit infestation.
Chemical control
Mango plantations account for major insecticide use in the tropics (Cunning-
ham, 1984). From the late 1960s to date, the conventional control of fruit flies
was through toxic bait sprays that combine proteinaceous bait (e.g. hydroly-
sed protein) with an insecticide (López et al., 1969; Soto-Manatiú et al., 1987;
Mangan et al., 2006; Mangan and Moreno, 2007). For many years the insecti-
cide of choice has been malathion (Peck and McQuate, 2000; Burns et al.,
2001). Fruit flies are highly susceptible to any insecticide, and other com-
pounds have also been widely used. For example, Singh (1991) reported that
5% aldrin dust, when mixed in soil, provided the highest residual toxicity to
falling mature larvae (23.4% after 15 days), compared to BHC endosulfan
and quinolphos. Vergherse et al. (2004), working on the control of B. dorsalis
in India, used a rotation of fenthion (0.05%), deltamethrin (0.0028%), carbaryl
(0.2%) and dimethoate (0.06%) to reduce the risk of resistance development.
Yee (1987) concluded that weekly applications of malathion for 3 months can
also provide effective control.
Since the late 1990s, there has been a concerted effort to find environmen-
tally friendly alternatives to malathion (Peck and McQuate, 2000). Cyromaz-
ine, imidacloprid (organochlorinated compound), spinosad (bacteria-derived
insecticide) and phototoxic dyes (Phloxine B) have been successfully tested
against various fruit fly species (Díaz-Fleischer et al., 1996; King and Hen-
nessey, 1996; Peck and McQuate, 2000; Vargas et al., 2002; Liburd et al., 2004;
McQuate et al., 2005). Despite their success, and as is typical with insecticides
intensively applied on a large scale, resistance has already been documented
in the case of spinosad (Wang et al., 2005; Hsu and Feng, 2006) or collateral
damage (i.e. negative impact on natural enemies) (Stark et al., 2004). In
Pakistan, application of pesticides caused reduction of fruit fly infestations,
but their use has created scale insect problems by eliminating their natural
enemies (Mohyuddin and Mahmood, 1993). Such an effect had been reported
by Ehler and Endicott (1984) with pests of olive, citrus and walnut. Another
recent development with respect to chemical control of fruit flies has been the
refinement of the bait-station concept (Mangan and Moreno, 2007).
Pests 325
Biological control
PARASITOIDS. Classical biological control and repeated augmentative releases
of mass-reared parasitoids have been used to suppress Anastrepha, Ceratitis
and Bactrocera populations (Wharton, 1978; Sivinski, 1996; Sivinski et al.,
1996, 1997, 2000; Montoya et al., 2000). In Florida USA, Mexico, Costa Rica,
Brazil, Colombia and Peru, parasitoid species (i.e. Diachasmimorpha longicau-
data (Ashmead), Fopius vandenboschi (Fullaway) and Aceratoneuromyia indica
(Silvestri)) have been imported and released for the control of A. suspensa, A.
ludens and A. fraterculus (Ovruski et al., 2000). Despite the widespread use of
exotic parasitoids over the past 80–100 years, the current trend is to resort to
native species as they pose less of an environmental threat to local fauna
(García-Medel et al., 2007; Aluja et al., 2009).
Use of parasitoids with mango is hindered by the fact that fruit are very
large and therefore provide larvae a refuge from parasitism (López et al.,
1999). As a consequence, Aluja (1993) and Montoya et al. (2000) recommended
that parasitoids should be released outside the mango orchards to attack fly
larvae in their much smaller native hosts and thereby significantly reduce the
size of populations entering mango orchards.
Several parasitoids, for example Opius fullawayi (= Diachasmimorpha
fullawayi (Silvestri)), Diachasmimorpha kraussi, D. Fullaway, D. tryoni (Cam-
eron), Opius bellus Gahan, Biosteres longicaudatus Ashmead (= D. longicaudata),
326 J.E. Peña et al.
Microbial control
Use of pathogens/disease agents (fungi, bacteria, nematodes) has been
attempted with varying degrees of success. For example, Metarhizium
anisopliae has been evaluated in small-scale mango orchards in Kenya using
bait stations laced with the pathogen. Results do not show differences
between use of pathogens and use of insecticides (malathion) (Lux et al.,
2003). Lezama-Gutierrez et al. (2000) also evaluated isolates of M. anisopliae
against larvae of A. ludens. They suggested that M. anisopliae can cause a
22–43% reduction in adult emergence, depending on the soil where the lar-
vae pupariates. De la Rosa et al. (2002) evaluated the fungus Beauveria bassi-
ana (Bals.) under laboratory conditions and concluded that highest mortality
was achieved at the adult stage, while Dimbi et al. (2003) reported on the
pathogenicity of M. anisopliae and B. bassiana on different species of Ceratitis.
Poinar and Hislop (1981), Lindegren and Vail (1986) and Toledo et al. (2006)
have investigated the use of various nematodes, Heterorhabditis bacteriophora,
Heterorhabditis heliothidis (Khan, Brooks and Hirschmann) and Steinernema
feltiae Filipjev, against Anastrepha, Bactrocera and Ceratitis. Finally, Robacker
et al. (1996) and Toledo et al. (1999) have tested various strains/isolates of
Bacillus thuringiensis (Berliner) against larvae of A. ludens, A. obliqua and A.
serpentina. For additional details on microbial control of pestiferous fruit flies,
we recommend the recent review by Dolinski and Lacey (2007).
Predators
In addition to parasitoids, pathogens and nematodes, ants have been used to
control fruit flies in mango orchards. Peng and Christian (2006) used the
weaver ant, Oecophylla smaragdina (Fabricius) for control of the Jarvis fruit fly,
B. jarvisi, in mango orchards in Australia. Van Mele et al. (2007 and references
Pests 327
Host resistance
Yee (1987) reported that B. dorsalis does not attack all mango cultivars to the
same extent. The most susceptible cultivars in Hawaii USA are ‘Hawaiian’,
‘Pirie’ and ‘Sandersha’. Singh (1991) indicated that the frequency of Bactro-
cera injury to physiologically mature fruit of ‘Dashehari’ ranged from 3.6 to
10%, while in fully ripe fruit the frequency of injured fruit ranged from 10 to
25.9%. Highest damage was reported in fully ripe fruit of ‘Mallika’ followed
by ‘Totapari’.
Susceptibility of different mango cultivars to attack by A. obliqua was
measured by Carvalho et al. (1996) who observed that ‘Espada’ showed no
infestation by A. obliqua, whereas ‘Carlota’ was highly infested. In this study,
the survival of adults of A. obliqua was lower when the larvae were fed on
‘Espada’ compared to ‘Carlota’. Furthermore, ‘Espada’ had an adverse effect
on the longevity of A. obliqua females, possibly due to the presence of toxic
substances (Carvalho and De Queiroz, 2002) or absence of essential nutri-
ents. Jirón and Soto-Manitiu (1987) also observed that susceptibility of
328 J.E. Peña et al.
Quarantine treatments
Quarantine treatments have been reviewed by Johnston and Hofman (Chap-
ter 15, this volume). Several quarantine treatments have been developed for
harvested mangoes: irradiation, hot water or hot water followed by immer-
sion cooling are widely used (Sharp et al., 1988, 1989a, b, c; Hallman and
Sharp, 1990; Nascimento et al., 1992; Mangan and Sharp, 1994; Mangan and
Hallman, 1998; Shellie and Mangan, 2002a, b; Bustos et al., 2004; additional
references in reviews by Mangan and Hallman, 1998 and Follet and Neven,
2006).
The mango seed weevil, Sternochetus mangifereae (Fabricius), and the mango
pulp weevil, Sternochetus frigidus (Fabricius) (Coleoptera: Curculionidae) are
important pests of mango (Plates 57–59). Quarantine restrictions prevent the
export of fresh weevil-infested mangoes into uninfested areas. The flesh of
ripe fruit is damaged when mango seed weevil adults emerge from the seed,
and weevil-damaged seed may limit plant propagation in nurseries and
orchards (Johnson, 1989). Early fruit drop may be caused by severe weevil
infestations (Subramanian, 1925). The mango seed weevil occurs from India
through South-east Asia to Australia, on tropical Pacific Islands, in parts of
Africa, in the Caribbean region and in northern South America (Balock and
Kozuma, 1964; Shukla and Tandon, 1985; Johnson, 1989; Schotman, 1989).
Biology
Srivastava (1998) reports that S. mangiferae is a greyish brown weevil, 8 mm
long and about 4 mm wide; its habits are nocturnal, usually feeding and ovi-
positing at dusk. After emergence, adults enter diapause and the duration
varies with the geographic range (Schotman, 1989). According to Shukla and
Tandon (1985), all adults emerging in southern India during June enter dia-
pause from July until late February in the following year. The beginning and
the end of diapause seem to be associated with long-day and short-day
Pests 329
Sampling
Shukla et al. (1988) reported the intra-tree distribution of eggs of S. mangiferae
on ‘Baganpalli’ mango; the highest number of eggs per fruit occurred on fruit
in the lower region of the tree. With increasing tree height, egg deposition on
fruit decreases. No statistical differences on fruit infestation were observed
on north, south, east or west directional quadrats of the tree. Eggs were
deposited in the lower region of the fruit rather than the pedicel. Weevils
enter diapause in crevices in the tree trunk. Most of the weevils (87%) are at
a height of 0–2 m in the trunk compared to 7% at 2–4 m and only 4% above
4 m. Emery (2002) considered that since both the mango pulp weevil (S. frigi-
dus) and the mango seed weevil (S. mangiferae) infest fruit at an early stage,
any fruit is a viable sample; however, as infested fruit all ripen precociously,
the sensitivity of surveys is enhanced by seeking out nearly ripe and fallen
fruit prior to harvest. If the survey coincides with the mango harvest, rejected
or fallen fruit should be inspected. Fruit should be sampled by longitudinal
dissection of fruit through the seed to expose the kernel. If the fruit is ripe, it
should be struck along the longitudinal axis with a hammer, and the seed
should be opened with pliers. The random sampling of 600 fruit from ran-
domly selected properties in each area provides a 9% chance of detecting a
330 J.E. Peña et al.
0.5% infestation of fruit. Sample size can be determined from the following
formula:
Probability of > 1 infested fruit = 1 – probability of no infected fruit in
total sample = 1 – (1 – 0.5%)600 = 1 – (1 – 0.005)600 = 1 – (0.995)600 = 95%
Economic damage
Follett and Gabbard (2000) report that germination rates for infested seed of
polyembryonic ‘Common’ are equal to those of uninfested seed. Germina-
tion is significantly reduced for infested seed of monoembryonic ‘Haden’
compared with uninfested control seed, although germination of infested
seed was >70%. Direct feeding damage to the pulp was found in only 0.11%
of 3602 mango fruit, which suggests that S. mangiferae is a less serious pest of
mangoes than previously considered.
Cultural control
Field sanitation, i.e. the removal of all fallen fruit and seed, is very labour
intensive, and demands complete removal and disposal of fallen fruit. This
procedure has been inconsistent in demonstrating pest control. In India, field
sanitation reduced infestation of the mango nut weevil, Sternochetus gravis
(Fabricius), by only 22% (De and Pande, 1987). In Hawaii USA, field sanita-
tion failed to reduce infestation rates (Hansen and Amstrong, 1990).
Chemical control
Various insecticides have been evaluated for controlling adult weevils, par-
ticularly during oviposition (Balock and Kozuma, 1964; Shukla and Tandon,
1985). The most effective control was provided by the organophosphate fen-
thion, which reduced infestation to <17%. In another field test, the pyrethroid
deltamethrin and the carbamate carbaryl were most effective, both resulting
in <15% infestation rates. Spot application of diazinon on tree trunks was
recommended based on cost, efficiency and least environmental damage.
Verghese et al. (2004) reported that commercially available azadirachtin was
not effective for management of S. mangiferae in India.
Resistant cultivars
Mango cultivars resistant to the mango weevil would be beneficial. Potential
mechanisms of resistance are seedless cultivars, those that form seed early or
those that fruit off-season. Most cultivars grown in Hawaii and India are
equally susceptible (Bagle and Prasad, 1984; Hansen et al., 1989), although
‘Itamaraca’ has shown some resistance (Balock and Kozuma, 1964).
Biological control
The mango weevil has few natural enemies. Parasitoids are unknown, prob-
ably because of the concealed nature of most life stages. Adults may be sus-
ceptible to predation by ants, rodents, lizards and birds (Hansen, 1993). A
baculovirus has been reported that affects the larvae of S. mangiferae (Shukla
et al., 1984).
Pests 331
Mango fruit infested with seed weevil do not show any visible external
symptoms and cause considerable quality control problems and economic
loss to the mango-processing industry as well as restriction in export of fresh
fruit. A non-destructive X-ray inspection method has been developed to
detect weevil-infested fruit. X-ray radiographs of infested mangoes show
dark areas in the seed corresponding to disintegrated kernel tissue as a con-
sequence of feeding by developing grubs. Non-infested mangoes show a uni-
formly light-grey area representing healthy kernel. There is a close agreement
between fruit showing weevil infestation based on their X-ray images and
physical examination of cut fruit, indicating the reliability of the technique.
X-ray imaging has good potential for application in the processing industry
and the export trade as a quality control measure (Thomas et al., 1995).
Damage
Mango fruit in all stages of development are susceptible to attack (Water-
house, 1998). The first larval instars feed on tissues beneath the skin, and
bore through the mango pulp to the seed, which is consumed. Up to 11 larvae
have been recorded in a single fruit, but usually there is only one. Infested
fruit split and rot, and fall to the ground (Anonymous, 1984). In the Guimaras
Islands, the Philippines, Golez (1991) recorded 12.5% fruit infestation and in
serious outbreak years, 40–50% yield reductions are possible. Waterhouse
(1998) considered that since D. sublimbalis is capable of causing such levels of
damage, it might be a more important pest of mangoes than has generally
been realized. It may have been overlooked as a pest or has recently spread
to new areas and has become evident as a pest there. Van Mele et al. (2001)
suggested that damage caused by D. sublimbalis in the Mekong Delta has
been wrongly attributed to fruit flies; however, Waterhouse (1998) states that
soon after boring by D. sublimbalis, secondary infestations with fruit flies
332 J.E. Peña et al.
Biological control
According to Waterhouse (1998) no parasitoids were detected in Java, Indo-
nesia. However, in the Guimaras Islands of the Philippines, the vespid wasp,
Rychium attrisimum, preys on the larvae as they exit the fruit to pupate. Lar-
vae are used to stock the wasps’ nests as food for their young. The egg para-
sitoids Trichogramma chilonis Ishii and Trichogramma chilotreae attack the pest
in Luzon (Golez, 1991). Leefmans and van der Vecht (1930) reported that an
entomopathogenic fungus infected the larvae in Indonesia.
The yellowish green coreid bugs, Amblypelta lutescens lutescens (Distant) and
Amblypelta nitida Stål occur along the coast of Queensland, Australia, and
attack most of the tropical and subtropical fruit crops there (Waite and Huwer,
1998). They prefer to feed on young, green fruit, but A. l. lutescens also dam-
ages the terminals of a number of hosts. In tropical north Queensland, A. l.
lutescens is the dominant species and feeds on young fruit causing black
lesions to develop and the fruit to fall. It also feeds on the terminals and leaf
petioles, causing wilting and dieback (Cunningham, 1989). In the subtropical
south, both species attack mango, but A. nitida confines its attention to the
fruit, while A. l. lutescens also attacks fruit and terminal growth (G.K. Waite,
1995, unpublished results). The bugs breed in natural rainforest areas, and
fly into the orchards to feed on the fruit and terminals. Female bugs lay
individual, opalescent green eggs under the leaves. There are five nymphal
instars and a generation takes c.40 days.
Pests 333
Thrips
Grove et al. (2001) reported that in South Africa, the citrus thrips, Scirtothrips
aurantii Faure and the red banded thrips Selenothrips rubrocinctus (Giard) are
the only thrips that caused lesions on fruit (Plates 60–63). However, Grove
et al. (2001) considered S. aurantii to have more economic importance than S.
rubrocinctus. Grove and Pringle (2000), using a two-stage sampling system to
determine population levels of S. aurantii, showed that S. aurantii has a clumped
distribution, and recommended that 50 fruit per orchard should be examined
in order to obtain accurate population estimates for pest management.
334 J.E. Peña et al.
Blossom pests
Midges, caterpillars, hoppers, thrips and mites are the most important pests
attacking mango inflorescences.
Midges
The mango gall midge or mango blister midge, Erosomya mangiferae Felt, is a
major pest, destroying flowers and up to 70% of set fruit (Plate 64). It was
first described by Felt (1911) in St Vincent (West Indies). Barnes (1948) recog-
nized nine gall midges from mango; two of these, Asynapta sp. and E. man-
giferae, are from the West Indies. Butani (1979) reported five cecidomyiid
species on mango blossoms, including Erosomya indica (Grover and Prasad).
Dasyneura mangiferae (Felt) was reported in Hawaii USA (Vannière et al.,
2004). In recent times, Gagne and Etienne (2006) reported the species Gephy-
raulus mangiferae (Felt), n.comb. infesting mango flowers on the island of
Guadeloupe, French West Indies. Male adults of E. mangiferae are 1.61 mm
and females 1.32 mm long. Eggs are deposited in folds between sepals and
petals of flower buds. The larval stage has four instars. Young larvae are
cream coloured and late instar larvae are yellowish. Larval feeding prevents
flower opening and consequently development of the fruit does not occur.
Infested buds develop as long pointed galls, in which pupation occurs (Van-
nière et al., 2004). Studies of population fluctuation of Erosomya sp. have been
conducted in India by Grover (1986a), who reported that emergence of adults
was higher at 24°C and 60–82% RH compared to lower temperatures and
RH. Abbas (1985) described systematic surveys to determine the percentage
of infestation of E. indica, and showed that infestation follows a negative
binomial infestation. The midge infests the newly emerged panicles by ovi-
positing at bud burst stage, and the first instar maggots bore into the grow-
ing panicle. The second generation then infests very young fruit, which drop
before the marble stage. Sampling of mango midges needs to include affected
tissue, different trapping devices, pheromones, etc. On citrus, use of coloured
sticky traps placed in the tree canopy provides a more efficient method of
sampling the citrus midge, Prodiplosis longifila Cagné, than ground emergence
traps and collection of larval samples (Peña and Duncan, 1992).
In a survey of parasitoids of cecidomyiid pests of mango in India, Grover
(1986b) reported that Platygaster sp., Systasis sp. and Eupelmus sp. were asso-
ciated with Dasineura sp., and Tetrastychus sp. was associated with E. indica.
An external parasitoid, the pteromalid, Pirene sp., attacked Procystiphora
mangiferae (Felt). Predators of the cecidomyiids include Formicai sp., Oeco-
phila sp. and Camponotus sp.
Mango hoppers
Approximately 18 species of leaf hoppers have been reported as pests of
mango. Of these, Idioscopus clypealis Leth., Idioscopus niveosparsus Leth., Idiosco-
pus magpurensis Pruthi and Amritodus atkinsoni Leth., are important (Virakta-
math and Viraktamath, 1985; Viraktamath, 1997; Fletcher and Dangerfield,
2002) (Plate 65). The females deposit their eggs in panicles or midribs of tender
Pests 335
leaves. The adults and nymphs preferentially feed on young leaves and flow-
ers or shoots, and excrete honeydew upon which sooty mould develops
(Ahmed et al., 1981). This interferes with photosynthesis, adversely affecting
plant growth and yield (Godase et al., 2004). Affected inflorescences turn
brown, become dehydrated and fruit set does not occur.
There has been no systematic study of the biology of most of the leaf
hoppers that attack mango; however, biology of A. atkinsoni, I. clypealis and I.
niveosparus has been studied by Sohi and Sohi (1990). Both A. atkinsoni and I.
niveosparsus are multivoltine. In A. atkinsoni, the egg, nymphal (five instars)
and adult stages require 7–9, 15–17 and 3–4 days, respectively (Patel et al.,
1977). Development from egg to adult is normally complete in 25–30 days.
There can be between one and six generations of A. atkinsoni in different areas
of India. In Pakistan there are four to five generations in the Central Punjab
(Mohyuddin, unpublished data). In the Philippines, I. clypealis is reported to
have one to four generations, whereas it has five or six generations in India.
Idioscopus nagpurensis is univoltine. In Pakistan, it normally oviposits in
mango inflorescences during March. Nymphs feed on inflorescences during
March and April. From May to February of the following year, only aestivat-
ing adults are found (Mohyuddin, unpublished data). Most of these species
are quite fecund. Amritodus atkinsoni reportedly lays 200 eggs during its life-
time (Rahman, 1940) and I. clypealis lays 100–190 eggs in the Punjab (Husain
and Pruthi, 1924). Amritodus atkinsoni eggs are laid in the midribs of tender
leaves, flower buds and inflorescences (Babu et al., 2002). Idioscopus niveospar-
sus lays c.238 eggs in 9 weeks under laboratory conditions (Mohyuddin,
unpublished data). Mohyuddin and Mahmood (1993) reported that A. atkin-
soni and I. niveosparsus occur in upper portions of mango trees during differ-
ent times of the year. Amritodus brevistilus and I. niveosparus populations
increase from February to peak in March–April in Sri Lanka, while peaks of
I. clypealis occur in March and September (Kudagamage et al., 2001). Idiosco-
pus clypealis populations peak in south-eastern India during March and April
(Tandon et al., 1983). Idioscopus nivesoparus and I. clypealis peaks coincided
with major and minor flowering periods while population peaks for A. bre-
vistilus coincide with the occurrence of vegetative flushing (Kudagamage
et al., 2001).
Azizur Rahman and Singh (2004) demonstrated that A. atkinsoni popula-
tions on panicles of ‘Langra’ mango were negatively correlated with high
RH; whereas no significant relationships were observed with rainfall, sun-
shine and wind velocity.
SAMPLING. Very few sampling studies have been reported for hoppers on
mango. A sequential sampling plan for mango hoppers was recommended
by Verghese et al. (1985) in India. Mohyuddin and Mahmood (1993) reported
sampling by direct visual examination: A. atkinsoni and I. niveosparsus were
found on upper portions of mango trees during different times of the year.
They moved to the lower parts of the stems and the leaves during summer.
Tandon et al. (1989) reported that distribution of I. niveosparsus was aggre-
gated and was best explained by Iwao’s patchiness regression. To assess
336 J.E. Peña et al.
BIOLOGICAL CONTROL. Several natural enemies have been described from west
and South-east Asia. Mohyuddin and Mahmood (1993) reported the egg par-
asitoids, Gonatocentrus sp., Miurfens sp. nr. mangiferae Viggiani and Hayat,
Centrodora sp. nr. scolypopae Valentine, Aprostocetus sp. and Quadrastichus sp.,
and the adult ectoparasitoid Epipyrops fuliginosa Tames in Pakistan. Fasih and
Srivastava (1990) reported that Aprostocetus sp., Gonatocerus sp. and Polynema
sp. parasitize eggs. Five species of predators, including Chrysopa lacciperda
(Kimmins), Mallada boninensis (Okomote), Bochartia sp. and two unindenti-
fied species (one each of Mantidae and Lygaeidae) prey on nymphs (Fasih
and Srivastava, 1990). In India, Sadana and Kumari (1991) studied the effi-
cacy of the lyssomanid spider, Lyssomanes sikkimensis on I. clypealis. Classical
biological control of mango hoppers has not been attempted. Whitwell (1993)
described four genera of parasitoids from Dominica, the most common being
Aprostocetus sp., followed by Platygaster sp., Synopeas sp. and Zatropis sp.
Peng and Christian (2005a, b) reported that the weaver ant, Oecophylla smarag-
dina (Hymenoptera: Formicidae) is an efficient biocontrol agent of I. nididulus
in northern Australia. The entomopathogens, Verticillium lecanii (Zimmer-
man) Viegas, Beauveria bassiana Balsamo (Vuillemin) and Isaria tax, infect
I. clypealis in India (Kumar et al., 1993; Srivastava and Tandon, 1986) while
the effectiveness of Metarhizium anisopliae var. anisopliae was tested under
laboratory conditions against A. atkinsoni (Vyas et al., 1993).
CHEMICAL CONTROL. Several pesticides have been tried for controlling mango
hoppers (Tandon and Lai, 1979; Yazdani and Mehto, 1980; Shah et al., 1983;
Shukla and Prassad, 1984; Islam and Elegio, 1997; Kudagamage et al., 2001).
Khanzada and Naqvi (1985) reported that six sprays of fenitrothion/year
were effective for controlling mango hoppers in Pakistan. Nachiappan and
Baskaran (1986) tested eight insecticides: phasalone, endosulfan, carbaryl,
penthoate, fenitrothion, monocrotophos, quinalphos and phosphamidom.
Endosulfan provided the best control when spraying was done 1 week after
flowering and then 14 days later. Mohyuddin and Mahmood (1993) reported
that monitor applied at 5 m on tree trunks and leaves in May provided control
of mango hoppers. Jhala et al. (1989) considered that sprays of carbaryl during
the off-season maintained the hopper population at low-density levels.
Godase et al. (2004) demonstrated that sprays of 0.05% monocrotophos at
the first panicle emergence and a second spray 15 days later are essential to
prevent yield loss. Kudagamage et al. (2001) found that imidacloprid (Admire
SL 200) controlled mango hoppers if applied just after flowering and again
10 days later.
Lepidoptera
The lepidopteran flower feeders are the second most important inflorescence
pests of mango. Geometrids, for example Chloropteryx glauciptera Hampson and
Pests 337
Thrips
The western flower thrips, Frankliniella occidentalis (Pergande) damages flow-
ers and fruit in Israel (Wysoki et al., 1993). The developmental time of F. occi-
dentalis from egg to egg at 25°C occurs between 14.8 and 16.65 days. The
duration of development of F. occidentalis from egg to adult is closely related
to environmental conditions, especially temperature. Frankliniella (possibly
cubensis) is present in mango flowers during the dry season in Costa Rica,
requiring several applications of systemic insecticides (Jirón, 1993). In Florida,
338 J.E. Peña et al.
midges, mites, scales, whiteflies, mealybugs, weevils, ants, locusts and cater-
pillars (Jeppson et al., 1975; Bhole et al., 1987; Jadhav and Dalvi, 1987; Tigvatt-
nanont, 1988). Formation of leaf galls in India is caused by the Eurytomidae,
Mangoma spinidorsum (Subba Rao, 1986), but there is little information on
their importance as foliage pests.
Thrips
The Mediterranean mango thrips, Scirtothrips mangiferae Priesner, is a severe
pest of mango in Israel, causing the young leaves to curl along the midrib,
distorting their shape and leading to premature drop (Wysoki et al., 1993).
The twigs of infested shoots are much shorter than those of uninfested ones.
The population of the thrips is low during winter, increases in early spring
and reaches its peak during summer (Wysoki et al., 1993). Yellow sticky traps can
be used for monitoring thrips densities. Ganz et al. (1990) established that an
average population of ten Mediterranean mango thrips per young shoot was the
threshold above which chemical control is required. Efficient control has been
achieved by spray application of fluvalinate or acephate (Ganz et al., 1990).
The red banded thrips, Selenothrips rubrocinctus (Giard), is an important
pest of cacao in the Caribbean islands and attacks mango and avocado in
Australia and Florida and Hawaii USA. The adults feed on the underside of
leaves, causing necrosis and subsequent leaf drop. According to Hill (1975),
S. rubrocinctus is only a pest in mango nurseries, and rarely damages mature
trees. Its biology was reviewed by Moznette (1922). Adult thrips are dark
bodied with a red band on the first abdominal segment. The immature stages
are light orange with abdominal segments one and two and the anal seg-
ments bright red. The population of this species peaks during the dry season
and declines during the rainy season. According to Yee (1987), the thrips are
controlled by malathion (25% v/w).
The weaver ant, O. smaragdina (Hymenoptera: Formicidae) is considered
an effective biological control of S. rubrocinctus in the Northern Territory of
Australia (Peng and Christian, 2004).
Midges
Mango leaves are attacked by different Cecidomyiidae species, especially in
Asia, but also in the Caribbean region. Two genera, Procontarinia Kieffer and
Cecconi and Erosomyia Felt, are particularly associated with mango and all
known species have been reared from mango (USDA, 1981; Schreiner, 1991;
Harris and Schreiner, 1992; Uechi et al., 2002; Gagne and Medina, 2004).
Prasad (1971) described the biology of the main species attacking mango in
India. A new species of gall midge, Procontarinia schreineri Harris, which
attacks mango foliage in Guam, lays eggs on young mango leaves and larvae
develop rapidly over c.5 days and induce blister galls. According to Harris
and Schreiner (1992), the main factors affecting populations of this midge are
rainfall and location. More galls are present during rainy periods, possibly
because RH improves larval and pupal survival. No differences were
observed in gall densities collected from lower and top portions of the tree.
Askari and Radjabi (2003) observed overlapping generations of Procontarinia
340 J.E. Peña et al.
matteiana in Iran related to the different leaf flushing patterns and found that
the optimum pest temperatures were 10–26°C. Differences on susceptibility
of mango cultivars to P. matteiana might indicate that susceptible cultivars
should not be grown in areas infested by this gall midge (Jhala et al., 1987;
Githure et al., 1998). Daneel et al. (2000) suggested that products to control P.
matteiana should be applied after harvest, coinciding with the first major
flush and a second spray 6 weeks later. Austin (1984) and Sankaran and Mjeni
(1989) have reported several platygastrid species parasitizing Procontarinia
spp. and their prospects for biological control of the pest.
Mites
The mango bud mite, Aceria mangiferae (Sayed) (Acari: Eriophyidae), attacks
buds and inflorescences (Keifer et al., 1982; Ochoa et al., 1994) (Plates 69–71).
According to Jeppson et al. (1975) this mite stunts and brooms twigs, causing
bud proliferation and appears to be responsible for necrosis of bud tissue
cells (Varma et al., 1974). In Hawaii, Tegonotus mangiferae (Acari: Eriophyidae)
feeds on the underside of leaves (Jeppson et al., 1975), while another species,
Metaculus mangiferae (Attiah) (Acari: Eriophyidae) causes russeting of termi-
nal leaves, buds and inflorescences. The latter is an important pest in Egypt,
India, Palestine and Angola (Jeppson et al., 1975). The puncture wounds of
several acarines (Acari: Tetranychidae) cause serious damage to leaves, which
may dry and fall. The main pest in Mauritius, India, Egypt, Israel and Peru is
Oligonychus mangiferus (Rahman and Sapra); in Israel, the spider mite Tet-
ranychus cinnabarinus (Boisduval), which lives on the underside of the leaves,
causes bronzing around the puncture wounds. The adult life span of O.
mangiferus is 10.11 days for females and 4.21 days for males (Rai et al., 1988).
The avocado red mites, Oligonychus yothersi McGregor and Oligonychus puni-
cae (Hirst), feed on the upper surface of leaves and cause considerable sti-
pling around the midrib at high population densities (Andrews and Poe,
1980). If the tetranychid mites are sufficiently abundant, infested leaves may
drop. There is little or no information on sampling techniques or for their
economic thresholds.
Aceria mangiferae occurs wherever mango is grown (Denmark, 1983;
Doreste, 1984). There has been controversy regarding a possible association
between this mite and floral and foliar galls, i.e. mango malformation (Sayed,
1946; Narasimhan, 1959; Summanwar and Raychoudhury, 1968; Denmark,
1983; Ochoa et al., 1994). However, A. mangiferae does not cause mango mal-
formation, but may be a carrier of Fusarium mangiferae, which is recognized
as the causal agent of mango malformation (Varma et al., 1974; Freeman et al.,
2004). Aceria mangiferae life history has been described by Abou-Awad (1981);
it is completed in 15 days at 25–27°C.
bud tissue cells, which is initiated externally at the edge of the bud and pro-
gresses toward the centre and internal areas of the bud (Peña et al., 2005).
CHEMICAL CONTROL. Osman (1979) reported that applications of four full cov-
erage sprays of dichlorvos were effective for controlling A. mangiferae in
Egypt. Rai et al. (1966) cautioned that chemical control should be directed to
apparently healthy and not malformed tissues. In Florida USA, agrimek plus
citrus oil, fenproximate and fenpropathrin resulted in the lowest mite densi-
ties 12 days after application. Agrimek plus citrus oil, and acequinocyl
resulted in the lowest mite densities 26 days after treatment (Peña et al., 2005).
In Brazil, Nascimento et al. (2002) recommended sulfur applications.
Scales
ARMORED SCALES. At least 26 species of diaspidids attack mangoes worldwide
(Chua and Wood, 1990). In India, Aspidiotus destructor Signoret causes serious
damage, while Parlatoria pergandii Comstock and Lepidosaphes gloverii (Pack-
ard) damage 3-year-old plants (cited in Chua and Wood, 1990). Radionaspis
indica (Marlatt) (= Leucaspis indica) encourages growth of black mould, which
covers young branches (Dekle, 1976). Several diaspidids, e.g. Aulacaspis
mangiferae (tubercularis) Newstead, attack shoots and leaves; the oleander
scale (Plate 72) in Florida USA (Miller and Davidson, 2005) and the mango
scale in Ghana (van Halteren, 1970) cause similar damage. They are damag-
ing not only because they feed on sap, but also because of the toxicity of their
saliva (Singh, 1991). Scales inhabit both leaf surfaces and also are on fruit.
Van Halteren (1970) concluded that A. mangiferae development is completed
in 35–40 days for females and 23–28 days for males.
SOFT SCALES. Other species of Coccidae, Coccus viridis (Green), Coccus longu-
lus, Ceroplastes actiniformis, Philephedra tuberculosa Nakahara and Gill and the
mango shield scales Milviscutulus mangiferae (Green) and Viusonia stellifera
(Westwood) in Asia, Africa, Australia, Israel and the Americas cause similar
damage. These coccids are generally polyphagous, attacking different genera
and species. They are mobile and injure mango because of the production of
honeydew and the subsequent accumulation of sooty mould on the honey-
dew (Escalante, 1974; Silva and Cavalcante, 1977). Most of these scales can be
suppressed at sub-economic levels, either by application of selective pesti-
cides (i.e. oils) or by biological control agents.
CONTROL. Various control methods, including banding tree trunks with vari-
ous materials to prevent D. stebbingi nymphs from climbing (Lakra et al.,
1980; Srivastava, 1981) and dusting chlorinated hydrocarbons on the soil
(Srivastava, 1981), have been tried with little success. In Pakistan the mango
mealybug was controlled by hoeing or ploughing the soil and conservation
of the predator, Sumnius renardi Weise, by wrapping burlap around the trunks
of the trees (Mohyuddin and Mahmood, 1993).
Termites
Mango orchards are becoming more common in dry and semi-arid areas
with vast termite populations. Mango growing in infested areas often results
in plant growth suppression as a result of reduced root establishment, inva-
sion and pruning of roots (Rogers et al., 1999). For example, six termite spe-
cies (Odontotermes indicus Thakur, O. lokanandi Chatterjee and Thakur, O.
obesus (Rambur), O. giriensis (Roonwal and Chhotani), O. bhagwatii Chatter-
jee and Thakur and Microtermes obesi Holm) were recorded from mango
orchards in India (Srivastava and Singh, 2004). More species of termites were
observed in winter than during the summer and rainy season. Veeresh et al.
(1989) observed that O. wallonensis, O. horni and O. obesus constructed earthen
sheeting on the stem of small mango trees. Singh (1960), cited by Srivastava
(1998), reports Eutermes (Nasutitermes) costali, Calutermes (Cryptotermes) bre-
bis, Heterotermes tenuis, Coptotermes gestroi, Neotermes (Kelatermes) bosei (Gard-
neri) Synder, Microcereoutermes peroffinis, Calotermes (Neotermes) greeni and
Coptotermes curvignathus also affecting mango in India. According to Srivas-
tava (1998), the most important termite species affecting mango are O. obesus
and O. wallonensis; O. wallonensis nests in the root zone in Uttar Pradesh,
India. The workers feed on roots, stems and branches.
Colonies of the subterranean termite, Coptotermes curvignathus Holmgren,
were monitored in Malaysia using bait matrices containing 0.5% hexaflu-
muron (Said Sajap et al., 2000). In Florida, urban dwellings infested with the
Formosan termite, C. formosanus Shiraki, were treated with baits containing
0.5% weight/weight noviflumuron (Cabrera and Thoms, 2006). These baits
might be useful when mango orchards are planned for areas infested with
termites. In India, termite infestations are controlled with a combination of
monthly irrigation, hoeing and application of neem cake (Singh and Singh,
2003).
10.3 Discussion
In general, most mango pests also occur on other fruit crop species. Fruit
flies, scales, mites, thrips, lepidopteran flower feeders, mirids, weevils and
beetles are mostly generalists, and some of their management schemes need
to be implemented with this in mind. In the case of fruit flies, Aluja (1996)
suggests surveying vegetation adjacent to infested mango orchards as
346 J.E. Peña et al.
populations are sustained and multiplied in these locations and from them
adult flies move into commercial orchards to attack ripening fruit (Aluja et
al., 1996). Management of key pests (i.e. fruit flies, seed weevils, etc.) must be
mandatory, in order to have an effect on a large region. The use of some mea-
sures (i.e. quarantine, etc.) must involve neighbouring producing countries
in order to have a positive effect on sanitation. The most progressive exam-
ples in management of mango pests are in Australia, Mexico and Israel, while
other producing countries, such as South Africa, Brazil and Ecuador, are tak-
ing serious steps to reconcile the opposing forces of globalization of markets
and sustainibility. For other areas where maximizing yields and blemish-free
fruit is not a priority, the emphasis should be biological control. Management
tactics that can be improved include the following:
Acknowledgements
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11 Crop Production: Propagation
11.1 Introduction
Sexual Asexual
Monoembryonic Polyembryonic
Grafting Rooting Micropropagation
seed seed
Somatic
Approach Cleft or wedge Ground
embryogenesis
Veneer
Rind or crown
Budding
Monoembryonic seed
Polyembryonic seed
Fig. 11.2. Grafting methods of mango propagation: (a) approach grafting; (b) tongue
grafting; (c) saddle grafting; (d) root grafting.
Crop Production: Propagation 371
Fig. 11.3. Grafting methods of mango propagation: (a) cleft grafting; (b) soft wood
grafting; (c) epicotyl grafting; (d) splice or whip grafting.
372 S. Ram and R.E. Litz
Fig. 11.4. Grafting methods of mango propagation: (a) side grafting; (b) notch grafting
or inlaying; (c) veneer grafting; (d) rind or crown grafting.
nucellar seedlings. Eiadthong et al. (1999), using single sequence repeats (SSRs)
could distinguish among mango cultivars, and separate them according to
their geographic origin; however, monoembryonic and polyembryonic geno-
types could not be differentiated. Amplified fragment length polymorphism
(AFLP) markers have been used for cultivar identification (Kashkush et al.,
1
Plate 1. Mango flowers: polygamous (left) and monoecious (right) flowers. Monoecious flowers are
functionally staminate.
Plate 2. Longitudinal section through the micropylar end of the ovule of a polyembryonic mango cultivar.
Arrows indicate the presence of globular stage nucellar embryos.
Plate 3. A mango tree in a relief on the Buddhist temple at Borobodur, Indonesia. The temple is thought to
have been built between the end of the 7th and beginning of the 8th century AD.
4
6
7
Plate 4. ʻAlphonsoʼ.
Plate 5. ʻArumanisʼ. Plate 7. ʻCalypsoʼ.
Plate 6. ʻAtaulfoʼ. Plate 8. ʻBombay Greenʼ.
9 10
11
12
13
14
16
17
18
20
21 22
23
26
27 28
31 32
33
35
36
37
39
40
43 44
45
46
Plate 41. A flooded mango orchard in south Florida where, following the seasonal rains, roots may remain
submerged for several days following periods of high rainfall.
Plate 42. Hypertrophied lenticels on the trunk of a mango tree in which the roots and basal part of the trunk
were submerged in water. Development of these structures helps confer flood tolerance to trees, probably
providing sites for O2 diffusion and transport to the roots and/or excretory sites for waste products of anaerobic
respiration in the flooded roots.
Plate 43. Rust-coloured thallus of Cephaleuros virescens, cause of algal leaf spot (Photograph courtesy of
R.C. Ploetz).
Plate 44. Sooty blotch thalli on the surface of a light-coloured mango fruit. These fungi grow on mango leaves,
panicles and fruit. They are often inconspicuous against dark backgrounds, but can be significant postharvest
blemishes on fruit (Photograph courtesy of R.C. Ploetz).
Plate 45. Stem bleeding on trunk and scaffold limbs of a ʻTommy Atkinsʼ mango tree in Florida. The tree was
exposed to temperatures of -4ºC the previous winter (Photograph courtesy of R.C. Ploetz).
Plate 46. Powdery mildew on a developing panicle in Israel. The white, powdery symptom is due to masses of
conidia of the causal fungus, Oidium mangiferae (Photograph courtesy of S. Freeman).
47a
47b
Plate 47. Sudden wilt in Oman and Pakistan is caused by a newly described ascomycete species,
Ceratocystis manginecans. (a) Vascular streaking and galleries of the scolytid beetle vector, Hypocryphalus
mangiferae, are typically found in affected trees, as is (b) unilateral death of portions of the canopy;
ultimately trees are killed (Photographs courtesy of R.C. Ploetz).
48 49
50 51
52 53
Plate 48. Internal fruit breakdown: jelly seed (Photograph courtesy of A. Cracknell Torres).
Plate 49. Internal fruit breakdown: soft nose (Photograph courtesy of A. Cracknell Torres).
Plate 50. Internal fruit breakdown: stem end cavity (Photograph courtesy of A. Cracknell Torres).
Plate 51. Internal fruit breakdown: spongy tissue (Photograph courtesy of A. Cracknell Torres).
Plate 52. Internal fruit breakdown: necrotic tissue (Photograph courtesy of A. Cracknell Torres).
Plate 53. Black tip disorder.
54
55 56
59
60
Plate 57. Damage to mango fruit caused by the mango seed weevil Sternochetus mangifereae.
Plate 58. Damage to mango fruit pulp caused by the mango seed weevil S. mangifereae.
Plate 59. Adult mango seed weevils S. mangifereae.
Plate 60. Damage to foliage caused by mango red banded thrips Selenothrips rubrocinctus.
62
61
64
63
65 66
69
70
71
72
74
Plate 73. Effect of nitrogen (N) on fruit colour. (a) Excessive N suppresses mango fruit colour in the field.
(b) Ripe ʻKeittʼ fruit grown on (front) low N (0 applied N) and (rear) grown on high N (417 g/tree).
Plate 74. Effect of N on postharvest disease. Ripe ʻKeittʼ fruit grown on (left) high N (417 g/tree) and (right)
low N (0 g/tree).
75
76
77
79
Plate 78. Boron (B) deficiency symptoms showing (top left) stunted growth and sickle-shaped leaves;
(top right) shot holes in leaves; (bottom left) bent tip of flower panicle; and (bottom right) gummosis from
cracking trunk.
Plate 79. B toxicity symptoms: black staining leading to necrotic leaf margins
80a
80b
81a
81b
81c
Plate 80. Internal (a) and external (b) colour development of ʻTommy Atkinsʼ mango fruit during ripening.
Plate 81. (a) Lenticel damage on ʻB74ʼ mango. Plates 81(b) and (c) are higher magnifications of (a).
82a
82b
82c
Plate 82. Harvest aids used in Australia. Desapping using racks in the field based on placing the fruit onto a tarpaulin
(a and b) or basket constantly sprayed with detergent and the fruit remaining wet with detergent for c.90 s.
(c) A cherry picker is used to harvest high fruit.
82d
82e
83
Plate 82. (d) Desappingusing racks in he field and (e) desapping conveyor used for desapping in the pack-
house.
Plate 83. Processed mango in the marketplace, including yoghurt, nectar, reconstituted drink (from powder),
fruit leather, dried mango slices and preserved fruit.
84
85a 85b
88
90
89
Fig. 11.5. Budding methods of mango propagation: (a) shield budding; (b) patch
budding; (c) modified patch budding; (d) forkert budding.
374 S. Ram and R.E. Litz
(b) H-budding
Fig. 11.6. Budding methods of mango propagation: (a) modified forkert budding;
(b) H-budding; (c) chip budding.
Fig. 11.7. Budding methods of mango propagation: (a) modified chip budding;
(b) flap budding; (c) window budding.
Fig. 11.8. Top working (a) and double working (b) of mango.
Preparation of rootstock
stress tolerance, and behaviour of the scion cultivar on clonal rootstock is highly
predictable. Monoembryonic seedlings are generally used as rootstocks in
India. Polyembryonic ‘Turpentine’ and ‘Kensington Pride’ seedlings are used
as rootstocks in Florida and Australia, respectively, whereas polyembryonic
‘Sabre’, ‘13-1’ and ‘4-9’ seedlings are used for rootstocks in Israel. Throughout
South-east Asia, polyembryonic seedlings are used for rootstocks, e.g. poly-
embryonic ‘Saing’ and ‘Thalapt’ in Myanmar (Grant and William, 1949). In
Senegal, polyembryonic ‘Amelioree’, ‘De Cameron’ and ‘Madame Francis’ are
used as rootstocks (Furon, 1966). In Brazil, seedlings of polyembryonic
‘Carabao’, ‘Mangua d’Agua’ and ‘Pico’ are used for rootstock; they are consid-
ered to have resistance to Ceratocystis fimbriata (Neto et al., 2002); the IAC series
(IAC100-104) are used to control ‘seca de mangueira’ (Neto et al., 2002).
Freshly extracted mango seeds from ripe fruits germinate with higher
frequency (76–91%) than those from overripe, firm or green fruits (Shant and
Saproo, 1974). Seeds with large cotyledons from seedling trees germinate
earlier, store better and seedling growth is more vigorous than seeds with
small cotyledons (Naik, 1949; Simao, 1960). Naik (1941) noted differences in
germination and vigour in monoembryonic seedling progenies of different
parentages. Seedlings of ‘Chausa’ reportedly produced better height and
girth than ‘Langra’ and seedling vigour was closely related to the weight of
the seed stone (Giri and Chaudhery, 1966). Mango seeds are recalcitrant, and
lose viability (Ledin and Ruehle, 1954; Ruehle and Ledin, 1955; Singh, 1960)
within 30 days. Therefore, seed should be collected and sown within 1 week
after collection. About 80% of seed germination occurs if they are sown
within 1 month of extraction (Stephens, 1960; Sturrock, 1968; Chacko and
Singh, 1971). Parisot (1988) recorded that seeds that are free of pulp could be
stored for up to 84 days at 15°C on sterile cotton with deionized water. Husk-
ing or decortication of the hard endocarp of the seed improves germination
(Paul and Guneratnam, 1938; Simao, 1960; Chauran et al., 1979; Abdel-Galil,
1992). At the time of sowing, treatment with fungicides also improves germi-
nation, particularly of dehusked seeds (Chauran et al., 1979). Seeds should be
sown without injuring the cotyledons and preferably early in the season.
Seedbed preparation
The standard practice for seed germination varies. In India, seedbeds are
used, whereas in most other countries, seeds are planted individually in pots
or plastic bags. Nurseries are usually under partial shade to prevent exces-
sive evaporation from the plant and soil and to protect seedlings from inclem-
ent weather; shade is particularly useful in areas with dry, hot climates.
Seedbeds can also be used in the open provided the soil is adequately moist.
Highly sandy or loamy soils are generally unsuitable for mango seed germi-
nation. Soil should be well drained with plenty of organic matter, and seeds
should be sown c.15 cm apart in a special seedbed in which 25 cm top soil of
the bed is excavated and the bed bottom is hardened with concrete or lined
with an iron or polyethylene sheet to prevent elongation of the taproot and
to encourage fibrous root development (Stephens, 1948). Germination can be
expedited if seeds are soaked in 20–100 mg/l gibberellic acid (GA3) for 24–48 h
378 S. Ram and R.E. Litz
or sprayed with 50–300 mg/l GA3; however, these seedlings are usually
unsatisfactory for grafting. Seeds should be planted with convex edge up at
a depth of 5–7 cm (Singh, 1960; Bakhshi, 1963) or with a small portion exposed
above the ground level (Pope, 1929; Ruehle and Ledin, 1955; Bakhshi, 1963;
Anonymous, 1975). Soil moisture should be maintained. Seeds germinate
2–3 weeks after planting (Ruehle and Ledin, 1955; Ahmed and Rashid, 1961;
Singh and Jawanda, 1962). Seedlings that are 1-month-old with coppery
leaves and with their stones attached survive transplanting better than older
plants without much damage to their root system (Ruehle and Ledin, 1955).
The taproot should be slightly pruned at the time of planting.
Nursery
Seedlings should be transplanted without disturbing the roots to nursery
beds or pots at least 1 month before grafting. Either the crown of the seed-
lings should be pruned or the distal half of the leaves should be removed to
reduce transpiration. Seedbed-grown plants generally have better germina-
tion rates and lower production costs than those grown directly in nursery
beds. Seedlings should be actively growing at the time of grafting. Success in
grafting depends on several factors, such as age and vigour of rootstock and
scion, diseases, insect pests, environment and the skill of the grafter.
Attached methods
Approach grafting
A scion shoot, while still attached to the parent plant, can be grafted onto the
seedling rootstock by making a long cut (5–7 cm) of similar length and depth
on one side of both rootstock and scion through the cambium and slightly
into the wood (Fig. 11.2a). The cut surfaces of rootstock and scion are pressed
firmly together and secured with waxed string, raffia or grafting tape. After
union, the scion is severed below the graft union and the rootstock above the
union. Several years ago, approach grafting was standard practice for propa-
gating mango in many countries, but has been largely replaced by detached
grafting methods, except for parts of India. For approach grafting, seedlings
45–60 cm long with a diameter of 1.25 cm are utilized (Asadullah and Khan,
1960; Singh, 1960). They are planted in pots or plastic bags containing soil
mixture, and are kept beneath the mother tree. Alternatively, the ball of earth
around the root system of the seedling grown for rootstock is tied in ‘jutties’
made of dried grass, paddy straw or plastic bags and planted under the
mother tree or raised to the shoot height of the mother tree on a raised plat-
form or directly suspended from branches of the mother tree. The seedling
rootstock can be a few months- to 2-years-old (Naik, 1949; Garg, 1954; Ahmed,
1960; Singh, 1960). Juvenile seedlings whose leaves are copper coloured and
Crop Production: Propagation 379
with the seed still attached to the seedling are often used. The root system is
covered with moist sphagnum moss or similar material and wrapped in
polyethylene to prevent drying. Production costs are less with young seed-
lings than with older material.
In the subtropics, approach grafting in spring can achieve 88–100% suc-
cess with some cultivars (Asadullah and Khan, 1960; Majhail and Singh,
1962a, b; Talukdar and Ahmed, 1965). Majhail and Singh (1962b) observed
that seedling size and age do not affect grafting success in spring, but that
larger seedlings yielded better success from July to September. Approach
grafting should be avoided during winter months when plants are dormant.
In the tropics, mangoes can be approach grafted year-round (Ahmed, 1960;
Singh, 1960). Approach grafting should be done when the trees are in active
growth (Singh, 1960; Rao, 1967). In low rainfall areas, approach grafting
should be done at the time of the onset of rains, while in regions of heavy
rainfall, it should be done soon after the rainy season is over, provided tem-
peratures are not <15°C, the critical temperature for the growth of mango
trees (Whiley, 1993). Healthy trees should be used for scion wood. The suc-
cess of approach grafting is cultivar dependent, for example ‘Langra’ scions
establishes better than either ‘Dashehari’ or ‘Chausa’ (Asadullah and Khan,
1960). Better success has also been reported with seedlings of 1.3–1.6 cm
diameter than with smaller shoots (Giri, 1966). Variations of approach graft-
ing include multistock, inflorescence, top working and adjuvant grafting. At
one time, south Indian nurserymen would approach graft a single large scion
with two to five rootstocks (Patwardhan and Deshmukh, 1931; Singh, 1960).
Grafts with large and old scions having several branches become top heavy, and
fail to thrive. New growth from inflorescences that are approach grafted is veg-
etative. Approach grafting has also been used to top work old seedling trees with
cultivars (Singh, 1960). Adjuvant grafting refers to approach grafting of diseased
or damaged trees in order to rejuvenate them on new rootstocks.
Tongue grafting
Tongue grafting is practised with thicker rootstock than scion. The rootstock
is first cut as in approach grafting; a second cut is made into the wood of the
stem approximately halfway down the first cut in a sloping direction point-
ing tongue upwards. A similar cut is made in the scion shoot. Both cuts are
made in such a manner that one fits into the other tightly (Fig. 11.2b). Root-
stock and scion are tied together with grafting tape and a graft union is com-
plete in c.1–2 months. The scion is then severed from the mother tree and the
rootstock is decapitated above the graft union. Tongue grafting is slightly
more complicated than simple approach grafting; however, the success rate
is better because three surfaces of the cambium layer are involved in the graft
union (Burns and Prayag, 1921).
Saddle grafting
With saddle grafting, the rootstock is decapitated c.25 cm above ground level,
and two upward sloping cuts are made on the sides of the rootstock to form
a 5–7 cm long wedge on its cut end. On the stem of the scion shoot, a tongue
380 S. Ram and R.E. Litz
is made and the outer surface of the tongue is not pared. The wedge of the
rootstock is then fitted into the groove of the tongue (Fig. 11.2c). Subsequently,
the joint is secured with grafting tape and the scion shoot is separated from the
mother tree after the union is complete (Singh, 1960). Saddle grafting is easier
to perform than tongue grafting but is more difficult than approach grafting.
Root grafting
Root grafting is similar to bench grafting. A seedling plant (c.l year old) is
potted in a U-shaped pot with a notched edge (7–8 cm deep and 5 cm wide)
(Singh, 1960). The 7–10 cm taproot is projected through the notch and the pot
itself contains the root. The seedling above the collar is staked, and placed in
the shade for 1–1.5 months for establishment. Seedlings are approach grafted
with the scion (Fig. 11.2d), and the graft is separated from the mother tree
after the union is complete. Grafts are maintained in the shade and watered
regularly. Naik (1941, 1949) reported 100% success with root grafting.
of various ages can be used; however, success decreases with age, i.e. 80%
with 1-year-old rootstocks and less with 6-month-old seedlings (Gaur, 1984;
Reddy and Melanta, 1988; Kulwal and Tayde, 1989; Panickar and Desai,
1989; Gandhoke, 1993). This technique is easy and is effective in dry, hot
weather and in areas with low precipitation.
Side grafting
Side grafting, also known as the ‘Nakamura method’ (Fig. 11.4a), was for-
merly popular (Burns and Prayag, 1921; Pope, 1929; Pope and Storey, 1933;
Walters, 1932; Tanaka, 1939; Lynch and Mustard, 1950; Thrower, 1954; Ruehle
and Ledin, 1955; Lynch and Nelson, 1956; Ahmed, 1960; Singh, 1960; Serpa,
1964; Rao, 1967). This method is supposed to be highly effective in mild
382 S. Ram and R.E. Litz
weather in the absence of strong winds, intense heat and heavy rain (Rao,
1967), and success has also varied (50–100%) with different cultivars (Veera-
raghavan, 1945). In India, side grafting is generally used in humid, coastal
areas with 1.0–1.5 cm diameter rootstock. Scions should be c.10 cm long and
defoliated c.1 week prior to grafting. The rootstock age has varied from 4 to
36 months with 72–90% success (Mulat, 1959; Kashyap et al., 1972; Faruque
and Fakir, 1973; Kanwar and Bhajwa, 1974; Ben-ya’acov, 1976). The scion is
inserted into the wedge and tied with grafting tape. The union is complete
after 2–3 months. The top of the rootstock can be removed above the graft
union after new scion growth occurs.
Veneer grafting
This grafting technique was first described by Lynch (1941), and has been
widely adopted (Cooper and Furr, 1945; Ruehle and Ledin, 1955; Mukherjee
and Majumder, 1961, 1962, 1964; Wolfe, 1963; Ahmed, 1964; Serpa, 1964; Jagirdar
et al., 1968; Bhambota et al., 1971; Prasad et al., 1973; Ramirez Diaz, 1973; Gun-
jate and Limaye, 1978; Ram and Bist, 1982; Pinto et al., 2004). Veneer grafting
is usually more effective than other methods of grafting (Bhambota et al.,
1971; Prasad et al., 1973), with approximately 90% success (Ram and Bist,
1982). For veneer grafting, 3- to 6-month-old 1.0–1.5 cm diameter scion shoots
with lush green leaves should be defoliated 3–15 days prior to grafting
(depending upon the season), leaving the petioles attached (Mukherjee and
Majumder, 1961; Singh and Srivastava, 1979a, b; Ram and Bist, 1982; Rajan
and Sinha, 1987; Bajpai et al., 1989). Prior defoliation may not be required under
humid conditions (Gunjate et al., 1976) or when extremes of temperature and
RH do not occur. The scion should be 10–15 cm long, although smaller scions
(5–10 cm) have also been used (Ram and Bist, 1982). Older shoots can also be
used (Mukherjee and Majumder, 1961; Jagirdar et al., 1968; Ram and Bist,
1982). Scions stored for 8 days at ambient temperature (25–35°C) in moist
sphagnum moss covered with polyethylene can still be grafted with a 50%
success rate during March through to July (Ram and Bist, 1982). Cultivar
differences may be important.
The use of seedling rootstock at different ages (3 months to 2 years) has
resulted in 40–100% success, depending upon the season, scion maturity,
predefoliation period, storage condition of scion, etc. (Zill, 1951; Subra, 1954;
Mukherjee and Majumder, 1961, 1962, 1964; Ahmed, 1964; Serpa, 1964;
Jagirdar and Bhatti, 1968; Bhambota et al., 1971; Prasad et al., 1973; Ramirez Diaz,
1973; Gunjate and Limaye, 1978; Ram and Bist, 1982). Dipping or smearing
Crop Production: Propagation 383
growth regulator onto the cut surfaces of the scion and rootstock has been
ineffective (Kulkarni et al., 1989). The rootstock is prepared for veneer grafting
by making a slanting cut (5 cm long); an oblique cut is then made at the base of
the first cut, so that a piece of wood along with bark is removed (Fig. 11.4c).
The base of the scion wood is then fitted into the rootstock so that the cut
surfaces with the cambium layers of scion and rootstock are in contact with
each other. The rootstock and scion are secured together with grafting tape.
The union takes place after 1.5–2.0 months. After scion growth begins, the
rootstock is removed above the graft union. Some workers have recom-
mended first ringing and then removing the rootstock shoot 1–2 weeks later,
while others have partially cut through the rootstock shoot before severing
the shoot completely a few days later. After new leaves of the scion turn
green, the rootstock may be removed completely together with grafting tape
(Mukherjee and Majumder, 1961; Ram and Bist, 1982).
Budding
Budding involves the grafting of a single bud, with or without wood attached,
with the rootstock. Budding methods are referred to by the shape of the bud,
flap of the rootstock covering the bud before insertion, etc. The various meth-
ods for budding are illustrated in Figs 11.5–11.7. The inserted bud eventually
develops as the crown. Rootstock above the bud is severed after bud estab-
lishment. Budding provides a stronger union and the graft union occurs ear-
lier than with other grafting methods. Both rootstock and scion should be
actively growing, and excision of the bud is easy. Stimulation of the bud prior
to its excision by predefoliation, application of growth regulators and gir-
dling can improve the efficiency of the procedure. A bud is normally selected
from a 3- to 4-month-old scion shoot, and is budded onto rootstock stems of
1.0–1.25 cm diameter c.25–30 cm above ground level where the rootstock is
greyish green, except in cases of very juvenile rootstock. Generally 1- and
2-year-old rootstocks are better for grafting than younger ones. The bud is
384 S. Ram and R.E. Litz
completely covered by grafting tape for a few weeks. Bud growth can be
stimulated by removing grafting tape c.2 weeks after grafting.
Patch budding
Patch budding (Figs 11.5b and c) (Verma, 1942; Ullah and Ali, 1955; Singh,
1960; Moreuil, 1963; Teaotia, 1963; Jauhari and Singh, 1970; Teaotia and
Maurya, 1970; Prasad and Singh, 1972; Prasad et al., 1973) is used as an alterna-
tive to approach grafting (Garner and Chaudhri, 1976). Under subtropical
north Indian conditions, March and May through to July is the optimum period
for patch budding. In tropical Malaysia, patch budding is successful from
December through to February. The scion is defoliated 2 weeks prior to budding.
Rootstocks are severed above the budding point 1–2 weeks after budding.
Crop Production: Propagation 385
Forkert budding
Over 90% grafting success has been reported with forkert budding (Fig.
11.5d) in tropical South-east Asia and Sri Lanka (Paul and Guneratnam,
1937a, b). In Maharashtra, India during July and August, success with this
technique using 1-year-old rootstock was 60–70% (Gandhi, 1942), and 100%
with ‘Langra’ and ‘Dashehari’ (Singh and Srivastava, 1962; Teaotia, 1963). A
modified forkert budding method (Fig. 11.6a) can be more effective than
shield budding (Garner and Chaudhri, 1976). H-budding is another modifi-
cation of the forkert method (Singh, 1960), and derives its name from an
‘H’-shaped cut made in the rootstock (Fig. 11.6b).
Chip budding
For chip budding, 2- to 3-month-old seedlings are used (Fig. 11.6c). The bud
is excised with a piece of wood attached to it. Graft union can occur 3–4
weeks after budding because rootstock tissues are partially undifferentiated
and possess a broadly exposed cambium. The method is very efficient, and is
widely used (Lynch and Nelson, 1949, 1956; Soule, 1954; Subra, 1954; Ruehle
and Ledin, 1955; Ahmed, 1960; Bhan et al., 1969; Rajput and Haribabu, 1971),
often with modifications (Lynch and Mustard, 1950; Singh, 1960) (Fig.
11.7a).
Flap budding
The excised bud shape is boat-shaped instead of rectangular (Fig. 11.7b). This
procedure has been used in South-east Asia (Naik, 1949; Garner and Cha-
udhri, 1976).
Window budding
This technique is similar to flap budding except that a window is created in
the flap for the bud so that flap does not press the bud while being tied (Fig.
11.7c). This method has been used in Queensland, Australia. The bud union
occurs in c.4 weeks (Stephens, 1948).
Effect of rootstock
the onset of spring. The new shoots are budded or grafted in the following
summer or autumn, usually by shield budding or veneer grafting. Singh
(1990) suggested that half of a tree should be top worked in the first year and
the other half in the second year, although top working of a complete tree in
a single year has been highly successful (Lynch, 1941; Ruehle and Ledin,
1955; Ahmed, 1960; Singh, 1960). Cutting back of limbs in October or early
November and veneer grafting in March–April has been recommended in
Florida (Carmichael, 1957/58; Miner, 1962); in Uttar Pradesh (India), top
working on 8-year-old mango trees was more successful during June (Singh,
1960). After top working, trees come into bearing within 2 years (Furon and
Plaud, 1972).
Grafting on an already grafted or budded tree is referred to as double
working (Singh, 1960), and double-worked trees therefore consist of three
genetically different parts, i.e. the rootstock, interstock and crown (Fig. 11.8b).
In Florida, old plantings of ‘Haden’ have been double worked with ‘Tommy
Atkins’ (Mitchell, 1971). Double working can also be used to repair trees
affected by disease or cold injury and to develop a strong framework. ‘Kala-
pady’ has reportedly been used as an interstock in order to dwarf ‘Langra’
(Sen, 1939). In one trial, 12 interstocks were grafted on two rootstocks for
‘Dashehari’; rootstock-interstock combinations significantly affected tree
height and vigour. Fruit yield was also influenced by rootstock in all the com-
binations. ‘Kurukkan’ interstock on ‘ST-9’ rootstock yielded more fruit in
comparison with ‘Ambalvi’ on the same rootstock. The maximum yield was
obtained with ‘Nakkare’/‘Chandrakaran’ and ‘Ambalvi’/‘Chandrakaran’;
whereas the lowest yield was obtained with ‘Ambalvi’/‘ST-9’. Iyer et al.
(1990) reported varying success (20–73%) with double-worked ‘Langra’.
Rooting
Mango air layers have been induced to root either while they are still attached
to the parent tree or by cutting shoots into pieces containing one to several
buds. Several rooting methods have been successful with mango.
Layering
Layering is used chiefly to propagate monoembryonic mango cultivars on
their own roots. Rooting methods involve the initial training of the main
stem. Shoots close to ground level are ringed and covered with soil, while
other shoots are covered with soil mixture or wet sphagnum moss at the
ringed portion and wrapped with polyethylene to maintain moisture (Singh,
1960; Majumder and Mukherjee, 1961; Mukherjee and Majumder, 1963).
Auxins such as E-naphthalene acetic acid (NAA) and indole butyric acid
(IBA) are generally essential in order to induce rooting even after ringing.
Ground layering
Vigorous, 1- or 2-year-old shoots are selected near the base of the parent tree.
The greyish-green bark portion of the shoot (3–5 cm length) is ringed without
388 S. Ram and R.E. Litz
injuring the wood. On the proximal end of the shoot just above the ring,
growth regulators are applied and the ringed portion is buried in the soil.
Depending upon the growth regulators and their concentration, ringed
shoots (or layers) generate roots within 2–3 months. The rooted shoot can
then be detached from the mother tree and planted.
Pot layering
With pot layering, the ringed portion of the shoot is maintained in a soil mix-
ture in special pots at a convenient height within the tree canopy. This can be
done only during periods of high RH, but the success is low (c.15–20%)
(Singh, 1960).
Air layering
Although air layering or marcottage is one of the oldest methods for propagat-
ing mango, good success has been elusive (Grove, 1947; Garg, 1954; Rangacha-
rlu and Rao, 1956; Rao and Rao, 1956; Choudhury, 1959; Jauhari and Nigam,
1962; Rao et al., 1963; Mukherjee and Majumder, 1963; Srivastava, 1963a, b;
Mukherjee and Bid, 1965; Sen and Bose, 1966; Bid and Mukherjee, 1969; Basu
et al., 1972; Prasad and Singh, 1972; Núñez-Elisea et al., 1992). Juvenility, high
RH and moderate temperature are important factors for air layering mango
(Singh, 1954; Rao et al., 1963; Rao, 1967; Chhonkar and Singh, 1972; Singh
et al., 1979; Ram and Sirohi, 1982a). Cultivar effects on air layering are also
pronounced; and vigorous cultivars root better than dwarf cultivars (Garg,
1954; Rao and Rao, 1956; Rao, 1967; Basu et al., 1972; Ram and Sirohi,
1982b).
Layering success can be improved with auxins, such as 2–3% indole ace-
tic acid (IAA) and 0.5–2.0% NAA (Thakurta and Dutt, 1941; Singh and Teotia,
Crop Production: Propagation 389
Cuttings
The potential of mango cuttings to form roots depends on maturity of the
cutting, rooting medium, temperature, RH, etc. Girdling or etiolation of
stems for a few weeks and use of growth regulators have improved rooting
frequency (Thakurta and Dutt, 1941; Gardner and Piper, 1943; Van Overbeek
and Gregory, 1945; Said and Shoushan, 1945; Khalifa et al., 1964; Dijkman,
1950; Ledin and Ruehle, 1954; Gowda, 1962; Ahmed, 1964; Sen and Bose,
1964; Mukherjee et al., 1966, 1967; Maurice, 1969; Sen et al., 1969; Ali and
Nazir, 1970; Prasad and Pathak, 1970; Bid and Mukherjee, 1972; Bid et al.,
1972). Rooting is improved with cuttings from the lower bole of the tree
rather than from the upper bole. Mukherjee et al. (1966) reported that cut-
tings from 4-year-old trees root more readily than those from 10-year-old
trees, alhough Sen et al. (1968) had better success with older shoots. Hard-
wood cuttings reportedly root better (Hussain, 1953; Benincasi, 1970) than
semi-hardwood cuttings (Singh and Teaotia, 1951). Mango cuttings should
be c.15 cm long and 1.2–1.8 cm girth with between three and five buds (Garner
and Chaudhri, 1976). Retention of one to two leaves at the apex of the cut-
tings has helped rooting, and is attributed to the presence of rooting cofactors
in the leaves.
In India and Pakistan, cuttings are generally made during the spring
months (Hussain, 1953). Cuttings can be stored for a short time before apply-
ing the rooting treatment. Khalifa et al. (1964) rinsed cuttings for 24 h with
running tap water, which increased their shelf life. Longevity of cuttings can
390 S. Ram and R.E. Litz
cuttings survived in the nursery. Sadhu (1979) and Sadhu and Bose (1980a, b)
found that 10,000 ppm cycocel pretreatments of 10-year-old ‘Langra’ resulted
in 41% rooting with 2.2 roots/cutting.
Micropropagation
11.5 Conclusions
Standard methods are widely utilized for propagating mango scion cultivars
with increasing efficacy. In many regions, including India and Mexico, scion
cultivars are still being propagated on heterogeneous monoembryonic seed-
ling rootstocks (see Crane et al., Chapter 13, this volume), despite the demon-
strated advantages of clonal nucellar rootstocks. The potential of clonally
propagated monoembryonic mango rootstock has not been properly investi-
gated. The genetic heterogeneity of monoembryonic mangoes has been
explored neither for stress tolerance nor for their effects on scion growth and
development. Somatic embryogenesis could play an important role in such
investigations as an alternative propagation method (see Litz et al., Chapter
18, this volume). Other Mangifera species also have interesting attributes, and
should be screened for graft compatibility with mango (see Bompard, Chap-
ter 2, this volume). The species that could be tested as rootstock for mango
might extend mango cultivation to areas where abiotic and biotic stresses
currently limit production and could provide a better source for dwarfing
rootstock for high-density orchards. Mango species growing in swamps or
seasonally inundated areas (i.e. Mangifera decandra, Mangifera gedebe, Mangifera
inicarpoides, Mangifera griffithii and Mangifera quadrifida) represent a promis-
ing source of rootstock for the development of mango cultivation on poorly
drained soils and inundated lands. In West Kalimantan, Mangifera laurina is
occasionally used as a rootstock for commercial mango cultivars on periodi-
cally inundated riverbeds, and is now being tried at Sabah (Bompard, 1993).
Mangifera zeylanica has been tried as a rootstock for several mango cultivars
in Sri Lanka (Parsons, 1931). Interspecific hybridization of other Mangifera
species with the common mango could also increase the available genetic
variability for rootstock development.
The potential for using species in other genera in the Anacardiaceae as
rootstock has scarcely been explored. Burns and Prayag (1921) investigated
the use of Semecarpus anacardium, Spondias mangifera (Spondias pinnate), Spondias
acuminata and Horigarna grahami as rootstocks for common mango without
any success. Anacardium occidentale (cashew) seedlings have been reported to
be graft compatible with mango scions and mango fruit on cashew rootstock
were reportedly almost double the size of normal fruit with smaller seeds and
free from fibre (Garner and Chaudhri, 1976). Mango rootstock development
should receive as much attention as breeding of scion cultivars.
Classical breeding and grafting studies should be greatly expanded to
include the enormous genetic variability within the genus. Emerging bio-
technological approaches also should not be overlooked as alternative prop-
agation methods and as the means to engineer specific horticultural traits in
candidate rootstocks.
Crop Production: Propagation 393
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12 Crop Production: Mineral Nutrition
I.S.E. Bally
Department of Primary Industries and Fisheries, Queensland, Australia
12.1 Introduction
Assessment of the mineral status of mango trees is not without its challenges.
Like many other tropical woody perennial tree crop species, mangoes have
complicated and variable phenological cycles that influence the trees’ uptake
and translocation of minerals. Their extensive root system enables them to
exploit unevenly distributed minerals throughout the soil profile; these min-
erals are often not assessed during routine soil analysis. The leaves, trunk,
bark and roots act as mineral reserves that buffer many short-term mineral
shortages (Robinson, 1986b). Soil and leaf mineral analyses used as short-
term indicators of tree mineral status, tree productivity or fruit quality are
therefore difficult and unreliable (Catchpoole and Bally, 1995).
Leaf mineral analysis is commonly used to assess mango tree mineral status,
and is useful for developing and monitoring tree fertilizer programmes.
Leaves often display visual symptoms of toxic and deficient concentrations
of many minerals. Sampling mango leaves for mineral analysis should be
done when the tree is at its most phenologically quiescent stage, i.e. when
leaf mineral concentrations are most stable. One of the most stable periods in
the mango phenological cycle is the dormant phase, which occurs after the
completion of summer flushing and approximately 2 weeks before the emer-
gence of flower panicles. The common practice of withholding irrigation
water leading up to flowering makes this period the most inactive of the year
and an ideal time for leaf sampling. At other times of the year, leaf mineral
concentrations sharply decrease when the tree is flowering and fruiting and
increase in the months following harvesting (Catchpoole and Bally, 1995;
Oosthuyse, 2000b). Sampling at an inactive growth stage reduces variability
between leaf samples and provides a stable reference point for annual com-
parisons. Guides with respect to the most appropriate leaves for sampling
have been variously reported (Kumar and Nauriyal, 1979; Chadha et al., 1980;
Smith, 1992; Catchpoole and Bally, 1995), and generally concur that the most
appropriate leaves to sample are the third or fourth leaf behind the apical
bud, or the first full-size leaf of the most recently matured dormant flush
where leaves are fully expanded and hardened off.
Leaves recently sprayed with foliar nutrients or fungicides should be
avoided or noted, as analyses of manganese (Mn), zinc (Zn), boron (B) and
copper (Cu) are commonly affected by mineral residues on the outside of
leaves or imbedded in the cuticle, and which are not available to the tree.
Some publications recommend that sampled leaves should be washed with
deionized water to reduce residues from sprays and dust from the operator’s
hands (Reuter et al., 1986; Shu et al., 1992).
et al., 1999) (Table 12.1). Most of these standards are in reasonable agreement
with respect to the optimal ranges of individual minerals (Samra and Arora,
1997). Soil and leaf analyses should be interpreted by comparing with these
standards and with previous soil and leaf analyses and past seasons’ crop-
ping and fertilizer application history. The recommended optimal ranges of
leaf mineral concentrations should be considered as general guides only, as
high-yielding trees producing good quality fruit have been found to vary
widely in leaf mineral composition (Catchpoole and Bally, 1995; Oosthuyse,
2000b; Medeiros et al., 2004).
Positive relationships between leaf minerals and tree productivity have
been reported. Oosthuyse (1997) determined that leaf concentrations of nitro-
gen (N), phosphorus (P), potassium (K), magnesium (Mg) and Zn influenced
the number of fruit retained, and that leaf Zn and Mg also influenced fruit
size. Rao and Mukherjee (1989) observed positive correlations between tree
yields and leaf N and K from non-fruiting terminals in five Indian mango
cultivars with generally low N and K concentrations. Although these rela-
tionships have been observed, there are many factors that can influence pro-
ductivity. Therefore, predicting productivity based on leaf analysis alone is
difficult. However, soil and leaf analyses are useful for identifying major
mineral imbalances and long-term trends in tree nutrition, and can be used
to adjust fertilizer programmes.
Leaf age can affect the mineral concentration in assays. Minerals that are
mobile within the plant, for example N, P, K and Mg, generally decrease during
leaf aging and the less mobile minerals, such as calcium (Ca), sulfur (S), B and
Mn, accumulate in leaves with age (Chadha et al., 1980; Medeiros et al., 2004).
Mango leaves that have been sampled from fruiting terminals generally
display increasing concentrations of N, K, Ca, Mn, iron (Fe), Cu and Zn dur-
ing early fruit development and decreasing concentrations during late fruit
development and maturation. These minerals, with the exception of Mg and
P, also vary greatly among fruiting terminals (Oosthuyse, 1997, 2000b).
The Diagnosis and Recommendation Integrated System (DRIS) for inter-
preting leaf mineral analysis, uses ratios of mineral concentrations rather
than the absolute mineral concentration to identify limiting minerals in order
of their effect on the tree (Beaufils, 1973). With mango, DRIS has been used
with varying success. Raghupathi et al. (2004) observed that DRIS was unable
to diagnose the nutrient imbalance of a particular nutrient in isolation; how-
ever, others have used the technique more successfully. Schaffer et al. (1988)
used DRIS to identify Mn and Fe as the most deficient elements in orchards
with tree decline, a disorder of unknown aetiology, and Hundal et al. (2005),
Raj and Rao (2006) and Wadt et al. (2007) utilized DRIS to identify yield-
limiting mineral combinations in mangoes grown in India and Brazil.
12.5 Nitrogen
Nitrogen is one of the most important elements for crop production, and has
a significant role in mango growth, yield and fruit quality. Nitrogen is an
408
Table 12.1. Suggested optimal mango leaf mineral concentration ranges according to different sources.
References
Young and
Smith and Koo (1969, Crane Catchpoole Kumar Pimplasker Stassen Bhargava
Robinson Scudder 1971), Young and et al. and Bally and Nauriyal and Bhargava et al. and Chadha
Minerala Unit et al. (1997) (1951) Sauls (1981) (1997) (1995) (1977) (2003) (1999) (1988)
I.S.E. Bally
K % 0.3–1.2 0.97 0.5–1.0 0.4–0.8 0.4–2.5 0.50 1.02–2.01 0.1 0.54
Ca % 2.0–3.5 0.91 3.0–5.0 2.0–3.5 1.5–2.8 1.50 – 0.8–1.05 1.71
Mg % 0.15–0.4 0.26 0.15–0.47 0.25–0.35 0.2–0.4 0.15 – 2.8 0.91
S % 0.5–0.6 – – – 0.1–0.23 0.50 0.11–0.17 0.3 0.12
B % 50–80 – 24–84 – 20–140 – – 50 –
Fe mg/kg 7–200 – 38–120 70–100 30–120 – – 80 1.71
Mn mg/kg 60–500 – 92–182 – 160–980 – – 80 66
Zn mg/kg 20–150 – 10–119 20–40 20–63 – 11–26 40 25
Cu mg/kg 10–20 – 28–35 – 10–150 – – 20 12
Mo mg/kg – – – – 0.2–0.4 – – 50 –
a N, nitrogen; P, phosphorus; K, potassium; Ca, calcium; Mg, magnesium; S, sulfur; B, boron; Fe, iron; Mn, manganese; Zn, zinc; Cu, copper;
Mo, molybdenum.
Crop Production: Mineral Nutrition 409
Nitrogen in the soil occurs in many forms, but for the most part as large and
complex organic molecules that comprise the organic matter. These mole-
cules are too large for roots to absorb, and are broken down to nitrate (NO3–)
and ammonium (NH4+). The concentration of N in soil is dependent on the
concentrations of soil organic matter and mineral N. Nitrogen that is avail-
able to the plant is determined by the processes of mineralization, immobili-
zation, de-nitrification, volatilization and leaching, which are influenced by
temperature, moisture, pH and aeration. Nitrogen is easily leached from the
soil by rain and irrigation, and consequently, soil N often limits tree growth,
especially in sandy soils (Strong and Mason, 1999).
Common N fertilizers include: urea (CO(NH2)2, 46% N), potassium
nitrate (KNO3, 13% N, 38% K), calcium nitrate (CaNO3, 15.5% N, 19% Ca),
ammonium nitrate (NH4NO3, 35% N), sulfate of ammonia ((NH4)2SO4, 21%
N, 23.6% S). Nitrogen is taken up by mango trees primarily through the roots
as NO3– and NH4+; NO3– is the preferred source. Nitrogen can also be adsorbed
through leaves as ammonia (NH3), urea and amino acids. Nitrate, after being
adsorbed by the roots, is either reduced to NH4+ in the roots or translocated
to the leaves, stems or other tissues, where it is reduced to NH4+ in a two-step
process in which NO3– is reduced to nitrite (NO2–) which is in turn reduced to
NH4+ (Marschner, 1995). Ammonium metabolism is complex, and several
pathways are necessary to produce amino acids, proteins, enzymes and hor-
mones. Within the tree, N concentrations vary among tissues and are depen-
dent on N availability in the soil and demand within the tree. If N is limiting,
the plant can translocate N from older tissues to new growing tissues where
it is required (Marschner, 1995).
Nitrogen has a major effect on mango tree vigour, stimulating both vegeta-
tive and floral growth. Increases in shoot length, leaves per shoot and leaf
area have been demonstrated by Singh et al. (1973), Tiwari and Rajput (1975),
Syamal and Mishra (1989), Reddy et al. (2000) and Sergent et al. (2000).
Yeshitela et al. (2005) reported that N in combination with K, as KNO3
and urea, improved the percentage of terminal shoots that flower; however,
excessive N can stimulate vegetative growth at the expense of flowering,
fruit set and fruit quality (Scholefield et al., 1986; Monselise and Goren, 1987;
Silva et al., 2002). Nitrogen can cause increased fruit set and retention (Singh
et al., 1973; Oosthuyse, 1997) and fruit weight and yield (Alvian, 1974; Tiwari
and Rajput, 1975; Young and Koo, 1975; Alvian and Figueroa, 1977; Syamal
and Mishra, 1989; Reddy et al., 2003). Kanwar et al. (1987) demonstrated that
N at 100 g/tree/year-of-age was sufficient for tree growth and Young and
Koo (1975) reported that maximum yields were increased by N applications
of 0.8–1.1 kg/tree/year. Many reports of the effect of N on increased mango
fruit size and yields are based on data of N in combination with other ele-
ments (i.e. P and K) and may be partly attributed to these combinations;
however, N has a major influence on productivity in mango.
High N application rates that stimulate yield increases can also have
negative effects on fruit quality. Nitrogen has been negatively associated
with fruit colour in mango (Oosthuyse, 1993; McKenzie, 1994, 1995). Nguyen
et al. (2004) demonstrated that high N applications during fruit growth
inhibited the de-greening of ripening fruit, causing green skin at ripeness.
Bally (2007) also reported a negative relationship between N and fruit colour,
demonstrating that high leaf N concentrations reduce the percentage of
yellow skin in ripe fruit, reduced the lightness and chroma (vividness) of the
yellow colour, the percentage of skin covered with blush and the intensity of
the blush colour (Plate 73). He also identified a significant exponential rela-
tionship between the severity of sunburn and skin N concentrations; the
effects of sunburn were reduced as N concentrations increased in the fruit
skin.
Early indications of the negative effect of N on postharvest rots in mango
were demonstrated by Weng and Chuang (1997), who observed that N posi-
tively affected the germination rate, hyphal growth and appressoria forma-
tion of Colletotrichum gloeosporioides. When Nguyen et al. (2004) investigated
the effect of N on fruit quality, they found that 300 g/tree applied as a
foliar spray significantly increased the severity of anthracnose caused by C.
gloeosporioides. These observations were confirmed by Bally (2007), who found
that the severity of fruit anthracnose during ripening increased with applied
N (Plate 74). He also demonstrated that the N effect was because N enhanced
the decline of antifungal resorcinol compounds in the fruit exocarp during
Crop Production: Mineral Nutrition 411
Nitrogen management
Leaf N concentrations between 1 to 1.5% dry weight (DW) are generally con-
sidered to be in the optimal range (Robinson et al., 1997). Optimum concen-
trations for fruit skin N have not been published, although Catchpoole and
Bally (1995) reported skin N concentrations of 0.069% DW in mature ‘Kens-
ington Pride’ fruit. Leaf N concentrations vary throughout the year, and are
influenced by the growth events during the phenological cycle. Nitrogen
concentrations are generally greatest at the end of the summer vegetative
flushing period and decrease during panicle growth, flowering and fruiting
(Catchpoole and Bally, 1995; Medeiros et al., 2004).
Because N is highly mobile within the tree and a primary driver of
growth and fruit quality, the general practice of monitoring leaf and soil N
annually is inadequate for assessing tree N status at all stages of tree phenol-
ogy. A cheap, rapid test for N is needed to provide closer monitoring of
changes in leaf N status and allow the rates and timing of supplementary N
applications to be matched to tree and fruit demands. Bally and Still (per-
sonal communication) have calibrated the Konica-Minolta Soil Plant Analy-
sis Diagnostic (SPAD-502) meter to measure the chlorophyll content of leaves,
which is directly related to their N status (Gonzalez et al., 2005).
12.6 Phosphorus
Phosphorus that is available for plants occurs in two forms in soil, primarily
as the monovalent phosphate anion (H2OP4–) and secondly as the divalent
anion (HPO42–) in the soil water solution. The balance of these anions is
dependent on soil pH, with HPO42– favoured in soils >pH 7 and H2OP4–
412 I.S.E. Bally
favoured in soils <pH 7 and most readily available in soils with pH between
6 and 7. In higher pH soils, calcium phosphate compounds tie up P from
plant availability, and in lower pH soils, P oxidizes with Fe, aluminium (Al)
and Mn to become unavailable to plants. Other forms of soil P include liable
P that is bound on clays and organic matter, which can become available to
trees when dissolved in water.
Phosphorus is most rapidly taken up by the tree as HPO42– and slower as
H2OP4– and often enhanced by chemical association with N. After entry into
the roots, the anions are either converted to organic phosphates immediately
or after transport to other tissues. Within the tree, P is easily translocated
between tissues, with redistribution usually occurring from older leaves to
younger actively growing tissues. Phosphorus concentrations within the
mango tree are highest in the roots, wood and bark and lowest in the leaves
(Vuuren and Stassen, 1997). Leaf P concentrations are generally at their low-
est during fruit development and at their highest in the vegetative growth
period (Medeiros et al., 2004).
12.7 Potassium
Potassium in the cytoplasm is an activator of enzymes involved in photosyn-
thesis, respiration and starch and protein synthesis (Bhandal and Malik, 1988;
Crop Production: Mineral Nutrition 413
Marschner, 1995). Potassium is important for cell growth due to its role in cell
expansion and development of thick epidermal cell walls that increase the
resistance of trees to pathogens and insect pests. Potassium is involved in
tree water status by regulating water uptake by the roots and water loss
through the leaf stomata (Salisbury and Ross, 1992).
Potassium deficiency first appears in older mango leaves as the tree redistrib-
utes K to young growing tissues. Sen et al. (1947) induced K deficiency by
growing mangoes in sand culture and observed brown necrosis on the leaf
margins extending from the leaf tip towards the base. Smith and Scudder
(1951) also generated K deficiency symptoms in sand culture and described
them as irregularly distributed yellow spots and necrotic areas along the
margins of small, thin attenuated leaves which are very persistent. Lim and
Khoo (1985) described the non-necrotic areas of the leaf as dull, yellowish
green to light green; symptoms usually develop in the dry season or when
irrigation has been stopped.
414 I.S.E. Bally
Potassium nitrate applications just prior to and at the flowering stage promote
flowering, increase fruit set and fruit retention (Sergent and Leal, 1989; Lyan-
naz, 1994; Ferrari and Sergent, 1996; Oosthuyse, 1997; Rojas and Leal, 1997;
Sergent et al., 1997; Saleh and El-Monem, 2003; Shinde et al., 2006). In the low
and mid-latitude tropics, KNO3 is used to stimulate out-of-season flowering;
however, this effect is lost in higher latitudes (Davenport and Núñez-Elisea,
1997; see Davenport, Chapter 5, this volume). Shongwe and Roberts-Nkrumah
(1997) suggested the KNO3 effect on flowering is a result of lowering the tran-
spiration rate and increasing water use efficiency. Protacio (2000) suggests the
KNO3 effect on flowering is primarily due to N stimulation rather than K and
he postulated that KNO3 overcomes the inhibitory effects of gibberellic acid
(GA3) on starch accumulation by elevating the N concentrations over the N
threshold to synchronize bud break from apices with an existing floral initial.
Potassium influences fruit quality of many species (Marschner, 1995);
however, in mango there are only a few studies that link K nutrition with
increased fruit quality. Shinde et al. (2006) observed that increased K fertiliza-
tion increased fruit weight (5.15%), ascorbic acid (26.99%), organoleptic score
for texture, flavour, colour and shelf life and reduced physiological weight loss
(22.79%) and spongy tissue (68.08%). Potassium in the form of monopotassium
phosphate (KH2PO4) suppresses powdery mildew on mango (Reuveni et al.,
1998; Oosthuyse, 2000a), but it is not clear if the effect is due to P or K.
12.8 Magnesium
Magnesium deficiency first appears in older leaves, due to its high mobility
within the tree, as pale green or yellow leaves with the inner vein areas of the
Crop Production: Mineral Nutrition 415
leaf lamina becoming mottled and necrotic while the leaf veins remain green.
Young trees are stunted by Mg deficiency. Smith and Scudder (1951) gener-
ated symptoms of Mg deficiency in sand culture and described them as a
green wedge pattern formed by the lateral intrusion of a bronzed chlorosis
along the leaf margin (Plate 75). Deficiency symptoms are more likely to
occur in textured, sandy or highly leached soils due to their low cation
exchange capacity or when heavy applications of liming products, K fertil-
izers or green manuring have been applied (Stassen et al., 1999). Magne-
sium deficiency has been associated with a browning skin discoloration in
‘Kensington Pride’ fruit.
12.9 Sulfur
12.10 Zinc
Zinc is chemically bound to Fe and Mn to form components of chlorophyll, and
is essential for photosynthesis (Weir and Cresswell, 1995). Zinc is essential for the
synthesis of proteins and hormones, including auxins, and is required for the
maintenance of biomembranes (Salisbury and Ross, 1992; Marschner, 1995).
Zinc deficiency is often referred to as ‘little leaf’ because leaves fail to reach
full size. Symptoms first appear in immature leaves in the coloured stage, as
Crop Production: Mineral Nutrition 417
a thickening of the leaf and failure to fully expand. Leaf expansion is often
stunted on one side of the blade, causing it to form a sickle shape (Dilly et al.,
1997) (Plate 76). Symptoms in fully mature leaves are prominent light-yellow
or olive-green veins on the upper surface that are thick and brittle. Internode
length is reduced, thereby causing a rosetting effect (Lim and Khoo, 1985;
Agarwala et al., 1988; Marschner, 1995). Zinc and Fe deficiency often occur
together because of their association with calcareous high pH soils.
Singh and Rajput (1977) reported that foliar sprays of 2.0–8.0% ZnSO4 prior
to flowering increased the number of perfect flowers in panicles and later
increased fruit yield, fruit sugar, ascorbic acid and total soluble solids (TSS).
Daulta et al. (1981) also observed that foliar Zn sprays increased fruit set and
improved fruit quality, but had no effect on sex ratio. Kumar and Kumar
(1989) demonstrated that 1% ZnSO4 sprays applied to ‘Dashehari’ trees
improved postharvest fruit life by reducing weight loss, spoilage, increasing
sugars, reducing acidity and slightly increasing vitamin A. Littlemore et al.
(1991) observed that 1% ZnSO4 foliar sprays applied quarterly to ‘Kensing-
ton Pride’ was sufficient to maintain leaf Zn concentrations above critical
concentrations and avoid symptoms of little leaf. Although leaf Zn concen-
trations were improved and symptoms cured, no effect on tree yields was
observed. Soil applications are often ineffective due to adverse soil conditions
making it unavailable to trees.
12.11 Manganese
Role of Mn
There are few reports on the effect of Mn on crop production or fruit quality.
Schaffer (1994) suggested that mango decline was associated with trees that
are deficient in Mn and Fe.
12.12 Iron
Iron is a trace element that is required in small quantities, and is a component
of several enzymes and molecules. It is a component of chlorophyll and leaf
Fe concentrations directly impact photosynthesis. Iron-containing enzymes
participate in oxidation processes that release energy from sugars and
starches, the conversion of nitrates to ammonia and the biosynthesis of eth-
ylene (Marschner, 1995). In proteins, Fe acts as an electron carrier by alternate
oxidation and reduction between Fe2+ to Fe3+ (Salisbury and Ross, 1992).
12.13 Calcium
A major role of Ca is membrane stability, which is achieved when Ca2+ binds
to phosphate and carboxylate groups of phospholipids and proteins on the
surface of the plasmalemma, and thereby preventing leaking of solutes to the
cytoplasm (Kirkby and Pilbeam, 1984). Calcium protects cells from toxins,
slowing the aging of plant tissues and promoting longer shelf life of many
fruits. Calcium is important for pectin polymers that strengthen cell walls
and provide defence from pathogens (Ferguson, 1984). Most Ca in trees is
fixed in cell walls and is not easily translocated. Calcium is essential for new
root hair and leaf development.
Calcium deficiency symptoms reflect the low mobility of Ca within the tree,
first appearing in young, actively growing tissues. Ca-deficient plants generally
display symptoms of membrane degeneration, which have been likened to
senescence and fruit ripening (Fallahi et al., 1977; Bangerth, 1979). Many symp-
toms of internal disorders in mango appear to be associated with premature
ripening or cell degeneration (Burdon et al., 1991, 1992). Raymond et al. (1998a)
observed that cell disruption and rupture of the cell walls were the first micro-
scopic indicators of soft nose, stem-end cavity and jelly seed. High Ca concen-
trations reduce the binding sites for enzymatic degradation (Rhodes, 1980). No
direct symptoms of Ca toxicity have been recorded in mango, but high concen-
trations of Ca in soils can displace other minerals such as Mg, Zn, B, Cu and P.
In mango, fruit Ca has been implicated as a factor in fruit quality, for example
with regards to internal physiological disorders, shelf life and disease resis-
tance. Many symptoms of internal physiological disorders appear to be asso-
ciated with low Ca-related membrane degeneration, i.e. premature ripening
(Burdon et al., 1992; Raymond et al., 1998a). Links between internal break-
down and Ca nutrition in mango were reported by Young (1957), who found
Crop Production: Mineral Nutrition 421
12.14 Boron
Boron binds as cis-diol borate complexes to mannose and certain other sug-
ars in cell wall polysaccharides and also has a role in sugar movement in the
tree. Boron also is important in nucleic acid synthesis (Salisbury and Ross,
1992), and is essential for pollen and flower development, pollen germina-
tion and pollen tube growth (Stanley and Lichtenberg, 1963; Gupta et al.,
1985) and is thus essential for mango fruit set. Boron also is important in the
synthesis of proteins that translocate sugars (Gupta et al., 1985).
Boron deficiency is expressed in tissues that are rapidly expanding and have
low transpirational rates, for example roots, fruits and shoots (Shelp et al.,
1995). In mango, B deficiency can result in poor flowering, pollination and
reduced fruit set. It is expressed in growing shoots by uneven cell division,
causing leaves to grow lop-sided with a curved sickle shape and deformed
lamina and margins (Plate 78). Leaves often have shot-holes that are sur-
rounded by a light-green halo and ragged margins. Apical dominance can be
lost with swelling of the internodes. The main raceme of panicles can develop
a slight bend or kink towards the tip and in some cultivars the bark splits and
oozes black gummy sap from the cracks, known as gummosis (Plate 78)
(Nartvaranant et al., 2002). Agarwala et al. (1988) generated B deficiency in
1-year-old seedlings using sand culture, and described mild deficiency symp-
toms, as mild chlorosis with a marked reduction in length and width of the
leaves. In more severe cases, older leaves become chocolate-brown at the base,
spreading to the tip before becoming completely chlorotic. Stems turned black,
lost apical dominance and eventually stopped growing. Boron deficiency can
be aggravated by high N status of trees (Ram et al., 1989; Raja et al., 2005).
In mango fruit, B deficiency causes cracks that split open; there is brown
discoloration of the mesocarp. Lumpy, deformed fruit may also be a symp-
tom of B deficiency. Meneses et al. (1994) examined normal and deformed
fruit using neutron capture radiography, and suggested that the deformities
were due to B toxicity. Boron toxicity is common in mango with typical symp-
toms appearing in leaves as dark spots on leaf margins that coalesce, eventu-
ally leading to marginal leaf necrosis in more severe cases (Plate 79). Boron
toxicity often occurs after excessive application of B fertilizers. Symptoms can
be amended through leaching of fertilizer from the root zone, raising soil pH
with applications of lime or stimulating growth through application of N;
however, these measures may have other implications for crop production.
25 g/m borax (11% B) during the summer wet season; however, the response
time and effect on gummosis of the two cultivars differed (Nartvaranant
et al., 2002). Foliar application of B solutions at the pre-flowering stage
increased yield and fruit quality in several studies. Dutta (2004) observed
that 3000 ppm boric acid is optimal for maximizing yield and quality of ‘Him-
sagar’ in West Bengal, India. Coetzer et al. (1991) reported that foliar applica-
tion of B at flowering raised leaf B concentrations to 60 mg/kg and increased
yields from 14 to 22 kg/tree as well as improving fruit quality. When Loría-
Meneses et al. (1992) applied boric acid to the skin of developing fruit they
found it was not significantly translocated into the mesocarp and remained
in the surface layers of the skin around secretary glands. The response of
mango trees to soil applications of B varies between cultivars. Rossetto et al.
(2000) observed that ‘Winter’ was least sensitive, whereas, ‘Tommy Atkins’,
‘Haden’ and ‘Van Dyke’ in declining order were more sensitive.
12.15 Conclusion
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Vuuren, B.P.H.J. and Stassen, P.J.C. (1997) Seasonal uptake of macro elements by young
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Wadt, P.G.S., Silva, D.J., Maia, C.E., Tome, J.B., Pinto, P.A.D. and Machado, P. (2007)
Modelling of functions in calculating DRIS indices. Pesquisa Agropecuaria Brasile-
ira 42, 57–64.
Weir, R.G. and Cresswell, G.C. (1995) Plant Nutrient Disorders. 2 Tropical Fruit and Nut
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Weng, F.Y. and Chuang, T.Y. (1997) Nutrient requirement of conidial germination and
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Yeshitela, T., Robbertse, P.J. and Stassen, P.J.C. (2005) Potassium nitrate and urea sprays
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South African Journal of Plant and Soil 22, 28–32.
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the Florida State Horticultural Society 70, 280–283.
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Extension Service Bulletin 189, 1–70.
13 Crop Production: Management
13.1 Introduction
Fruit
development
Flowering and
fruit set
Flower bud Postharvest
development Unwanted vegetative flush
vegetative
flush
Harvest
Amount of development
Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct
n
s May Jun July Aug Sept Oct Nov Dec Jan Feb Mar Apr
Fig.13.1. Theoretical mango phenological cycle (Source: after Cull, 1991). s, southern
hemisphere; n, northern hemisphere.
434 J.H. Crane et al.
The mango industry of Brazil is extensive due to favourable soil and climatic
conditions. Bahia and São Paulo are the most important production areas of
Brazil with 34,600 and 28,800 ha, respectively, of the estimated 70,000 ha of
mango production in 2001 (Table 13.1). The north and south of Brazil account
for only 6.0% of the total mango-production area (Souza et al., 2002). The
most recent mango census (2005 data) estimated mango production in Brazil
to be c.970,700 t (Gazeta, 2006; Agrianual, 2007), which is a slight increase
since 2001. Bahia and São Paulo also have the largest production, respec-
tively (Table 13.1). Brazil exports 133,300 t of mangoes, 13.7% of the total
mango production. Although pest and disease problems and the highly com-
petitive internal and external markets have limited the expansion of the
mango industry in some Brazilian regions, important technologies have been
Table 13.1. Regions of mango production in Brazil, Mexico, Taiwan and the USA
(Source: see table footnotes).
developed to improve and support the mango industry (Santos Filho et al.,
2002). This has resulted in an increase of 58% in mango exports from 1998 to
2005 (Souza et al., 2002; Agrianual, 2007).
There are c.181,525 ha of mangoes in 23 of Mexico’s 32 states (SIIAP,
2007); there are four large mango-producing regions (Table 13.1), ranging
from 14°33N to 27°N latitude. They are distinguished by differences in cli-
mate, soils and cultivars. There are 106,740 ha of mango in the Northern and
Central Pacific region; the major cultivars are ‘Tommy Atkins’, ‘Ataulfo’,
‘Kent’, ‘Haden’ and ‘Keitt’. The Southern Pacific region has 42,180 ha and
produces ‘Manila’ (‘Carabao’), ‘Ataulfo’, ‘Tommy Atkins’, ‘Haden’ and
‘Kent’. The leading cultivars of the Gulf of Mexico region (28,778 ha) are
‘Manila’ followed by ‘Tommy Atkins’. Mango production in Mexico was
about 1.7 million t in 2006 and the states of Sinaloa, Guerrero, Nayarit, Oax-
aca, Chiapas, Veracruz and Michoacán accounted for 92% of the crop (SIIAP,
2007). All these states except Veracruz (which abuts the Gulf of Mexico) are
situated along the Pacific coastal and inland areas.
In 2006 Mexico exported 196,120 t to the USA valued at >US$132 million
(EMEX, 2007). Mexico supplies 87.5% of the mangoes consumed in the USA
and mango exports in 2006 were 17.9% higher than in the 2005. The major
export cultivars are ‘Tommy Atkins’ (33%), ‘Kent’ (23%), ‘Ataulfo’ (19%), ‘Keitt’
(14%) and ‘Haden’ (11%). None the less, most mango production (c.85%) in
Mexico is for the domestic market. The most important mango warehouses
and distributors are in Mexico City, Guadalajara and Monterrey.
The 7-month mango season in Mexico is due to differences in latitude,
phenological cycles, rainfall and soil moisture patterns and the use of growth
regulators. Chiapas, along the South Pacific coast, is usually the first to har-
vest mangoes (‘Ataulfo’). Full bloom of ‘Ataulfo’ occurs from mid-November
to mid-February and fruit is harvested from the end of January to the end of
May. Mango harvesting continues northwards from Chiapas to the states of
Oaxaca, Guerrero, Michoacán, Colima, Jalisco, Nayarit, Sinaloa and Sonora.
Sinaloa is a major mango producer with the latest harvest season for ‘Manila’
(July), ‘Haden’ (July), ‘Tommy Atkins’ (July), ‘Kent’ (July–August) and ‘Keitt’
(August–September). The ‘Manila’ mango is produced mainly in Veracruz
and Guerrero. Full bloom occurs in January–February and fruit is harvested
in May–June.
The production area in Taiwan increased from 17,000 ha in 1987 to 21,000
ha in 1992 and stabilized at c.20,000 ha for c.10 years when the yield was
17 kg/tree; however, as production has increased to c.27 kg/tree, the plant-
ing area has decreased to c.18,200 ha in 2006 (Table 13.1). There are six
important mango-growing prefectures in Taiwan: Nantou, Taichung, Chiayi,
Tainan, Kaohsiung and Pingtung from north to south in western and eastern
Taiwan. Tainan and Pingtung are the most important production areas, com-
prising about 90% of the total harvested area.
Mangoes are grown commercially in four areas of the USA: Florida
(25–28°N), Puerto Rico (18–18°5´N), Hawaii (22–30°N) and California
(33–33°5´N) (Table 13.1). Puerto Rico has the largest area (1079 ha) (Alvarado-
Ortiz and Acin, 2004), followed by Florida (324 ha) (FASS, 2003; Anonymous,
436 J.H. Crane et al.
Mangoes are mainly cultivated in the tropics (25°N, 25°S) and subtropics
(35°N, 35°S), although limited production also occurs in warm temperate/
subtropical, i.e. Mediterranean-type areas, for example Israel, southern Spain
and the Canary Islands and California. The ideal areas for mango production
have a cool and/or dry period prior to flowering followed by moderate soil
moisture and moderately hot temperatures (30–33°C) (Chacko, 1986). The
diversity of climates and soils in mango-production areas reflects the adapt-
ability of the species and improvements in cultural practices.
Temperature and availability of water are the most significant environ-
mental factors that influence commercial mango production by affecting the
frequency, intensity, duration and time of vegetative growth and flowering
(Chacko, 1986, 1989; Whiley, 1993; Schaffer et al., 1994; Núñez-Elisea and
Davenport, 1995; Davenport, 2006) and disease incidence (Johnson et al.,
1989; Ploetz, 1994, 2003; Ploetz et al., 1994). Temperatures <15°C and >30°C
inhibit pollen tube germination, thereby impeding fertilization and resulting
in embryo abortion and fruit abscission.
Mango production in Brazil extends from 3°N, close to the Amazon
region, where a humid, hot tropical climate predominates almost year-round
to approximately 25°S in Paraná, which has a subtropical climate. Between
these two extremes, there are various soil and climatic conditions where
mango trees are grown intensively. The semi-arid tropical climate of the
north-east region of Petrolina in Pernambuco (21°13´N and 47°50´W) is more
appropriate for high-quality mango production than Votuporanga in São
Paulo (21°13´S and 47°50´W), which has a subtropical climate (Table 13.2).
Table 13.2. Climatic parameters for Brazil, Mexico, Taiwan and the USA.
Yearly Yearly
Country State Region Climatic category mean Mean Mean mean Wet Dry
Brazilb Pará North region Tropical hot humid 26.0 31.5 22.0 2893 436.2 111.8
Goiás Central region Savannah dry tropics 29.8 31.9 17.7 1576 270.3 6.2
Bahia North-east region Tropical semi-arid 27.9 33.8 22.0 545 139.6 1.7
437
(Continued)
438
Table 13.2. Continued
Yearly Yearly
Country State Region Climatic category mean Mean Mean mean Wet Dry
USAe Florida Southern coasts Marine subtropics – 23.2 26.8 (26–27) 18.9 (18–19) 1643 1313 333
east coast
precipitation is averages for wettest (May–October in Pintung; April–September in Tainan and island average) and driest (October–March) months and yearly
annual. Available at http://www.cwb.gov.tw/
e Source of data on USA: Butson and Prine (1968); Getz (1979); E.E. Toro, personal communication; C.L. Chia, personal communication; Quayle et al. (1995);
Some cultivars, for example ‘Haden’ in north-eastern Brazil, have poor fruit
set when temperatures are >35°C.
Solar radiation is very important for fruit development, since its dura-
tion and intensity are directly related to photosynthesis and carbohydrate
production (Mukherjee, 1953). Light incidence depends on the season (Allen
et al., 1998). According to Lima Filho (2000), mango production in the semi-
arid region of Brazil is where maximum solar radiation occurs in summer
(October southern hemisphere; 528 cal/cm2/day) and the minimum in win-
ter (June southern hemisphere; 363 cal/cm2/day), which corresponds to the
flowering and fruit development stages, respectively.
Mango production in Mexico is in the tropics. Most rainfall occurs dur-
ing the summer and may be accompanied by hurricanes (Table 13.2). Climate
in the Northern and Central Pacific region (17–27°N) ranges from warm sub-
humid to warm/very warm semi-arid tropics with a 7-month dry season
(García, 1973). In Colima and Michoacán, mangoes are irrigated due to low
annual precipitation (semi-arid condition with a 6- to 8-month dry season;
the mean annual temperature is 26.5–28.3°C). The Southern Pacific and Gulf
of Mexico coastal regions have isothermal (< 5°C monthly temperature oscil-
lation) warm, subhumid to humid climates with a 6–8-month dry season
(García, 1973).
The Gulf of Mexico coastal region is affected by winds from the north
(11–28 m/s) from October to April. These winds cause direct damage, such as
limbs breaking and flower and fruit drop, and also increase plant respiration
and transpiration rates that stress the trees. In the Pacific coastal regions hur-
ricanes usually occur from August to October. Recent, unusually warm win-
ters have resulted in undesirable late autumn or winter vegetative flushes and
poor flowering. Normal bloom occurs in late January and February; how-
ever, late autumn or winter shoots delay anthesis during hot spring tempera-
tures (April to early May) which result either in low fruit set or parthenocarpic
fruit.
The primary production areas in Taiwan are from 22°N (Fungshan, Ping-
tung) to 24°N (Nanto and Taichung prefectures). The average temperature of
these areas is c.23°C, with a mean of 20°C during flowering (December–
March) and c.18.6°C during development (April–August) (Table 13.2). Flower
induction in mango is not a problem in Taiwan because of its subtropical
climate; however, fruit set can be affected by low temperature, rainfall, etc. A
monsoon-type climate prevails in southern Taiwan with precipitation
occurring mostly from May through to September. There is little rainfall
(usually <50 mm) during flower bud formation and flowering (autumn-winter)
(Table 13.2).
The production areas of the USA have different climates. South Florida
has a marine subtropical climate (Table 13.2). South-east Florida has a mean
annual temperature of 23°C, a mean annual rainfall of 1643 mm and 62%
relative humidity (RH) (Butson and Prine, 1968; Getz, 1979; Barrick and
Black, 1980). Constant ocean-spawned winds of 4–9 km/h buffet the region
from February through to October. The wet season (two-thirds annual rain-
fall) occurs during the late spring into autumn (May–October, northern
440 J.H. Crane et al.
hemisphere) and the dry season occurs during winter and early spring
(November–April). Lowest temperatures (-4 to -6°C) occur from December
through to February with a 70% probability of 0°C at least once each year
(Bradley, 1975).
The Puerto Rican industry is mostly along the south and south-west
coast at elevations between sea level and 50 m (Table 13.2) (Aponte-Morán
et al., 1977). The La Cordillera central mountain range divides the island from
north to south and has a major impact on the island’s climate, with annual
rainfall of 1550 mm in the north and 904 mm in the south (Espenshade, 1992;
SERCC, 2007). In the main production areas, the dry season is from Decem-
ber to May, although July and August are also dry (Aponte-Morán et al.,
1977). Mean maximum and minimum temperatures vary with altitude; how-
ever, along the coastal production areas, 24–29°C is normal (Espenshade,
1992; SERCC, 2007). The lowest temperatures (18–20°C) occur along the coast
during January and February.
The Hawaii mango industry is located mostly on the leeward coasts of
Oahu, Hawaii and Maui islands; however, limited production occurs on the
windward side of Kauai Island (Table 13.2). Each commercial planting has a
distinct climatological niche that varies with altitude and location with
respect to mountains and predominant north-east trade winds. Annual rain-
fall is 531 mm on Maui, 559 mm on Oahu and 3279 mm near Hilo (east coast
of Hawaii) (Table 13.2; C.L. Chia, personal communication). Temperatures,
like rainfall, vary with elevation and location but generally range between 21
and 27°C (Bahr and Johnston, 1993b). Lower (13°C) and higher temperatures
(32°C) occur occasionally.
The Coachela Valley of California is in the Salton Trough Desert area
(Bahr and Johnston, 1993a) (Table 13.2). The valley ranges from 80 m below
sea level to 488 m above sea level (Espenshade, 1992; Aslan et al., 1993) and
the climate is arid. Temperatures in the valley vary considerably depending
upon season, elevation and exposure. Some data reported herein represent
the average for the area and may not accurately reflect the particular micro-
climate of the mango orchards in the valley (Garczynski, 1995). The mean
annual temperature is 22.5°C; the average summer temperature is 31.8°C,
and the average winter temperature is 13.3°C (Aslan et al., 1991; Garczynski,
1995). Winter temperatures range from 22.1°C to 4.5°C (Garczynski, 1995);
however, Schacht (1992) reported a low of -4.4°C for 2 h. Summer tempera-
ture extremes range from a low of 11°C to 50°C (Garczynski, 1995). The aver-
age annual rainfall is 79.8 mm with two-thirds of this occurring during the
winter (December–March) (Aslan et al., 1991; Garczynski, 1995). The valley is
buffeted by wind and sandstorms during late spring, and these can cause
severe damage (Schacht, 1992; Aslan et al., 1993).
in deep, fertile, moderately acid to neutral pH, loam-type soils. They tolerate
infertile sands, volcanic ash and limestone-based soils, excessively drained
and or periodically flooded soils and soils with acid (pH 4.5–7) to alkaline pH
(pH 7–8.5). Mangoes are sensitive to saline and sodic soil conditions and
proper irrigation practices and the use of salt-tolerant rootstocks is imperative
for successful crop production in some areas (Schaffer et al., 1994).
Land preparation includes clearing virgin or existing orchard trees, disk-
ing, destruction of subsurface hardpans (slip ploughing), mixing of upper
and lower soil profiles, crushing superficial bedrock and mixing rock and soil
layers, formation of land contours to facilitate drainage and/or flood irriga-
tion, bedding and amending soils with organic matter and inorganic chemi-
cals, for example hydrated lime (Ca(OH)2), dolomite and preplant fertilizers.
Mango is grown on a wide range of soil types, from latossols with a high
percentage of sand in the north-east of Brazil to loamy oxysols in the south-
east. Some areas in the north-east have shallow soils that need improved
drainage. Soils of the central region are chemically poor and acid (pH 3.7–4.7)
and require lime and/or gypsum application prior to orchard establishment.
Soil analysis is used for determining the suitability of soil for production and
potential fruit quality, especially in areas cultivated for export fruit. Typically
soil analysis is conducted prior to land clearing and ploughing to determine
nutrient levels and the need for lime and/or gypsum application. Soil sam-
ples are taken from 0–20 cm and 20–40 cm depth of the soil profile. Soil sam-
ples are usually taken at a 0–30 cm and 30–60 cm soil depth for mature and
established orchards. Ploughing, harrowing and lime and/or gypsum incor-
poration are recommended at 30 cm depth and at least 30 days prior to rainy
weather (Pinto and Ramos, 1998). Liming is very important for acid soils
(<pH 4.0) in the central region of Brazil in order to increase soil pH to 6.0–6.5.
It also improves the soil base saturation to 60–70% and results in better
soil conditions for growth and production (Pinto, 2000). The amount of lime
to be applied is based on a basis saturation equation (where hydrogen (H),
aluminium (Al), calcium (Ca), magnesium (Mg) and potassium (K)):
Lime rate (t/ha): (T × 0.5) – S where T = (H + Al) + S and S = Ca + Mg + K
Gypsum applications are recommended for acid subsoils with >20% Al
saturation and <0.5 cmol/dm3 Ca at soil depth of 60 cm (Andrade, 2004).
This has been shown to significantly reduce fruit physiological disorders,
such as soft nose.
Production areas of Mexico vary in soil characteristics, and include flat
coastal areas and steep mountain slopes from sea level to >600 m above sea
level. Orchards on sloped land commonly utilize individual tree terraces.
Soil depth ranges from 30 cm to >3 m (alluvial soils). Shallow and eroded
soils are common on hilly terrain. The presence of medium to large (10–50 kg)
boulders can impede the use of machinery but they reduce erosion and
increase soil-water storage capacity. In general, poor soil drainage is not a
problem; however, areas with clay soils have drainage problems during peri-
ods of heavy rain, especially if there is a shallow water table. A range of soil
types is used for mango production in Mexico. In the Northern Pacific region,
442 J.H. Crane et al.
mangoes are planted in cambisols (pH 6.0–7.0), luvisols (pH 6.5–7.5) and
feozem (pH 5.0–6.0) soils with loamy textures (Anonymous, 1970, 1982).
These soils are well drained, with 2–3% organic matter, moderate to high
water holding capacity and cation exchange capacity (CEC) of 10–20+
meq/100 g soil. In the Central Pacific region mangoes are planted on cambi-
sols, feozem and regosols with light textures (Anonymous, 1970, 1982). The
regosols (pH 6.5–7.5) are well drained, with <2% organic matter content, low
water holding capacity and CEC <10 meq/100 g soil. Soils planted to man-
goes in the Southern Pacific region include nitosols, luvisols and feozem
(Anonymous, 1970, 1982). The nitosols (pH 5.0–6.0) are well drained, with
>3% organic matter content, low to moderate water holding capacity and
low CEC (<10–20 meq/100 g soil). Soils in the Gulf of Mexico region include
fluvisols, cambisols and vertisols of loamy and clayey textures (Anonymous,
1970, 1982). The fluvisols and vertisols are moderately and slowly drained
soils, respectively. Soil pH for fluvisols ranges from 5.5 to 7.5 with <2%
organic matter content. Soil pH for the vertisols is alkaline (7.5–8.0) and the
organic matter content is moderate (2.1–3%). The water holding capacity and
CEC is low to moderate for the fluvisols (<10–20 meq/100 g soil) and high
for the vertisols (10–20+ meq/100 g soil).
Soil tillage is used in flat lands before planting. Dimensions of planting
holes range from 40–60 cm depth and 30–50 cm diameter in light-textured
soils. Planting holes can be larger in heavy-textured or stony soils. On steep
hills only planting holes are made and individual terraces are built up to
hold soil, water, organic matter, fertilizers or soil amendments. Preplant soil
analyses are rarely taken; however, chemical or organic fertilizers are com-
monly applied at planting time. With rapid expansion of the mango industry,
orchards have been planted in shallow soils (0.75–1.2 m depth) underlain
with a hardpan. These soils are poorly drained and root growth and exten-
sion are limited. Trees growing in such soils are prone to drought stress dur-
ing dry periods, flooding stress after rains and nutritional deficiencies. The
weakened trees appear to be more susceptible to pathogens, for example
Botryodiplodia theobromae, which causes cankers or stem dieback and eventu-
ally tree death (Ponce-González and Salazar-García, 1992). Cultural practices
are not available to ameliorate these problems and planting in such soils is
not recommended.
Mangoes are grown in Taiwan on sandy loam, loamy sands, clay and
coarse sandy soils. Soil pH ranges from 5.0 to >7.8. Most trees are grown on
sloping land. Silt clay loam with pH 5.0–6.0 is mostly found in the Pingtung
area, while the Tainan area has clay soils with pH 7.3–7.8. Acid soils are
amended with Ca(OH)2 or dolomite and alkaline soils are amended with
sulfur (S) or acid-based fertilizers.
The topography in the mango-production areas of Florida is flat and
ranges from sea level to c.6.1 m above sea level. Soils in Florida include vari-
ous sands, muck and oolitic limestone. In the main production area (Miami-
Dade County), mangoes are planted in extensively scarified (crushed) and
trenched oolitic limestone rock (Krome and Chekika very gravelly loam)
(Colburn and Goldweber, 1961; Noble et al., 1996). The limestone soil is very
Crop Production: Management 443
permeable (1.5–5.1 cm/h), with high pH (7.4–8.4), low organic matter con-
tent (3–10%), low water holding capacity (0.2–0.3 cm/cm of soil) and low
CEC (16.0–37.0 meq/100 g soil) (Calhoun et al., 1974; Anonymous, 1989). In
areas subject to flooding, crushed rock is formed into beds (0.6–1.0 m high
and 1.0–1.5 m wide) before planting. The sandy soils in other Florida produc-
tion areas are poorly to well drained with or without an undulation hard-
pan 0.5–3.0 m down in the soil profile (Henderson et al., 1984). The highly
organic muck soils in Palm Beach County are poorly drained and underlain
by dense limestone bedrock. Beds of varying dimensions are made in the
sandy and muck areas to increase the proportion of the root system above
flood levels. The sand and muck soils are characterized by an acid to alka-
line pH (3.6–8.4) and low cation exchange capacities (Carlisle et al., 1978;
Henderson et al., 1984).
In Puerto Rico, mangoes are grown on flat and gently sloping land con-
sisting of alluvial fans and terraces level with or slightly above the flood
plain (Bierbolini et al., 1979). Some orchards in western Puerto Rico are
located on moderately steep to very steep slopes (12–60% slope) and rounded
hill tops that are somewhat eroded and superficial (Bierbolini et al., 1975).
Soils in southern Puerto Rico are very deep, moderately well to well drained
and consist of fine-textured sediment of sandy and clayey loam over gravelly
fine-textured sediment. The pH ranges from moderately acid (pH 6) to alka-
line (pH 8 or above). Soils have good to very good native fertility and good
water holding capacity. Soils in the west are well drained and moderately
acid (pH 5.0–6.0).
Mango production in Hawaii occurs on volcanic ash soils varying from
recent to highly weathered (R. Yost, personal communication). They tend to
be well drained; some soils must be slip ploughed to break up the hardpans
in preparation for planting. The pH ranges from 5 to 8 and the fertility varies
substantially. Mangoes in California are on lacustrine deposits consisting of
fine-textured sediments that are highly stratified sandy loam soils with clay
lenses (S. Aslan, personal communication). The soil is alkaline (pH 7–8.4) and
calcareous (Aslan et al., 1991). Land is slip ploughed to 1.5–1.8 m depth to
break up and mix the stratified soil layers.
The main commercial cultivars in the USA, ‘Keitt’, ‘Tommy Atkins’, ‘Van
Dyke’, ‘Haden’ and ‘Brooks’, are monoembryonic. Orchards are planted with
grafted or budded trees (Aponte-Morán et al., 1977; Chia et al., 1988; Crane
and Campbell, 1991; Hamilton et al., 1992). The most common rootstocks in
Florida are seedlings of polyembryonic ‘Turpentine’, ‘Number 11’ and
‘Criollo’. In Hawaii, any mango seedlings are used. In California, ‘Turpen-
tine’ or ‘Criollo’ are used. In Puerto Rico, polyembryonic ‘Mangotino’ and
‘Pasote’ seedlings are used in the Ponce region and ‘Mayaguezano’, ‘Turpen-
tine’ and ‘Colombo Kidney’ in the Mayaguez region (Aponte-Morán et al.,
1977; Toro, 1988; E.E. Toro, personal communication). Regardless of the seed
source, roguing the zygotic seedlings is important for obtaining uniform tree
size, growth characteristics and production (Schnell and Knight, 1991, 1992;
Degani et al., 1993; Schnell et al., 1994). Seed is removed from mature fruit
along with the husk, and the seed is planted in well-drained container media
(Sauls and Campbell, 1980; Young and Sauls, 1989). Seedlings are large
enough to graft or bud after 2–6 months. The most common vegetative prop-
agation methods in Florida are veneer and cleft grafting (Sauls and Camp-
bell, 1980; Crane and Campbell, 1991) and chip and ‘T’ or inverted ‘T’
budding. The trees are usually ready for field planting in 6–12 months. Graft-
ing may be done at any time of year when suitable rootstocks are available,
but it is more successful during warm weather. In Puerto Rico the most
common propagation method is cleft grafting (Aponte-Morán et al., 1977;
Toro, 1988). Grafting is most successful during the spring and from September
through to November.
Topworking of established trees is common. Trees are cut back to several
major limbs or the trunk. In Florida, several sprouts emerging from the
pruned limbs are allowed to develop to 1.3–7.6 cm diameter, and are veneer
grafted with the new cultivar (Sauls and Campbell, 1980). In Puerto Rico,
several sprouts are allowed to develop, and larger sprouts (2.5–7.6 cm diam-
eter) are cleft grafted and smaller diameter sprouts (1.0–1.5 cm) are bark
grafted (Toro, 1988).
Country Cultivar Origin Seed type Production seasona Tree growth habitb
Florida selections (i.e. ‘Haden’, ‘Keitt’, ‘Tommy Atkins’, ‘Van Dyke’ and ‘Palmer’)
are grown for the export trade. Most Florida cultivars produce from Decem-
ber to January except ‘Palmer’, which is harvested between January and Feb-
ruary and ‘Keitt’, which is harvested between February and March. ‘Tommy
Atkins’ represents 80% of the commercial export volume in Brazil (Pinto,
2004); ‘Palmer’ is increasing in demand in the national market.
Mango production in Mexico relies on the following cultivars (from early
to late beginning of harvest season): ‘Manila’, ‘Ataulfo’, ‘Kent’, ‘Haden’,
‘Tommy Atkins’ and ‘Keitt’ (Table 13.3). ‘Zill’, ‘Irwin’, ‘Sensation’, ‘Diplomático’,
‘Manililla’, ‘Oro’, ‘Ovo’ and criollos are also grown. Mango harvest seasons
are no longer inflexible as they may be modified by pruning, water manage-
ment and growth regulators. The most widely planted cultivar in Mexico is
‘Manila’ (45,396 ha); no export figures have been reported. Although ‘Manila’
is grown all over the country it dominates in Veracruz (24,908 ha) and
Guerrero (9198 ha). It is an alternate bearer and the harvest season is from
May to June in Veracruz and July in Sinaloa (De los Santos and Mosqueda-
Vázquez, 1988–89; C. Guzmán, personal communication). There are 22,890
ha of ‘Tommy Atkins’ and it is the major export cultivar to the USA (33% of
total exports). Fruit is harvested from February (Michoacán) to August
(Colima, Jalisco and Sinaloa) (V. Medina and C. Guzmán, personal commu-
nications). ‘Kent’ is cultivated on 13,366 ha and accounts for 23% of exports
to the USA. Sinaloa is the major producer of ‘Kent’ (9710 ha) and the begin-
ning of harvest season is similar to ‘Tommy Atkins’ (Campbell, 1992); however,
it can be harvested as late as September in Colima, Jalisco and Nayarit.
‘Ataulfo’ is from Chiapas and is now cultivated nationwide on >34,000
ha. The major producing states are Chiapas (south) and Nayarit (north) with
18,334 ha and 7156 ha, respectively. The harvest season is February–May in
Chiapas and June–July in Sinaloa (V. Palacio and C. Guzmán, personal com-
munications). In some years this cultivar is the third most important export
to the USA, and accounted for 19% of exports in 2006 (EMEX, 2007). ‘Keitt’
(6828 ha) is the fourth most important export cultivar (14% exports). Most
‘Keitt’ orchards are located in Sinaloa (5198 ha). ‘Keitt’ is harvested from May
(Nayarit) to September (Colima, Jalisco and Sinaloa). Large fruit size and
fungal lesions sometimes cause marketing problems because harvest occurs
during the rainy season. ‘Haden’ (27,768 ha) is still an important mango
although exports to the USA have decreased to 11% (EMEX, 2007); the major
producing states are Michoacán (15,573 ha), Guerrero (5053 ha) and
Sinaloa (3518 ha). Fruit are harvested from February/March (Michoacán and
Guerrero) to August/September in Colima, Jalisco and Michoacán. Alternate
bearing and mango malformation are problems and ‘Haden’ orchards are
being replaced by or topworked to cultivars like ‘Ataulfo’.
The mango was introduced into Taiwan from South-east Asia by the
Dutch in the 16th century (Chen, 1991). The introductions were all polyem-
bryonic and included ‘Tsar-swain’, ‘Pung-swain’, ‘Va-swain’ and ‘Kee-gway-
swain’. ‘Tsar-swain’ is still very important (Table 13.3). From 1954 to 1973, 35
monoembryonic cultivars were introduced from the USA, and ‘Irwin’ and
‘Keitt’ have been major cultivars in Taiwan since 1960. ‘Jin-hwung’, which is
448 J.H. Crane et al.
derived from a chance seedling of ‘White’ and ‘Keitt’, was selected in 1980.
‘Tainong No. 1’ and ‘Tainong No. 2’ are derived from controlled pollinations
and were released in 1985 by the Fengshan Tropical Horticultural Experiment
Station, Taiwan Agricultural Research Institute. Only ‘Tsar-swain’, ‘Jin-hwung’,
‘Irwin’ and ‘Keitt’ and ‘Tainong No. 1’ are commercially important today.
Fruit in the southern prefectures or in lower elevation orchards are har-
vested earlier than the northern prefectures or orchards at higher elevations.
‘Tsar-swain’ comprises about 40% of total production; its season is from
March to August, depending on the prefecture or location of the orchard. The
fruit of ‘Tsar-swain’ is 154 g, 100 mm long and 64 mm wide, total soluble
solids (TSS) value is 17° Brix, total titratable acidity 2.2% and seed/pulp
weight ratio 0.84 g. ‘Irwin’ comprises 40% of the production and is harvested
from Pingtung in April and in Tainan in August. ‘Jin-hwung’ comprises
about 10% of production and is harvested from June to September. The fruit
of ‘Jin-hwung’ is 965 g, 144 mm long and 99 mm wide, TSS value is 15° Brix,
total titrateable acidity 2.3% and seed/pulp weight ratio 0.94 g. ‘Keitt’ makes
up c.5% of production and is harvested from August to October. The fruit of
‘Tainong No. 1’ is 237 g, 99 mm long and 69 mm wide, TSS value is 20° Brix,
total titrateable acidity 4.0% and seed/pulp weight ratio 0.85 g.
The major cultivars in Florida are ‘Keitt’ and ‘Tommy Atkins’, which
account for c.70% and 20% of the hectarage, respectively (Table 13.3). Small
commercial hectarages of ‘Van Dyke’, ‘Palmer’, ‘Irwin’, ‘Raposa’ and ‘Kent’
are also grown. In Puerto Rico, the major export cultivars are ‘Keitt’, which
makes up c.60% of the hectarage, ‘Parvin’ (20%), ‘Irwin’ (10%), ‘Tommy
Atkins’ (5%) and ‘Haden’ (< 5%). Other cultivars (i.e. ‘Davis-Haden’, ‘Palmer’,
‘Kent’, ‘Mayaguezano’, ‘Poste’ and ‘Cubano’) are grown on a small scale. The
local cultivars are grown for the domestic market (Toro, 1988). The commer-
cial hectarage of California is mostly ‘Keitt’, although other cultivars have
been evaluated (Scott, 1990; Linden, 2006).
Immature ‘Keitt’ is the major early season cultivar (picked green) and
mature ‘Tommy Atkins’ is the major early season cultivar in Florida (Table 13.3)
(J.H. Crane, personal communication). Mature ‘Keitt’ and ‘Kent’ mangoes
are the major late season cultivars. Six- to 8-year-old ‘Tommy Atkins’ trees
produce 75–150 kg/tree and older trees may produce up to 300 kg/tree.
Internal breakdown varies from year to year and may be aggravated by
over-fertilization with nitrogen (N). Fruit are considered moderately
resistant to anthracnose. The harvest season is June through to August.
‘Keitt’ trees are precocious and produce large crops regularly during July
through to September.
‘Palmer’, ‘Irwin’ and ‘Van Dyke’ are grown commercially on a limited
scale in Florida and Puerto Rico (except ‘Van Dyke’). In Florida, ‘Palmer’ is
harvested from July to early September, ‘Irwin’ from June to early July, and
‘Van Dyke’ from July to August. In the recent past, ‘Bailey’s Marvel’, ‘Brooks’
and ‘Haden’ were grown commercially; however, their importance has
declined due to natural disasters and susceptibility to anthracnose. ‘Haden’
is no longer grown commercially in Florida (Crane and Campbell, 1991;
Campbell, 1992) but, ‘Glenn’, a seedling of ‘Haden’, has been recommended
Crop Production: Management 449
Plant spacing and density are influenced by climate, soil type and depth,
rootstock and scion vigour, growth habit and the targeted tree size. Cultural
practices including tree-size control, fertilizer and irrigation availability,
methods, rates, timing and frequencies; current technology and the necessity
for orchard access by farm machinery also influence plant spacing and con-
figuration. The cost of borrowed capital, land, water, irrigation, orchard
maintenance and net returns will dictate what cultivars are grown and how
they are managed (see Evans and Mendoza, Chapter 16, this volume).
Initial plant spacing and density should be planned to maximize early
yield and returns from young orchards and to maintain high yields from
mature orchards. Trees in overcrowded orchards compete for water, nutri-
ents and light and eventually lose production in the lower canopy. The effi-
cacy of foliar sprays (nutrients, pesticides and growth regulators) is reduced
and harvest is more difficult in crowded orchards.
Plant spacing among trees and rows has decreased in recent years to
optimize returns on investments in land, equipment and orchard infrastruc-
ture. This has been possible because of more insight into the physiology of
mango trees, improvements in irrigation system design and efficiency, and
better fertilizer delivery systems and pruning (i.e. intense hand and/or
mechanical pruning) and the use of plant growth regulators. Closer plant
spacings require more tree-size control and expertise on the part of the pro-
ducer. Various training systems are advocated for new trees to force complex
branching and increase bearing surface volume and fruit production poten-
tial (Fig. 13.2). Topping and hedging and/or hand pruning (selective prun-
ing and/or heading back) is used to maintain mature tree size and fruit
production and improve fruit colour (Fig. 13.3).
In Brazil, the traditional spacing of 10 m × 10 m in a rectangular or qua-
dratic format, with a density of 100 plants/ha, has been replaced by plant-
ings of 8 m (between rows) × 5 m (within rows) to 5 m × 5 m with higher plant
densities, which vary from 250 to 400 plants/ha (Cunha and Castro Neto,
2000; Mouco et al., 2002). In general, growers use two types of pruning: for-
mation and production pruning (Albuquerque et al., 2002). Formation prun-
ing is used to establish the intial tree architecture and trees are pruned five
times for several years leaving c.243 branches prior to starting mango pro-
duction (Fig. 13.2). Pruning systems include cleaning, skirting of the lower
canopy, lateral, central and top-canopy pruning, and pruning to correct
poor canopy development and maintain adequate canopy after production
commences (Albuquerque et al., 2002).
450 J.H. Crane et al.
Fig. 13.2. Tree training system for young trees to increase branching and productive
canopy volume (Source: after Oosthuyse, 1995; Campbell and Wasielewski, 2000).
Fig. 13.3. Various mechanical topping and hedging schemes that may include
selective hand pruning or shoot tipping to maintain tree size and regular bearing.
(Mouco et al., 2002). Windbreaks, consisting of pine trees, elephant grass and
three rows of banana plants, are often used during the first 2 years of orchard
establishment.
In Mexico, orchards were originally established at 10 m × 10 m to 16 m ×
16 m, and trees were not pruned, which resulted in enormous and very pro-
ductive trees; however, care and harvest from very large trees is problematic.
Plant spacing is based on cultivar vigour (‘Ataulfo’ and ‘Manila’ are most
vigorous), land slope (wider spaces as slope increases), farm machinery, cli-
matic conditions and soil fertility (wider distances for warmer climates and
more fertile soils) and water availability (Cruzaley-Sarabia et al., 2006). Irri-
gated orchards may handle closer spacings if pruning is practised. Under
rainfed conditions, soil moisture availability may have an important impact
on tree size. For example, 10-year-old orchards may have a few big trees
(c.69–100 trees/ha) or if trees are spaced more closely (300–600 trees/ha), tree
size is reduced.
Currently, both square and rectangular planting patterns are common in
Mexico although hexagonal planting systems are also used. For square
designs, tree spacing ranges from 5 m (400 trees/ha) to 10 m (100 trees/ha).
Rectangular planting systems have more options and the most common
arrangements are 6 m × 5 m (333 trees/ha), 8 m × 5 m (250 trees/ha), 8 m × 6 m
(205 trees/ha) and 10 m × 5 m (200 trees/ha) (Chávez-Contreras et al., 2001).
In Taiwan, plant spacing for monoembryonic cultivars (i.e. ‘Tsar-swain’)
ranges from 6 m × 6 m to 10 m × 10 m (100–256 trees/ha). In contrast, plant
spacing for polyembryonic cultivars ranges from 4 m × 5 m to 5 m × 6 m,
depending on the topography and soil fertility. Orchards on sloped lands
and infertile soils are less vigorous and are planted at higher densities than
orchards on lowlands and fertile soils.
Recommended plant spacings in Hawaii and Florida are similar. The
cooler climate of south-eastern California allows close spacing, i.e. 3 m
in-row by 5.4 m between-rows (617 trees/ha). Traditional plant spacings in
Florida were as high as 11 m in-rows and 11–14 m between-rows (64–82
trees/ha) (Young and Sauls, 1989). Plant spacings in Florida currently include
3.5 m × 6.1 m (468 trees/ha), 4.5 m × 6.1 m (364 trees/ha), 4.5 m × 7.6 m (292
trees/ha), 6.1 m × 6.1 m (268 trees/ha), 6.1 m × 7.6 m (215 trees/ha) and 7.6
m × 7.6 m (173 trees/ha) (Young and Sauls, 1989; Crane and Campbell, 1991).
In Puerto Rico, plant spacings are 3.7 m × 5.5 m (491 trees/ha), 4.6 m × 9.1 m
(238 trees/ha), 6.1 m × 9.1 m (180 trees/ha), 7.6 m × 7.6 m (173 trees/ha) and
9.1 m × 9.1 m (120 trees/ha) (Toro, 1988). Interplanting mango trees at moder-
ate to wide plant spacings (i.e. 5.4–7.5 m in-row) with banana or plantain
(Musa sp.), and papaya (Carica papaya L.) in Florida and plantains in Puerto
Rico is widely practised. Mango trees have also been planted at close spacing
(e.g. 3.0–4.5 m) and every other tree is removed if crowding becomes a prob-
lem. Controlling tree size and maintaining crop productivity is important,
otherwise, competition among trees will reduce yields and fruit quality (Toro,
1988; Crane and Campbell, 1991; Oosthuyse, 1995). Annual or biennial hand
pruning and/or mechanical hedging and topping is necessary and should
begin several years before trees begin to crowd and should continue after
452 J.H. Crane et al.
trees grow to their desired size based on plant spacing. Mature trees are
topped at 3.5–5 m, and hedged trees are cut on a slight angle (5–10°) to leave
a 2.5–3.5 m row middle (J.H. Crane, personal communication). Trees can be
maintained at in-row spacings of 2–3 m; however, this involves intense tree
training and hand pruning, which most producers have been unwilling to
adopt (Fig. 13.3) (Oothuyse, 1995; Oosthuyse and Jacobs, 1995; Stassen et al.,
1999; Campbell and Wasielewski, 2000). Severe hedging is utilized to increase
light penetration and re-establish inner productive canopy but this can result
in little to no production in the following year. Some producers utilize a com-
bination of mechanical pruning followed by selective pruning to open the
inner canopy to light and air movement, improving fruit colour and reducing
disease pressure.
Table 13.4. Standard leaf nutrient content ranges for mature mango trees in Brazil,
Mexico, Taiwan and Florida USA.
Table 13.5. Soil phosphate and potash corrective fertilization rates based on soil
analysis in Brazil.
(a) Rate of P2O5 (kg/ha) application.
≤15 60 30 0
16–35 100 50 0
36–60 200 100 0
>60 280 140 0
Table 13.6. Quantity of P2O5 and K2O applications based on young tree plant age, and
P and K soil content of oxisoils for São Paulo and the Brazilian central regions.
0–1 30 0 0 0 0 40 0 0 0
1–2 60 160 120 80 0 80 40 0 0
2–3 120 240 160 100 0 160 120 80 40
3–4 160 320 240 120 0 240 180 120 80
(Table 13.5); however, the quantities of N, P and K for mango production are
based mainly on soil and leaf analysis. Leaves used for analyses are 6–8
months old, from the mid-canopy and from branches with fruits and from all
four sides of the canopy to reduce variation in analysis results. The proce-
dure for sampling leaves is: (i) divide the orchard into separate areas of no
more than 10 ha with trees of the same age and productivity and growing on
similar soil; (ii) collect healthy leaves from the middle of the tree canopy,
from the four cardinal points on normal branches with recently matured
leaves from the previous flush of growth, with leaves not less than 4 months
old. Remove four leaves per plant, from a total of 20 plants selected ran-
domly, and take the leaves prior to the application of nitrates or other foliar
fertilizers that are applied to break the dormancy of the floral buds.
There are two distinct periods of mango fertilization in Brazil: pre- and
postharvest fertilizations (Alves et al., 2002). In the preharvest fertilization of
non-irrigated orchards, P should be applied in a single dose, before flower-
ing, and incorporated with a medium-weight plough. At the beginning of the
rainy period 40% of the N and K should be applied and the remainder after
flowering, during fruit development. In irrigated orchards c.40% of the P
should be applied before flowering and 60% postharvest. For N, 50% is
applied preharvest (i.e. after the start of fruit set) and 50% postharvest. Potas-
sium applications should be distributed throughout the production cycle but
more during fruit development and c.25% postharvest. In São Paulo and cen-
tral Brazil c.40% and 20% of the N and K should be applied after harvest and
at the end of the rainy season (i.e. beginning of March), respectively.
When the productivity of orchards in north-east Brazil is <10 t/ha and
leaf N concentrations exceed 16 g/kg and the P and K concentrations in the
soil profile are >40 mg/dm3 and 0.45 cmol/dm3, respectively, application of
N, P and K is unnecessary (Table 13.7). On the other hand, if the expected
productivity is >50 t/ha and leaf N concentrations are <12 g/kg and the P
and K concentrations in the soil profile are <10 mg/dm3 and <0.16 cmol/
dm3, respectively, 120 kg/ha of N, 150 kg/ha of phosphate (P2O5) and 250 kg/
ha of K2O should be applied.
Table 13.7. Quantity of N, P2O5 and K2O applications based on fruit productivity, leaf N content, and P and K soil content for the semi-arid
N leaf content (g/kg) Soil P content (mg/dm3) K soil content (mmolc dm-3)
Fruit <12 12–14 14–16 >16 <10 10–20 21–40 >40 <1.6 1.6–3.0 3.0–4.5 >4.5
production
(t/ha) N application rate (kg/ha) P2O5 application rate (kg/ha) K2O application rate (kg/ha)
15–20 60 40 20 0 45 30 15 0 80 40 20 0
20–30 75 50 25 0 65 45 20 0 120 60 30 0
30–40 90 60 30 0 85 60 30 0 160 80 45 0
455
456 J.H. Crane et al.
Table 13.8. Standard leaf nutrient content values for selected mango cultivars in Mexico
(Source: Guzmán-Estrada, 2001, 2004, 2006).
Cultivar
Deficiencies of zinc (Zn) and boron (B) are common in orchards in Brazil.
Zinc sulfate and borax are normally used to correct these deficiencies; the
rates depend upon leaf analyses (Silva et al., 2002). Lime is applied when base
saturation is <60%. Gypsum at a rate of 2 t/ha for sand and 3 t/ha for clay
soils is recommended to reduce the incidence of internal breakdown (Silva
et al., 2002). Gypsum (280 g/m2) incorporated at a 30 cm soil depth before
orchard establishment under Cerrados conditions (pH 3.7 and very poor Ca
levels) reduces internal breakdown from 60 to 3% in ‘Tommy Atkins’ (Pinto
et al., 1994).
Fertilization is not a common practice in most mango-producing regions
of Mexico. In Colima (Central Pacific region) <30% of orchards are fertilized.
However, in the Soconusco area of Chiapas (Southern Pacific region), 86% of
‘Ataulfo’ orchards are fertilized; 56.7% orchards are fertilized once a year,
39.5% twice a year, and 2.7% receive three applications per year. In fertilized
orchards, there is a high variation in amounts, materials, timing and forms of
fertilizer applications. Recommendations are empirical since there are few
published guidelines. Standard leaf nutrient levels have been proposed for
several cultivars grown in Sinaloa (Table 13.4, Table 13.8). Only a few growers
use leaf or soil analyses. Fertigation is rarely employed. In the Soconusco area
of Chiapas, ‘Ataulfo’ mango trees have K, Ca, Mg, Zn and B deficiencies. In
southern Sinaloa, most mango cultivars (‘Ataulfo’, ‘Haden’, ‘Tommy Atkins’,
‘Kent’, ‘Keitt’) are deficient in N, P, K, Mg, S, copper (Cu), Zn and B, normal in
Ca and iron (Fe) and high in manganese (Mn) (Guzmán-Estrada, 2001). In
Colima, where mango is cultivated on sandy soils with pH 7–8.4, Fe and Zn are
the most common deficiencies. In Nayarit, irrigated and non-irrigated ‘Haden’
and ‘Tommy Atkins’ trees are deficient in K, P and Ca (in this order) and have
excess Mg (Salazar-García et al., 1993). No Al toxicity has been reported.
Crop Production: Management 457
Leaf nutrient levels in mature mango trees on calcareous and sandy soils
have been investigated (Young and Koo, 1969, 1971; Koo and Young, 1972).
Significant variation in mineral content was due to cultivar, leaf age, position
of sample leaf on the twig, presence or absence of fruit and soil type. Ranges
of mineral nutrient levels for maintaining productive trees under Florida
conditions were developed (Table 13.4). Fertilizer frequency, rates and timing
in Florida are based on observation, leaf and soil nutrient content and experi-
ence. Leaves are sampled at least once a year between December and Febru-
ary; mature leaves are collected from at least 30 trees at 0.9–2.4 m and from
all sides of the trees, and before the trees have been sprayed with nutrients
(Young and Koo, 1971; Koo and Young, 1972; Young and Sauls, 1989). Sam-
ples should be taken for trees showing nutrient deficiencies, different culti-
vars and from groves under different cultural programmes and growing in
different soil types.
Nitrogen has the greatest effect on tree growth and yield and is used as
the basis for determining the amount of fertilizer to apply; however, some
caution must be used in recommendations from the past because of the
change from highly to less soluble fertilizer ingredients that has occurred
over the last 30 years (J.H. Crane, personal communication). The aim of the
programme for the first 2 or 3 years is to produce a strong, healthy canopy so
that when production begins, trees will bear regularly and heavily. Young
non-bearing trees (1–3-years-old) are fertilized with 100–225 g/tree of a
2–10% N and K (K2O) source (e.g. 4-2-8, 4-8-12, 8-2-8-2, 8-3-9; N-P2O5-
K2O-Mg) or similar material also containing P (P2O5) and Mg, at 6–8 week
intervals during the first year (Young, 1974; Young and Sauls, 1989). About
25–50% of N should be in an organic or slow-release form. The amount of
fertilizer is gradually increased (up to 1.4 kg application/tree) and the fre-
quency decreased (2–4 times/year) during years 2 to 4. Sources of N recom-
mended for mangoes in Florida soils include ammonium nitrate (NH4NO3)
and potassium nitrate (KNO3), some in a slow-release and/or organic form.
Urea is not recommended for calcareous soils because it volatilizes as ammo-
nia gas, but S-coated urea is used. Currently, low N analysis fertilizers are
recommended because excessive vegetative growth and increased fruit
physiological disorders (e.g. internal breakdown) are caused by moderate to
high N applications (Young and Miner, 1960, 1961; Nguyen et al., 2004). Cal-
cium applications may be necessary to raise the pH of acid sandy soils in
Crop Production: Management 459
trees are maintenance of tree health, avoidance of severe drought stress and
enhancement of vegetative dormancy. Mangoes are drought tolerant (Schaffer
et al., 1994); however, in some production areas under non-irrigated condi-
tions, fruit production and quality may be reduced. Excessive irrigation can
cause reduced tree growth and tree decline (Larson et al., 1989a, 1991d; J.H.
Crane, personal communication). Mango production has been displaced
onto marginal lands that possess low water and/or nutrient holding capacity
and high pH. Saline water for irrigation is a concern. Irrigation methods
include flood and furrow, high-pressure volume guns, high-volume under- and
over-tree sprinkler and low-pressure microsprinkler and drip systems.
In north-eastern Brazil, under semi-arid tropical conditions, irrigation is
essential throughout the hot and dry season (Albuquerque et al., 1999). Sev-
eral irrigation systems are used, with c.41% of orchards using microsprinkler
systems. About 21% use other types of irrigation (e.g. furrows, drip, basin,
etc.) and 33% of orchards use no irrigation (Gomes et al., 2002). Mean produc-
tivity of irrigated orchards may be as high as 25 t/ha compared with only 12
t/ha for non-irrigated orchards (Coelho et al., 2002). Irrigation is used on
14,500 ha (74% of the cultivated area) of commercial mango in the north-east.
Microsprinkler irrigation is the most common method (30.3% of the irrigated
area) (Gomes et al., 2002). Commercial mango orchards in the south-east,
mainly in São Paulo, are not irrigated.
Fertigated orchards require well-trained people. Some nutrients can
cause corrosion of irrigation pipes, and proper management is essential to
prevent environmental damage. Proper selection of nutrients is critical due
to problems of solubility and compatibility. Urea, (NH4)2SO4 and KNO3 are
the primary N fertilizers; they are highly soluble and are compatible with
most nutrients. However, the SO42– is incompatible with Ca and their mixture
causes precipitation and clogging of emitters.
The oldest commercial orchards in Mexico were rainfed and established
in areas with deep soils and annual rainfall >900 mm (subhumid and humid
tropics). However, growth of the mango industry has forced the planting of
semi-arid tropical areas (annual rainfall 600–700 mm) (Colima and Micho-
acán) (Table 13.2). Currently, 67% of orchards are rainfed (SIIAP, 2007) with
annual rainfalls that fluctuate from 900 to 3700 mm; 80% of the rainfall occurs
in June–October. Irrespective of the amount of annual rainfall, since the late
1980s a significant proportion of the new and old mango orchards (60,000 ha)
have installed irrigation systems. This is to prevent water deficits and to
increase yield and fruit size and quality. Sources of irrigation water include
rivers or deep wells and irrigation may be applied either by gravity or by
electrical or diesel engines.
Irrigation management involves either furrow or pressurized systems.
The furrow soil surface system (FSSS) can be simple, crossed, furrow-basin
and fish spine. Low-pressure drip and microsprinkler systems are most com-
mon, particularly the latter. Two microsprinklers (90–120 l/h) or 10–15 drip
emitters (8–12 l/h) are used per mature tree in a low-pressure system. In
semi-arid regions, mango potential evapotranspiration from October to June
each year is 16,000 m3/ha. The FSSS system applies >25,000 m3/ha of water;
462 J.H. Crane et al.
however, low-pressure systems can reduce water use to 5000 m3/ha with no
negative effect on yield or fruit quality (L.M. Tapia, personal communica-
tion). Water requirements in the semi-arid region of Michoacán are calculated
by using evapotranspiration measured with a Class A pan and multiplied by
the crop coefficient Kc, which for practical purposes is considered as 0.4 for
vegetative growth and 0.8 for fruit growth and development (Chávez-
Contreras et al., 2001). The FSSS water schedule for obtaining maximum
yield and fruit quality in Michoacán is: September (pre-bloom), one 20 cm
irrigation; October–December (bloom), two 10 cm irrigations spaced 20 days
apart; January–April (fruit growth), eight 10 cm irrigations at 15–17 day inter-
vals; May–July (vegetative growth), three 50 cm irrigations at 21–25 day inter-
vals. The amount of water used for drip irrigation in mature mango trees
during fruit growth and development is 1350 l/tree/week and for microsprin-
klers it is 1560 l/tree/week. After the first irrigation, growers water every 8
days in low-pressure irrigation systems and every 15–20 days in FSSS.
Soil-water content influences the frequency of flowering, the number
and length of panicles, yield and quality of mango fruits in Taiwan (Chang
and Lu, 1995). Two types of irrigation system are used: (i) furrow flooding in
the low lands; and (ii) microsprinklers, overhead sprinklers and drip (trickle)
irrigation on sloped lands. Microsprinklers have become the most important
irrigation system in recent years and fertigation is utilized by many produc-
ers. Irrigation management is based on producer experience and observation
of the soil, current weather and tree phenology.
In Florida, rainfall is not evenly distributed through the year, with the
dry season occurring during the autumn–winter (October–April). During the
wet spring–summer season (May–September), dry periods of 3 days or more
can occur. Most orchards in Florida are irrigated only during prolonged dry
periods. Several systems are common. High-volume overhead or under-tree
sprinklers which run at high pressures (28,121–70,303 kg/m2) and distribute
large amounts of water (0.51–0.89 cm/ha) and microsprinkler systems which
run at low pressure (7030–28,121 kg/m2) and distribute lower volumes of
water (37.9–1113.6 l/tree/ha). Some growers use both systems: the microsprin-
kler system for irrigation and fertigation and the high-volume system for
cold protection during freezing weather.
Well-established trees require little to no irrigation. During prolonged
drought conditions (>30 days with no significant rainfall), irrigation may be
applied. Irrigation is not recommended during the cool autumn and winter
months, to enhance the vegetative dormancy period induced by cool tem-
peratures, to enhance synchronization of apical stems and to intensify the
flowering response after growth commences.
Currently, irrigation recommendations and practices are based on obser-
vation and experience with c.6.35 cm water/application/ha. Preliminary
research suggests that irrigation at 7-day intervals during the period of fruit
development increases fruit size, earliness and yield (Larson et al., 1989b);
however, this has not been implemented by growers. Most Florida producers
rarely irrigate their mango orchards during the spring and summer as rain-
fall during this period coincides with fruit development and is sufficient for
Crop Production: Management 463
Mango trees require new vegetative growth in order to produce fruit each year.
The optimum temperature for vegetative growth is 24–30°C (Krishnamurthi
et al., 1961; Shü and Sheen, 1987; Whiley et al., 1989) and when levels of essen-
tial plant nutrients and water are not limiting, vigorous growth results.
Mature leaves and a period of cessation of vegetative growth (i.e. mature api-
cal and subtending meristems) are required for the transformation from veg-
etative to reproductive growth (Núñez-Elisea and Davenport, 1992; Kulkarni,
2004; Davenport, 2006).
Canopy management and reproductive manipulation vary according to
climatic conditions, cultivar and available technology (Davenport, 1993).
Ultimate mango-tree size depends upon the climactic and edaphic condi-
tions and cultural practices. Under optimum conditions, trees can reach
heights and canopy diameters of 30 m or more (Kostermans and Bompard,
1993); however, it is difficult to protect large trees from insects, diseases and
strong winds, and harvesting is difficult and costly. With the increase in costs
for orchard establishment and maintenance, the number of trees per unit
area has increased and tree size has decreased, whereas production of high
quality fruit per unit area has increased. Growers in some production areas
manipulate the period of flowering and fruit production. Tree size can be
controlled to maximize the number of trees per unit area and maintain
productive tree canopy and yields.
The natural period of mango production in Brazil was originally from
October to January. Production has increased from September through to
April through the use of precocious and late-bearing cultivars along with
floral induction techniques. There is potential that whole-year harvesting
and mango supply can be achieved, although there are some months with
low mango yields. There are three types of floral induction and their use
depends upon phenology and season. In general, the steps are: (i) pruning of
apical internodes after harvesting; (ii) application of paclobutrazol (PBZ) to
stimulate flowering by inhibiting gibberellin biosynthesis; (iii) spraying with
K2SO4 (2–2.5%) to increase carbohydrate levels in the tree; (iv) drought stress
to facilitate growth cessation; (v) applications of ethylene (optional); and
464 J.H. Crane et al.
Pacific region. In the Central Pacific region, ‘Haden’, ‘Manila’ and ‘Ataulfo’
trees are treated with one or two sprays of 2–4% KNO3 or 1–2% NH4NO3 at
any time during the first half of November. Similarly, ‘Haden’ and ‘Manila’
are treated during November with 8% KNO3 or 4% NH4NO3 in the Northern
Pacific region. ‘Tommy Atkins’ does not respond to foliar nitrate treatments
for promoting early flowering. This is due to delayed floral initiation in this
cultivar so that when nitrate treatments are applied the buds are not irrevers-
ibly committed to flowering (Pérez-Barraza et al., 2000). Consequently, veg-
etative growth is produced in response to treatments that stimulate bud break
(Pérez-Barraza et al., 2006a). However, application of PBZ promotes early
bloom in ‘Tommy Atkins’ (Salazar-García and Vázquez-Valdivia, 1997).
Currently, soil applications of Cultar® (25% a.i.) close to the tree trunk is
used for most cultivars in dosages that range from 1 ml/m canopy diameter
for ‘Manila’ in Veracruz to 2–4 ml for ‘Tommy Atkins’, ‘Haden’ and ‘Ataulfo’
in Michoacán, applied at 1–2 year intervals. The response to PBZ treatment is
enhanced by 30–45 days water stress and canopy sprays with nitrates
(Chávez-Contreras et al., 2001). In Nayarit and Sinaloa late bloom and harvest
are profitable because they are the last two production areas to be harvested
in Mexico. Two canopy sprays of 50 mg/l GA3 (15 and 30 November) cause
delayed bloom and shift 86% of the harvest to 1 month later (Pérez-Barraza
et al., 2006b).
In Taiwan, cultivar, latitude, elevation and cultural practices are used
for off-season production (Shü et al., 2000). Several strategies are utilized to
stimulate flowering and fruit set:
1. After harvest the last one or two vegetative flushes on each shoot are cut
back to control tree size. Subsequently, two flushes of healthy shoots are
allowed to grow to serve as fruiting shoots for the next year. Weak or crowd-
ed shoots are removed to facilitate ventilation and light penetration.
2. In general, flowering is not a problem in subtropical Taiwan; however,
poor fruit set due to bad weather and lack of pollinators occurs occasionally.
A recent programme to increase the population of pollinators, mainly the
greenbottle flies (Chrysomyia megacephala Fabricius) in mango orchards has
been very successful. Increased yield has been noted and the practice has
been exploited commercially throughout the island.
3. Most of the mango fruit harvest goes to the domestic market within a
short time period, causing the price to decline rapidly. Off-season fruit pro-
duction is thus very important to avoid this sharp drop in price.
4. Physical trunk damage by girdling and ringing and application of ethrel
is used to promote early flowering of the early season cultivar ‘Tsar-swain’
(Liu, 1996). Foliar applications of KNO3 can promote early flowering but
only if applied to flower-bud-induced shoots; PBZ is not recommended for
use on edible crops in Taiwan.
5. Panicle removal has been used to postpone flowering and fruiting (Shü
and Sheen, 1987; Shü, 1993). Emerging terminal panicles are removed by
hand. Chemical removal of terminal panicles with hydrogen cyanamide
(CH2N2) or calcium cyanamide (CaCN2) causes leaf damage and is not as
466 J.H. Crane et al.
effective as pruning (Hwang et al., 2004). Axillary panicle induction has some
advantages, because flowering can be timed to avoid frost or cold tempera-
tures or a period of excessive rainfall during the normal flowering period
(Singh et al., 1974; Shen and Huang, 1980). Axillary panicle removal also
reduces mango malformation and reduces alternate bearing (Majumder et al.,
1976; Pal and Chadha, 1982).
Control of flowering and tree size in the USA varies with respect to differ-
ent climatic, edaphic and soil conditions as well as cultural practices (Daven-
port, 1993). In Florida, cool temperatures during the winter months (December
through to February) are usually adequate to arrest vegetative growth and
induce flower bud differentiation. In some years, the duration of cool tempera-
tures prohibits floral expression until late winter/early spring (February/
March) and when continuous warm temperatures begin (March), profuse, syn-
chronized flowering occurs. In some years, warm and cool periods may occur
for a few days to weeks during the winter and partial flowering may occur 2–4
times during the winter. This prolongs the flowering and harvest season.
Flowering and fruiting of trees are not actively manipulated in Florida.
For young trees, vegetative growth is encouraged during the first 2–3 years
and panicles may be removed by hand or natural pathogens, i.e. powdery
mildew or anthracnose are allowed to kill panicles and flowers. Tipping and
selective pruning of young trees is recommended to improve tree structure,
control tree size and enhance early fruit production (Oosthuyse and Jacobs,
1995; Campbell and Wasielewski, 2000); however, selective pruning and
mechanical topping and hedging are used to control mature-tree size. Selec-
tive pruning usually involves removal of selected scaffold limbs to open up
the tree canopy to light and to remove dead wood. Mature trees may or may
not be allowed to grow together in the tree row to form hedgerows. Periodic
mechanical topping at 3.5–5 m and hedging to leave a 2.5–3.5 m row middle is
common (Crane and Campbell, 1991; J.H. Crane, personal communication).
Limiting the between-row spread of the trees to 2.5–3 m improves light pen-
etration into the tree canopy. In some orchards, hedging the inner sides of the
canopy of adjacent rows every 2–4 years and/or topping every third or fourth
row every 2–4 years is recommended. Trees are mechanically pruned immedi-
ately after harvest. Timing the pruning to selected rows each year ensures that
most of the planting will always be productive if continuous flushing of
pruned parts of the canopy prohibits reproductive growth the following
spring. Pruning trees shoots of 2–10 cm diameter immediately after harvest
and then tip pruning 3–4 times to force multiple lateral growths has been
advocated (Davenport, 2006). This strategy: (i) reduces or controls tree size;
(ii) shapes trees to facilitate subsequent tip pruning; (iii) synchronizes the veg-
etative flushing; and (iv) inhibits continuous vegetative flushing and prolongs
the period of vegetative dormancy. After vegetative growth has ceased for 5 or
more months, trees will synchronously flower when regrowth occurs.
In Puerto Rico, continuous vegetative growth of 1–2-year-old trees is
promoted. During this time panicles may be removed by hand to promote
vegetative growth, and this encourages rapid development of large trees.
Crop Production: Management 467
Early season flowering when cold temperatures and dry windy conditions
prevail (December–February) results in poor fruit set and abnormal fruit.
Therefore, pruning of mature trees just before April (early spring) delays
flowering and induces synchronous axillary flowering after the danger of
cold has passed.
mangoes are irrigated in low rainfall areas. In Veracruz and Chiapas most
orchards are adjacent to riverbanks and the deep soils provide root access to
the water table. No data are available supporting the benefit of irrigation in
medium-high precipitation areas.
Most mango-producing regions in Mexico are frost free. The only report
of frost damage to 2–4-year-old mango orchards was in Sonora where freez-
ing events may occur every 6–8 years (E. Sánchez, personal communication).
Low temperatures (≤10°C) during bloom reduce fruit set, especially in
‘Ataulfo’. Treatment with GA3 to delay the bloom has been suggested as a
method to avoid low-temperature damage during flowering but flowering
under high-temperature conditions is also detrimental to fruit set and crop
yield (Pérez-Barraza et al., 2006b).
In Taiwan, damage from typhoons occurs periodically and trees are reset
and either pruned to recover or replanted. In general, freezing temperatures
are not a problem in the mango-production areas but cool temperatures can
reduce fruit set. To avoid cool or cold temperatures during the normal flow-
ering period, shoot tips may be pruned to delay and force flowering from
lateral buds. Flooding in production areas is uncommon, and drought stress
is not an issue because most producers irrigate during dry periods.
The production areas of Hawaii and Puerto Rico are frost free; however,
Florida and California may experience temperatures at and below freezing
during the winter months (December through to February). In Florida, freez-
ing temperatures (0 to −6°C) may occur for a few hours for 1–4 nights/year,
although freezing temperatures for 13–15 h within a 24 h period have been
reported (Johnson, 1970; Campbell et al., 1977). In California, temperatures as
low as −6.6°C are common and frosts may occur 10–15 times/year during
January and February (Aslan et al., 1993). Mango trees do not acclimate to
cold temperatures (McKellar et al., 1983), although differences in cold toler-
ance and recovery from cold damage have been observed (Carmichael, 1958).
In Florida, high-volume overhead and under-tree irrigation is used to protect
trees during freezing weather. At least 0.6 cm of water/ha is distributed.
Overhead systems are designed for complete coverage (overlapping spray
patterns) of the trees and under-tree systems are designed to spray 0.9–2.4 m
into the tree canopy. Irrigation commences before freezing temperatures are
reached (usually c.2–3°C) and continues until ice has melted. These systems
are powered by diesel or gas engines as electrical power is unreliable during
freezing weather conditions. In California, microsprinklers, wind machines
and helicopters are used to raise the air temperature of plantings (Schacht,
1992).
Mango trees are relatively tolerant of wind stress (Schaffer et al., 1994;
Crane and Balerdi, 2005); however, newly planted trees are commonly staked
at planting in the calcareous soils of Florida to prevent damage to the bark
and cambium caused by constant movement and rubbing against the rocky
soil. Staking also stabilizes the tree against toppling during hurricanes. In
contrast, tolerance to mature trees depends on tree size, with larger trees
being more vulnerable to wind damage than pruned trees (Crane et al., 1993,
1994, 2001; Crane and Balerdi, 1996; NASS, 2006).
470 J.H. Crane et al.
Harvesting is done by hand in Brazil, Mexico, Taiwan and the USA (Evans,
2007) and is one of the most expensive operations in mango production because
fruit do not mature synchronously, and trees require multiple pickings. The
technology to mechanize mango harvest is difficult because of differences in
fruit colour, size and weight among cultivars, difficulty in determining fruit
maturity, requirement for multiple pickings, lack of tree-size-controlling
rootstocks and tree training and moderate to large tree canopies.
Prior to harvesting in Brazil, excessive set fruit and injured and diseased
fruit are removed, part of the rachis is removed to prevent scarring and bruis-
ing of the fruit, and some leaves that shade fruit may be removed to allow for
better peel colour development (Alves et al., 2002). Fruit in the lower part of
the canopy are harvested from the ground; however, ladders are required for
picking fruit high in the tree canopy. Peel damage due to latex exudation at
the stem end of the fruit during harvest is a very common problem, and
>50% of harvested fruit may be affected. This problem occurs mainly when
fruit are picked high in the canopy with a picking pole by severing the fruit
Crop Production: Management 471
near the stem end of the pedicel. An improved picking pole that cuts the
petiole c.2 cm above the pedicel reduces latex burn to <10%. Before trans-
porting to the packing house, fruit containers are placed in the shade to avoid
increasing the pulp temperature.
In Mexico, harvesting is done by hand when fruit have reached physio-
logical maturity. Picking poles with bamboo baskets (Gulf of Mexico and
Southern Pacific regions) or nylon net bags or cotton bags (Central and
Northern Pacific regions) and a cutting blade at the distal end are used to
reach fruit high in the canopy. Ladders may be used, although climbing trees
is more common. Fruit is usually placed in 20–30 kg wooden or plastic boxes.
The inner walls of the picking crates are covered by newspaper to absorb
latex and decrease sap peel damage.
Several fruit maturity indexes exist for mango: pulp TSS content and/or
acidity, pulp firmness, skin or pulp colour, calendar days or heat units from
bloom to harvest. No index alone is successful because of differences among
cultivars and significant variability in fruit maturity within trees and orchards
and among orchards. Pickers are trained to distinguish the following charac-
teristics: (i) size, form and fruit colour; (ii) shoulder development (higher
than the base of peduncle); (iii) cavity formation at the base of the peduncle;
and (iv) increased lenticel size (Chávez-Contreras et al., 2001). The Associa-
tion of Mango Packers and Exporters (EMEX; Empacadoras de Mango de
Exportación) have established some minimal physical and chemical stan-
dards to define maturity for ‘Haden’, ‘Tommy Atkins’, ‘Kent’, ‘Keitt’ and
‘Ataulfo’. A photographic maturity index chart is used by growers, pickers,
companies and packing houses. Mango fruit-peel damage by latex is common
when tree sap pressure is high. To minimize this damage, irrigation is stopped
at least 2 weeks prior to harvest; fruit are also picked with a long peduncle
and are washed immediately after the harvest bin is full.
Mature-green mangoes are the first fruit to be harvested and all the cri-
ollo, Oro and Florida cultivars fit this category. These mangoes are for the
domestic market and usually sell for high prices. ‘Manila’ mangoes are
picked when their colour changes from pale green to greyish green. ‘Ataulfo’
is harvested when the green peel shows a yellow colour break. The Florida
mangoes are picked ‘mature-green’ and at colour break.
The harvest season in Taiwan begins in March with ‘Tsar-swain’ and
ends in September or October with ‘Keitt’. Depending on the cultivar and
market, fruits are harvested at 70–80% maturity. ‘Tsar-swain’ is harvested
either at the green stage for pickles or at a mature ripe stage for domestic
markets. ‘Irwin’ fruit are harvested at either 80% maturity for export or at
>90% maturity for local markets. Green mature fruits may be triggered to
ripen with calcium carbide, ethephon or ethrel at 30–40°C, depending on the
cultivar. Mature fruits are stored at at 8–12°C for several days to several
weeks. Fruit destined for export are disinfested using the vapour heat method;
the fruit core temperature must reach and be held at 46.5°C for 30 min.
There are two markets for mango producers in the USA: the ‘green’ mar-
ket for non-ripe fruit and the ‘tree-ripened’ market. The green and tree-ripened
markets are speciality, niche markets where the green fruit are used as a
472 J.H. Crane et al.
13.13 Conclusions
Differences in mango culture are due to the climatic and edaphic conditions,
available information and technology, and tradition in each production area.
The interaction of climate and cultural practices, for example irrigation, fer-
tilizer, pruning, etc., that are essential for optimizing crop yields and quality
is not completely understood. None the less, horticultural systems can be
developed and tested that can impact fruit production and quality. It is
important that area- and, in many cases, site- and cultivar-specific cultural
information and practices need to be developed to optimize production and
fruit quality. This chapter has addressed the current state of mango culture in
four different production areas, and has emphasized improvements that are
essential for a prosperous industry.
Ackowledgements
The Mexican co-author acknowledges the following INIFAP researchers,
based at several Research Stations (CE) and states: Ernesto Sánchez-Sánchez,
CE-Valle del Yaqui (Sonora); Camerino Guzmán-Estrada, CE-Sur de Sinaloa
(Sinaloa); R. Mosqueda-Vázquez (deceased) and Enrique N. Becerra-Leor,
CE-Cotaxtla (Veracruz); Fulgencio M. Tucuch-Cauich, CE-EDZNA (Campeche);
Crop Production: Management 473
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14 Postharvest Physiology
14.1 Introduction
Successful postharvest handling of mangoes requires knowledge of the post-
harvest physiology of the fruit and how the fruit physiology determines the
best handling practices to maintain and develop high fruit quality. For example,
mango, like banana, tomato and avocado, is a climacteric fruit, which means
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
484 (ed. R.E. Litz)
Postharvest Physiology 485
that it may be picked when mature but before ripening has commenced, and
subsequently ripened postharvest. As mango fruit mature on the tree and
begin to ripen, eating quality improves, but potential marketable life de-
creases due to the difficulty of controlling the ripening changes once they
have been initiated, increased bruising susceptibility and increased decay.
Susceptible mango cultivars tend to develop more internal breakdown (jelly
seed, soft nose and stem-end cavity) the longer that harvesting is delayed
(Raymond et al., 1998; see Galán Saúco, Chapter 9, this volume). As a tropical
species, mangoes are subject to chilling injury (CI), which limits the use of
refrigeration to maintain postharvest quality. Mangoes are also subject to
other physiological disorders, physical damage and decay, the symptoms of
which may make the fruit unmarketable (Yahia et al., 2006a).
Mangoes harvested at a mature but unripe stage of development (‘mature-
green’) can be stored in the unripe state as long as the initiation of ethylene
production and hence ripening is avoided. The initiation of ripening can be
avoided by prompt cooling and storage at a low temperature at which ripen-
ing does not occur or, more effectively, by changing the composition of the
storage atmosphere so that the oxygen (O2) level is reduced and carbon diox-
ide (CO2) level is raised. This latter approach is called either modified atmo-
sphere (MA) or controlled atmosphere (CA) storage, depending on the degree
of control. These technologies slow fruit metabolism and specifically inhibit
the initiation of ethylene production. With MA or CA transport or storage,
mangoes can typically be maintained in a firm, green condition for several
days longer than can be achieved with normal refrigerated air storage. How-
ever, there are limits to the levels of O2 and CO2 that can be tolerated by
mangoes and these limits are affected by several factors, including cultivar,
maturity or ripeness stage, storage temperature and storage time (Yahia,
1998).
Mango postharvest physiology and technology have been described in
previous reports, book chapters and reviews (Subramanyam et al., 1975; Lak-
shminarayana, 1980; Ledger, 1986; Peacock, 1986; Lizada, 1991; Coates and
Johnson, 1993; Johnson and Coates, 1993; Lizada, 1993; Heather, 1994; Jacobi
et al., 1994; Johnson et al., 1997; Mitra and Baldwin, 1997; Tharanathan et al.,
2006).
Consumers are becoming aware of the nutritional and health benefits of fresh
fruits and vegetables. Mango fruit are a rich source of vitamin C (Table 14.1),
although the content decreases during ripening (Thomas, 1975; Vinci et al.,
1995). ‘Raspuri’ mango is rich in vitamin C (300 mg/100 g fresh fruit) during
the early stages of development, but the concentration is less (39.1–69.5
mg/100 g) at maturity (Siddappa and Bhatia, 1954). The content of vitamin C
was between 13 and 178 mg/100 g in the ripe fruit of 50 cultivars surveyed
by Singh (1960). The vitamin C content in fully grown mango fruit of culti-
vars in Puerto Rico ranged between 6 and 63 mg/100 g (Iguina de George
486 J.K. Brecht and E.M. Yahia
Table 14.1. Composition of the edible portion of mango fruit (Source: USDA/ARS,
2007).
Water g 81.71
Energy kcal 65
Energy kJ 272
Protein g 0.51
Total lipid (fat) g 0.27
Ash g 0.50
Carbohydrate, by difference g 17.00
Fibre, total dietary g 1.8
Sugars, total g 14.80
Minerals
Calcium mg 10
Iron mg 0.13
Magnesium mg 9
Phosphorus mg 11
Potassium mg 156
Sodium mg 2
Zinc mg 0.04
Copper mg 0.110
Manganese mg 0.027
Selenium Pg 0.6
Vitamins
Vitamin C (total ascorbic acid) mg 27.7
Thiamine mg 0.058
Riboflavin mg 0.057
Niacin mg 0.584
Pantothenic acid mg 0.160
Vitamin B6 mg 0.134
Folate, total Pg 14
Folic acid Pg 0
Folate, food Pg 14
Vitamin B12 Pg 0.00
Vitamin A IU 765
Retinol Pg 0
Vitamin E (D-tocopherol) mg 1.12
Vitamin K (phylloquinone) Pg 4.2
Lipids
Fatty acids, total saturated g 0.066
4:0 g 0.000
6:0 g 0.000
8:0 g 0.000
10:0 g 0.000
12:0 g 0.001
14:0 g 0.009
16:0 g 0.052
18:0 g 0.003
Fatty acids, total monounsaturated g 0.101
(Continued)
Postharvest Physiology 487
et al., 1969). Vitamin C content was 105.2, 65.7 and 17.3 mg/100 g in ‘Langra’,
‘Ashwini’ and ‘Fazli’ mangoes, respectively (Gofur et al., 1994), and decreased
rapidly 5–7 weeks after fruit set, and when ripe fruit were stored at room
temperature. Vitamin B1 (thiamine) in two mango cultivars was 35–60 Pg/100 g,
488 J.K. Brecht and E.M. Yahia
3.5
All-trans-violaxanthin
3.0 9-cis-Violaxanthin
All-trans-β-carotene
Pulp carotenoid content (mg/100 g)
2.5
2.0
1.5
1.0
0.5
0.0
‘Haden’ ‘Ataulfo’ ‘Tommy ‘Manila’ ‘Criollo’ ‘Kent’ ‘Paraíso’
Atkins’
Cultivar
Fig. 14.1. Content of selected carotenoids in pulp of several mango cultivars. Data
represent the mean of eight individual observations for each cultivar ± standard error
(Source: Ornelas-Paz et al., 2007).
490 J.K. Brecht and E.M. Yahia
600
500
α-Tocopherol (μg/100 g)
400
300
200
100
0
‘Haden’ ‘Ataulfo’ ‘Tommy ‘Manila’ ‘Criollo’ ‘Kent’ ‘Paraíso’
Atkins’
Cultivar
Fig. 14.2. The content of D-tocopherol in the pulp of several mango cultivars. Data
represent the mean of eight individual observations for each cultivar ± standard error
(Source: Ornelas-Paz et al., 2008).
Postharvest Physiology 491
Climacteric behaviour
6 50
Hard green
5 Sprung green
40
Ethylene (mmol/kg/h)
Half ripe
CO2 (mmol/kg/h)
4 Ripe
30
3
20
2
Fig. 14.3. The climacteric pattern of respiration and ethylene production during mango fruit
ripening (Source: Lalel et al., 2003).
ethylene production occurs 110 days after flower initiation, and declines
as fruit approached full maturity (Cua and Lizada, 1990). The content of
1-aminocyclopropane-1-carboxylic acid (ACC), the immediate precursor of
ethylene, increases in different tissues (peel, outer and inner mesocarp) dur-
ing ripening in both cultivars, ‘Amrapali’ and ‘Deshehari’, while ACC oxi-
dase (ACO; EC 1.14.17.4), which catalyse the conversion of ACC to ethylene
and ethylene production, decline (Reddy and Srivastava, 1999). Fruit peel
has the highest levels of ethylene and ACO and less ACC accumulation than
the outer and inner mesocarp at the mature-green stage. The inner mesocarp
has less ACO activity and high ACC accumulation during the ripening pro-
cess compared to peel; levels in the outer mesocarp are intermediate between
those in the peel and inner mesocarp. Changes in the ability to convert ACC
to ethylene in the peel are not related to changes in ripening parameters in
the fruit pulp (Lederman et al., 1997). Mango seed also produces ethylene
(Reddy and Srivastava, 1999). Fruit slicing has no measurable effect on ethyl-
ene production in ‘Julia’ and ‘Graham’ mangoes (Allong et al., 2001).
Treatment of mango fruit with acetaldehyde or ethanol (0.1, 0.5 or 1%
ethanol or acetaldehyde vapour) has concentration-dependent inhibitory
effects on ethylene production (Burdon et al., 1996). Application of ACC to
acetaldehyde- or ethanol-treated fruit discs indicates that acetaldehyde can
completely eliminate increased ACO activity, whereas ethanol cannot (Bur-
don et al., 1996). Accordingly, Burdon et al. (1996) suggested that acetalde-
hyde can either inhibit ACO activity directly or prevent the increase in the
enzyme, thereby providing a possible mechanism for retarding fruit ripening.
Several important metabolic changes occur during the maturation and ripen-
ing of mangoes, and some of those are useful as maturity indices (Ketsa et al.,
1991). The ripening changes are irreversible senescence processes that are
related to degradation of organelles or changes in chemical constituents, and
thus relate to the quality and postharvest life of the fruit. Natural senescence
is aggravated and promoted by ethylene, mechanical injury and high tem-
perature, and can be delayed by low temperature, elimination of mechanical
damage and reduction of ethylene production.
Organic acids
Organic acids are important for respiratory activity and as flavour constitu-
ents. During maturation and ripening, mango fruit experience a substantial
loss of organic acids. The predominant acids in mature mango fruit are citric,
succinic, malic and tartaric acids; citric acid has the highest concentration
and tartaric acid the lowest (Shashirekha and Patwardhan, 1976; Sarker and
Muhsi, 1981; Medlicott and Thompson, 1985). Citric acid content increases
steadily during fruit development in ‘Irwin’, reaching a maximum at the
beginning of the endocarp-hardening period, and then decreases steadily
Postharvest Physiology 495
(Ito et al., 1997). In ‘Keitt’ the predominant organic acids are citric and malic
acids, but tartaric, oxalic, ascorbic, and D-ketoglutaric acids also are present,
and the initial loss in acidity is due to a substantial loss in citric acid and a
small loss in malic acid (Medlicott and Thompson, 1985). In ‘Badami’ man-
goes, citric acid is the major organic acid, but malic and succinic acids are
also present (Shashirekha and Patwardhan, 1976). In ‘Fazli’ mangoes, oxalic,
citric, malic, pyruvic and succinic acids have been detected and tartaric acid
has been detected in ‘Zardalu’ mangoes (Kumar et al., 1993). In general, citric
and succinic acids decrease during ripening while malic acid shows different
changes with different cultivars (Lizada, 1993).
Mango fruit contain organic acids involved in tricarboxylic acid cycle
reactions (i.e. oxalic, succinic, pyruvic, oxaloacetic and D-ketoglutaric acids).
In ‘Pairi’ mangoes, maximum concentration of D-oxoglutaric and pyruvic acids
occur before the climacteric peak. Aspartic and glutamic acid concentrations
increase for c.3 days after harvest and then decrease as the climacteric maxi-
mum is reached (Krishnamurthy et al., 1971). Malic enzyme (EC 1.1.1.40), which
catalyses the oxidative decarboxylation of L-malic to pyruvic acid, occurs in the
three-quarter-ripe and ripe stages and the activity pattern during ripening is
similar in ‘Alphonso’, ‘Banganpalli’, ‘Dasheri’, ‘Fazli’, ‘Langra’ and ‘Suvar-
narekha’ (Selvaraj and Kumar, 1994). In ‘Alfonso’ (sic), the levels of malic dehy-
drogenase (EC 1.1.1.37) and succinic dehydrogenase (EC 1.3.5.1) increase with
the onset of ripening; whereas, the level of citrate synthase (EC 2.3.3.1) increases
several-fold during maturation but decreases markedly during ripening (Baqui
et al., 1974). The activity of malic enzyme increases during ripening, reaching a
maximum immediately after the climacteric peak, and then declines (Dubery
et al., 1984). The activity patterns of phosphoenol pyruvate carboxylase (PEPC;
EC 4.1.1.49) and pyruvate decarboxylase (EC 4.1.1.1) during ripening vary
among different cultivars, while malic enzyme activity increases during ripen-
ing. PEPC activity is relatively high in ‘Alphonso’ and ‘Langara’, but low in
‘Dashehari’ and ‘Totapuri’ during ripening (Selvaraj and Kumar, 1994).
Soluble sugars
The increase in soluble sugars is a major change during mango fruit ripening,
and sweetness is the most important compositional change related to mango
flavour. While starch content increases in chloroplasts during mango fruit
development, it is almost completely hydrolysed to simple sugars during
ripening (Medlicott et al., 1986; Selvaraj et al., 1989; Kumar et al., 1994; Ito
et al., 1997). In ‘Alphonso’, starch content is 14% (by weight) at the immature
stage and c.0.3% at the ripe stage. Similarly, starch is almost undetectable in
‘Irwin’ mangoes after ripening, whereas sucrose increases significantly and
fructose increases slightly (Ito et al., 1997). Starch content decreases slightly
during ripening of ‘Haden’, but is insufficient to account for the observed
increase in the level of sucrose (Castrillo et al., 1992).
Ripe mango contains up to 10–20% total sugars on a fresh weight (FW)
basis, depending on the cultivar and the stage of ripeness. At the beginning
496 J.K. Brecht and E.M. Yahia
of ripening, reducing sugars make up most of the sugar content, while there
are more non-reducing (c.17%) than reducing (3%) sugars in completely ripe
fruit. Sucrose contributes 57% of the total sugar in ripe ‘Keitt’ mangoes, with
fructose and glucose making up 28% and 15%, respectively (Medlicott and
Thompson, 1985). Although Krishnamurthy et al. (1971), Lakshminarayana
(1973, 1975) and Shashirekha and Patwardhan (1976) reported a simultane-
ous increase of glucose, fructose and sucrose during ripening, Vazquez-Salinas
and Lakshminarayana (1985) observed a gradual reduction in glucose and
fructose and a continuous increase of sucrose during ripening in ‘Haden’,
‘Irwin’, ‘Kent’ and ‘Keitt’. Medlicott and Thompson (1985) and Vazquez-Salinas
and Lackshminarayana (1985) identified the main reducing sugar as fruc-
tose, while Selvaraj et al. (1989) reported that glucose is predominant. Con-
flicting reports on the relative concentrations of individual sugars in mango
fruit during ripening is cultivar-dependent and due to different storage and
handling conditions (Medlicott and Thompson, 1985).
Sucrose content increases during ripening as a result of starch hydrolysis
from increased amylase (EC 3.2.1.1) activity (Mattoo and Modi, 1969a; Fuchs
et al., 1980; Tandon and Kalra, 1983). The high activities of sucrose synthase (EC
2.4.1.13) and invertase (EC 3.2.1.26) in the mesocarp during ripening indicate
active sucrose metabolism (Kumar et al., 1994). Hexoses and hexose phosphates
can be formed from pyruvate by gluconeogenesis (Selvaraj and Kumar, 1994).
The activity of glucose-6-phosphatase (EC 3.1.3.9) reportedly increases up to
the three-quarter-ripe stage; whereas, fructose-1,6-diphosphatase (EC 3.1.3.11)
activity increases as the fruit ripens from the three-quarter-ripe to full-
ripe stage (Kumar and Selvaraj, 1990). The glycolytic enzyme hexokinase
(6-phosphofructokinase; EC 2.7.1.11) has maximum activity at the ripe stage,
while pyruvate kinase (EC 2.7.1.40) activity increases until the three-quarter-ripe
stage and declines at ripening (Selvaraj and Kumar, 1994). The pattern of
activity changes in hexokinase/phosphofructokinase and pyruvate kinase
demonstrates that glycolysis is activated during mango fruit ripening.
Reducing sugars, mainly fructose, increase slightly during ripening, and
sucrose synthase (EC 2.4.1.13) activity increases approximately ten times
during the phase of rapid sucrose accumulation (Castrillo et al., 1992). This
activity accounts for the maximum rate of sucrose synthesis. The proportion
of sucrose phosphate synthase (EC 2.4.1.14) activity that is sensitive to inhibi-
tion by inorganic phosphate changes during ripening (Castrillo et al., 1992).
Maximum catalytic activity of sucrose synthase is constant throughout the
ripening period and contributes significantly to sucrose metabolism. The
activities of neutral and acid invertases (EC 3.2.1.26) are very low in com-
parison with the other enzymes of sucrose synthesis. Acid invertase activity
increases and later decreases during ripening.
Structural polysaccharides
Pulp firmness is important for the evaluation of fruit maturity potential for
transport and storage, and as a quality characteristic. Fruit softening and cell
Postharvest Physiology 497
wall changes are principal changes associated with fruit ripening. Fruit tex-
ture changes are due to changes in cell walls and pectic substances in the
middle lamella, and these are cultivar-related (Selvaraj and Kumar, 1989).
Softening of mango fruit is characterized by increased solubility of cell wall
pectins (Roe and Bruemmer, 1981; Tandon and Kalra, 1984; Lazan et al., 1986;
Nasrijal, 1993). In general, water-soluble polysaccharides increase during
ripening (Lazan et al., 1986; Brinson et al., 1988), but water- and alkali-soluble
pectins decline in ‘Keitt’ mangoes, and ammonium oxalate-soluble pectins
increase as the fruit become soft (Roe and Bruemmer, 1981). There is an over-
all loss of galactosyl and deoxyhexosyl residues during mango fruit ripen-
ing, the latter indicating degradation of the pectin component of the wall
(Muda et al., 1995). The loss of galactose appears to be restricted to the chela-
tor soluble fraction of the wall pectin, while loss of deoxyhexose seems to be
more evenly distributed among the pectin.
Pectinesterase (PE; EC 3.1.1.11), which catalyses the deesterification of
methyl groups from acidic pectins, has been detected in ripening mangoes
(Tahir and Malik, 1977; Roe and Bruemmer, 1981; Ali et al., 1990, 1995; Abu-
Sarra and Abu-Goukh, 1992). Physiological maturity in ripened mangoes is
associated with lower PE activity (van Lelyveld and Smith, 1979) and peel
has higher PE activity than pulp (Ashraf et al., 1981). Endo-polygalacturonase
(PG; EC 3.2.1.15), which is responsible for degrading the 1-4-linked galactur-
onic acid residues, occurs in ripening fruit (Lazan et al., 1986, 1993; Abu-Sarra
and Abu-Goukh, 1992). Enzymatic and/or non-enzymatic processes, in addi-
tion to PG activity, are involved in the extensive softening of fruit (Mitcham
and McDonald, 1992). Other cell wall hydrolases can be detected in ripening
fruit, including cellulases (EC 3.2.1.4; Lazan et al., 1986; Abu-Sarra and Abu-
Goukh, 1992), E-galactosidase (EC 3.2.1.23; Ali et al., 1990, 1995; Lazan et al.,
1993), galactanase (EC 3.2.1.145; Ali et al., 1990) and xylanase (EC 3.2.1.8; Ali
et al., 1990).
Ripening in mangoes, as characterized by decreased tissue firmness, is
initiated in inner mesocarp tissue close to the seed, and progresses outwards
(Lazan et al., 1993). Pectin solubilization in inner and outer mesocarp tissues
is comparable, but pectin solubilization begins earlier in the inner than in the
outer mesocarp (Lazan et al., 1993). The outer mesocarp of ‘Keitt’ remains
firm longer than ‘Tommy Atkins’, and the inner is softer than the outer meso-
carp at each stage of ripening in both cultivars (Mitcham and McDonald,
1992). Cell wall neutral sugars, particularly arabinosyl, rhamnosyl and galac-
tosyl residues, decrease with ripening in both cultivars. ‘Keitt’ has more
loosely associated, chelator-soluble pectin, accumulates more soluble poly-
uronides and retains more total pectin at the ripe stage than ‘Tommy Atkins’.
Both cultivars have similar PG activity, which increases with ripening. The
amount and molecular weight (MW) of cell wall hemicellulose decreases
with ripening in both cultivars. Galactose is the only cell wall neutral sugar
to show a significant decrease during ripening of ‘Sensation’ mangoes (Sey-
mour et al., 1990). Losses of neutral sugars can be due to hydrolysis of galac-
tans and arabinogalactans by E-galactosidase having galactanase activity.
E-Galactosidase activity shows a parallel increase to tissue softening during
498 J.K. Brecht and E.M. Yahia
Mango skin colour is important for its role in the perception of overall qual-
ity (González-Aguilar et al., 2001) and can be important for determining the
appropriate maturity for harvesting (Cocozza et al., 2004; Jha et al., 2007), pro-
cessing (Mahayothee et al., 2004) and consumption (Cocozza et al., 2004; Jha
et al., 2007). The loss of green colour is an obvious sign of fruit ripening in
many mango cultivars. The development of the optimum skin colour usually
defines mango quality. Some mango cultivars retain green colour in ripe
fruit. Depending on the cultivar, skin colour can change from dark to olive-
green; sometimes reddish, orange-yellow or yellowish hues appear from the
base colour. Some cultivars develop a reddish blush, which has been attrib-
uted to anthocyanins. Colour changes in mango fruit are due to the disap-
pearance of chlorophyll and the appearance of other pigments (Fig. 14.4).
Chloroplasts are transformed to chromoplasts containing yellow or red pig-
ments (John et al., 1970; Lakshminarayana, 1980; Parikh et al., 1990; Lizada,
1993). Well-arranged grana and osmiophilic globules occur in chloroplasts of
cells in the peel of unripe mangoes (Parikh et al., 1990), and lose integrity
during ripening. Osmiophilic globules appear, indicating the transformation
Postharvest Physiology 499
15
5
8
LSD
(P = 0.05)
Anthocyanin
4
Anthocyanins (μg/cm2)
Chlorophylls (μg/cm2)
Carotenoids (μg/cm2)
10 6
3
LSD
Carotenoid (P = 0.05)
4
2
5
2
1 LSD
Chlorophyll (P = 0.05)
0 3 6 9 12 15
Storage time (days)
1998). However, John et al. (1970) detected 15, 14 and 17 carotenoids in ‘Bad-
ami’ mangoes at mature-green, partially ripe and fully ripe stages of fruit,
respectively. Variation with respect to pigment types and quantities is due to
cultivar differences, geography and climate, different maturity stages and
treatments after harvest; discrepancies in results are probably due to differ-
ent analytical procedures.
Mango skin colour can be used to estimate the content of all-trans-E-
carotene (Vázquez-Caicedo et al., 2004), the most important provitamin A
carotenoid (Wolf, 1984). Ornelas-Paz et al. (2007) demonstrated that the val-
ues of external and internal colour are similar in ‘Manila’ and ‘Ataulfo’ man-
goes (non-blushed) in contrast to blushed cultivars (‘Criollo’, ‘Paraíso’ and
‘Kent’). The carotenoids in fruit skin of some mango cultivars can be corre-
lated with some non-destructive colour measurements (Table 14.2; Figs.
14.5–14.8 (Ornelas-Paz et al., 2008).
The most abundant carotene of mango is all-trans-E-carotene, while the
most important xanthophylls are violaxanthin and its isomers (Wilberg and
Rodriguez-Amaya 1995; Chen et al., 2004). Mercadante et al. (1997) quantified
many cartenoids of ‘Keitt’ mangoes and concluded that the most predomi-
nant xanthophylls were all-trans-violaxanthin and 9-cis-violaxanthin, account-
ing for 38% and 18% of total carotenoid content, respectively, although other
xanthophylls are important in other cultivars (Ben-Amotz and Fishler, 1998;
Setiawan et al., 2001).
Modi and Reddy (1967) reported an increase during mango ripening of the
carotene precursors, mevalonic acid (MVA) and geraniol, with a concomitant
increase in carotene content. The geraniol concentration of unripe ‘Alphonso’
Table 14.2. Correlation coefficients (R) for the relationships between the content of the
main carotenoids in mesocarp and the internal/external colour values in ‘Ataulfo’ and ‘Manila’
mango fruit. The correlation analysis was performed using D = 0.5 (Source: Ornelas-Paz
et al., 2008).
space, where L* indicates lightness on a scale of 0 (black) to 100 (white), a* indicates chromaticity on a
green (–) to red (+) axis, and b* indicates chromaticity on a blue (–) to yellow (+) axis. Numerical values
of a* and b* were converted into chroma (C) and hue angle (h°), which represent colour purity and the
shade of colour, respectively.
40 Mesocarp Mesocarp Mesocarp
Pigment content
30
(10–3 g/kg)
20
10
Postharvest Physiology
0 5 10 15 20 25 30 40 45 50 55 60 60 65 70 75 80 85
30
(10–3 g/kg)
20
10
0
–5 0 5 10 15 20 25 30 35 40 45 60 62 64 66 68
a* value b* value L* value
Fig. 14.5. Relationships between the content of all-trans-E-carotene (▲), all-trans-violaxanthin (as dibutyrate, z), 9-cis-violaxanthin (as dibu-
tyrate, {) in mesocarp and the a*, b* and L* values, measured in mesocarp or peel of ‘Ataulfo’ mango fruit during ripening. Each point repre-
sents the mean of two independent measurements ± the standard error (vertical bars). The continuous line represents an exponential regression
(Source: Ornelas-Paz et al., 2008).
501
502 J.K. Brecht and E.M. Yahia
40
Mesocarp Mesocarp
Pigment content
(10–3 g/kg)
30
20
10
0
40 45 50 55 60 65 55 60 65 70 75 80 85
40 Peel Peel
Pigment content
(10–3 g/kg)
30
20
10
0
30 35 40 45 55 60 65 70 75 80 85 90 95
C* value h° value
mangoes varies from 0.5 to 3.0 Pmol with 0.0 to 0.5 Pmol MVA; in ripe mangoes
the corresponding levels are 5–10 and 1–5 Pmol, respectively. The increase in
free geraniol and MVA indicates that these compounds are dephosphorylated
during ripening. Acid phosphatase (EC 3.1.3.2) may regulate carotenogenesis
in ripe mangoes (Mattoo et al., 1968). Mangoes stored at low temperatures and
then ripened at room temperature fail to synthesize as much carotenoids as
fruit held at room temperature (Krishnamurthy and Subramanyam, 1973;
Thomas, 1975). Hot water treatments increase the colour intensity of the pulp
(Medlicott et al., 1986) and the peel (Esguerra and Lizada, 1990).
‘Tongdum’ mangoes, which ripen without changing colour, have three-
fold more chlorophyll and slightly more E-carotene in the peel and have higher
rates of ethylene production compared with ‘Nam Dok Mai’ mangoes, which
change from green to yellow upon ripening (Ketsa et al., 1999). Activities of
chlorophyllase (EC 3.1.1.14) and peroxidase (EC 1.11.1.7) in the peel of ripe
‘Tongdum’ fruit are about half of that in ‘Nam Dok Mai’ fruit. Changes in the
peel of ripe green mangoes are due to either or both a lower activity of chloro-
phyllase or peroxidase activity and are not a result of low ethylene production.
Phenolic compounds
30
(10–3 g/kg)
20
10
Postharvest Physiology
-5 0 5 10 15 20 25 20 30 40 50 60 50 55 60 65 70 75 80 85
40
Peel Peel Peel
Pigment content
30
(10–3 g/kg)
20
10
0
-10 -5 0 5 10 15 20 20 25 30 35 40 55 60 65 70 75
a* value b* value L* value
Fig. 14.7. Relationships between the content of all-trans-E-carotene (▲), all-trans-violaxanthin (as dibutyrate, z), 9-cis-violaxanthin
(as dibutyrate, {) in mesocarp and the a*, b* and L* values, measured in mesocarp or peel of ‘Manila’ mango fruit during ripening. Each point
represents the mean of two independent measurements ± the standard error (vertical bars). The continuous line represents an exponential or
second order polynomial regression (Source: Ornelas-Paz et al., 2008).
503
504 J.K. Brecht and E.M. Yahia
40
Mesocarp Mesocarp
Pigment content
30
(10–3 g/kg)
20
10
0
20 25 30 35 40 45 50 55 60 65 60 65 70 75 80 85 90 95
40
Peel Peel
Pigment content
30
(10–3 g/kg)
20
10
0
20 25 30 35 40 45 60 65 70 75 80 85 90 95 100 105 110
C* value h° value
1970). This is associated with loss of astringency (Selvaraj and Kumar, 1989).
The peel of mango fruit has a higher phenolic content than the pulp at all
stages of fruit development (Jain, 1961; Lakshminarayana et al., 1970).
Polyphenol oxidase (PPO; EC 1.14.18.1) catalyses the oxidation of mono-
and diphenols to o-quinones, which polymerize to produce brown pigments.
PPO activity increases slightly from harvest maturity to the half-ripe stage and
then declines in ‘Banganapalli’, ‘Dashehari’, ‘Fazli’ and ‘Langra’ mangoes, and
decreases in ‘Alphonso’, ‘Suvarnarekha’ and ‘Totapuri’ mangoes (Selvaraj and
Kumar, 1989). The PPO isolated from ‘Haden’ mango is active towards the
o-diphenolic compounds, showing higher activity in the presence of catechol,
followed by chlorogenic acid, but not with monophenols (Park et al., 1980).
Sugar changes are very important for organoleptic attributes in the mango
fruit. Fruit flavour is mostly a balance between the content of sugars and
organic acids (Medlicott and Thompson, 1985) as well as aromatic volatiles.
Kapse et al. (1989) determined that increasing TSS and decreasing acidity
Postharvest Physiology 505
Table 14.3. Characteristic aromas in ‘Alphonso’ and ‘Totapuri’ mangoes and their possible
chemical causes (Source: Bandyopadhyay, 1983).
mangoes, and at 8°C compared to 12°C for tree-ripe fruit. However, aroma
volatile levels in tree-ripe mangoes from 25 kPa CO2 are equal to or greater
than those in mature-green fruit treatments. Atmospheres that prolong
mango shelf life by slowing ripening processes can allow tree-ripe mangoes
to be stored or shipped without sacrificing their aroma quality.
Quality enhancement has been used to determine properties critical to fla-
vour acceptability of mangoes, and focus group interviews have been conducted
to determine sensory attributes important to the purchase and consumption
of mangoes (Malundo, 1996). Sugars and acids enhance perception of specific
flavour notes in mango, including aromatics (Malundo et al., 2001).
Water loss lowers fruit weight, resulting in shrivelling, and may further reduce
quality by causing poor colour development and uneven ripening. Water is lost
from mango fruit through stomata, lenticels and other openings. Relative
humidity (RH) inside the fruit is 100% and water is lost when RH in the envi-
ronment surrounding the fruit is <100%. Water loss is also greatly influenced by
temperature. With constant RH and air movement, water loss increases signifi-
cantly with any increase in temperature. Transpiration rate is influenced by cul-
tivar and ripeness stage. It is correlated with skin thickness, morphological
structure, epidermal cells and surface wax coating. For example, waxes usually
develop on the epidermis of fruit in the later stages of development and thus it
is common for fruit harvested early to shrivel faster compared with those har-
vested at a more advanced stage of development (Yahia et al., 2006a).
Heat injury
Mango is highly tolerant of heat (Yahia et al., 2000; Jacobi et al., 2001b). Man-
goes that are not stored in refrigerated conditions after harvest may be
exposed to extremely high ambient temperatures in many production areas.
This may lead to heat injury, especially if the fruit are exposed to >30°C for
>10 days, but injury can also occur more rapidly at higher temperatures. The
heat disinfestation treatments of mangoes that are required for insect quar-
antine security may injure fruit that are not fully mature (Jacobi and Giles,
1997; Jacobi et al., 2001a).
External symptoms of heat injury include lenticel spotting and skin
browning (‘scald’) with secondary disease development, while internal
symptoms include mesocarp browning, tissue cavitation and ‘starch spots’
(Jacobi and Wong, 1992; Jacobi and Giles, 1997; Mitcham and McDonald,
1997; Jacobi et al., 2001a, b). Ripening of heat-injured mangoes may also be
inhibited (Jacobi et al., 2001a, b).
the different cultivars and maturity stages of mangoes used, different atmo-
spheres implemented and lack of experimental controls. Optimum condition
for prolonged shipping or storage is reported to be 3–5 kPa O2 plus 5–10 kPa
CO2, which can delay ripening, but the benefits are not very significant. Use
of CA and MA would most likely be beneficial in delaying fruit ripening
during marine transport for 2 weeks or more.
Bender et al. (2000b) determined the tolerance of preclimacteric ‘Haden’
and ‘Tommy Atkins’ to reduced O2 levels for storage times in typical marine
shipments. They reported that mangoes can tolerate 3 kPa O2 for 2–3 weeks
at 12–15°C and that tolerance of low O2 decreases as mangoes ripen. All low
O2 treatments reduced mature-green mango respiration; however, elevated
ethanol production occurred in 2 and 3 kPa O2 storage, with the levels two to
threefold higher in ‘Tommy Atkins’ than in ‘Haden’. ‘Haden’ fruit at the
onset of the climacteric accumulated ethanol in 4 kPa O2 and produced 10–20
times more ethanol in 2 and 3 kPa O2 than preclimacteric fruit. There were no
visible injury symptoms, but off-flavour developed in mature-green fruit at 2
kPa O2 and in ripening-initiated fruit at 2 and 3 kPa O2. Ethanol production
was not affected by storage in 25 kPa CO2. Ethylene production was reduced
slightly by low O2; however, ‘Haden’ fruit also showed a residual inhibitory
effect on ethylene production at 2 or 3 kPa O2 storage, while ‘Tommy Atkins’
fruit stored in 2 kPa O2 produced a burst of ethylene upon transfer to air at
20°C. Fruit firmness, total sugars and starch levels did not differ among treat-
ments, but 2, 3 or 4 kPa O2 and 25 kPa CO2 maintained significantly higher
acidity than 5 kPa O2 or air. The epidermal ground colour responded differ-
ently to low O2 and high CO2 in the two cultivars. Only 2 kPa O2 maintained
‘Haden’ colour better than air, while all low O2 levels maintained ‘Tommy
Atkins’ colour better than air. High CO2 was more effective than low O2 in
maintaining ‘Haden’ colour, but had about the same effect as low O2 on
‘Tommy Atkins’.
Properly selected atmospheres, which prolong mango shelf life by slow-
ing ripening processes, can allow tree-ripe mangoes to be stored or shipped
without sacrificing their superior aroma. Mature-green and tree-ripe ‘Tommy
Atkins’ mangoes were stored for 21 days in air or in a CA (5 kPa O2 + 10 kPa
or 25 kPa CO2) at 12°C (mature-green) and at either 8 or 12°C (tree-ripe)
(Bender et al., 2000a). Tree-ripe mangoes produced much higher levels of all
aroma volatiles except hexanal than mature-green fruit after ripening for 2
days. Both mature-green and tree-ripe mangoes stored in 25 kPa CO2 had
lower terpene (especially p-cymene) and hexanal levels than those stored in
10 kPa CO2 and air-stored fruit. Acetaldehyde and ethanol levels were higher
in tree-ripe mangoes from 25 kPa CO2 than in those from 10 kPa CO2 or air
storage, especially at 8°C. Inhibition of volatile production by 25 kPa CO2
was greater in mature-green than in tree-ripe mangoes, and at 8°C com-
pared to 12°C for tree-ripe fruit. Aroma volatile levels in tree-ripe mangoes
from the 25 kPa CO2 treatment equalled or exceeded those in mature-green
fruit treatments.
Mangoes have high tolerance of short-term elevated CO2 atmospheres
(Yahia, 1998). Mangoes can tolerate CO2 atmospheres of up to 25 kPa for
510 J.K. Brecht and E.M. Yahia
2 weeks at 12°C (Bender et al., 2000b). High (25 kPa) CO2 inhibits ethylene
production, but increases ethanol production. Aroma volatiles are reduced
following 25 kPa CO2 treatment, while 10 kPa CO2, low O2 atmospheres and
storage temperature did not significantly influence production of terpene
hydrocarbons, which are characteristic of Florida-type mangoes. Mature-
green ‘Tommy Atkins’ mangoes can be stored for 21 days in CA (5 kPa O2 +
10 kPa or 25 kPa CO2) at 12°C, while tree-ripe fruit can be stored for 21 days
in the same atmospheres at either 8 or 12°C (Bender et al., 2000a).
The quality of ‘Keitt’ mangoes was evaluated during storage for 6 days
at 20°C in an extremely low O2 (LO) CA (approximately 0.3 kPa) before stor-
age in modified atmosphere packaging (MAP) made from three, low-density
polyethylene (LDPE) films with different gas permeability characteristics
(González-Aguilar et al., 1997). Both LO and MA treatments delayed the
losses of colour, weight and firmness. Fruit maintained good appearance
with a significant delay of ripening. Mangoes are very tolerant of LO treat-
ment; however, some MAP fruit developed a fermented taste after 10 and 20
days at 20°C. Short duration (6-day) storage of mangoes in LO did not other-
wise have any deleterious effect on fruit quality during subsequent storage
under MA or normal atmosphere. Properly selected atmospheres, which pro-
long mango shelf life by slowing ripening, permit fruit to be shipped without
sacrificing superior aroma.
Beaulieu and Lea (2003) studied ‘Keitt’ and ‘Palmer’ mangoes without
heat treatment to assess volatile and quality changes in stored fresh-cut man-
goes prepared from firm-ripe (FR) and soft-ripe (SR) fruit, and to assess what
effect MAP may have on cut fruit physiology, overall quality and volatile
retention or loss. Subjective appraisals of fresh-cut mangoes based on aroma
and cut edge or tissue damage indicated that most SR cubes are unmarket-
able by day 7 at 4°C. Both cultivars stored in MAP at 4°C had almost identical
O2 consumption, which is independent of ripeness. The CO2 and O2 concen-
trations measured for cubes stored in passive MAP indicated that the system
is inadequate to prevent potential anaerobic respiration after 7 days storage.
Semipermeable coatings
Some fruit coatings can create an internal MA within the fruit due to semi-
permeable restriction of O2 and CO2 movement in and out of the fruit. Bald-
win et al. (1999) tested two types of fruit coatings – polysaccharide-based and
carnauba wax-based – for their effect on external and internal mango fruit
atmospheres and quality factors during simulated commercial storage at 10
or 15°C with 90–99% RH, followed by simulated marketing conditions at
20°C and 56% RH. The coatings exhibited markedly different O2 permeabil-
ity characteristics under laboratory conditions. Polysaccharide coatings were
less permeable to respiratory gases (i.e. O2 and CO2) and more permeable to
514 J.K. Brecht and E.M. Yahia
water vapour compared to carnauba wax. When applied to fruit under simu-
lated commercial conditions, however, the differences between the coatings
with regards to their permeability to respiratory gases were much reduced,
most likely due to high RH during cold storage. Both coatings created a MA
within the fruit, reduced decay and improved appearance by imparting a sub-
tle shine; but only the polysaccharide coating delayed ripening and increased
concentrations of flavour volatiles. The carnauba wax coating significantly
reduced water loss compared to uncoated and polysaccharide-coating treat-
ments.
‘Julie’ mangoes treated with 0.75% w/v aqueous solution of Pro-long
semipermeable fruit coating (a mixture of sucrose esters of fatty acids and
sodium salt of carboxy methyl cellulose) and stored at 25°C and 85–95% RH
exhibited reduced weight loss, retarded ripening and increased storage life
(6 days longer) without evidence of adverse effects on quality (Dhalla and
Hanson, 1988). Treatment with 1.0% Pro-long could increase ethanol concen-
tration in the pulp. Treatment with Pro-long (0.8–2.4%) also delayed ripening
of ‘Haden’ (Carrillo-López et al., 1996).
Insecticidal CA
14.9 Conclusions
Mango fruit have the potential to develop extremely desirable texture, taste
and aroma that make this fruit highly appreciated and desirable. Strategies
used to extend mango shelf life are based on control of ripening, ethylene
action and ethylene production. Therefore, fruit are usually harvested at the
mature-green stage, prior to ripening initiation, and stored and transported
at low temperatures at or near the threshold for induction of chilling injury.
These practices result in poor quality, immature and chill-injured mangoes
on the market. Successful handling of ripening-initiated mangoes is prob-
lematic due to the fruit’s short shelf life and the increased incidence of inter-
nal breakdown that accompanies delayed harvests makes international
transport of ripening mangoes almost impossible. Consequently, the export
market for fresh mangoes, which expanded rapidly in the 1990s, has not con-
tinued its rapid expansion in recent years.
Future expansion of mango consumption will require an understanding
of mango postharvest physiology in order to overcome the problems of CI,
internal breakdown, and premature and uneven ripening. This may involve
increased transport of tree-ripe mangoes in CA-equipped marine containers
or in MAP. It may involve development of improved procedures for storage
and ripening to offer preconditioned, ripening-initiated, ready-to-eat man-
goes to consumers. It may also involve genetic transformation of mango to
manipulate the progression and uniformity of ripening and softening.
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15 Postharvest Technology and
Quarantine Treatments
15.1 Introduction
Postharvest handling of mangoes is the last phase (from the tree to mouth) of
an agribusiness venture. To optimize productivity, profitable uses for all
grades of fruit should be sought, and stable employment provided for key
skilled staff. Sustainable land and water management, and compliance with
health, safety and financial obligations to employees are also necessary.
Increasingly, Good Agricultural Practice (GAP) protocols need to be observed
(GAP, 2003; FFTC-GAP, 2007).
This chapter reviews the technology and quarantine treatments that have
been developed for the postharvest handling of mangoes. Peacock (1986), Led-
ger (1986, 1991b) and Opara and Nguyen (1999) have previously reviewed
postharvest handling of mangoes, while several others have reviewed spe-
cific aspects of the topic (Johnson and Coates, 1993; Heather, 1994; Jacobi
et al., 2001b; Singh et al., 2004; Yahia, 2006). Less than 10% of total world
mango production is exported. Export markets for mangoes have expanded
because of social changes and rising demand, increased international air
cargo space for some sectors and promotion of export fruit production in
developing countries (Procter and Cropley, 1994; FAOSTAT, 2008). Expan-
sion of production to meet the supply requirements of export and distant
domestic markets has been possible because of successful integrated man-
agement strategies and disinfestation technologies to control diseases and
insects (Johnson and Coates, 1993; Johnson and Heather, 1995; Ploetz, 2007),
and increased land availability due to deforestation and diversification away
from rice. Market development has also been facilitated through harmoniza-
tion of the rules of trade between nations and regions at global and near
global levels (WTO, 2008), agreement on pest and disease risk management
under the International Plant Protection Convention (IPPC), various bilateral
and regional Free Trade Agreements (FTA, 2007), and the global expansion of
supermarkets. Simultaneously, knowledge of and concerns about exotic pest
risks to domestic fruit production, socio-political concerns about chemical
residues on food, environmental management and labour conditions, and
rising production and marketing costs, have impinged upon market access
and stimulated international dialogue and research initiatives which address
these concerns (Buchanan, 1994; Gullino and Kuijpers, 1994; Ploetz, 2003,
2007).
Postharvest Technology 531
Strategies and procedures for horticultural market research have been out-
lined by Minnis (1993, 1994), Hall et al. (2001) and Kitinoja and Kader (2003).
Increasingly, a supply chain approach is being taken (Johnson and Hofman,
2004). Hofman and Ledger (2006) proposed that the supply chain approach
should be used to guide research and development and that there needs to be
a champion in the supply chain with significant influence and a desire for
improving chain status and performance. Key features are identification of
market demand, through-put, price and profit flows, seasonal fluctuations in
availability and demands, supply competitors (and their commodity statis-
tics), importer-buyer requirements and relationships, options for value-adding,
and consumer expectations and sales promotions. Point-of-sale transaction
data from supermarkets and other sales outlets in target markets can be an
invaluable source of intelligence and can be purchased from marketing infor-
mation specialists. Detailed supply chain assessments can guide options for
innovation and improvement (Johnson and Hofman, 2004).
The volume, grade and quality of product available for export requires
analysis in relation to buyer specifications, retail customer and provedore
preferences, technological and regulatory requirements for supplying the
market, and the production, packaging, cooling and transportation proto-
cols/options that are needed/available or specified under agreed codes of
practice (NRI, 2008a). Procter and Cropley (1994), Mahendra et al. (2002) and
the Natural Resources Institute (NRI) (2008b) provide some perspectives on
these issues.
Table 15.1. Typical phytosanitary requirements for mangoes for some countries.
Australia Approved treatment for fruit fly, area free of pulp weevil
(Sternochetus gravis (F.))
Canada No phytosanitary certificate required
China Phytosanitary certificate required. Field management measures for
specific pests of quarantine concern to China plus approved
disinfestation treatment for fruit flies
EU Phytosanitary certificate required
Indonesia Phytosanitary certificate required plus grown in area free of Queensland
and Mediterranean fruit fly
Japan Phytosanitary certificate required plus disinfestation schedule approved
for nominated mango cultivars and fruit fly species and inspection of
approved quantity of fruit (2–5%)
Korea Phytosanitary certificate required, combined with field surveys
Malaysia Must be free of seed weevil on inspection
New Zealand Phytosanitary certificate required plus approved disinfestation schedule
for nominated fruit fly species
Saudi Arabia Phytosanitary certificate required. Require destructive test of 2% of
consignment for seed weevil, or field survey verification of block freedom
Singapore No restrictions
United Arab Phytosanitary certificate required. Require destructive test of 2% of
emirates consignment for seed weevil, or field survey verification of block freedom
USA Phytosanitary certificate required plus disinfestation approved for
nominated mango cultivars and fruit fly species
a General requirements: Prior approval to import is required to access the market of many countries.
Packhouse and disinfestation treatment/facility inspections may be required by exporting and importing
regulatory authorities. Import permits may cover multiple importations but usually require renewal every
3–12 months. Phytosanitary certificates must be issued by a government authority. Fruit must be free
of soil and debris and packed in clean, new containers. Timber packaging and pallets will be subject to
additional requirements. Consignments found to contain quarantinable pests will be rejected, and either
re-exported or destroyed.
Personnel
QA systems and GAP protocols must encompass the human component of
an organization or business (Bunt and Piccone, 1994; Rolle, 2006; Sonneveld,
2006; FFTC-GAP, 2007). Market agents, exporters, farm/packhouse suppli-
ers, finance providers and transport personnel and companies need to be
selected and worked with from the quality perspective. Human resource
development, training and education have been of major significance in the
success of many industries.
Postharvest Technology 535
Maturity
Peacock (1986) considered that fruit maturity referred to its stage of ontog-
eny, with fruit of different maturities being at different stages of ontogeny.
Fully mature mango fruit are strictly those which have produced a fully
developed seed and which have reached their full physiological potential in
536 G.I. Johnson and P.J. Hofman
relation to size increase and dry matter accumulation within the constraints
of the growth environment. When fruit size and dry matter concentration
reach a plateau, climacteric fruit such as mango can undergo ripening, where
colour, texture, flavour and aroma may change (Watada et al., 1984). In these
fruit, a sharp rise, followed by a decline in respiration also accompanies the
transformation from not-ready-to eat (unripe), to edible (ripe), to senescent
(overripe). Ripening signals the completion of seed ontogeny, and encour-
ages dispersal of the seed by attracting vertebrate fructivores (Cipollini and
Stiles, 1992). If fruit are not harvested, maturation and ripening occur on the
tree. Ripe fruit fall to the ground, or are consumed by bats, primates, pha-
langers, birds or humans, either on the tree or after detachment.
Softening and sweetening of fruit flesh and colour changes can occur at
any stage of ontogeny, even in pea-sized fruit (Oosthuyse, 1995). Fruit drop
at any stage of development is preceded by these events, and the likelihood
of their occurrence increases as fruit size and dry matter levels approach
their maxima (Singh et al., 2004). Although the changes constitute some of
the components of ripening, they can only be regarded as such if the fruit
have attained physiological maturity, i.e. ‘the stage of development when a
plant or plant part will continue ontogeny even if detached’ (Watada et al.,
1984; Yashoda et al., 2006).
When fruit are removed from the tree several days before the onset of
ripening, they are initially hard and green. The fruit progressively soften,
change colour and develop aroma at a rate determined by cultivar, storage
environment and at-harvest maturity. Ideally, fruit are picked, treated, packed
and transported while hard-green, and arrive at retail markets at some pre-
determined stage of colour development (usually more yellow or red, than
green, and ‘sprung’, but still firm). The rate at which ripening will occur
under particular storage conditions depends upon the stage of ontogeny at
harvest. More mature fruit will ripen more rapidly than less mature fruit.
Accurately estimating when the fruit are ready for harvest is critical to
consistently meet customer expectations. This is called horticultural matu-
rity, and several criteria of horticultural maturity are possible. One is a legal
minimum or buyer-specified standard of maturity, which confirms that the
fruit would be acceptable for consumption or processing when ripe or ready
to eat for green-eating types. Maturity estimation often relies on visual or
calendar-based (days from flowering) assessment or in some cases the appli-
cation of a simple test (e.g. dry matter or flesh colour assessment). In more
exacting cases, more accurate estimates of horticultural maturity may be
required to assess product suitability for more stringent or narrower quality
specifications, as may be required for contract sales or sea-export consign-
ments.
In both cases, easy to assess harvest indices relying on visual, chemo-
sensory or fruit-age attributes are needed, and they must correlate with the
commercially relevant fruit characteristics measured in prescribed tests. Pea-
cock (1986), Harvey (1987) and Askar and Treptow (1993) reviewed methods
for assessing fruit maturity. In mango, dry matter, flesh colour, skin colour,
fruit shape, Brix, specific gravity or days from flowering or heat accumulation
Postharvest Technology 537
7
7 Early harvest
Mid-harvest
6 Late harvest
6
Flavour (1–9)
Flavour (1–9)
5
5
4
4
3
r² = 0.86** 3 r² = 0.83
Fig. 15.1. Relation between flesh colour (1 = white; 15 = very yellow) and accumulated
heat units (>10°C) with flavour of ripe ‘B74’ mango grown under Australian conditions.
**= Significant to P = 0.01. (Source: Hofman and Marques, unpublished data).
units (e.g. degree-days) have been used (Baker, 1986; Hofman and Ledger,
2006). In South Africa flesh colour is favoured, while in Australia, skin colour,
dry matter and accumulated heat units are considered as well (Fig. 15.1).
Using several maturity indicators will usually increase the accuracy of pre-
dicting the first acceptable harvest date. Information would be cultivar-
specific. In some cultivars there may be few reliable visible indicators of
maturity that allow picking of the most mature fruit on the tree, particularly
when flowering has occurred over many weeks. In these instances it is very
difficult for pickers to spot pick the more mature fruit in a cost-effective way.
Variation in maturity between fruit can also be influenced by where fruit
develop on the tree. In the cooler subtropical areas of the southern hemi-
sphere, fruit on the northern or western side mature more quickly than fruit
on the southern side (Hofman et al., 1995; Oosthuyse, 1995; Hofman et al.,
1998). In these situations, it may be more feasible for growers to map the
maturity of their blocks or orchards to identify groups of trees that have more
mature fruit, and/or identify parts of the tree (e.g. the northern/western side
in the southern hemisphere) that generally hold more mature fruit. Pickers
can then be instructed to pick all the fruit on a specified canopy position and
from specified areas of the orchard in order to harvest more mature fruit.
This can be more cost effective than selectively picking from individual trees.
New technologies such as portable near infrared spectroscopy (NIRS) units
may assist in non-destructively mapping maturity profiles across orchards
(Subedi et al., 2007).
The maturity of fruit at harvest is important for determining fruit quality
at out-turn in overseas markets (see 15.8 Pre- and Post-shipping Storage sec-
tion, this chapter). If fruit in a carton or pallet are of uneven maturity, it may
be impossible to find an effective storage regime(s) which will ensure good
quality of all fruit on arrival. One fruit of more advanced maturity in a box or
pallet can accelerate the ripening of other fruit, which can then arrive with
538 G.I. Johnson and P.J. Hofman
disease symptoms and with little or no shelf life. Variable maturity within
treatment lots can also adversely affect product quality after heat disinfesta-
tion (Jacobi et al., 1995).
On-farm record keeping and analysis of date of flowering, seasonal prod-
uct maturity, orchard management schedules, environmental data, transport
regimes, market destinations and out-turn problems may enable some pre-
diction of at-market quality and better selection of the appropriate destina-
tion market. Recent research is focusing on non-destructive methods that
could be used for checking fruit maturity in automated grading systems
(Joyce et al., 1993; Subedi et al., 2007). Improvements in product quality and
performance resulting from the effective use of such systems over several
seasons, can provide considerable competitive advantages when developing
buyer relationships or consumer ‘brand’/country-of-origin loyalty.
Skin colour can affect sales, with markets preferring colour (green, yellow,
orange, red blush) familiar to past purchase experience, known use and cul-
tivar knowledge, or ethnic-group preferences. Fruit position on the tree af-
fects red colour development, since sun exposure is important for anthocyanin
development. Likewise, bagging of fruit can decrease red and green skin
colour on ripe fruit (Hofman et al., 1997a). Nitrogen (N) can increase the pro-
portion of the ripe fruit with green skin (Fig. 15.2) by retaining skin chloro-
phyll (McKenzie, 1994; Nguyen, 2003; Nguyen et al., 2004; Bally, 2007). In
cultivars susceptible to green skin at ripeness, N fertilization rates should be
Postharvest Technology 539
40
30
20
Green colour (%)
10
2
1
0
0 75 150 300 Fol. 0 150 300 450 Fol. 0 150 300 450 Fol.
HG orchard LG1 orchard LG2 orchard
Total N application (g/tree)
Fig. 15.2. Percentage of green skin colour on ripe ‘Kensington Pride’ mango fruit
from trees in three different orchards (coded HG, LG1 and LG2) following application
of N (0–450 g/tree). (Source: H. Nguyen et al., 2004). Fol. = foliar N sprays to a total of
50 g/tree. Bars represent least significant difference (LSD) at 5%.
Table 15.2. Effects of leaf:fruit ratios on quality of ‘Kensington Pride’ mango fruit after ripen-
ing at 22°C (Source: Simmons et al., 1998).a
Disease
Leaf:fruit Fruit Dry Lenticel spotting
ratio mass (g) matter (%) (1–5) Severity (1–5) Incidence (%)
mechanism against foreign particles and infection entering through the len-
ticels (Bezuidenhout et al., 2005; du Plooy et al., 2006).
The severity of lenticel damage is often difficult to control, but strategies
for minimizing damage exist. There are cultivar differences in susceptibility
(Oosthuyse, 1999), possibly related to differences in lenticel, wax and/or
cuticle structure or composition (du Plooy et al., 2004). Strong negative cor-
relations have been found with maximum and minimum temperature and
Class A pan evaporation, and strong positive correlations with maximum
relative humidity (RH) and rain at harvest (Oosthuyse, 1998). These results
suggest that cool, humid and wet conditions around harvest increase the risk
of lenticel damage. Increased damage following excess irrigation during the
latter stages of fruit growth (Simmons, 1998) support the above conclusions.
Lenticel damage can also be more severe in larger fruit obtained from
branches with higher leaf:fruit ratios (Table 15.2), possibly because of greater
damage to the lenticels during fruit growth (Simmons et al., 1998).
Physiological disorders include a range of symptoms that affect shelf life and
marketability (Johnson et al., 1996). These generally result in either prema-
ture ripening of parts of the fruit (e.g. soft nose and jelly seed) or tissue break-
down (i.e. stem-end cavity) (Winston, 1986; Mead and Winston, 1991; Whiley,
1999) and tissue breakdown in ‘Keitt’ (Bally, 2007). These disorders are more
severe in more mature fruit (Young, 1957; Katrodia, 1989; Mead and Winston,
1991) and are often evident on the tree or after ripening without storage.
Mango disorders are affected by growing conditions (Young, 1957; Young
and Miner, 1961). Production away from the coast, and higher altitude and/
or lower temperature are associated with lower incidence of spongy tissue
(Subramanayam et al., 1980; Katrodia, 1989), and susceptibility to the disor-
der is also affected by rootstock (Joshi and Roy, 1985). Stem-end cavity ap-
pears to be more severe in wet conditions near harvest (Wainwright and
Postharvest Technology 541
Burbage, 1989; Mead and Winston, 1991). Incidence of spongy tissue has been
reduced by mulches that decrease radiated and reflected field heat (Katrodia
and Sheth, 1989). Severity of watery pulp breakdown in ‘Keitt’ is lower with
higher crop loads from similar size trees (Bally, 2007), possibly because of
smaller fruit size at high crop loads.
Several reports suggest links between low fruit Ca and mango disorders.
High leaf Ca has been related to reduced soft nose (Young and Miner, 1961)
and reduced stem-end cavity (Mead and Winston, 1991). Soil Ca applications
can reduce stem-end cavity incidence (Whiley, 1999), but these responses are
not always consistent. Applications of Ca to ‘Keitt’ from just before flowering
onwards did not increase leaf or fruit Ca during fruit growth or at commer-
cial harvest (Bally, 2007). However, soil characteristics may also affect responses
to soil Ca applications. In the sandy soils typical of Australian orchards,
and even in the heavier clay subtropical soils with low cation exchange
capacity, Ca can be rapidly removed from the top soil profile, resulting in
little long-term increase in soil solution Ca (Hofman and Mullen, 2005;
Bally, 2007). Regular (two/week), small applications are required to consis-
tently increase solution Ca (Hofman and Mullen, 2005). Other factors (e.g.
vegetative vigour and water status) can influence fruit Ca uptake (Hofman
and Smith, 1994).
Foliar Ca treatments have produced inconsistent effects, in some cases
having little or no effect on fruit Ca concentrations or quality (Singh et al.,
1987; Simmons et al., 1995), and in other instances reducing internal disor-
ders (Chitarra et al., 2001). Postharvest dips have also had mixed results, with
both positive (Wills et al., 1988; Singh et al., 2000) and nil or very small effects
(Joyce et al., 2001) reported. As a result there is little commercial use of foliar
or postharvest Ca treatments in mango. Future development of more labile
Ca formulations may provide more consistent results.
Other nutrients have also been associated with fruit quality (Hofman
and Smith, 1994). There are relatively few reports of potassium (K) and mag-
nesium (Mg) effects on mango fruit quality. Higher Ca and Mg, and a ten-
dency towards lower K in mango fruit, have been noted in fruit from orchards
with no incidence of soft nose, compared with fruit from orchards with high
incidence of the disorder (Burdon and Moore, 1991). Conversely, K levels are
positively correlated with disease resistance; Karunanayake (2007) reported
reductions in disease on mango fruit from trees receiving additional K.
Application of triple the recommended level of K significantly reduced stem
end rot caused by Lasiodiplodia theobromae, while the severity of anthracnose
was most reduced by application of the recommended level of K compared
to nil and triple rate treatments.
Excess N can reduce storage life and quality, and excessive levels cause
deterioration of avocado fruit quality (Wolstenholme, 2004) where its effect
may be mediated through vegetative:reproductive balance and crop load
(Hofman and Mullen, 2005). High N has been associated with increased dis-
orders in mango (Young and Miner, 1961; Mead and Winston, 1991), possibly
through the dilution effect of increased fruit size on Ca concentrations; how-
ever, soil N applications later during fruit growth do not affect watery pulp
542 G.I. Johnson and P.J. Hofman
breakdown in fruit ‘Keitt’ fruit (Bally, 2007). Heavy rain late during fruit
development can release soil N previously unavailable to trees due to low
soil moisture, potentially resulting in high fruit levels. Boron (B) deficiency
has been related to abnormal fruit development (Lahav and Whiley, 2002)
and increased fruit storage disorders (Yogaratnam and Johnson, 1982; Smith
et al., 1997). It may also be important in mango (Coetzer et al., 1991). The effect of
larger fruit size and maturity on shelf and storage life (Seymour et al., 1990) can
also be mediated through production factors influencing fruit set and leaf:fruit
ratio. Fruit position in the canopy may also play a role here (see above).
Weather conditions
Rain before harvest, and high RH and temperatures can increase disease lev-
els, fruit susceptibility to heat and brush damage, lenticel damage and reduce
storage life (Dodd et al., 1991a; Estrada et al., 1993; Prusky et al., 1993a, b; Cooke
and Johnson, 1994; Oosthuyse, 1998; Jacobi et al., 2001b). Disease risk predic-
tion based on the monitoring of environmental variables to determine fungi-
cide application frequency, can reduce pesticide residues (Fitzell and Peak,
1984; Fitzell et al., 1984; Peak et al., 1986; Dodd et al., 1991a, b, 1992; Prusky et al.,
1993b). Irrigation frequency and water availability for tree growth can signifi-
cantly impact postharvest diseases and disorders (Simmons, 1998).
Flavour is largely determined by sugars and volatiles in the ripe fruit, both of
which increase in more mature fruit. The aroma produced by ripening and
ripe mango can help attract customers, and provide some indication of fla-
vour development. In mango fruits, more than 280 different aroma volatile
compounds have been reported (Singh et al., 2004). Variation in the constitu-
ent aromatic compounds in mango cultivars results in aroma and flavour
diversity (MacLeod and Snyder, 1985; MacLeod et al., 1988; Torres et al., 2007).
The high fruit levels of D-terpinolene contribute to the characteristic flavour
of stronger-flavoured cultivars such as ‘Kensington Pride’ (Bartley and
Schwede, 1987; MacLeod et al., 1988). ‘Kensington Pride’ harvested at the
green-sprung stage have higher concentrations of total aroma volatiles com-
pared with fruit harvested at the hard-green or coloured stages (Lalel et al.,
2003b). Most of the glycosidally bound aroma compounds increase in the
pulp as the fruit matures, which contribute to improved flavour. During the
first 7 days of ripening, D-turpinolene is the major volatile compound, but in
the later stages of ripening ethyl octanoate dominates (Lalel et al., 2003a).
Timing
this chapter). Fruit water potential fluctuates diurnally, and can affect fruit
quality. The water potential of fruit at harvest can affect susceptibility to han-
dling, heat damage and product storage potential (Joyce and Patterson, 1994).
In hot weather, fruit should be harvested in the coolest part of the day to
reduce fruit overheating and energy requirements for postharvest cooling,
and to minimize worker discomfort. Harvest during rain can reduce fruit
quality (see Weather conditions section under 15.3 Preharvest Management,
this chapter).
Sapburn
Severing the stem from the fruit causes relatively large volumes of latex to
spurt or ooze from the cut stem. The sap is of low pH and high oil content
and can burn the surface of the fruit (Bagshaw and Brown, 1989). The oil frac-
tion contains terpinolene and resorcinols and is the fraction of the latex that
causes the damage. Skin damage is particularly severe with ‘Kensington
Pride’ (O’Hare, 1994), and less serious in Florida cultivars. In Pakistan,
‘Chausa’ is more susceptible than ‘Sindhri’ and ‘Dashehari’ (Maqbool et al.,
2007). O’Hare (1994) observed that latex levels are lower and less phytotoxic
in ‘Nam Doc Mai’, ‘Nang Klang Wun’, ‘Tong Dum’ and ‘Keow Savoey’ (0.16–
0.48 ml/fruit), than in ‘Kensington Pride’ (1.67 ml/fruit). The oil compo-
nent of the latex of Thai cultivars is much lower than that of ‘Kensington’
(O’Hare, 1994; Hassan, 2007). The concentrations and ratio of the two main
resorcinols, 5-n-pentadecylresorcinol and 5-n-heptadecenylresorcinol, differ
among cultivars (Hassan, 2007).
Factors affecting the potential of latex to cause sapburn are not well
understood. It appears that both the terpene in the oil fraction of the sap, and
adequate polyphenol oxidase (PPO)/peroxidise concentrations in the skin
are required to develop sapburn; PPO and resorcinols in sap are less signifi-
cant (Loveys et al., 1992; Robinson et al., 1993; John et al., 2002). Rain near
harvest and high N in fruit result in more severe sapburn in ‘Kensington
Pride’. However, negative relationships have been observed between exo-
carp N concentration and alkyl resorcinols (Hassan, 2007). Sap from fruit
harvested early in the day causes less sapburn than sap from fruit harvested
later in the day, although early-harvested fruit exude more sap than late-
harvested fruit (Maqbool et al., 2007).
Latex may provide protection against infestation by fruit fly larvae (Joel,
1978, 1980) and may also contribute to disease tolerance. The 5-substituted
resorcinols have antifungal properties (Cojocaru et al., 1985; Droby et al., 1986,
1987; Prusky and Plumbley, 1992). Karunanayake (2007) extracted a resorci-
nol and a resorcinol derivative from the dichloromethane phase of Sri Lankan
mango peel extracts that had antifungal properties. Differing relationships
occur between resorcinol levels and relative susceptibilities of different
mango cultivars to anthracnose (Karunanayake, 2007; Hassan et al., 2007).
Strong positive relationships occur between resorcinol concentration in the
peel and latex, and fruit resistance to artificially inoculated anthracnose.
546 G.I. Johnson and P.J. Hofman
Rough handling at harvest can cause skin damage and internal fracturing or
bruising. Using hooked sticks or shaking the tree to detach fruit causes skin
damage and flesh fracturing (Ledger, 1991a; Abu-Goukh and Mohamed,
2004) and sapburn. Mechanical damage during harvest also causes soft,
darkened areas and bruises on fruit following hot water treatment. Mangoes
should be handled as if they were eggs. Long-handled secateurs cut and grip
the stem, allowing the fruit to be carefully lowered to the picking bin. Contact
with soil and soilborne pathogens should be avoided (Johnson et al., 1993).
In cultivars where sapburn is a problem, latex should be drained from
the fruit (desapping or bleeding) to minimize the incidence and severity of
sapburn. Several systems have been assessed for reducing damage (Brown
et al., 1986; Ledger, 1991b; Holmes et al., 1993; Lim and Kuppelweiser, 1993;
O’Hare and Prasad, 1993; O’Hare, 1994; Shorter and Joyce, 1994). In Austra-
lia, the main commercial practices (Plate 82) are:
● Desapping in the field with harvest aids using detergent. The basic de-
sign characteristics include detergent spraying onto a tarpaulin, a trough
with the same detergent and a final spray before fruit are placed in a field
bin. The fruit are either hooked from the tree in the direction of the harvest
aid and onto the catching surface or the fruit are snapped directly off the
tree and placed onto the tarpaulin. The fruit roll from the tarpaulin into
Postharvest Technology 547
the trough containing detergent and then into 300–400 kg field bins.
Alkaline detergents that deactivate damaging sap components are most
effective; high concentrations of surfactant in the detergent are not re-
quired. The crucial factors are that fruit should be exposed to detergent
for at least 90 s, the detergent is either not recycled, or replaced before
sap accumulation in the detergent causes other damage such as skin
browning (Bally et al., 1997; O’Hare et al., 1999).
● Another design includes a motorized hydraulic ladder (cherry picker)
with the fruit desapped for 1–2 s before placing in a basket containing a
spray of alkaline detergent. This system is particularly effective for tall
trees, but care must be taken that fruit are covered by the detergent for at
least 90 s.
● Picking fruit with long stems into small 18 kg crates and desapping in
the shed. The fruit are dipped into detergent before desapping and plac-
ing on a long conveyor system that holds the fruit inverted and provides
detergent/water sprays for a few minutes. The fruit are inverted for c.20
min before drying and packing.
In these systems, the detergent is not strongly alkaline, but the surfactant
should be of sufficient concentration to provide a protective coating around
the fruit before desapping. Stem breakage must be minimized in the crates,
as this can cause sapburn and quality loss (Holmes et al., 1993; Holmes and
Bally, 1994). Latex must not spray or drip onto fruit being desapped. Workers
who are sensitive to mango sap should wear hand protectants, aprons and
footwear to minimize skin contact. The detergent must reduce sapburn and
skin browning without causing other damage (i.e. lenticel spotting) (Fig.
15.3). Desapping in the field by inverting fruit directly onto racks without
detergents has been used for sensitive cultivars and when particular growing
conditions have increased fruit susceptibility to lenticel spotting or other
damage from detergents; however, labour costs are becoming prohibitive,
requiring compromises between cost and quality. Desapping by inverting
and placing on the ground significantly increases the incidence and earlier
appearance of stem end rot caused by soilborne L. theobromae, and is not
recommended (Johnson et al., 1993).
Harvest aids have reduced in-shed desapping. Holmes et al. (1993) found
9–16% of fruit in field crates were affected by sapburn when harvest aids
were not used. Harvest aids provide the greatest reduction in total sapburn
(from 69% to 15–18%). While harvest aids can significantly reduce sapburn,
inappropriate use can increase some forms of skin browning (Bally et al.,
1997). Underhill and Dahler (1995) described four types of skin browning
which produce symptoms distinct from the sapburn caused by the oil phase
of latex. Several forms of skin browning involve tissue reactions with sap/
detergent mixtures. Symptoms vary if latex enters fruit through micro-cracks
in the cuticle or the lenticels (O’Hare et al., 1999). Holmes (2003) developed
guidelines for the use of harvest aids.
Cost savings associated with the use of harvest aids can be lost if fruit-
to-fruit or fruit-to-ground impact is not minimized during harvest. Rough
548 G.I. Johnson and P.J. Hofman
C
l
lo
us
ic
ar
te
h
tro
as
LO
ag
G
ra
le
Pl
on
W
C
nt
go
go
C
ce
go
go
an
go
an
on
an
an
M
an
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Su
Detergent
harvesting can increase the incidence of bruising and internal fracturing, and
lower wholesale returns. Thorough training of picking crews and supervi-
sion of their performance is required to maintain good practice.
Fig. 15.4. Packhouse and marketing activities for mango. Waxes are not applied in some
countries because of abnormal ripening and off-flavour development. When disinfestation is
not required, steps 11, 12 and 15 would be omitted. When fruit are heat disinfested, they may
be packed into an inner box prior to cooling. For some markets accessible by air, the fruit may
be treated with ethylene and stored until near ripe. The boxed fruit may then be packed into
an outer carton prior to palletizing. Pre-ripening allows fruit to be rechecked prior to despatch,
with fruit of unsatisfactory appearance (e.g. skin damage) redirected to domestic market or
processing.
550 G.I. Johnson and P.J. Hofman
Harvested fruit in field crates should be treated, packed and cooled as soon
as possible. Quality and contaminant management systems may require that
a record system tracks the block or trees from which each bin of fruit is har-
vested. This enables records to be kept of tree or block yield, quality perfor-
mance and defect levels as well as labour performance rates and pesticide
residues. Individual fruit in a tray can be traced back to the tree or block from
which it was harvested. Block/tree-to-tray traceability systems and pack-out
records allow problems (i.e. excessive or unapproved pesticide detections) to
be traced to the site of the problem and relevant action taken. They can also
be used to motivate producers or picking teams to deliver high quality
produce.
At the packhouse, samples of fruit should be evaluated immediately for
maturity, blemishes and disease and pest incidence and recorded in an appro-
priately designed computerized system. Preharvest orchard inspections can
reveal the defects that can be anticipated in the packhouse. Some degree of
in-field sorting can occur at the point of harvest, and soft or damaged fruit
collected separately and discarded.
Disease control
harvest (Johnson et al., 1989a, b, 1991, 1992; Kernot et al., 1999; Poffley et al.,
1999; Plan et al., 2002). Suslow (2000) provides generic recommendations for
the postharvest handling of produce. Postharvest disease treatment efficacy
varies with infection level, cultivar, ripening status and storage regime. Hot
water treatment also cleans fruit, but can contribute to increased skin dam-
age (Cooke and Johnson, 1994). Anthracnose caused by Colletotrichum gloeo-
sporioides Penz. and Colletotrichum acutatum Simmonds, is controlled more
readily than stem end rots (or soft brown rot) caused by anamorphs of Botry-
osphaeria spp. (Fusicossum spp., Neofusicoccum spp., L. theobromae (Pat.) (Griff.
and Maubl.)) (Johnson, 1994; Slippers et al., 2005; Crous et al., 2006) and Pho-
mopsis mangiferae Ahmad, and alternaria rot caused by Alternaria alternata
(Fr.) Keissler. The latter is generally only a problem in fruit from dry regions
or in fruit from more humid areas during storage for 3 weeks or more (Prusky
et al., 1980, 1993a, b, and Chapter 7, this volume; Johnson et al., 1990b).
Fruit are moved through a water bath for 5 min at 48–50°C for less mature
fruit and hot-water-damage-susceptible cultivars (e.g. ‘Zill’ and ‘Irwin’) and
at 50–55°C for mature fruit and less susceptible cultivars (Anonymous, 1994b).
Treatment for 3 min may be adequate for control of anthracnose, while
immersion for up to 7 min may enhance control of stem end rot (Muirhead
and Grattidge, 1986; Sepiah, 1986; Johnson et al., 1989b). In large-scale facili-
ties, dip tanks may range from 3000 to 5000 l, with fruit immersed and moved
through the tank by a series of paddles. In tank construction, non-corrodible
materials such as stainless steel and fibreglass are preferred, and the con-
veyor system that contains the paddles should travel along the bottom of the
tank to reduce damage to fruit that sink. Accuracy in temperature control,
efficacy of the heating unit and timing of fruit flow through the bath are
critical. Temperature probe placement at pump inlet and outlet and thorough
water circulation to ensure accurate temperature reading and to minimize
hot spots are critical. Impurities (e.g. minerals, sediment and debris) in dip
water can affect fungicide performance and stain or damage the fruit. In-line
filters in the inlet and pump circulation systems should be installed and
cleaned regularly.
Where acceptable, carbendazim can be added to the hot water at the rec-
ommended rate, and topped up and replaced regularly, to provide improved
control of stem end rot and anthracnose at lower temperatures (52°C). Beno-
myl has been withdrawn for postharvest use, but much of the benomyl use
information (from earlier research) is relevant for carbendazim (Johnson et al.,
1997). Also, hot thiabendazole (TBZ) is generally as effective as hot benomyl
for controlling stem end rot, but may provide inferior control of anthracnose
(Coates et al., 1993). The active component of benomyl and TBZ in plants,
carbendazim (MBC), is identical (Erwin, 1973; Muirhead, 1976); however,
TBZ also contains sulfur (S) which affects its rate of breakdown and spec-
trum of activity (D. Guest, personal communication, Melbourne, 1995). Beno-
myl penetrates plant tissue more effectively than TBZ, carbendazim or
thiophanate methyl (Eckert, 1983).
Dipping fruit in hot, dirty, latex-contaminated water can increase phyto-
toxicity and lenticel damage. Hot fungicide dips lose efficacy due to sap
552 G.I. Johnson and P.J. Hofman
build-up in the dip tank and stripping out of fungicide (Wells and Littlemore,
1989). Ledger (2004) optimized dip:fruit ratio and dipping practices. Prochlo-
raz provides good control of anthracnose and alternaria rot, but does not pro-
vide control for stem end rot (Johnson et al., 1990b; Johnson and Coates, 1993).
In South Africa, for local markets prochloraz is added at 405 ppm of active
ingredient (ai) of a 45% emulsifiable concentrate (ec) formulation; for export
markets, 810 ppm prochloraz is used. Fruit are immersed for 20 s. In Australia,
prochloraz at 250 ppm is applied by overhead spray, and fruit require 15–20 s
to pass through the prochloraz spray race on a roller conveyer system.
A maximum residue level (MRL) of 7.0 mg/kg for prochloraz for assorted
tropical and subtropical fruits with an inedible peel is recommended (CODEX-
MRL, 2008). This group MRL replaces individual fruit commodity MRLs,
and takes into account the lower residues in the flesh compared to the skin
(Muller and Burt, 1989). Some registered use-rates for postharvest applica-
tion of prochloraz are listed in Table 15.3.
Hot water sprays over brushes (55°C for 15–20 s) is an effective alterna-
tive to hot water dips containing prochloraz for controlling alternaria rot
(Prusky et al., 1999, 2006). Application of hot water spraying and brushing for
15–20 s (HWB) followed by a spray of 50 mM hydrochloric acid (HCl), alone
or in combination with prochloraz, also improved control of alternaria rot
(Prusky et al., 2006). These treatments have not been tested for anthracnose.
Using 2,4-dichlorophenoxyacetic acid (2,4-D) diluted in wax after HWB and
prochloraz reduces stem end rot (Kobiler et al., 2001). A hot water and beno-
myl combination treatment followed by a prochloraz spray provides effec-
tive control of anthracnose, stem end rot and alternaria rot during longer
storage (Johnson et al., 1989a, 1990b). Similar benefits are now attributed to
hot TBZ dip and cold prochloraz spray (Ledger, 2004).
When fungicides are used in the packhouse, spent dip suspensions and
fungicide containers must be disposed using approved methods, often
included with supplier recommendations. Carbendazim suspensions can be
drained into a trench filled with stones, but runoff must be avoided. Car-
bendazim and other benzimidazole fungicides are toxic to earthworms (Wright
and Stringer, 1973).
Postharvest Technology 553
Heat
Pest disinfestation treatments involving heat provide some control of anthra-
cnose, but do not adequately control mango fruit pathogens for export. Tem-
perature and time combinations suitable for non-deleterious fruit disinfestation
are sublethal to a significant percentage of quiescent infections beneath the
fruit cuticle and pedicel tissues (Coates and Johnson, 1993). In many regions,
fruit skin temperatures frequently approach the mid-40°C range during pre-
harvest development, a natural selection pressure favouring heat-tolerant
fungal infection structures. For ‘Kensington Pride’, hot benomyl in combina-
tion with either prochloraz or vapour heat at 46.5°C for 20 min controls stem
end rot more effectively than hot benomyl alone and TBZ, alone or in combi-
nation with vapour heat, during storage at 23°C for 15 days (Coates et al.,
1993). Disease control in combination with heat disinfestation has been
reviewed by Coates and Johnson (1993) and Jacobi et al. (1994, 2001a).
Future options
Heat is an ideal disease control treatment, since it is environmentally safe and
non-chemical. Its effectiveness would be enhanced if fruit tolerance could be
increased by genetic manipulation or the development of pre-conditioning
treatments. Pre-treatments to render quiescent structures of pathogens more
susceptible to heat would also improve disease control. Measures to increase
efficacy could include other energy sources, chemicals, adjuvants, fumigants
or microorganisms to damage or soften fungal wall structures.
Treatments to delay fruit ripening also limit or reduce disease losses. Storage
quality would benefit from the development of cultivars or pre-conditioning
treatments to improve tolerance of fruit for cool storage or controlled atmo-
sphere (CA) and modified atmosphere (MA) storage (Brecht and Yahia,
Chapter 14, this volume). With increasing concerns about the use of chemi-
cals on food (Gullino and Kuijpers, 1994), and in view of current limitations
on heat treatment and storage regime disease control efficacy, non-deleteri-
ous alternatives to synthetic fungicides are required. Alternatives to fungi-
cides for controlling postharvest diseases have been reviewed by Johnson
and Sangchote (1994) and Korsten (2006). Options include: (i) biological con-
trol, i.e. the use of microorganisms to control pathogens (Wilson and Pusey,
1985; Jeffries and Koomen, 1992; Korsten et al., 1993, 1994; Korsten 2006); (ii)
enhanced exploitation of naturally occurring antifungal compounds in fruit
(Prusky et al., 1982; Johnson et al., 1998; Joyce et al., 1999; Zainuri et al., 2003;
Hassan et al., 2007); (iii) application of fruit coatings such as chitosan with
both MA and antifungal effects (El Ghaouth et al., 1992a, b; Wilson et al., 1994;
El Ghaouth and Wilson, 1995); (iv) exposure to UV-C light (wavelength <280
nm) (Chalutz et al., 1992; Wilson et al., 1994). Zainuri (2006) reported some
promise in the use of UV-C radiation for control of anthracnose, but fruit
damage risks and treatment dose accuracy were critical; (v) containment of
fruit in atmospheres containing high levels of carbon dioxide (CO2) for 24–48
h after harvest (flushing) (Prusky et al., 1992, 1993c); (vi) regulation of fruit
ripening (Brady, 1994); and (vii) application of naturally occurring plant
products (Fallik and Grinberg, 1992; Wagner and Flores, 1994). Many of these
554 G.I. Johnson and P.J. Hofman
Brushing
Brushing on mango packing lines can occur after, or at the same time as, hot
water and fungicide treatments. Hot water treatment washes sap away, and
loosens superficial debris, scale insect carapaces and sooty mould, which
are removed as the fruit pass over rotating brushes. Brushing also removes
superficial deposits of fungicides that accumulate on fruit from orchard
application of Cu fungicides (Lonsdale, 1993) and incorrect mixing or sedi-
mentation of benzimidazole fungicides resulting from sap accumulation in
dip tanks (Wells and Littlemore, 1989). Soft, non-damaging brushes should
be used, washed every day and replaced seasonally.
For ‘Kensington Pride’ mangoes harvested after rain, skin marking, fruit
shrivel and weight loss increase significantly on fruit treated with a hot water
and fungicide dip or a hot water and fungicide dip followed by treatment with
prochloraz, when fruit brushing followed either or both treatments relative to
untreated and untreated/brushed fruit. Prochloraz before brushing resulted
in fruit quality similar to untreated or brushing only (Cooke and Johnson, 1994).
Brushing can increase lenticel spotting (Oosthuyse, 1999). Therefore, the num-
ber and type of brushes must remove foreign matter and polish the fruit, while
not increasing risk of brush and lenticel damage, especially during wet weather
and with heat treatments (see Weather conditions and Skin colour and lenticel
damage sections, both under 15.3 Preharvest Management, this chapter).
The purposes of grading are to sort fruit into defined categories of unifor-
mity and to divert out-of-grade fruit from the pack line to either a second
Postharvest Technology 555
grade, processing or reject line. Mangoes with defects outside acceptable lim-
its as defined in a grade schedule or chart are manually removed and trans-
ferred (by conveyer belt) to seconds or processing lines as appropriate. The
purpose of sizing is to categorize fruit into size or weight groups for packing.
Fruit must be sized prior to disinfestation with hot water or vapour heat to
ensure consistent treatment responses. Typical systems include automatic
graders that separate fruit by weight into groupings that correspond to pre-
determined categories (Schoorl and Holt, 1982, 1985). Camera vision systems
can separate for colour, defects and shape. Fruit usually accumulate in sepa-
rate bins for packing into cartons or into bulk containers for processing or
disinfestation. The fruit are packed manually into single-layer trays, with
plastic or cardboard liners that have depressions designed to accommodate
fruit of a particular size. The depressions provide some support for individ-
ual fruit during packing, while the cardboard liners also provide some
additional buffering against impact damage. The pattern of the depressions
facilitates most efficient utilization of carton space. Mango tray liners com-
monly accommodate 12–25 fruit for 6.5 kg trays. Some tray liners may be
inappropriate for sea export due to interference with vertical airflow.
Organic materials (i.e. paper, leaves or shredded wood) have been
used to cushion individual fruit in cartons. These materials can harbour
pathogens, for example Rhizopus stolonifer (Ehrenb. Fr. Lind.), which
causes transit rot of mangoes and has been detected in shredded wood
used in mango packaging. Shredded wood creates micro-wounds in the
fruit skin, providing points of entry for hyphae growing on the wood.
Losses are more severe when fruit have been removed from cold storage,
allowing condensation to develop on the fruit and shredded wood
(Muirhead and Grattidge, 1986).
Grade standards
Packing-line QA inspections
Packing-line control inspections are used to monitor grading efficacy and
packing-line damage. Packing-line inspection samples are taken soon after
fruit pass points in the line at which defects are most likely to be overlooked
and/or induced. For start and end-of-line pack-out checks, and out-turn
inspections, randomly selected cartons are unpacked, and all fruit are checked
for compliance with preset quality parameters. Most value is gained from
quality control checks if records are kept and evaluated, with feedback/trou-
ble shooting as necessary, to constantly improve the system (Ledger and Bag-
shaw, 1994; Ledger and Premier, 2006). Computer analysis of such information
provides a seasonal benchmarking record of QA improvement, and high-
lights areas for attention in packing-line improvement and personnel train-
ing. Record keeping is mandatory under GAP certification systems.
Future options
Greater automation of grading and packing will become necessary as pro-
duction and labour costs increase, and as customers become more demanding
(Hilton, 1994). Recent advances in computing have made possible high-speed
sorting using visual systems for colour, shape and externally visible defects.
Also, NIRS systems can now be used in-line to sort for flesh characteristics
that influence flavour. In mango, percentage dry matter and flesh colour are
related to ripe fruit flavour, and can be estimated using NIRS (Saranwong
et al., 2004; Subedi et al., 2007). Estimation is sufficiently accurate to allow
acceptable separation into several categories for final flavour. NIRS may also
be useful for predicting ripening behaviour and weight loss during ripening
(Mahayothee et al., 2004). Given the influence of weight loss in chilling injury
(CI) development during cold storage (Bower et al., 2003), NIRS may also be
able to estimate potential for CI during cold storage.
There is interest in other non-destructive quality assessment for the pack-
house. Joyce et al. (1993) noted that future innovation could lead to proton
magnetic resonance imaging (MRI) technology suitable for packing-line
applications to allow non-destructive detection of internal disorders and pest
infestations. X-ray imaging may have potential for detecting seed weevil
Postharvest Technology 557
damage in mangoes (Thomas et al., 1995; Reyes et al., 2000), but recent inves-
tigations suggest that neither X-ray imaging nor MRI is sufficiently reliable
for quarantine purposes, particularly where larvae are small (R.A. Jordan,
personal communication, 2007). New methods of nuclear magnetic reso-
nance (NMR) (Marigheto et al., 2008) may distinguish internal disorders such
as jelly seed. Acoustic/ultrasonic methods can sort for fruit firmness (Miz-
rach, 2008) and may help identify softening fruit with internal disorders and
reduced storage life. Robotics in sorting and packing will be used increas-
ingly where labour costs and availability are high.
Disinfestation
Required for
Import Permit No No
Phytosanitary Certificate Yes Yesa
Additional Declaration No Yesb
Post Entry Quarantine No No
EX188 No No
EX46 No No
Radiation Statement No No
a Treatment details, including date of treatment, are to be endorsed on the Phytosanitary
Certificate in the treatment section. The treatment is to be shown as: irradiation at minimum
250 gray.
b The mangoes in this consignment have been treated in accordance with Appendix 12 of the
Bilateral Quarantine Arrangement between New Zealand Ministry of Farming and AQIS.
Target pests
A pest of regulatory concern that could become established in an area where
it is not found is a quarantine pest risk, and requires quarantine action.
Mango fruit pests include internal pulp feeders (i.e. fruit fly immatures), seed
and fruit pulp pests (i.e. mango weevils and fruit caterpillars) and external
pests (i.e. scales, mealybugs, thrips and mites) (see Peña et al., Chapter 10,
this volume). External pests pose detection risks as surface hitchhikers that
Postharvest Technology 559
Fruit fly disinfestation
Tephritidae are the most important mango pests and occur wherever man-
goes are grown (Waite, 2002). Eggs are oviposited below the peel. The wound
provides an opening for microorganisms and scars the peel. Larvae feed and
tunnel throughout the pulp. Fruit flies infest tropical and temperate fruits. It
is the risk to temperate climate fruits and commodities produced in fly-free
areas that has prompted the development of quarantine restrictions and
treatments for fruit fly hosts.
At present, quarantine treatments against fruit flies are not required for
fruit entering the European Union (EU), despite the large production of tem-
perate fruit in fruit-fly-free regions. Fly infestation has not been perceived as
a threat because winter temperatures throughout much of the region effec-
tively prevent establishment of the flies, despite geographical continuity
with the distribution range of the Mediterranean fruit fly (Ceratitis capitata
(Wiedemann)). Canada does not require fruit fly disinfestation of tropical
produce for the same reason. Exotic fruit fly pests could become established
in southern Florida, Texas and California because of their subtropical climate.
The USA requires that mangoes be disinfested by vapour heat, irradiation,
hot water or hot air.
560 G.I. Johnson and P.J. Hofman
Vapour heat
Vapour heat treatment (VHT) involves heating air that is nearly saturated
with moisture, and passing the air stream across the fruit (Jacobi et al., 2001b).
When the temperature of the mango fruit is at or below the dew point of air,
condensation occurs on the fruit surface and rapidly heats the fruit by con-
ductive energy transfer. The core of the fruit next to the seed is heated to
c. 45°C for the required time before cooling. Fruit have to be sorted for size
before treatment because of different rates of attaining the required core
temperature.
Vapour heat is used worldwide to disinfest mangoes of fruit flies. Jacobi
et al. (2001b) list the VHT protocols approved for importation of mangoes
into Japan from the Philippines, Taiwan, Thailand, Australia and Mexico.
Conditions range from 43–47°C pulp core temperature for 10 min to 6 h;
however, the most common treatment conditions are 46–47°C for 10–30 min.
Melon fruit fly (Bactrocera cucurbitae Coquillett) immatures in mangoes from
Okinawa were killed at 44 ± 0.3°C core temperature for 3 h (Sunagawa et al.,
1987). Taiwanese mangoes infested with melon fly can be disinfested with
vapour heat at 47.5°C until the centre pulp is >46.5°C for 45 min (Kuo et al.,
1987).
A VHT schedule was approved against Queensland fruit fly (Bactrocera
tryoni), in ‘Kensington Pride’, ‘R2E2’, ‘Keitt’, ‘Palmer’ and ‘Kent’ from Aus-
tralia for the Japanese market (AQIS, 2008b), which consists of a core tem-
perature of 47°C for 15 min. The United States Department of Agriculture,
Postharvest Technology 561
Animal and Plant Health Inspection Service Plant Protection and Quarantine
(USDA-APHIS PPQ) approved VHT as a quarantine treatment for Mexican
fruit fly (Anastrepha ludens (Loew)) and other Anastrepha species in ‘Manila’,
and for mangoes from Taiwan infested with oriental fruit fly (Anonymous,
1994a). Generic guidelines for use of VHT in treating commodities for the
USA market are provided by the USDA-APHIS PPQ manual on vapour heat.
Mangoes from Taiwan imported into Australia must be treated until the pulp
temperature has been 46.5°C for 30 min (AQIS, 2008a).
Hot air
Hot or forced hot air systems also heat the air to 40–50°C, but at a lower RH.
Relative humidity usually remains >50%, depending on ambient RH, but is
never high enough to produce condensation. Heat is transferred to the fruit
by convection, with no condensation of water on the skin (Gaffney and Arm-
strong, 1990; Jacobi et al., 2001b). Relative humidity should be high enough to
prevent fruit desiccation during treatment. Transfer of heat from the air to
the skin is slow compared with VHT. Mangan and Ingle (1992) reported that
a mean centre pulp temperature of >47°C killed all stages of West Indian fruit
fly, Anastrepha obliqua (Macquart), in Mexican mangoes, and Sharp (1992)
found a centre pulp temperature of >46°C killed all stages of Caribbean fruit
fly, Anastrepha alletis (Loew), in Florida-grown mangoes.
Hot water
Provided that fruit are not damaged, hot water immersion is environmen-
tally safe and efficient for killing mango pests. Use of hot water to kill fruit
fly eggs and larvae intensified in the USA when the Environmental Pro-
tection Agency (EPA) removed ethylene dibromide from the market as a
chemical fumigant because of health concerns (Anonymous, 1983). Sharp and
Spalding (1984) showed that mangoes could be disinfested of Caribbean fruit
fly using hot water. The work led to more studies in Haiti and a disinfestation
method for West Indian fruit fly (Sharp et al., 1988), as well as Mediterranean
fruit fly and other Anastrepha spp. in Texas and Mexico (Sharp et al., 1989a, b),
Puerto Rico (Segarra-Carmona et al., 1990) and Peru (Sharp and Picho-Martinez,
1990). Nascimento et al. (1992) developed a hot water treatment for fruit flies
in mangoes in Brazil. Hot-water-treated mangoes may be imported into the
USA from Mexico, Central America, South America and the West Indies
(Anonymous, 1994a). Typical treatments include 46.1°C for 65 min for smaller
fruit to 90 min for larger fruit (Jacobi et al., 2001b). Large commercial hot-
water-treatment facilities have been constructed, certified by the USDA-
APHIS PPQ, and used in Mexico, Central and South America, and the West
Indies. Generic guidelines for the use of hot water are provided by the USDA-
APHIS PPQ manual for hot water treatment.
In Australia, Smith (1992) showed that immersing five Australian mango
cultivars in 48°C water for 30 min killed eggs and larvae of Bactrocera aquilo-
nis; however, ‘Kensington Pride’ is more sensitive to hot water than to vapour
heat, so the latter has been adopted for disinfestation of mangoes in Australia
(Jacobi et al., 1994). Grové et al. (1997) found that treatment of several cultivars
562 G.I. Johnson and P.J. Hofman
in hot water at 46.1°C for 90 min followed by refrigeration for 24 h did not
damage fruit, although some cultivars showed severe lenticel damage.
Refrigeration of ‘Tommy Atkins’ fruit immediately after treatment resulted
in scald development. Weevils in ‘Alphonso’ mangoes from India were not
killed when infested mangoes were immersed in water at 48–52°C for up to
90 min and 54–70°C for up to 5 min (Shukla and Tandon, 1985).
Compared with hot air treatments, hot water treatments can damage the
skin, partly because of rapid heat transfer from the water to the skin com-
pared with from the skin to the centre of the fruit. Damage includes skin scald-
ing, lenticel damage, cavities, white starchy areas in the flesh and delayed
ripening (Jacobi et al., 2001b). Several factors influence damage severity after
heat treatment, for example cultivar, temperature and duration (Jacobi et al.,
2001b). Immature fruit have low heat tolerance, and small fruit are damaged
by heat more readily than large fruit. Conditioning treatments (i.e. 37°C core
temperature, for at least 12 h in air) can reduce injury, and preharvest condi-
tions, especially rainfall before harvest, can increase skin damage (Esguerra
and Lizada, 1990; Esguerra et al., 1990; Jacobi and Wong, 1992; Jacobi et al.,
1994, 1995; Jacobi and Giles 1997). Better understanding of these influences
could increase the commercial potential for hot water disinfestation.
Hot water dips could pose human health risks. Sivapalasingam et al.
(2003) reported that an outbreak of Salmonella enterica that infected 72 patients
from 13 USA states may have been due to contamination of hot-water-dipped
mangoes from a single farm in Brazil. No outbreaks were reported among
consumers in the EU of mangoes from the same farm, and the EU does not
require hot water disinfestation.
Irradiation
Irradiation involves γ rays (at <1000 Gy), X-rays, electrons and microwaves
(Thomas, 1986; Velasco and Medina, 2004; Follett et al., 2007; Moreno et al.,
2007). A 2005 FAO/International Atomic Energy Agency (IAEA) report indi-
cated that >20 irradiation facilities have been planned, constructed or reno-
vated in ten countries, some of which are mango exporters (Eustice, 2007).
Radiation treatments have been developed for fruit flies in mangoes from
Florida, Mexico, India and Australia. Von Windeguth (1986) treated mangoes
with 76 Gy and disinfested them of Caribbean fruit fly eggs and larvae. Third
instar Mediterranean fruit fly larvae in Mexican mangoes irradiated with 250
Gy did not emerge from pupae, and 60 Gy applied to third instar Mexican
fruit fly, and West Indian fruit fly in Mexican mangoes prevented adult emer-
gence (Bustos et al., 1992). Bustos et al. (2004) recommended a generic dose of
150 Gy for control of Mexican fruit fly (A. ludens), the West Indian fruit fly (A.
obliqua), the sapote fruit fly (Anastrepha serpentina) and the Mediterranean
fruit fly (C. capitata) in mango. ‘Kensington Pride’ mangoes infested with
eggs and larvae of Queensland fruit fly and Bactrocera jarvisi (Tryon) are
disinfested with 74–101 Gy (Heather et al., 1991).
International guidelines for the use of irradiation as a phytosanitary
measure are available (ISPM, 2003), and recently a fast track process has been
proposed as an Annex to ISPM 28 (ISPM, 2008), which endorses irradiation
Postharvest Technology 563
Table 15.5. Minimum absorbed dose of gamma irradiation required by USDA for specific
pests (Source: adapted from EPA, 2002).
many of which can infest mangoes. The USDA-APHIS PPQ manual on irradia-
tion provides generic guidelines. Irradiation was approved for the USA market
as a phytosanitary treatment for all fresh fruits and vegetables from all coun-
tries in 2002. Effects of J-irradiation on mango fruit quality and disease control
have been reported (Mitchell et al., 1992; Moreno et al., 2006; Reyes and Cisneros-
Zevallos, 2007; Wall, 2008). Only marginal disease control was obtained with
‘Kensington Pride’ at the highest non-deleterious doses for mature-green fruit
(300 Gy), with additive effects of disease control treatments and irradiation on
disease reduction (Johnson et al., 1990a). Disease control may be more effective
in cultivars with greater tolerance of irradiation (van der Linde and Thord-Gray,
1986; Johnson et al., 1990a). Other types of irradiation have been evaluated
for mango disinfestation but none has been adequately suitable.
Quick freezing
Quick freezing of mango, lowering the temperature to –17°C and holding at
–6°C or below for 48 h is used to disinfest mangoes for processing (Anony-
mous, 1994a; PPQ, 2007). The process is not approved for importing man-
goes with seeds from most of the West Indies, French Guiana, all countries
outside of North, Central and South America, Oceania, Hawaii, South-east
Asia, the Philippines and the Republic of South Africa into continental USA
because mango weevil could be present (Anonymous, 1994a; PPQ, 2007).
Fumigation
Fumigation is an ideal methodology for ensuring effective control when the
fumigant is effective and safe to use. Until 1994, New Zealand required fumi-
gation of mangoes from Australia, the Cook Islands and the Philippines
using 33, 29 or 22 g/m3 ethylene dibromide at 10–15, 15.5–19.5, or 20°C and
above, respectively, at normal atmosphere pressure (NAP) to disinfest man-
goes of fruit flies before entry. As part of the international phase-out of ozone-
depleting substances, the process was banned in 1994 (Anonymous, 1992;
N.W. Heather, personal communication, Brisbane, 1994) and most applica-
tions as a fruit fumigant have ceased worldwide. Methyl bromide was phased
out completely in the USA in 2005, but some emergency uses for quarantine
applications may be permitted, e.g. to destroy a serious quarantine pest in an
imported consignment or to meet official requirements of an importing coun-
try (EPA, 2008). Mangoes imported into Australia from countries where fruit
flies occur must be fumigated with 16–35 g/m3 ethylene dibromide for 2 h at
21–26°C or above (Anonymous, 1985, 1988).
Phosphine is widely used as a fumigant of durable produce (grains and
tobacco). It provides effective control of fruit fly larvae and other pests in
temperate fruits under experimental conditions (Horn and Horn, 2004).
However, phosphine when mixed with water is highly explosive and the
vapour is toxic to humans, so prospects for utilization are not strong.
Miscellaneous treatments
CHEMICAL TREATMENTS. Postharvest chemical treatments using dimethoate are
effective against Queensland fruit fly with ‘Kensington Pride’ (Swaine et al.,
Postharvest Technology 565
NATURAL PRODUCTS. The short shelf life of mango and the high level of insect
mortality required obviates the use of natural products for disinfestation.
Suhaila and Halim (1994) reported the potential of low toxicity, insecticidal
compounds from edible plants that may be effective for topical application to
harvested fruit. Extracts of black pepper (Piper nigrum) were particularly active
in laboratory tests against vinegar fly (Drosophila melanogaster (Meigen)).
PACKAGING. Some markets, for example Japan and the USA, require that fruit
must be packed into insect-proof packages following disinfestation to preclude
566 G.I. Johnson and P.J. Hofman
Surface coatings
Surface coatings are used to improve fruit appearance and to alter gas per-
meability to reduce moisture loss or retard ripening. Commercial use of sur-
face coatings on mango fruit needs to be considered carefully because of the
fine balance between beneficial and undesirable effects on fruit quality. Neg-
ative effects of coatings include reduction in chlorophyll loss (Fonseca et al.,
2004a), anaerobic conditions and off-flavours (Amarante and Banks, 2001)
and skin damage, possibly due to cytotoxic reactions with other components
in the coating formulation (Bower et al., 2003). Generally, coatings have less
effect on delaying ripening during cold storage, compared with extending
the shelf life at typical ripening temperatures (Amarante and Banks, 2001).
Less significant effects are observed in more mature and in ripening fruit.
Coatings often delay skin colour change rather than softening, which increases
the risk of soft, green fruit with less consumer appeal.
Coatings are generally emulsions of synthetic (e.g. polyethylene) or nat-
ural (e.g. polysaccharides, carnauba, beeswax, etc.) origin. Surface coatings
containing waxes, oils (e.g. carnauba, beeswax, etc.) and resins (e.g. shellac)
have a greater effect on limiting water loss then reducing O2 and CO2 perme-
ability, compared with those containing polysaccharides, (e.g. those based on
cellulose) (Amarante and Banks, 2001). Formulations based on shellac result
in a shinier appearance than those based on carnauba wax and polysaccharide-
based waxes (Baldwin et al., 1999; Hoa and Ducamp, 2008).
Factors other than coating formulation can affect fruit gas permeability,
i.e. cultivar, variations in skin permeability between fruit, inconsistency in
coating thickness during application, interference from water during appli-
cation causing coating cracking and coating thickness and evenness-of-
spread over the fruit surface. The effect of coating on fruit quality can vary
with holding conditions because of larger temperature effects on respiration
rate than on coating permeability.
Shorter and Joyce (1994) found commercially formulated Avocado and
Passionfruit Wax, a polyethylene and shellac emulsion, and Technimul 9122
Wax, a polyethylene-based emulsion, were acceptable with ‘Kensington Pride’
mango, while Peach Wax, a polyethylene-based emulsion and starch solution
was unacceptable. With Peach Wax, deleterious modified atmosphere effects
on colour development, softening and flavour were obtained (El Ghaouth
et al., 1992b; Shorter and Joyce, 1994). Coating ‘Tommy Atkins’ mango with a
carnauba-based coating and BeeCoat (based on beeswax) reduced water loss,
shrinkage, chlorophyll breakdown, CI and decay after cold storage, and Bee-
Coat also reduced red lenticel discoloration (Feygenberg et al., 2005). With
Postharvest Technology 567
Packaging
sides of the cartons. Storage and product use information can also be printed
on the cartons. Many QA systems require adequate labelling linked to appro-
priate record keeping for plate-to-farm traceability. Clear labelling facilitates
correct delivery, allows immediate buyer recognition of product profile and
ensures maintenance of accurate sales records. An exporting country may
find it of value to identify individual packers by barcoding or numbers
stamped on cartons, so that sources of faulty packaging can be traced. Some
countries also use date codes which enable exporters to determine the fresh-
ness of the produce at the point of export and evaluate an importers’ capacity
to achieve adequate turnover of the fruit without prolonged storage. It also
provides invaluable feedback on the efficiency of the total distribution chain.
Cartons used for export should be clean, strong, unbroken and new. The
water absorption capacity of the material should be evaluated as excess
absorption will lead to collapse on the pallet. The cartons’ strength will
depend on the starch used by the manufacturer, the outer liner and the direc-
tion and numbers of fluting in the carton (Anonymous, 1994b). There is
increasing pressure in the EU for recyclable packing material. Cartons that
are recyclable should be marked with the appropriate international symbol.
Returnable plastic crates are increasingly being used for domestic trade, but
the return cost would make this less profitable for international trade.
Inspection
Palletizing
Precooling
Precooling removes field heat from the product and lowers the temperature
to that required for ripening, transportation or storage. Precooling also
reduces the cooling demand on any in-transit cooling system. Precooling
concepts and systems are described by Thompson et al. (2002). Forced-air
cooling systems efficiently and rapidly remove field heat, and are preferred
for bringing fruit to storage temperature. High RH systems are preferred as
they reduce fruit water loss. Hydrocooling can increase the risk of infection
by wound pathogens (i.e. Rhizopus spp.) and are less effective with large fruit.
Kitinoja and Kader (2003) describe low-cost cooling facilities for use in devel-
oping countries.
Table 15.6. Days for fruit to reach the eating soft stage (days to ripe at 20°C),
percentage weight loss/day and percentage of fruit surface area affected by stem
rots in ‘Kensington Pride’ mango fruit treated with 25 Pl/l 1-methylcyclopropene
(1-MCP) for 14 h at 20°C followed by exposure to 100 Pl/l ethylene for 24 h at 20°C.
Fruit were then ripened at 20°C. Means followed by the same letter in each column
are not significantly different (P >0.05) (Source: Hofman et al., 2001).
70
24 h
72 h
60
50
40
Green colour (%)
30
20
10
5
2
1
0
0 10 100 1000 0 10 100 1000 0 10 100 1000
Treatment at 15°C Treatment at 20°C Treatment at 25°C
Ethylene concentration (μl/l)
Fig. 15.5. Effect of ethylene concentration and time in ethylene and ripening
temperature on the percentage skin surface area with green colour of ‘Kensington
Pride’ mangoes at eating soft; least significant difference = 5.16 (P <0.05) (Source:
Nguyen et al., 2002). Note the increased green colour on the skin of ripe fruit with
lower ripening temperature and higher ethylene concentrations and duration.
the start of ethylene treatment; otherwise, skin spotting can develop (Ledger,
2003a). Ripening at 18–22°C is recommended for maximum yellow skin colour,
less disease and higher flavour volatiles (Hofman, 1997; Lalel et al., 2004).
The relatively high respiration rate of ripening mangoes can result in
CO2 accumulation in the ripening room, particularly if the room is full and
there is poor ventilation. Carbon dioxide concentrations up to 5.3% have
Postharvest Technology 571
been recorded in ripening rooms (Ledger, 2007), which can cause more green
colour and a dull appearance on the ripe fruit (Nguyen, 2003). Ripening room
CO2 concentrations should be maintained at <1% with adequate ventilation
to minimize fruit quality loss (Kernot et al., 1999; Ledger, 2007).
Accidental exposure of mangoes to ethylene and its analogues from adja-
cent ripening rooms, exhaust fumes from internal compression engines or
wound ethylene produced from damaged/ripening fruit can cause prema-
ture ripening. Various systems can remove unwanted ethylene, for example
oxidizing mechanisms such as potassium permanganate either in sachets or
in ethylene scrubbing units in storage rooms, catalytic oxidizers or ozone-based
systems (Reid, 2002). Smartfresh™ (active ingredient 1-methylcyclopropene;
1-MCP) is a relatively new approach for preventing undesirable ethylene
effects. 1-MCP is a structural analogue of ethylene and irreversibly binds to
the ethylene receptors in the plant, thus preventing ethylene-initiated ripen-
ing. Ripening re-commences as additional ethylene-receptor sites are pro-
duced in the fruit (Blankenship and Dole, 2003). Generally, Smartfresh™
treatment is applied in well-sealed cold rooms or plastic tents as soon as pos-
sible after packing. 1-MCP concentrations of 250–200,000 Pl/l for 12 h are
optimum for delaying ripening (Jiang and Joyce, 2000; Hofman et al., 2001;
Adkins et al., 2002; Penchaiya et al., 2006), although most reports state 250–
1000 Pl/l. 1-MCP treatment completely negated any effect of subsequent eth-
ylene on ripening, and can almost double the days to eating soft compared
with ethylene-treated fruit ripened at 20°C (Table 15.6) (Adkins et al., 2002).
However, the 1-MCP effects were less in more mature fruits (Alves et al.,
2004), and ethylene exposure before 1-MCP will negate any 1-MCP benefit
(Adkins et al., 2002). Any beneficial effects of 1-MCP also appear to be less
with longer-term storage (Hofman, unpublished results). 1-MCP treatment
can cause more disease on ripe fruit, because the longer days to ripen allows
more disease development (Hofman et al., 2001). Sourcing fruit from well-
managed orchards can help minimize this effect (Adkins et al., 2005).
For some domestic markets, on-farm treatment of mangoes with ethyl-
ene is used to ensure that fruit have more attractive colour when they are
displayed at the wholesale market 48–72 h after dispatch from the farm. This
practice improves returns as fruit can be delivered to retail outlets ready-to-
eat. Ethylene induction of ripening is undesirable for more distant markets
because fruit arrive at the market too ripe for sale, with greater risk of bruis-
ing and disease.
Cool storage
Cool storage is important when delivery time from harvest to the consumer
exceeds the typical ripening time (5–10 days). The ideal storage temperature
is dictated by the risk of CI, fruit ripening and disease development during stor-
age, and storage time. CI is first noted as greying of the skin, which intensifies
572 G.I. Johnson and P.J. Hofman
Decreasing the O2 and/or increasing the CO2 concentration can have several
advantages with respect to storage (i.e. reduced ethylene production, better
flavour retention, slowing softening and green skin colour loss and reduced
CI) (Thompson, 1998; Yahia, 2006). However, if the O2 concentration is too
low (dependent on cultivar, storage temperature, fruit maturity and ripeness
stage) anaerobic respiration will commence, with associated production of
ethanol and acetaldehydes, leading to off-flavours and physiological disor-
ders (Bender et al., 2000). Atmosphere modification generally has less benefit
for tropical fruit compared with temperate fruit, but does have commercial
potential for sea freight to distant markets. Atmosphere control can be active
or passive, or combinations of the two. Surface coatings (see Surface coatings
section under 15.7 Preparing Fruit for Market, this chapter) also provide
modified atmosphere inside the fruit.
With CA systems, O2 and CO2 concentrations are actively monitored and
controlled by injecting N2 and CO2, or bleeding air into the container as
required. In more passive systems, such as the MaXtrend® system (Maxtend,
2008) fruit respiration directly lowers O2 concentrations, and its concentra-
tion is monitored and manipulated by venting as required. In some cases, the
container is flushed with N2 at the start of storage to rapidly establish the
desired atmospheres. Excess CO2 is absorbed with hydrated lime. In MA sys-
tems, atmospheres are modified by placing a semi-permeable membrane
around the fruit (usually plastic film), and relying on fruit respiration to
modify the atmosphere.
574 G.I. Johnson and P.J. Hofman
15.9 Transport
Transportation of tropical fruit and vegetables has been reviewed by McGregor
(1987) and Thompson (2002). For local markets (<3 h access), transportation
of fruit in non-refrigerated carriers is feasible, particularly if the fruit has
576 G.I. Johnson and P.J. Hofman
been precooled and transported at night with few stops. Fruit must be shel-
tered from direct sun and rain. For sea export, fruit must be cooled to the
required vessel carrying temperature (or within 2°C thereof) and the cold chain
must be maintained until the fruit is displayed for purchase.
Some retailers prefer fruit that are ready to eat within 1–2 days of receipt. In
these situations, the ideal scenario is to ripen the product as close as possible to
the retail outlet to minimize physical damage to the soft fruit. However, where
end-market location and transport arrangements allow delivery to market
within 3 days, ripening on-farm has advantages by reducing costs for growers,
and extending ownership of the product. Transport time is a major consideration
for determining optimum systems (Ledger, 2003b) (Table 15.7).
When the export dispatch facility is >1 h away from the packhouse, the
following road transport recommendations apply:
● The refrigerated truck should be clean and in good mechanical condi-
tion. The insulation and floor should be in a sound condition, the door
seals must be intact and the doors must close very tightly.
Table 15.7. Ripening and transport recommendations for ‘Kensington Pride’ mango within
Australia, to cater for the ‘ripe-for-tonight’ programme of major retail chains (Source: Ledger,
2003b).
Table 15.8. Typical packing and shipping schedules for mangoes consigned by sea
to the EU from South Africa.
15.10 Marketing
15.11 Conclusions
Mango production has been based almost entirely on Mangifera indica, albeit
a variable meld of thousands of cultivars which may be derived from inter-
specific hybrids of a few closely related species (Kostermans and Bompard,
1993). Given its perishable nature, capitalizing more on the diversity of exist-
ing germplasm to develop cultivars with superior storage traits linked to
customer appeal could deliver major benefits.
Future improvements in postharvest technology and quarantine treat-
ment will come from refinement of preharvest management, for example
reducing disease inoculum and increasing fruit resistance to disease, reduc-
ing harvest costs and fruit damage, improving postharvest treatments and
systems, and supply chain approaches to enhance fruit longevity and quality
and reduce the risks of product damage. Improvements will also accrue from
the provision of user-friendly information for supply chain personnel, but
only if the information is utilized and implemented. Increases in throughput
via the automation of harvesting and treatment systems for fruit will increase
as production and marketing costs escalate. Labour saving and work effi-
ciency will also become more critical. Innovative transport arrangements
may become necessary as regional development places greater pressures on
transport systems. International, collaborative joint-marketing ventures will
ensure year-round supplies of uniform quality fruit, and per capita consump-
tion of mangoes will increase (Johnson, 1995).
Acknowledgements
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Postharvest Technology 605
16.1 Introduction
Worldwide mango production occurs in over 90 countries. Although only a
relatively small proportion of total mango production enters international
trade (<4%), the volume traded has increased substantially since the late
1990s. Among the factors responsible for the increased mango production,
trade and consumption are lower prices, year-round availability, fewer
horticultural trade barriers, changes in food consumption preferences,
longer shelf life for perishables and consumer interest in healthier foods.
Although not a major mango producer, the USA has developed most of the
popular cultivars traded on the international market, and is the largest
single-country mango importer. The costs and returns and general practices
of establishing and maintaining orchards vary considerably from coun-
try to country and within each country (different regions and production
systems).
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
606 (ed. R.E. Litz)
World Mango Trade and Economics 607
16.2 Recent Trends in World and USA Mango Production, Trade and
Consumption
World situation
Asia accounts for approximately 77% of global mango production, and the
Americas and Africa account for approximately 13% and 9%, respectively
(FAOSTAT, 2007). In 2005, world production of mango was estimated to have
reached 28.51 million t, an increase from the 27.82 million t recorded in the
previous year. Between 1996 and 2005, production grew at an average annual
rate of 2.6%. Table 16.1 shows the world’s top ten mango-producing coun-
tries, which account for about 85% of the world’s production.
India is the largest producer, accounting for 38.58% of global production
from 2003 to 2005. During that period, the Indian mango crop averaged
10.79 million t, followed by China and Thailand at 3.61 million t (12.90%) and
1.73 million t (6.20%), respectively. Other leading mango-producing coun-
tries and their respective shares of world production during the 2003–2005
period include Mexico (5.50%), Indonesia (5.29%), Pakistan (4.48%), Brazil
(4.30%), the Philippines (3.53%), Nigeria (2.61%) and Egypt (1.28%).
Although currently only 3.3% of the world production of mango is traded
globally, this represents a noticeable increase over the quantities traded since
the late 1980s. In terms of distribution, Mexico, Brazil, Peru, Ecuador and
Haiti supply the majority of North America’s imports. India and Pakistan are
the predominant suppliers to the West Asian market. South-east Asian coun-
tries get most of their supplies from the Philippines and Thailand. European
Union (EU) buyers source mango mainly from South America and Asia.
In 2005, global exports of mango reached 912,853 t, a slight decrease of
0.73% compared with the previous year, and were valued at US$543,100,000
(FAOSTAT, 2007). Table 16.2 shows the top ten major mango-exporting coun-
tries. India is the largest producer but only recently has overtaken Mexico as
the number one exporter of the fruit. For the 2003–2005 period, Mexico and
India dominated the export trade with shares of 22.64% and 20.25%, respec-
tively, followed by Brazil (13.18%) and Pakistan (6.94%). Other major export-
ers include the Netherlands (major re-exporter), Peru, Ecuador, the Philippines,
Thailand and China.
World imports of mango increased from 397,623 t in 1996 to 826,584 t in
2005. The USA is the number one importer of mango. During the 2003–2005
period, the USA imported 271,848 t, or approximately a third of the total
mango imports (Table 16.3).
The Netherlands imported 88,300 t of mangoes (10.62%), although most of it
is redistributed throughout the EU. Other prominent importing countries that
are also major redistributors are the United Arab Emirates (6.82%) and Saudi
Arabia (5.32%). Most of these imports are redistributed to other countries within
the Middle East. Although China (4.91%) appears as a major importer, the quan-
tities imported have been declining. For example, China imported 57,000 t in
2004 and only 19,000 t in 2005. This could be due to increases in domestic pro-
duction in response to an increase in domestic demand driven by rising per
608
Table 16.1. World’s ten major mango producers, 1996–2005 (1000 t) (Source: FAOSTAT, 2007).
Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2003–2005 (%)
Mexico 148 187 209 204 207 195 195 216 213 195 22.64
India 27 45 47 38 39 46 42 179 156 223 20.25
Brazil 24 23 39 54 67 94 104 138 111 114 13.18
Pakistan 18 25 39 41 48 52 48 60 82 49 6.94
Netherlands 21 25 17 37 34 43 33 58 51 69 6.42
Peru 11 6 11 20 21 27 35 40 60 58 5.71
Ecuador 0 2 7 0 26 34 30 38 41 40 4.31
Philippines 40 45 53 35 40 39 36 38 36 25 3.61
Thailand 8 9 10 10 9 11 9 8 33 2 1.55
China 12 7 9 10 5 5 15 22 10 4 1.31
Others 80 104 87 103 132 121 127 126 127 135 14.08
World total 391 478 529 552 628 666 673 923 920 913 100.00
609
610
Table 16.3. World’s top ten major mango-importing countries, 1996–2005 (1000 t) (Source: FAOSTAT, 2007).
Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2003–2005 (%)
The USA is not a major mango producer even though most of the commer-
cially traded varieties have been developed in Florida. USA production,
mainly in Florida, remains fairly stable at a little under 3000 t/year.
The USA is currently the world’s leading importer of fresh mangoes,
accounting for 32.70% of the total imports during the 2003–2005 period
(FAOSTAT, 2007). Figure 16.1 shows the trend of mango imports into the
USA between 1997 and 2006. Overall, the graph indicates a steady increase
in the volume of mango imports. Between 1997 and 2006, imports increased
from 187,193 t to 298,088 t, an average annual growth rate of 5.46%. Mango
imports were valued at about US$233,100,000 in 2006 (USDA, Foreign Agri-
cultural Service, 2007).
The main sources of USA imports of mango are Mexico, Peru, Ecuador
and Brazil. Mexico is the main supplier of mango to the USA (60.78% share
in 2006) (Fig. 16.2). In recent years, Brazil, Peru and Ecuador have become
significant exporters to the USA, competing with Mexico at the beginning
and the end of the season. The USA exports very few of its mango imports,
mainly to Canada and the UK.
612 E.A. Evans and O.J. Mendoza
350,000
300,000
250,000
USA total imports (t)
200,000
150,000
100,000
50,000
0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
Fig. 16.1. USA total imports of mango (t), 1997–2006 (Source: USDA Foreign
Agricultural Service, 2007).
200,000
180,000
160,000
140,000
USA total imports (t)
120,000
100,000
80,000
60,000
40,000
20,000
0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
Fig. 16.2. USA total imports of mango (t) by country, 1997–2006 (Source: USDA
Foreign Agricultural Service, 2007).
Table 16.4. Average cost, insurance and freight (CIF) prices for selected varieties from
main suppliers to the USA, 1998–2006 (US$/kg) (Source: USDA Agricultural Marketing
Service, 2007).
Country of origin
16.3 Sample Costs and Returns Associated with the Establishment and
Production of Mango Orchards
In this section, we have estimated the per hectare costs to establish an 18 ha
commercial mango orchard in Florida USA, as well as the annual estimated
costs and net returns per hectare after establishment. We have not provided
too much detail on production practices (since such information is readily
available elsewhere), but rather focus on the methodology of estimating the
costs and returns of establishment and production. Although somewhat hypo-
thetical, the data presented are based on a combination of information
obtained from the growers, economic engineering using recommended prac-
tices, and discussions with industry experts. We begin by briefly describing
the general methodology, followed by a listing of some of the major assumptions
used in the analysis. Estimates of the establishment costs are then discussed,
and the chapter closes with a discussion of profitability of a mango operation.
614 E.A. Evans and O.J. Mendoza
The general approach for estimating the cost of production of perennial crops
(orchards and vineyards), which usually take more than a year to begin pro-
duction, is to develop two separate budgets: one for the establishment
phase and the other for the production phase. The establishment phase bud-
get reflects the sum total of all expenses (expressed on a yearly and per unit
basis) that are incurred over the years to bring an orchard into meaningful
(mature trees) production. Once this amount is determined, it is treated as if
it were an expense incurred in the purchase of a ‘capital item’, for example
machinery to be used in the production phase of the orchard. As in the case
of a capital item, an equal annual amount (amortized amount) is charged to
the production operation as an expense spread over the estimated future life
of the orchard. In other words, the amortized amount is included as a non-
cash overhead expenditure in the production phase budget when determin-
ing net returns from the enterprise. The production phase budget estimates
the costs and returns on an annual per unit basis associated with the mainte-
nance of the orchard after it has been established. Although the procedure
seems daunting, it is made much easier by using spreadsheet software with
built-in formulas for calculating amortization.
Main assumptions
Land
The hypothetical farm consists of 20 ha. A mango orchard is being estab-
lished on 18 ha, with roads, irrigation system and farmstead occupying 2 ha.
The 18 ha orchard is considered large enough to use machinery and equip-
ment efficiently. The orchard is farmed by the owner with the help of hired
part-time labour.
Site preparation
It is assumed that the land is relatively clear, with no costs being included for
major land preparation such as timber clearing, rock removal or land level-
ling. However, if these operations are required, they should be included. The
main operations considered here are associated with fencing and road con-
struction for travelling and harvesting. These operations are usually done 1
year prior to planting, but costs are shown in the first year.
Year 1 2 3 4 5 6 7+
Hedging and topping operations are carried out immediately after fruit har-
vesting.
Fertilization
During the first 3 years, an N-P-K fertilizer (6% nitrogen) is spread by hand
six times each year. After year 3, the frequency of application decreases to
four times each year. Table 16.5 shows the annual fertilizer rates assumed. In
addition, the trees are given annual nutritional sprays of copper, zinc, man-
ganese and boron. Iron is applied in chelated form as a soil drench two to
three times each year.
Irrigation
Total irrigation costs include the cost of pumping water and irrigation labour.
Water for our irrigation system is supplied from a well; therefore, the cost of
the water is zero. Irrigation costs for individual orchards vary, depending on
the amount of water pumped, pumping system, energy source and irrigation
district.
Harvest
Mango fruits are best harvested using clippers and hand-carried harvesting
bags (about US$0.11/kg). With large trees, fruits are harvested using picking
poles, with or without attached clippers, which are equipped with bags into
which the fruit falls. After harvesting, the fruits are usually piled under a tree
on the ground. The fruit is then loaded onto trucks in the field and hauled to
packing houses for US$0.11/kg.
Year 1 2 3 4 5 6 7+
Labour
Hourly wages for workers are US$10.00 for skilled workers and US$7.00 for
field workers. Adding 34% for the employer’s share of federal and state pay-
roll taxes, insurance and other possible benefits, yields a labour rate of US$13/h
for skilled labour and US$9/h for field labour.
Cash overhead
Cash overhead consists of numerous cash expenses paid during each year
that are assigned to the entire farm, not to a particular operation. These costs
include property taxes, office expenses, capital interest, liability and property
insurance, equipment repairs, sanitation services and crop insurance.
PROPERTY TAXES. In the USA, most counties charge a base property tax rate of
approximately 1% on the assessed value of the property.
SANITATION SERVICES. Sanitation services offer portable toilets for orchards and
cost the farm US$1900/year (US$106/ha). This cost includes delivery and
servicing of a single toilet and washing unit.
Non-cash overhead
Non-cash overhead is calculated as the capital recovery cost for equipment
and other farm investments. Capital recovery cost is the annual depreciation
World Mango Trade and Economics 617
and interest costs for a capital investment. It is the amount of money required
each year to be set aside so that a capital item can be fully replaced at the end
of its useful life. It can be viewed as the value of the portion of the capital
item that was utilized in the production process during that year. The for-
mula for the calculation of the annual capital recovery costs is the PMT Excel
formula:
Capital Recovery = –PMT (I, N, (PP − SV)) + (I)*(SV)
Where:
● PMT = the formula that calculates the payment for a loan based on con-
stant payments and a constant interest rate.
● I = the interest rate used to represent the cost of borrowed capital (in this
case, 5%).
● N = the number of years.
● PP = the purchase price for the capital item.
● SV = an estimate of the remaining value of an investment at the end of its
useful life. In this case, the percentage of remaining value is calculated
from equations developed by the ASAE based on equipment type and
years of life.
Farm equipment on a mango orchard in the region is purchased either
new or used. The study shows the current purchase price for new equip-
ment. The new purchase price is adjusted to 60% to indicate a mix of new and
used equipment.
The orchard is irrigated using a sprinkler irrigation system. The life of
the irrigation system is estimated at 20 years. The irrigation system is installed
before the orchard is planted and includes costs of a pumping system, instal-
lation labour and design and materials.
Although it is assumed that the grower owns the land, the going rental
rate in South Florida of approximately US$2500/ha is used to reflect the
opportunity (economic) cost of land.
Table 16.7 shows the layout of the establishment phase budget. The budget
reflects the annual per hectare costs incurred in establishing a mango orchard.
Here it is assumed that a mango orchard reaches maturity in the seventh
year. As such, the budget reflects all associated cash and non-cash expenses
incurred over 6 years.
The information required for each year is similar except for the first 2 or
3 years. With reference to Table 16.7, year 1 includes pre-planting costs (site
preparation and orchard layout), which in this case amounts to US$5000/ha.
Most of the pre-planting costs most likely would occur in the year before, but
the practice is to include these costs in year 1. Planting costs include new
trees, digging holes for trees, setting trees, and wrapping and staking trees.
Both pre-planting and planting costs appear only in year 1. Total per hectare
618
Table 16.7. Sample costs to establish a mango orchard in south Florida (source: compiled by the authors).
Year (US$/ha)
Pre-planting costs
Site preparation 4,500
Orchard layout 500
Total pre-planting costs 5,000
Planting costs
Mango trees (175 trees/ha) US$25.00 4,375
619
Total net cost for the year 17,228 6,037 5,318 5,334 3,295 –361
a FOB, freight on board.
620 E.A. Evans and O.J. Mendoza
Table 16.8 provides the estimates of the annual growing costs (after establish-
ment) for mango in south Florida. The budget is similar to the one used in the
establishment phase except that there are no provisions for pre-planting,
planting and replanting expenses, and the estimates are for a typical year.
Also of importance, and shown in italics in Table 16.8, is the amortized estab-
lishment cost (US$2615/ha) obtained from Table 16.7. This reflects the annual
amount that must be recovered for the investment made in establishing an
orchard.
Based on Table 16.8, it can be seen that the total annual cash cost to pro-
duce a hectare of mango, assuming a yield of 22,000 kg/ha (Table 16.6) is
622
Table 16.8. Sample cost per hectare to produce mango in south Florida.
Costs (US$/ha)
Non-cash overhead
Land rent 2,500 2,500 2,500
Machinery and equipment 3,648 405 405
Building 2,964 185 185
Tools (shovels, picking bags, saws, etc.) 217 20 20
Shop tools 464 42 42
Irrigation system 4,817 380 380
Sprinklers 20 20 20
Amortized establishment cost 2,615 2,615
Total non-cash overhead costs 14,630 6,167 6,167
Total cost/hectare 18,774
623
624 E.A. Evans and O.J. Mendoza
Profitability analysis
Table 16.9 utilizes the information presented in Tables 16.6 and 16.8 along
with information obtained elsewhere to analyse the profitability of mango
production in South Florida based on 2006 data. Based on an average freight
on board (FOB) price of US$0.90/kg and marketable yield of 22,000 kg/
ha (actual yield would be slightly more), the gross returns are calculated as
US$19,800/ha. Subtracting total operating costs of US$10,962 (cultural, and
harvesting and marketing costs) from this amount gives a gross margin of
US$8838/ha. In other words, based on these assumptions, the enterprise is
profitable in the short run since gross margin is positive. This implies that the
returns from crop sales are more than sufficient to cover the variable costs
incurred in the production and marketing of the crop and there are still funds
left over to go towards paying overhead costs.
To be viable in the long run, the amount remaining should be able to
cover overhead (cash and non-cash) and provide a reasonable return to the
operator. Subtracting cash overhead costs (US$1645) and non-cash overhead
costs (US$6167) from gross margin (US$8838) gives net returns of US$1026/
ha. Viewed slightly differently, from total revenue of US$19,800 obtained from
selling a hectare of the crop, after paying all costs (variable and fixed) the
amount of US$1026/ha remain as payment to management. This results in a
return of approximately 7% on cash and non-cash investment.1
Given that prices and yield can vary, Table 16.10 shows a sensitivity anal-
ysis for changes in price and yield variables. The sensitivity analysis is cal-
culated on net returns but could be done on gross margin. As seen in Table
16.10, a 10% price increase combined with a 10% yield increase would result
in over a 335% net return per hectare increase (from US$1026 to US$4460/
ha). A 10% price increase would have a greater impact on net return than
would a 10% yield increase. In the case of the former, the net returns would
increase from US$1026/ha to US$2984/ha (191%). In the case of the latter, net
returns would increase to US$2324 (127%). This underscores the importance
of doing whatever is necessary to improve crop prices such as improving the
quality of the product and engaging in direct marketing where possible.
Assuming marketable yields of 22,000 kg/ha, the break-even price, or
the price at which net returns (economic profits) would be zero, is computed
as US$0.85/kg. In other words, at this price the grower is just able to cover
both variable and fixed costs. Any FOB price above this would result in pos-
itive net return and vice versa. Likewise, if prices were to remain at US$0.90/
kg, growers would have to ensure that they obtain yields in excess of about
20,000kg/ha (break-even volume) to generate positive returns.
World Mango Trade and Economics 625
Table 16.9. Costs and returns to produce mango in south Florida, 2007.
Price or Value or
cost per cost/ha
Unit Quantity/ha unit (US$) (US$)
(Continued)
626 E.A. Evans and O.J. Mendoza
Price or Value or
cost per cost/ha
Unit Quantity/ha unit (US$) (US$)
16.4 Conclusions
The major trends and developments occurring within the world and USA
mango market have been discussed and the methodology used to compute
the costs of establishing and maintaining an orchard has been demon-
strated. Several trends were highlighted, including the increasing levels of
production, trade and consumption of mangoes and the declining or stag-
nating prices. Although the case for establishing an orchard was hypotheti-
cal, the statistics are based on information obtained from growers, economic
engineering using recommended practices, and discussions with industry
experts.
Acknowledgements
The authors are indebted to many farmers and agricultural researchers who
provided us with innumerable advice and relevant comments about this
study. We wish to thank those who kindly offered suggestions for improve-
ment, including Dr Jonathan Crane, Dr Carlos Balerdi, Scott Smith, Luis D.
Roman, Sikavas Nalampang, Denys Benjamin and Valderez Gonzalez. A spe-
cial thanks to Carol Fountain for editing the manuscript.
World Mango Trade and Economics 627
Note
1This is obtained by adding the interests charged on operating and cash overhead costs
to the net returns to give an adjusted net income. Next, subtract these interests from the
total costs per hectare to give an adjusted total cost per hectare. Finally, express the
adjusted net income as a percentage of the adjusted total cost per hectare.
References
FAOSTAT (2007) Food and Agriculture Organization of the United Nations database.
Available at: http://faostat.fao.org/ (accessed 30 November 2007).
United States Department of Agriculture (USDA) Agricultural Marketing Service (2007)
USDA, Agricultural Marketing Service: Fruit and Vegetable Market News Website.
Available at: http://www.marketnews.usda.gov/portal/fv (accessed 31 March 2008).
United States Department of Agriculture (USDA) Economic Research Service (2007)
Fruit and Tree Nut Yearbook, 2005. Available at: http://www.ers.usda.gov/Data/
FoodConsumption/FoodAvailSpreadsheets.htm (accessed 31 March 2008).
United States Department of Agriculture (USDA) Foreign Agricultural Service (2007)
USDA Foreign Agricultural Service: Market and Trade Data. Available at: http://
www.fas.usda.gov/markettradedata.asp (accessed 3 November 2007).
17 Fruit Processing
17.1 Introduction
Processing is a value-adding step that fills the gap between farm production
and marketing. The objective of processing is to convert highly perishable
but prime quality farm commodities to more stable forms. When the process
is accomplished with the least alteration in the nutritional value and aes-
thetic properties of the food, high consumer acceptance is assured. Com-
pletely new product lines can likewise be created out of fresh raw materials
through processing. In other instances, the raw material may undergo such
extensive physical alteration during processing that the product is used dif-
ferently by consumers. Culinary experts devise new uses for these products
to fit the changing lifestyles of present-day consumers. The availability in the
market, for example, of pre-mixed condiments and various meat powders
used for flavouring foods such as instant noodles and similar convenience
© CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
628 (ed. R.E. Litz)
Fruit Processing 629
foods, has simplified the life of working women who do not have the time
or the skill to prepare food the way their mothers and grandmothers did
at home. The products are affordable and easy to prepare; hence, consumer
acceptance is high. In addition, these foods are found everywhere, from
neighbourhood stores in the suburbs and countryside to supermarkets in
urban centres. The product and process research and development work of
food scientists and food technologists, the business acumen of entrepreneurs
and the marketing expertise of product distributors ensure the availability of
the food products.
The global competitiveness of agricultural produce of a country can be con-
siderably enhanced by utilizing appropriate technologies to produce high-
quality processed foods. Research focusing particularly along the areas of
processing equipment, product design and packaging has made the latest
techniques available for the manufacture of new generations of food prod-
ucts. Tropical fruits have had an excellent record of breaking into the world
market because of their exotic flavours (Plate 83). When processed into purée,
juice, nectar or simply dried fruit slices, they can be shipped long distances
with only minimum changes in quality.
The popularity of fruits may be attributed to consumer perception of
their health benefits. In the case of fruit juices, many consumers are now
looking for healthy alternatives to the traditional carbonated beverages. Fruits
are rich in dietary fibre as well as phytonutrients, especially antioxidants,
and have no cholesterol. The availability in the market of natural fruit juices
derived from fresh fruits is a welcome alternative to synthetic juices that are
being passed off as fruit juices but are, in reality, sugar-based, fruit-flavoured
beverages prepared from artificial ingredients.
Mango fruits are usually eaten fresh as dessert or as relish depending on
their stage of maturity when picked. In the Philippines and elsewhere, fresh
mangoes are available throughout the year because the flower-induction
technology first described by Barba (1974) allows growers to produce fruit
out-of-season. The off-season fruits, however, are still expensive. Most mango
growers have not put the technology into practice due to the high cost of
additional farm inputs necessary for its successful implementation during
the rainy season.
At the peak of the harvest season, on the other hand, oversupply of
fresh mangoes depresses the market price to the detriment of the growers.
Moreover, high temperatures combined with high relative humidity (RH)
and intense sunlight during the harvest season accelerate the metabolic
processes associated with ripening in fresh mangoes, rendering them sus-
ceptible to microbial attack, particularly by Colletotrichum gloeosporioides
Penz., the cause of anthracnose. The physico-chemical changes that occur
during ripening also lead to fruit deterioration and loss of quality. Thus,
fresh mangoes are processed to facilitate better distribution and to stabilize
prices.
Mango processing was previously reviewed by Nanjundaswamy (1997),
who described the status of processing technologies and products in India.
This review discusses current trends in mango processing.
630 L.C. Raymundo et al.
Dried mango
Dehydrated or dried mango slices are among the first products manufac-
tured commercially from ripe mango fruits that caught the attention of con-
sumers in the international market for processed tropical fruits (Plates 83 and
84). The product was developed in Cebu, the Philippines, from where it
spread to the neighbouring islands of Panay and Negros. It is now produced
in many regions of the Philippine archipelago where mango is abundant. In
addition, it is a popular product in Thailand and elsewhere in South and
South-east Asia.
In the Philippines, the ‘Carabao’ mango is the preferred variety for dehy-
dration or drying. The fruit should be at the firm-ripe stage. When over-ripe
fruit is used as raw material, a dark-coloured product will invariably result.
Although the dried pieces from over-ripe mangoes have a more distinct ripe
mango flavour that attracts customers, the shelf life is considerably shorter.
Fruit Processing 631
The fruit is washed thoroughly. The cheeks are sliced with a sharp
stainless-steel knife. Each slice is then cut into two or three pieces length-
wise. The flesh is scooped from the skin with a stainless-steel scoop or ladle.
These operations are done manually; however, a simple and novel peeling
and slicing machine for ripe mangoes has been developed and patented in
Australia (as cited by Nanjundaswamy, 1997).
Sugar syrup is prepared by adding 175 kg sugar, 1.7 kg citric acid and
0.85 kg sodium metabisulfite in 175 l H2O to make a 45 Brix sugar concen-
tration and heating to 90°C to dissolve the metabisulfite. The prepared
mango slices (1 t) are added to the syrup. The preparation is heated to 90°C
and then allowed to stand for at least 6 h. Subsequently, the sugar concen-
tration of the syrup is adjusted by dissolving more sugar until the concen-
tration reaches 50 Brix using a hand refractometer. After 6 h, the mango
pieces are again removed from the syrup and the sugar concentration is
adjusted to 60 Brix using a hand refractometer for the ‘plumping’ stage.
The syrup is reheated to 90°C; the mango slices are added to the syrup and
soaked for a further 6 h.
The final step in the process involves the removal of mango pieces from
the syrup. They are lightly rinsed with H2O to remove the excess sugar that
may crystallize on the surface of the mango during drying. The slices are
then loaded in drying trays and dehydrated at 40–50°C in a convection dryer.
Drying time usually requires 18 h. The dried mango pieces are unloaded
from the dryer and reconditioned at room temperature overnight. Each
piece is coated with confectioner’s sugar. The product is then packed in
aluminium-foil-laminated pouches and sealed. Recent improvements in
dryer design can reduce drying time to 8 h. As a result, up to three batches of
prepared mango slices can be processed daily instead of only two batches
every 36 h. The process for the production of dried mangoes has been
described by Raymundo et al. (1999).
The product is also commonly referred to as mango ‘leather’ (Plate 83). The
processing of mango fruit bar has also been described previously by Amor-
iggi (1992) and Raymundo et al. (1999, unpublished work, 2006). Purée pro-
cessed from ripe ‘Carabao’ mango is the preferred raw material for the
manufacture of mango fruit bar in the Philippines, although ‘Pico’ is also
suitable because of the orange colour of the purée.
The total solids content of 1 t of mango purée is adjusted to 25 Brix with
the use of a hand refractometer by adding sugar to the purée. The amount of
sugar required depends on the initial total solids content of the mango purée.
Then 2 kg each of citric acid and sodium metabisulfite are blended into the
purée. Juice of calamansi, also known as the Philippine lemon (× Citrofor-
tunella microcarpa Wij. (Bunge); Citrus mitis Blanco) may be used instead of
citric acid at the rate of c.20 kg per batch, although the resulting cost of the
product will be slightly higher. There is no real difference in the appearance
632 L.C. Raymundo et al.
and flavour of the finished product. Citrus juice is generally utilized if the
client prefers an all-natural product.
The prepared purée is heated for 2 min at 80°C with constant stirring to
avoid scorching. The hot mixture is carefully transferred to stainless-steel
drying trays. The trays are loaded into the dryer and dried for 14 h at 55°C.
At the end of the drying operation, the dried sheets of mango should be pli-
able and should not stick to the fingers when touched.
The sheets of mango are removed from the trays. Six layers of the dried
mango leather are stacked on top of each other. The edges are trimmed and
bars measuring 2 × 4 cm are cut from the stack. Each bar is coated with con-
fectioner’s sugar, wrapped in cellophane then packed in labelled plastic bags.
The processing of mango fruit roll has been described previously (UPLB,
1996; Raymundo et al., 1999, unpublished work, 2006). The product and pro-
cess are similar to mango fruit bar, and they only differ in the presentation.
The total solids content of 1 t of mango purée is adjusted to 25 Brix using a
hand refractometer by adding sugar to the purée. The amount of sugar
needed depends on the initial total solids content of the mango purée. Then
2 kg each of citric acid and sodium metabisulfite are blended into the purée.
As with mango fruit bar, citric acid may be replaced by calamansi juice at the
rate of c.20 kg per batch without affecting the overall sensory quality of
the fruit roll, if the client specifies an all-natural product.
The prepared purée is then heated for 2 min at 80°C with constant stir-
ring to avoid scorching. The hot mixture is carefully transferred to stainless-
steel drying trays. The trays are loaded into a convection dryer and dried for
12–16 h at 55°C.
The dried sheets of mango are removed from the trays. Confectioner’s
sugar is sprinkled over a stainless-steel-topped table to avoid sticking of
the sheets on subsequent rolling. Each sheet is rolled manually into 1 cm
diameter pieces. The ends are trimmed; and each roll is sliced into 5 cm long
pieces. The pieces are coated with confectioner’s sugar and wrapped with
cellophane. The rolls are then packed in appropriate containers.
The use of vacuum for puffing and drying mango and similar food materials
is not as widespread as explosion puffing (Eskew et al., 1963), high tempera-
ture short time (HTST) pneumatic drying and centrifugal fluidized bed (CFB)
drying (Brown et al., 1972), because the vacuum-puff dryer is still expensive.
With the increasing consumer demand for high-quality processed foods and
their willingness to pay a higher price for such products, vacuum-puff dehy-
dration could become an economically viable investment opportunity for
entrepreneurs, particularly in the mango processing industry.
Fruit Processing 633
Khalid and Sial (1974), Anonymous (1998) and Diaz (2000) have discussed
the recovery of fruit powder and production of instant mango juice powder
Fruit Processing 635
using the technology. Both products use mango purée as the raw material.
They differ only in the composition of the liquid feed. The liquid feed is
mango purée with the total solids adjusted to the right consistency, thereby
allowing the purée to be discharged through a high-speed nozzle in the form
of fine droplets into the drying chamber that quickly dries to a yellow free-
flowing powder. The patent application for the manufacture of spray-dried
mango powders is still pending at the Philippine Patent Office. The process
parameters used, therefore, cannot be discussed in detail in this chapter. The
parameters used are the same as reported previously, namely, the inlet tem-
perature, the feed rate, air speed and the outlet temperature (Welti and Laflu-
ente, 1983; Liang and King, 1991; Anonymous, 1998). The powder of instant
mango juice is comparatively dense so that it gets wet easily, enabling it to
sink immediately during reconstitution. The plain mango fruit powder, on the
other hand, does not disperse as readily; it has a tendency to clump on the H2O
surface. However, it will dissolve quickly in hot H2O or with the use of an
ordinary household blender.
A modest capacity spray-drying plant equipped with a NIRO SD 12.5R
model spray-dryer made in Denmark, valued at approximately US$207,860,
with an evaporative capacity of 25 kg H2O/h will require an initial invest-
ment of about US$313,860 to make it operational. In addition, if fresh man-
goes are be used as raw materials, a purée processing facility will cost around
US$98,000. The figure represents expenses for buying and installation of
equipment, and does not include the costs of the building, land and ancillary
expenses such as environmental pollution control system and spray-dryer
accessories.
At the evaporative capacity rate of 25 kg H2O/h, the facility can produce
124.8 t/year of plain mango powder and 202.8 t/year of instant mango juice
at a total cost of US$251,460 and US$467,355, respectively (Table 17.1). The
net profit for plain mango powder is US$439,983 and US$965,088 for instant
mango juice before taxes.
The purée of green mangoes can be converted to a powder (UPLB, 2005) just
like the purée of ripe mangoes by spray-drying (Plate 85a). The spray-dried
powder can be mixed with other condiments and used as a souring agent for
exotic or native dishes, or as the raw material in the manufacture of instant
green mango shake (see below). The powder may be dry-mixed with sugar,
powdered honey, caramel powder or powdered sugar syrup to instantize it.
During this process, the mixture can be fortified with vitamins (e.g. ascorbic
acid) and other nutrients.
The estimated cost of goods sold using a commercial spray-dryer with
an evaporative capacity of 25 kg H2O/h is US$245,226/year (Table 17.1). At
the rated capacity of the spray-drying plant of 141,221 kg/year, the net profit
before taxes for green mango fruit powder is US$600,216 at a selling price of
US$10/kg of powder.
636 L.C. Raymundo et al.
Green mango shake on cracked ice is a very popular thirst quencher in South-
east Asia and elsewhere (Plate 85b). Its origin can be traced back to the coun-
tryside where finely diced fresh green mango pieces are mixed with H2O,
sugar and ice to make a cheap, wholesome summer drink during the mango
season. The practice has since been modified and upgraded to cater to upscale
domestic markets and abroad.
The beverage is an excellent source of vitamin C. When the green mango
purée is spray-dried, a free-flowing white to greenish-white powder is pro-
duced that will dissolve instantly even in cold H2O. The powdering process
offers unprecedented convenience to the consumer, especially when the pow-
der is packed in sachets. Since no artificial or synthetic colours and flavour-
ing agents are included in the liquid-feed formulation, the natural taste and
aroma of green mango is retained in the powder.
Using a commercial spray-dryer with an evaporative capacity of 25 kg
H2O/h, the estimated cost of goods sold is US$209,468/year. At the rated
capacity of spray-drying plant of 124,800 kg, the net profit before taxes for
instant green mango shake is US$473,975/year at a selling price of US$10/kg
of powder (Table 17.1).
The technology for the production of instant green mango shake and
green mango powder was developed at UPLB (2005). If the proper feed for-
mulation and process parameters are applied, spray-drying is an efficient
and hygienic method for producing cheap but high-quality mango fruit
powder and instant mango juice. Table 17.2 demonstrates that the spray-
dried mango powders have much lower microbial load, i.e. 2.8–10.4 × 102
colony forming units (cfu)/g, compared to the industry standard of 1 × 104 to
5 × 105 cfu/g total plate count. Except for mango powder, the mould and
yeast counts are within the limit of 1 × 102 cfu/g. Locally refined sugar and
imported modified starch, on the other hand, have much higher mould
and yeast counts than the Heinz standard for powdered products (Shapton
Table 17.2. Microbial load of raw materials and spray-dried mango fruit powder and instant
mango juice (cfu/g).
and Shapton, 1991) at 11 × 102 and 63 × 102 cfu/g total plate count, respec-
tively. These materials are essential ingredients of the liquid-feed formula-
tion used for the production of the spray-dried powders.
The formulation and processing of numerous mango products popu-
lar in South Asian countries were reported previously (Nanjundaswamy,
1997).
Evaporative rate
Table 17.4. Estimated volume of raw material needed for the operation of a mango
spray-drying and a mango dehydration plant.
17.6 Conclusion
Processing of mango is a profitable business venture once economies of scale are
attained, i.e. when the processor has the proper proportions of raw materials,
labour and machinery to meet a given market demand. Fresh mangoes are now
available year-round. But the supply is still insufficient to satisfy the demand by
the fresh fruit market and the mango-processing sector. It is, therefore, essential
that a farming system for mango be designed in order to minimize the cost of
production of off-season fruits and ensure the sustainable operation of mango
processing facilities. Since raw material sourcing is the primary cause of the dif-
ficulties encountered by processors of dried mangoes, mango in syrup and
similar products, growers need more assistance for them to adopt the tech-
nology for off-season mango flower induction. The problem is not as serious
for product lines utilizing mango purée as starting material since the purée
can be processed during the peak of the harvest season and held in cold stor-
age for later use. Once the system is in place, the processed mango industry
is expected to develop further and become a major revenue generator.
In the near future, with the advances in the field of genetic engineering,
it may be possible to eliminate the biennial fruiting habit of many current
mango cultivars or, at the very least, minimize its influence on the perfor-
mance of the crop. Species of Mangifera and some non-cultivated, wild types
of mango can be the source of desirable traits that may be incorporated in the
next generation of mango cultivars, such as their innate ability to bear fruits
during the rainy season (see Bompard, Chapter 2, this volume).
The technologies for processing mangoes are readily available. Others
are being developed in research laboratories to cope with the changing needs
of consumers. The main problem is to ensure a continuous supply of high-
quality fresh mango fruit in order to produce the prime quality commodities
that consumers expect from the industry.
640 L.C. Raymundo et al.
References
Amoriggi, G. (1992) The marvelous mango bar. Ceres 24, 25–28.
Anonymous (1998) Instruction Manual for Spray Drying Plant. NIRO A/S Soeborg, Denmark.
Barba, R.C. (1974) Induction of flowering of the mango by chemical spray. Proceedings
of the Crop Science Society of the Philippines 5, 154–160.
Brown, G.E., Farkas, D.F. and De Marchena, E.S. (1972) Centrifugal fluidized bed.
Blanches, dryers and puff piece-form foods. Food Technology 26, 23–30.
Candelaria, N.M. (1993) Dehydration of vacuum-puffed fruits and vegetables. PhD dis-
sertation, University of the Philippines Los Baños, Laguna, the Philippines.
Candelaria, N.M. and Raymundo, L.C. (1994a) Comparative drying and reconstitution,
characteristics of some fruits and vegetables. Philippine Agriculturist 77, 321–326.
Candelaria, N.M. and Raymundo, L.C. (1994b) Vacuum puffing and dehydration of fruits
and vegetables. Philippine Agriculturist 77, 251–260.
Diaz, J.V. (2000) Development of spray-dried instant mango (Mangifera indica L.) juice pow-
der. MSc thesis, University of the Philippines Los Baños, Laguna, the Philippines.
Eskew, R.K., Cording, J., Jr and Sullivan, J.F. (1963) A gun for explosive, puffing of dehy-
drated fruits and vegetables. Food Engineering 35, 91–96.
Khalid, M.A. and Sial, M.B. (1974) Spray-drying of mango juice powder. Mesopotamia
Journal of Agriculture 9, 47–56.
Liang, B. and King, L.J. (1991) Factors influencing flow patterns, temperature and conse-
quence drying rates in spray-drying. Drying Technology 9, 1–27.
Nanjundaswamy, A.M. (1997) Processing. In: Litz, R.E. (ed.) The Mango: Botany, Pro-
duction and Uses. CABI International, Wallingford, UK, pp. 507–542.
Raymundo, L.C., Ombico, M.T. and De Villa, T.M. (1999) Establishment of integrated
mango processing plant. In: Namuco, L.O. and Andam, C.J. (eds) Mango Production
Manual. Philippine Council for Agriculture, Forestry and Natural Resources Research
and Development (PCARRD), Los Baños, Laguna, the Philippines, pp. 97–119.
Raymundo, L.C., Ombico, M.T., De Villa, T.M. and Jaen, R.M. (2006) Processes of Fruits,
Nuts, Vegetables and Other Tropical Foods. Fruit and Vegetable Laboratory, Food
Science Cluster, University of the Philippines Los Baños, Laguna, the Philippines.
(Unpublished)
Shapton, D.A. and Shapton, N.F. (1991) Principles and Practices for the Safe Processing
of Foods. Butterworth-Heinemann Ltd, Oxford, UK.
University of the Philippines Los Baños (UPLB) (1996) Standardization of Processing
Method and Evaluation of Different Packaging Material and Storage Conditions for
Mango Roll. Annual Report, Institute of Food Science and Technology, College of
Agriculture, University of the Philippines Los Baños, Laguna, the Philippines.
University of the Philippines Los Baños (UPLB) (2005) Spray-dried Instant Juice and
Powder from Green Mango. Annual Report, Food Science Cluster, College of Agri-
culture, University of the Philippines Los Baños, Laguna, the Philippines.
Welti, J.S. and Lafluente, B. (1983) Spray-drying of comminuted orange products. I. In-
fluence of air and temperature and feed rate on product quality. Revista de Agro-
nomica y Technologia de Alementos 23, 97–106.
18 Biotechnology
18.1 Introduction
embryo and organ development, including shoots, leaves, roots, flowers and
fruit. Genomic analysis and the identification of horticulturally important
genes that control these and other processes will ultimately enable us to under-
stand and control many aspects of mango fruit production (i.e. disease and
pest control, fruit quality, tree architecture, etc.). The development of molecu-
lar approaches will also facilitate mango breeding through the identification
of DNA markers for important traits.
This chapter is a discussion of the current state of mango cell culture,
gene cloning and the manipulation of cell cultures to address plant breeding
objectives by genetic engineering.
Organogenesis
Somatic embryogenesis
Genetic engineering of mango has been based upon the efficient recovery of
somatic embryos from embryogenic cultures. Efficient regeneration of woody
plants from cell cultures derived from mature phase materials is often difficult
to achieve, and the optimum procedure generally must be determined empir-
ically. Mangifera indica consists of two ecogeographic races: a monoembryonic
644 R.E. Litz et al.
Induction
The induction of embryogenic competence is associated with the develop-
mental stage of the nucellus at the time of explanting, and also can be af-
fected by the physiological status of the tree (Litz, 1987). Fruits that are
approximately 30–40 days after pollination contain seeds in which the nucel-
lus is at the ideal stage for explanting: relatively thick and intact and easily
removed. The embryo (mass) in fruit and developing seed of this age is usu-
ally no more than half the length of the immature seed. Mango fruit of the appro-
priate stage of development are surface-disinfested with 20–30% (v/v) domestic
Biotechnology 645
Country Reference
bleach containing Tween 20 for 30 min. The sterilant is rinsed from the fruit
with three changes of sterile deionized or distilled water, and each fruit is
bisected along its longitudinal axis without damaging the immature seed
under axenic conditions. The immature seed is removed from the bisected
fruit, which is also bisected carefully along its longitudinal axis. Manza-
nilla Ramirez et al. (2000) obtained optimum induction with polyembryonic
‘Ataulfo’, monoembryonic ‘Tommy Atkins’ and monoembryonic ‘Haden’
nucellar explants when the embryo (mass) to immature seed ratio was 1:3.
The zygotic embryo (of a monoembryonic cultivar) or polyembryonic mass
(of a polyembryonic cultivar) is excised and discarded. The nucellus can be
carefully peeled from the interior of the seedcoat using a sterile, flat spatula.
Following the transfer of the nucellus onto induction medium in sterile Petri
dishes, the cultures are incubated in darkness at 25°C. Thereafter, it is essen-
tial to subculture the nucellar explants onto fresh induction medium at daily
intervals until the oxidation of the explant ceases; oxidation is associated
with darkening of the medium around the explanted nucellus.
A summary of those mango cultivars that have been successfully estab-
lished as embryogenic cultures is provided in Table 18.2. The efficiency of
646 R.E. Litz et al.
Maintenance
Embryogenic mango cultures are friable and cream to light brown in
colour, although the cultures rapidly darken on semi-solid medium, and
must therefore be subcultured at 3–4 week intervals. PEMs develop from
globular somatic embryos in the presence of the primary induction agent,
2,4-D. The PEMs originate as globular somatic embryos, but their devel-
opment as individual somatic embryos is arrested in the presence of 2,4-D.
The PEMs increase in diameter with cells of the protoderm dividing rap-
idly; secondary globular somatic embryos develop from these proliferating
embryogenic cells. This highly repetitive pattern of somatic embryogen-
esis in the presence of 2,4-D is the basis for maintenance of embryogenic
cultures.
Embryogenic mango cultures can be maintained as proliferating PEMs
either on semi-solid or in liquid induction medium. Maintenance of embryo-
genic cultures of many cultivars is optimal in liquid induction medium sup-
plemented with 4.8 PM 2,4-D (Litz et al., 1984; DeWald et al., 1989a) (Plate 87);
however, rapid proliferation of embryogenic suspension cultures is cultivar-
dependent (Litz et al., 1993). Embryogenic suspension cultures are initiated
by inoculating approximately 400 mg of PEMs into sterile 80 ml maintenance
medium in 250 ml Erlenmeyer flasks (or 200 mg of PEMs into 40 ml mainte-
nance medium in 125 ml Erlenmeyer flasks). The flasks are maintained on a
rotary shaker at 100–125 rpm in semi-darkness at 25°C with regular transfers
of PEMs into fresh medium at 10–14 day intervals. Regular subculturing is
essential to prevent loss of morphogenic potential and darkening of the tis-
sue. The typical embryogenic suspension culture consists of PEMs, embryo-
genic cells and multicellular complexes.
Maturation
Development of somatic embryos from embryogenic cultures maintained on
semi-solid maintenance medium occurs sporadically and without synchroni-
zation, due to the lack of direct contact of parts of a culture with medium
containing 2,4-D. Exposure to 2,4-D is necessary for embryogenic culture
proliferation, while at the same time, somatic embryo development is
648 R.E. Litz et al.
somatic embryos. The medium that has been utilized for development of
mango somatic embryos to maturity consists of B5 major salts, MS minor
salts and organic components, 400 mg/l glutamine, 20% (v/v) filter-sterilized
CW, 40 g/l sucrose and 2.0 g/l gellan gum (DeWald et al., 1989b). As the
somatic embryos enlarge, the sucrose concentration of maturation medium is
gradually reduced to 10 g/l.
During somatic embryo development certain developmental anomalies
become apparent, of which the most frequently observed are polycotyly, fas-
ciation, absence of bipolarity, secondary somatic embryogenesis from the
hypocotyl and precocious germination. Polycotyly and fasciation do not
affect subsequent plant development; however, failure to address problems
associated with absence of bipolarity, secondary embryogenesis and preco-
cious germination can seriously impact the recovery of plants. Control of
precocious germination of developing embryos is occasionally necessary,
since immature somatic embryos that germinate before they are physiologi-
cally mature cannot survive. Some of these developmental anomalies can be
eliminated by maintaining relatively high sucrose concentrations and/or by
incorporating 100 PM ABA in the maturation medium (Monsalud et al., 1995;
Pliego-Alfaro et al., 1996b). The cultures are maintained in darkness at 25°C
during somatic embryo maturation.
Germination
When somatic embryos begin to germinate, they are finally transferred to
light conditions. The radicle elongates, followed by growth of the taproot.
The shoot apical meristem remains quiescent for approximately 2 weeks
after germination, at which time the shoot elongates. Although many mango
somatic embryos germinate under these conditions, their survival or conver-
sion ex vitro is low, primarily due to apical shoot necrosis, a physiological
disorder that is associated with calcium ion (Ca++) deficiency. Different strat-
egies have been attempted to improve the conversion rate (i.e. survival of
somatic embryo-derived plants):
1. The period for embryogenic cultures in/on maintenance medium should
be minimal (Litz and Lavi, 1997).
2. Ara et al. (1998) rescued in vitro microshoots obtained from germinated
somatic embryos by pulsing them for 24 h with 24.6 PM IBA in liquid medi-
um followed by transfer to auxin-free medium in darkness for root develop-
ment.
3. Somatic embryo shoots have also been rescued by micrografting the
shoots on decapitated in vitro-germinated seedling rootstocks (Plate 90).
4. Enhanced recovery of mango plantlets can occur following the induction
of photoautotropism by transfer of small plantlets onto minimal plant growth
medium, containing <5% sucrose and 1% (w/v) activated charcoal (Litz et al.,
1993). A filter-sterilized air mixture consisting of 20,000 ppm carbon dioxide
(CO2) in a nitrogen gas carrier is introduced into the growing containers, and
the plantlets are exposed to a 16 h photoperiod at 180 Pmol/s/m2 provided
by cool white fluorescent tubes.
650 R.E. Litz et al.
microcarpa (Chen et al., 1980), Dimocarpus longan (Yang and Wei, 1984) and
Litchi chinensis (Fu and Tang, 1983). Mangifera indica is thought to be an auto-
tetraploid (2n = 4x = 40), and recovery of diploid (n = 2x = 20) plants would
reveal the nature of the ancestral species and simplify genomic analysis.
Chromosome doubling would restore autotetraploidy, and somatic hybrid-
ization of diploid mangoes with wild Mangifera species would allow interest-
ing genetic recombination.
Gene cloning
can be detected. For that reason, the molecular analysis of fruit ripening
requires construction of a gene library. In mango, cDNA libraries have been
constructed, mainly from ripe fruit, and screened using several approaches.
The mRNA for virtually all the ripening-related genes isolated so far have
been shown to be absent or at a low level in immature fruit, increase during
ripening and decline as ripening progresses (Giovannoni, 2001). Unlike other
fruits, none of the identified genes in mango code for enzymes involved in
the ripening process itself (see below).
Mango fruit are highly perishable commodities due to over-ripening,
which is mainly caused by the sharp increase in ethylene production, which
occurs simultaneously with the climacteric peak (Tucker and Grierson, 1987;
see Brecht and Yahia, Chapter 14, this volume). Mangoes have poor storage
quality and storage in controlled or modified atmospheres has been associ-
ated with physiological disorders (Chaplin, 1989). Genetic manipulation of
mango fruit ripening represents an attractive alternative to extend storage
life and, therefore, the isolation of mango genes coding for enzymes involved
in ethylene biosynthesis has been a target for research.
The two key enzymes in the ethylene biosynthetic pathway are those
catalysing the conversion of S-adenosylmethionine (SAM) to 1-aminocyclo-
propane-1-carboxylic acid (ACC) and ACC to ethylene, i.e. ACC synthase
and ACC oxidase or ethylene forming enzyme (EFE), respectively and cDNA
clones coding for mango ACC synthase and ACC oxidase have been identi-
fied (Gómez-Lim, 1993). Their expression during ripening was studied in
pulp and peel in northern blot analysis-type experiments. The ACC synthase
message is undetectable in unripe fruit and starts to appear in turning fruit,
reaching a maximum in ripe fruit (Gómez-Lim, 1993). This pattern of expres-
sion is similar in the peel and in the pulp; however, the message appears in
the pulp before the peel. The ACC oxidase message shows similar kinetics in
both types of tissue, but the message is clearly detectable before any ACC
synthase message becomes detectable (Gómez-Lim, 1993). These results sug-
gest that ACC oxidase is expressed before ACC synthase and that ripening
starts on the inside of mango fruit and proceeds outwards. Ethylene-treated
mango fruits show a different pattern of expression, with ACC oxidase and
ACC synthase appearing initially in the peel.
If ethylene is being actively produced, the gas must be clearly perceived
by plant cells and therefore ethylene sensitivity is important for the ripening
process. Major advances in understanding ethylene signal transduction have
come from a molecular genetic approach using ethylene responsive mutants
of A. thaliana and tomato. Several genes coding for ethylene receptor homo-
logues have been isolated from various plants (Johnson and Ecker, 1998).
Genetic manipulation of the ethylene receptor represents an interesting alter-
native to control ethylene production and delay ripening, particularly in
those fruits where other alternatives, such as storage in controlled atmo-
spheres, have not been effective (Wilkinson et al., 1997). A cDNA coding for a
mango ethylene receptor has been isolated (Gutiérrez-Martinez et al., 2001).
The message seems to be present at low levels in unripe fruit and to increase
Biotechnology 653
Genomics
The human genome project has been the catalyst for the development of sev-
eral high-throughput technologies that have made it possible to map and
sequence complex genomes. Many bacterial genomes and the genomes of
Sacchromyces cerevisiae, Caenorhabditis elegans, Drosophila melanogaster, A. thali-
ana, Oriza sativa and Populus trichocarpa have been fully sequenced. In addition,
the National Center for Biotechnology Information Entrez Genome Projects
web site reports that sequencing of several more plant genomes is in prog-
ress. Nevertheless, the completion of the entire genomic sequence of a par-
ticular organism represents the end of the structural genomics segment of the
project. It is clear, therefore, that the identification of every gene within the
genomes of model organisms is only the initial step to understand what these
genes do and how they interact to make up a living organism. Understand-
ing the functions of the 20,000–50,000 genes comprising plant (and animal)
genomes and the variations within a population and roles in normal devel-
opment will represent a possibly greater task than the mapping and sequenc-
ing efforts currently underway.
Understanding the function of genes and other parts of the genome is
known as functional genomics, and research has involved model organisms
such as A. thaliana and rice. Model organisms offer a cost-effective way to
follow the inheritance of genes through many generations in a relatively
short time. Functional genomics is characterized by high throughput or
large-scale experimental methodologies combined with statistical and com-
putational analysis of the results. The fundamental strategy in a functional
genomics approach is to expand the scope of biological investigation from
studying single genes or proteins to studying all genes or proteins at once in
656 R.E. Litz et al.
India has focused upon the development of mango cultivars that are largely
indistinguishable from traditional selections with respect to fruit size, appear-
ance, taste, flavour and overall quality.
Despite its impact on crop breeding (Maluszynski, 2001) and particularly
on banana improvement (Novak et al., 1990), mutation breeding has not been
successfully exploited for mango cultivar improvement by production of use-
ful off-types of existing selections. There is some anecdotal evidence that
somatic mutations can occur naturally in mango on the basis of variation that
occurs within seed-propagated polyembryonic cultivars. There are reported
to be marked phenotypic differences within polyembryonic ‘Kensington Pride’
trees in Australia, and among polyembryonic cultivars of South-east Asia
(e.g. ‘Arumanis’, ‘Golek’, etc.). Different phenotypes of polyembryonic ‘Aru-
manis’, for example, have been characterized as ‘Arumanis-1’, ‘Arumanis-2’,
etc. Gan et al. (1981) and Litz et al. (1993) described isozyme variation within
populations of South-east Asian polyembryonic mangoes.
The most important production and postharvest problem of mango in
the humid tropics and subtropics is anthracnose, caused by Colletotrichum
gloeosporiodes (Penz.) Penz. and Sacc. In Penz. The current strategies for con-
trol of this disease involve the use of moderately resistant cultivars (i.e.
monoembryonic ‘Calypso’, monoembryonic ‘Keitt’ and monoembryonic
‘Tommy Atkins’, etc.) and at least weekly applications of fungicides (i.e.
benomyl or maneb or mancozeb) from the time of flowering until harvesting
(Dodd et al., 1997). This can result in as many as 25 spray applications in
a season, and is considered an increasingly unsustainable agricultural
practice.
The effect of γ-irradiation on embryogenic cultures of polyembryonic
‘Hindi’, monoembryonic ‘Keitt’ and monoembryonic ‘Tommy Atkins’ was
reported by Litz (2001) as part of the Food and Agriculture Organization
(FAO)/International Atomic Energy Agency (IAEA) Co-ordinated Research
Project entitled Improvement of Tropical and Subtropical Fruit Trees through
Induced Mutations and Biotechnology. Embryogenic mango cultures on semi-
solid maintenance medium were exposed to 0–200 Gy irradiation provided
by 60Co. The lethal dose for 50% mortality (LD50) for each cultivar was deter-
mined: it was approximately 125 Gy for ‘Keitt’ and approximately 100 Gy for
‘Tommy Atkins’. The LD50 of embryogenic ‘Hindi’ cultures could not be
established within this dosage range, possibly due to its high rate of prolif-
eration relative to ‘Keitt’ and ‘Tommy Atkins’. This study was confirmed by
Manzanilla Ramirez et al. (2000). The main objective of these studies has been
to develop appropriate technologies for utilizing induced mutations in vitro
for recovery of useful off-types of existing cultivars.
Culture filtrates produced by pathogenic fungi and bacteria can be uti-
lized not only to select for resistance to the pathogen in vitro (Hammerschlag,
1992), but also to induce the host resistance response. In order for in vitro
selection to be used effectively, essential prerequisites include: (i) a highly
morphogenic suspension culture; and (ii) evidence that the culture filtrate or
phytotoxin can produce symptoms of the disease on plant organs as well as
with cells in suspension cultures. Litz et al. (1991) first reported that the C.
658 R.E. Litz et al.
Genetic transformation
General protocols
Mathews et al. (1992, 1993) first reported the genetic transformation of mango
using embryogenic cultures of polyembryonic ‘Hindi’ and of a monoembry-
onic ‘Keitt’ zygotic embryo-derived embryogenic line, respectively. These two
studies utilized two different disarmed, engineered strains of Agrobacterium
tumefaciens: (i) strain C58C1 containing the plasmid pGV 3850::1103 with the
selectable marker gene for neophosphate transferase (NPTII) which confers
resistance to the antibiotic kanamycin, both of which were driven by the
CaMV constitutive 35S promoter (Mathews et al., 1993); and (ii) strain A208
containing the plasmid pTiT37-SE::pMON9749, a co-integrate vector, with
genes for NPTII and the scorable marker E-glucuronidase (gus or uidA) with
the 35S promoter (Mathews et al., 1992). A report by Cruz Hernandez et al.
(1997) utilized A. tumefaciens strain LBA4404 containing NPTII, E-glucuronidase
(GUS) and genes that mediate a horticulturally useful trait in binary plasmid
pBI121 with the CaMV 35S promoter. Mathews and Litz (1990) earlier had
demonstrated that 12.5 Pg/ml kanamycin sulfate is toxic to embryogenic
suspension cultures; whereas, much higher levels (200 Pg/ml kanamycin)
are toxic to embryogenic cultures that are grown on semi-solid medium.
These genetic transformation reports have followed a similar two-step
selection (Mathews et al., 1992; Cruz Hernandez et al., 1997). Embryogenic
suspension cultures in their logarithmic phase of growth are separated by
passing them through sterile filtration fabric (1000 Pm pore size), and the
large fraction (>1000 Pm) is abraded with a sterile brush on sterile filter
paper. The abraded PEMs are then incubated with acetosyringone-activated
A. tumefaciens for 3 days in liquid maintenance medium, with subculture
into fresh medium at 24 h intervals. The PEMs are then transferred onto
semi-solid maintenance medium supplemented with 200 mg/l kanamycin
sulfate and 500 mg/l cefotaxime. After 10 months on this selection medium,
the PEMs are transferred to semi-solid maintenance medium containing
400 mg/l kanamycin sulfate. Proliferating cultures are subcultured in liq-
uid maintenance medium containing 100 mg/l kanamycin sulfate, and
somatic embryo development is initiated by subculture onto semi-solid
maturation medium. Mathews et al. (1993) regenerated transgenic mango
plants derived from a ‘Keitt’ zygotic embryo embryogenic culture and
which had been transformed with pGV 3850::1103 containing the selectable
marker gene nptII. Genetic transformation was confirmed by: (i) growth in
selection medium containing inhibitory levels of kanamycin sulfate; (ii)
positive histochemical reaction for GUS with X-GLUC (Jefferson, 1987); and
(iii) Southern hybridization.
660 R.E. Litz et al.
Long-term storage
18.6 Conclusions
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constituents of the fruits of three mango cultivars, Mangifera indica L. Journal of
Essential Oil Research 11, 65–68.
Ara, H., Jaiswal, U. and Jaiswal, V.S. (1998) Rooting of microshoots of Mangifera indica
L. cv. Amrapali. Current Science 74, 240–242.
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Index
671
672 Index
Tahar 60
Salinity 196–197 Taimour 61
Sandersha 11, 327 Taiwan 435, 439, 442, 447–448, 457, 465–466,
Sapburn 545–548, 550 469, 471
Scab 274 Taxonomy 19–20, 22–30
Scale 342–343, 344–345 Temperature
Scaly bark 256–259 crop production 198–199
Sclerotium rot 285 flowering 111–113, 129, 134–135
Seed 4–5, 369–374 growth 192–193
cultivar description 44–63 internal fruit breakdown 306
germination 377–378 photosynthesis 179–181
hormones 130, 132–133 Termites 345
mango seed borer 331–332 Thailand 88
weevil 78, 328–331 Thiabendazole (TBZ) 551–553
Seedbed 377–378 Thrips 333, 337–338, 339
Sensation 60 Tissue culture 643–651
Sex ratio Tommy Atkins 11, 61, 323, 328, 448, 498–500
flowering 134–136 Top working 386–387
fruit set 140–141 Totapuri 61, 327, 506
Shoot Trade 34–35, 530–535, 606–611
chimeric 107–108 Transport 533–534, 548, 575–578
development 100–104 Tree growth 190–198, 463–468
formation 128–129 Triazoles see Paclobutrazol (PBZ)
initiation 105–106, 110–111, 129–131 Turpentine 11, 62, 78, 377
reproductive 104–105
Smartfresh 571
Smith 11, 328 USA 462–463, 466–468, 469–470, 471–472,
Soil 611–613
analysis 405–407 costs 613–626
crop production 440–443 cultivars 78, 448–449
flooding 182–184, 194–195 cultivation 10–11, 70
temperature 192–193 fertilizer 457–460
Soluble sugars 495–496 production 435–436, 439–440, 442–443,
Somatic embryogenesis 643–650 451–452
680 Index