Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
Dr Ratikanta Maiti
Dr Pratik Satya
Dasari Rajkumar
and
Allam Ramaswamy
QK641.C8653 2012
580--dc23
2012005599
Preface vii
Index 307
v
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Preface
The field of plant anatomy is one of the oldest life science disciplines. Starting its journey
from the inquiries of Greek philosophers, this branch of science has provided a strong
support to plant taxonomy, physiology and more recently plant developmental biology and
molecular biology. Not very surprisingly, anatomy of crop plants has attracted less attention
to botanists and taxonomists, since the major food crops belong to only few taxonomic
families. In certain model crops such as rice and maize, the progress in developmental
biology can largely be credited to the field of anatomy. However, relating structural anatomy
with crop adaptation and productivity has received comparatively less attention from
either agriculturists or botanists and few books give proper importance to this field of
agricultural science. However, while searching literature, one can find overwhelming sup-
port provided by anatomical investigations towards drawing meaningful conclusions from
research experiments on crop plants, particularly in the area of physiology and adaptation.
Still, the significance of anatomical variation in explaining the adaptation of crop plants
towards different abiotic stresses needs to be further investigated. In comparison to many
other modern techniques, which target cellular or nucleic acid variation, anatomy provides
low cost phenotypic screening options at organ, tissue or cellular level, further helping
emerging fields of life sciences such as transcriptomics and system biology as well as
providing rapid and reliable phenotypic screening procedures for selection of better
performing genotypes.
Increasing crop productivity is the most immediate concern of mankind in today’s
world. Crop anatomy is intrinsically linked with better genetic potential for productivity of
all crops, be it a derivative of primary or secondary metabolic pathway. To a large extent,
anatomical structures determine the synthesis of food by crop plants, absorption of mineral
nutrients for maintaining the process and channelling of reserve metabolite in certain
organs, which are harvested by humans as economic product. Crop plants have thus been
selected and manipulated during the course of evolution and selection for storing higher
reserve elements. The clue to further improving productivity thus lies in the process of
improvement of these physiological processes, which obviously need modifications of
structural anatomy at cellular, tissue, organ or whole plant level. Thus in future crop
science, anatomy is certain to play a pivotal role in helping crop improvement approaches,
both at phenotype and genotype level.
vii
viii Preface
In presenting this book the authors hope to fill a need for a textbook in crop plant
anatomy with an applied approach towards crop adaptation and productivity. Not only,
however, in our opinion, is there a need for a book for class study and guidance, but also
for one which shall serve as a reference text for workers in fields of agricultural science,
applied botany, and for teachers and students in other fields of crop ecology and botany.
This book first discusses the significance of anatomy in modern plant science followed
by an outline of the basic anatomical structures of angiosperms to give readers a basic idea
about plant anatomy. In the second section, structural anatomy of major crop plants has
been discussed in detail with an objective to delineate the significance of variations in the
anatomical features under different environments. These are followed by two sections, one
emphasizing the role of anatomy in adaptation of crop plants and the other on signifying
impact of the variations in structural anatomy on crop productivity, both of which are very
important for increasing agricultural productivity. Throughout the book we also show how
simple, low-cost light microscopy of hand sections can be used for rapid identification of
anatomical features and be used for selection of genotypes under different environments,
along with citing examples from research publications for further justification. Almost all
the anatomical figures presented in the book have been prepared from live samples by the
authors using simple light microscopy and low cost digital cameras, which is expected to
encourage students and new researchers in the field of agricultural science to explore the
tremendous possibility of utilizing anatomical techniques in their research fields. Training
on anatomy is mainly undertaken only by laboratory practice. On such practice the authors
believe emphasis must be given not only on lectures and text study, but also on extensive
practical training. For laboratory teaching the present book should provide a background of
facts, terms and history; it may, indeed, be used, in part, as a laboratory guide. The sequence
of subjects adopted mainly on the long experience of the first author is giving emphasis of
possible application of anatomical traits for adaptation of the particular crop to biotic and
abiotic stress factors.
In the treatment of subject matter, emphasis has been placed on adaptability to
classroom use from the standpoint of the student beginning anatomical study. Thus the
book is, first of all, a textbook in the elements of crop plant anatomy and their significance,
providing an introduction to the field. It presupposes an acquaintance only with the funda-
mental structure and activities of plants, an acquaintance such as is ordinarily obtained
from a first course in botany. However, we also understand that while dealing with crop
anatomy, which has vast and numerous research applications in the field of plant and crop
science, some areas may be neglected, which may not be on purpose. We welcome any
suggestions or critical input for further improvement of this book.
We are highly thankful to P. Vidyasagar, Chairman Vibha Agrotech Ltd, for provid-
ing facilities in Hyderabad as well as UNALA, ICRISAT, JRI UANL MEXICO, DLAA
Poebla, Mexico.
1
Origin and Development
of Crop Anatomy
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 1
2 Chapter 1
Crop anatomy is not therefore only anatomi- 1.3 Resurgence in Anatomical Studies
cal studies for the generation of basic bio- after Invention of the Microscope
logical information about the structure and
function of internal organs of crop plants; it After a long dormant period in the develop-
is an interdisciplinary science with highly ment of science, plant anatomy resurged as
significant applied value for the betterment a major technique in plant science largely
of agricultural science. due to the discovery of microscopic tech-
niques. During the 17th century, major con-
tributions in the field of plant anatomy were
1.2 Plant Anatomy in Ancient made by Marcello Malpighi (1628–1694)
Civilizations and Nehemiah Grew (1641–1712). However,
before their work a significant discovery
Although the subject of anatomy devel- was made by an English mathematician,
oped only after the discovery of micro- Robert Hook (1635–1703). He investigated
scopes, several ancient literatures reveal the internal structure of a thin slice of bottle
that visual anatomical studies have been cork under a crude microscope designed
carried out in different plant species. by himself, and described a honeycomb
Eames and MacDaniels (1925) reported like structure in it, and to each compartment
that plant anatomy study was initiated by he termed a cell (Latin, cellula = a small
Theophrastus of Eresus (c. 369–262 BC), apartment).
who is also regarded as the ‘Father of Malpighi is considered as one of the
Botany’. He described bark, wood and pith. forerunners in the field of animal and plant
However, even before the works of anatomy. He studied structural similarities
Theophrastus, anatomical studies using in plant and animal body plans and intro-
visual observations were initiated in differ- duced the words ‘epidermis’ and ‘stomata’
ent civilizations. in his book Anatome Plantarum published
Perhaps the earliest well documented in 1675. Grew, an English physician, is con-
studies on plant botany and anatomy were sidered as the father of plant anatomy. He
carried out in the early Indian civilization provided detailed anatomical description of
in the Vedic age. People during this period flower bud and vascular tissues and many
accumulated a great amount of knowledge other internal structures of plants that were
about botany and anatomy of plants, recog- earlier unknown. He first differentiated
nizing the values of different plants in agri- between the soft and hard parts of the plant
culture, medicine, fuel, construction and body and described the vertical and hori-
religious performances (Choudhury, 1971). zontal system in plants. He introduced the
In Rig Veda, the oldest Vedic literature, terms ‘parenchyma’, ‘pith’, ‘vesicles’ and
wood has been termed as ‘daru’ to differen- ‘cortex’ for the first time. He was also the
tiate it from the bark or ‘vakala’. Later, in pioneer in comparative anatomy of different
Brihadaranyaka Upanisada, wood anat- plant species, such as a comparison between
omy has been described mentioning xylem, the stem anatomy of apple, pear, plum or of
pith, bark and fibres. One of the earliest pine and oak. In spite of the limited resolu-
works in the field of plant science is the tion of microscopes during that time, Grew
Vrksayurveda (science of longevity of generated accurate and vivid descriptions
plants) by ‘Parasara’ (c. 400 BC), which of many plant anatomical features.
describes flower anatomy including ovary The 18th century witnessed some spe-
shape and also mentions sexuality in plants cific developments in the field of plant
(Kanjilal, 1999). It also describes leaf anat- anatomy, such as the description of vascular
omy, mentioning leaf structural compart- bundle elements (Sprengel, 1766–1833),
ments storing sap and being covered by a protoxylem (Treviranus, 1779–1864) and
boundary, which may be the cell wall in cambium (Du Hamel, 1700–1781). In the
today’s context. absence of good resolution, the earlier
Origin and Development of Crop Anatomy 3
microscopic observations were less specific. meristems. He also described the structure
However, a number of theories were devel- of starch granules in plant cells.
oped in this century to explain the function- The second development was progress
ality of vascular bundles and uptake of in understanding of the fertilization process.
water by plants. In this century, Carolus The anatomy of pistils and ovules helped a
Linnaeus established taxonomy based on lot to understand the process of pollen tube
structure, with plant anatomy in his epic growth, entry and fertilization. E. Strasburger,
work, Species Plantarum (1753). who was the first to describe fertilization
In the 19th century, two major develop- processes of plants in detail, extensively
ments took place that further established the used anatomical drawings to describe the
importance of plant anatomy as an inde- process of fertilization in his book Die
pendent science. The compound microscope Angiospermen und die Gymnospermen.
was further modified to obtain a better look During this period, a number of text-
at the cellular and subcellular structures. In books on anatomy appeared describing the
1831, with the help of a compound micro- basic anatomical features of root, stem,
scope, Robert Brown described the nucleus leaves, reproductive organs and seeds
in a cell. With the firm establishment of cell (Table 1.1). In 1884, the first book on stain-
theory by Schleiden (a botanist) and ing technique for plant anatomy was writ-
Schwann, (a zoologist) in the years 1838–39, ten by V.A. Poulsen. It described many
the functional significance of anatomical chemicals and their applications in staining
structures were recognized, followed by dis- various plant parts, such as application of
covery of cellular components such as pro- starch to identify and differentiate starch
toplasm (Von Mohl, 1846) and cytoplasm from cellulose, copper-ammonia to detect
(Kolliker, 1862). About the same time Carl pectic substances or the use of chromic acid
Wilhelm von Nägeli (1817–1881) exten- for study of the cell wall. This book served
sively studied the meristem and introduced as a very good description of several histo-
the terms ‘xylem’ and ‘phloem’. He studied chemical techniques that had been used
ontogeny of apical meristems and distin- in the 18th and 19th centuries for plant
guished between primary and secondary anatomy studies.
and its potential has not been well recog- question of providing food security to
nized either in basic science or in app- billions of people on the earth in the near
lied scientific fields such as agriculture. future is the most important question in
However, in the late 20th century, the phys- front of mankind, where crop anatomy
iological basis of plant productivity has can be used as a handy tool to generate criti-
been in the forefront of quests of agricul- cal inputs in basic as well applied plant
tural scientists and related disciplines. The sciences for enhancing productivity.
References
Chowdhury, K.A. (1971) Botany: Prehistoric Period. In: Bose, D.M., Sen, S.N. and Subbarayappa, B.V. (eds)
A Concise History of Science in India. Indian National Science Academy, New Delhi, pp. 371–375.
Eames, A.J. and MacDaniels, L.H. (1925) An Introduction to Plant Anatomy. McGraw-Hill Book Co.,
New York, pp. 321–342.
Kanjilal, D.K. (1999) A note on the Vrksayurveda of Parasara. Indian Journal of History of Science 34,
127–131.
2
Relevance of Anatomical Studies
in Modern Crop Science
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
6 (R. Maiti et al.)
Relevance of Anatomical Studies 7
When plant anatomy was born as a scientific has been used as secondary evidence after
discipline, it was immediately adopted as a the primary classes have been established
tool in plant classification. Wood anatomy based on morphological characters. At spe-
was used by Auguste Mathiew in 1858 for cies level characterization, morphological
description of forest plants in his book Florae and cytogenetic characters are used pre-
Forestiere (Naik, 2006). Bureau in 1864 first dominantly for systematics, followed by
used anatomical characters for plant system- confirmation through anatomical observa-
atics at taxa level in the family Bignoniaceae. tions of different plant parts such as root,
Along with the morphological descriptions stem, leaves or reproductive organs. In
of plants and cytogenetic behaviour of chro- many cases, the differences are not very
mosomes, anatomical descriptions (also prominent among closer relatives, although
referred as micromorphology) have been suc- their morphological and cytogenetic behav-
cessfully used in classification of several iours are quite distinct. Rice and wheat, the
land plants. The basis of classification was two major food crop species of the world
similar to morphology in that plants sharing from Gramineae exhibit remarkable dis-
common anatomical features are related. We similarity in vegetative morphology (many
can group these anatomical evidences in two tillers versus few tillers), reproductive mor-
distinct categories. In the first category, plant phology (panicle versus spike), chromo-
anatomy has been used as a primary basis for some number (2n = 18 versus 2n = 42), show
classification, being supported by other evi- no crossability with each other and are
dence. In most of these cases, anatomy has adapted to different climatic conditions
been used as a primary tool in higher order (tropical versus temperate). However, the
classification of land plants. One prominent anatomical features of root, stem, leaves and
example is the difference in the vascular seed are quite similar in rice and wheat,
system of monocotyledonous and dicotyle- suggesting remarkable similarity in struc-
donous plants. The use of anatomy as the tural mechanism albeit their other differ-
primary basis of classification is more com- ences. Such anatomical similarity persists
mon at genus, family or higher level. in the other members of Gramineae, estab-
However, in certain cases, anatomy has been lishing the importance of anatomy as a key
used as primary evidence in classification at descriptor at family level.
species or subspecies level, mostly when Anatomy is particularly helpful in tax-
morphological differences are insufficient onomic studies of the herbarium material
for classification. when morphological features are not suffi-
There are only a handful of cases where cient to identify the taxonomic status of the
crop anatomy has been used as primary plant material. This may happen when the
basis for the purpose of taxonomy and sys- morphological differences are insufficient,
tematic studies. A major reason is that or when only parts of plant are available
unlike non-crop plants, crop plants have (pieces of leaves or stem).
been in closer association with humans, Out of the major anatomical features of
thus their morphological characteristics had plants, anatomy of leaves has been utilized
been well studied. In many cases, plant most in plant classification. Leaf anatomy is
classification initiated from these crop the most utilized anatomical descriptor in
plants by noting and comparing their mor- the families Gramineae, Euphorbiaceae,
phological features. Prominent examples Zingiberaceae, Musaceae and Ericaceae.
are classification of rice, wheat, maize, sug- Root anatomy and root architecture are also
arcane and sorghum in the family Gramineae, utilized for taxonomic studies, although
based on their similarity in reproductive as root characters are less utilized for plant
well as vegetative features. Only later, anat- systematics (Scatena et al., 2005; Zobel and
omy was used to confirm the classification Waisel, 2010).
pattern. The anatomical variations of stems
The second category, which is more com- have been mostly utilized to differentiate
mon, includes cases where plant anatomy the structural aspects of vascular bundles of
8 Chapter 2
and weedy rice species. The thick noble (less water, higher temperature), a number
cane varieties of sugarcane, on the other of common anatomical modifications are
hand have been preferred for better vegeta- observed, such as a drastic reduction in the
tive growth and sugar production than the number of stomata on upper leaves, the den-
thin cane varieties and the wild Saccharum sity of trichomes on leaves forming a micro-
species, where food channelling occurs climate, development of cuticles on leaves,
from vegetative (leaf as source) to vegetative thickening of laminar cells, development of
(stem as sink) tissues. In both cases, robust- bulliform cells etc. On the other hand, under
ness in structural anatomy has helped in hypoxia (induced by waterlogging stress),
better adaptation of these species as pre- the formation of roots having aerenchyma
ferred crops. is a common mechanism for survival. Aeren-
chyma formation is under the control of
ethylene accumulation in the collar zones,
which is formed under hypoxic, anoxic or
2.4 Anatomy in Crop Ecology other stress conditions such as heavy soil
prohibiting penetration of roots. Similarly,
Ecology is defined broadly as the science of halophytic plants undergo special anatomi-
interaction of an organism or a group of cal modifications, such as enlargement of
organisms to its surrounding environment. cellular vacuoles, the formation of salt
Ecological adaptation is an essential require- glands for storing NaCl, thickening of hypo-
ment of survival of any species. Plant species dermis and surface hairs to remove excess
are adapted to different ecological conditions salt. Hydrophytic plants adapted to flooded
in nature. When a species is introduced in a conditions produce aerenchyma in tissue
new ecology, or the existing ecology changes, functioning as a supply of oxygen under
plants try to adapt to the new environment anoxic condition.
by modification of their own body plan to Crop species have been adapted to dif-
survive and reproduce in the altered envi- ferent environments mostly due to migration
ronment. Such changes are induced by mod- of human population. The major adaptations
ification of genome constitution through of crop plants under extreme environments
mutation and recombination, higher order are many times common, such as aeren-
chromosomal changes, or changes at pheno- chyma formation under submerged condi-
type level. Plants with genetic constitution tion in rice and maize. A very prominent
befitting to the environment survive to pro- example of adaptation to change in ecologi-
duce progeny adapted to the environment. cal conditions due to human migration is
The adaptation is manifested by changes in maize, which originated in the New World
morphology, anatomy, growth or reproduc- (Central and South America), but has been
tion or a combination of these. adapted to different environments of the Old
Plants when adapted under different World within a very short time period of few
environments show difference in anatomi- hundred years. Due to its special C4 photo-
cal structures. By interpreting such anatomi- synthesis system, it has adapted easily to the
cal changes, one can not only understand tropical as well as humid subtropical condi-
the specific responses of anatomical struc- tions of African and Asian countries.
tures of root, stem, leaf, flower and seed of a The study of ecological crop anatomy
species under different ecology, but also enables us to understand the basis of
study the common anatomical modifications adaptation of crop species in different eco-
taking place in several related and unrelated logical conditions, including extreme envi-
species under a particular ecology. For ronments such as drought, waterlogging,
example, mesophytic plants surviving under heat, cold stress, or biotic stress conditions.
optimum environmental conditions (light, In the third section of this book, we will
water, nutrient) exhibit several common discuss how adaptation of crop plants under
anatomical features. When many of these these conditions has affected the anatomi-
plants are adapted to xerophytic conditions cal structures of crop plants, or how anatomy
10 Chapter 2
has helped in adaptation of some of the crop cells, de-differentiate from specific cell
plants in extreme weather conditions. types to undifferentiated cellular mass and
Global warming and increase in CO2 again re-differentiate into specific cell types.
concentration is a major concern of present This makes the plant cell truly immortal,
day agriculture. The ecological conditions providing the conditions for growth and
for growth and survival of plants and ani- survival of the cells are met. In contrast, the
mals in general are undergoing rapid change animal cells differentiate into specific cell
and are predicted to worsen in near future. types, but cannot de-differentiate into non-
Crop species in the future generation are going specific cells once the cell fate is deter-
to face changed ecological conditions, which mined. The only exception to this category
will affect crop productivity. Understanding is the stem cells, which can be maintained
structural and physiological changes to sus- for indefinite period and can be subjected to
tain in the future environment is a major differentiate into other cell types. But the
research area in crop science. Experiments capability of re-differentiation is limited to
under artificially enriched CO2 and observa- plant cells only. This is the primary basis of
tion on plants that sustain better under vegetative reproduction of plants through
higher CO2 concentration show that plants stem cuttings, a very common form of prop-
undergo distinct morphological, anatomi- agation for horticultural plants. The ability
cal and physiological changes under such to re-differentiate comes very handy under
extreme conditions. Such modifications are stressed conditions, such as formation of
more obvious on leaf structures, such as aerial roots under submerged condition for
modification in stomatal density, alterations harvesting oxygen from the atmosphere.
in thickness of leaf palisade parenchyma or However, although all plant cells are
increase in leaf thickness. Chapter 12 of this totipotent, not all the cells in a growing plant
book describes possible effects of climate are actively dividing. The actively dividing
change on crop anatomy and consequences cells are located in the meristematic regions
on crop productivity. leading to primary growth of the plant. The
secondary growth results from the thicken-
ing of the differentiated tissues by formation
of new layers of differentiated tissues or by
2.5 Anatomy in Developmental maturation and thickening of the cells or
Studies deposition of specific materials in the inter-
cellular cavities. Anatomical observation of
Plant anatomy is an indispensable tool in the meristematic regions is the primary tool
studying growth, development and differen- to investigate the growth pattern of the
tiation. From a tiny seed, the angiosperms dividing tissues and pattern of formation of
develop into a full plant having root, shoot, specific tissues. Anatomy of growth pattern
leaf and reproductive organs. Even during of apical meristems (root and shoot) clearly
seed development, a single fertilized diploid differentiates the growing regions from the
cell develops and differentiates into a multi- differentiated cells and also identifies dis-
cellular embryo bearing a cellular structure tinct layers (L1, L2, L3 etc.) in the apical
dedicated for shoot and root development. meristematic regions. These layers are not
The anatomical features of the body parts of only structurally different, but also have
seed plants are well described in many plant different functional significance. Different
anatomy textbooks. A brief description of layers form different types of tissues.
anatomy of plant body parts is also provided
in Chapter 3 of this book, which will help
the readers to understand general plant
anatomy as well as anatomy of crop plants. 2.6 Anatomy in Plant Evolution
Plant cells have higher plasticity than
animal cells, i.e. the cells can differentiate Plants have emerged on the planet for
from uncategorized cell mass to specific millions of years. During the course of
Relevance of Anatomical Studies 11
evolution plants have advanced from Hibiscus rosa chinensis), or at species level
simple structures to more complex organ (stress tolerant versus susceptible genotypes).
differentiation. Land plants have evolved
from aquatic plants by changing their inter-
nal structures for adherence to the soil by
modifications in morphology and anatomy 2.7 Anatomy in Anthropology
of root structures. Likewise, the water and Palaeobotany
transport system of land plants evolved in
a different way from those of aquatic plants. Humans have used plant products as food,
Consequently, the evolution of xylem ves- fuel, fibre or for construction purposes
sel systems selected separate paths in since prehistoric times. Anatomical inves-
aquatic and land plants. The land plants tigations of fossil remains of plant parts
had to establish the plant stand by mechan- used for household purposes by ancient
ical supports, leading to higher root civilizations or early human establishments
number, root volume and anchorage. The provide important clues about the habitat
body plan of land plants has also been and culture of that time. Archaeological
modified to develop more structural tis- and anatomical investigations of wood and
sues in definite orientations to provide charcoal samples from the excavated sites
higher mechanical support to the aerial can also help in estimating the climatic
parts. Moreover, since these plants are not conditions based on the study of growth
in constant association with water, a rings. The cell walls of late-wood cells are
number of advanced water loss protection preserved better than that of early-wood
mechanism evolved in these plants under cells in remains of wood samples. Such
different adaptation conditions. Anatomical anatomical studies are also helpful in
investigations of terrestrial and aquatic understanding evolution of structural
plant body systems have generated numer- compartments of cells and tissues during
ous evidences for such modifications, evolution. For example, studies of car-
which help us to understand the course of bonized samples of Taxodixylon species
evolutionary progress in these two plant aged at 20 million years revealed that
systems. The basic evolutionary differ- the fibrillar structures of cellulose are well
ences are also very clear in aquatic, semi- conserved throughout the evolutionary
aquatic and terrestrial crop species. The period. Anatomical observation of charred
aquatic and semi-aquatic crop species such remains of fossil plants also helps in
as lotus or water lily have tender stem identification of nearest relatives of these
structures with little mechanical support prehistoric plants in the present-day plant
compared to land crop species. In contrast, kingdom.
both the monocotyledonous and dicotyle- Since crop species have been domesti-
donous crop species derive mechanical cated early in civilization, investigations on
support from collenchyma and scleren- remains of crop plants or seeds generate
chyma tissues, and many at times also from valuable information regarding the civiliza-
parenchyma tissues like tracheary ele- tion. An important area of anatomical inves-
ments. The variations in anatomical con- tigations is the assessment of remains of
figuration of these elements are observable clothes. Early humans primarily used grass
at higher order levels of crop classification species for the purpose of clothes. With
(difference in plant architecture of pulse advancement of civilization, human popu-
crops and beans), family level (growth lation learned to extract fibre from plants
habit of leguminous pulse crops like pea and make durable clothes. Fibres from flax
and lentil and leguminous green manure were used in producing clothes of very high
crops like Sesbania and Crotalaria, varia- value in the Egyptian civilization, which
tion in fibre formation in cotton and jute), was considered as a royal commodity and
genus level (variation in development of beyond the reach of general people. The
bast fibres in Hibiscus cannabinus and archaeological remains from the pyramids
12 Chapter 2
References
The minute internal structure of the plant For future anatomical study, the specific
body is identified mainly from thin sec- organs need to be fixed with reagents to stop
tions taken by hand or microtome and the life processes without disintegration of
from maceration in which the individual cell structures and organelles. Different flu-
cells are freed from one another. Plant ids are used for fixation, such as glacial acetic
microtechnique consists of three proce- acid, 1% chromic acid, formalin (30–40%);
dures (Fig. 3.1): (i) preparation of plant tis- the most common fixative is FAA (formalin
sues for microscopic study; (ii) use of the acetic alcohol).
microscope and related equipment for crit-
ical study and interpretation of the materi-
als; and (iii) detailed descriptions of each
3.4 Section Cutting
part. There are different steps to study the
anatomy of plant parts, which are men-
tioned below. Section cutting is carried out by hand with
the use of a sharp razor blade and the
assistance of a microtome. The section
must be very thin and should not be
oblique. Different types of section cutting
3.2 Collection of Desired Plant are used to study plant organs, such as a:
Materials and Subdivisions (i) transverse section (TS): the plant organ
root, stem, petiole, leaf etc. is cut trans-
For anatomical study, desired plant organs, versely (perpendicular 90°) to its axis;
such as stems, leaves, floral organs, should (ii) longitudinal section (LS): the plant
be cut with a sharp knife or scalpel and organ is cut longitudinally (180°) to its
transported between wet blotting paper. In axis; (iii) radial longitudinal section: the
the case of dried herbarium specimens, the plant organ is cut transversely, then cut
specimen can be softened for sectioning to through the radius; and (iv) tangential lon-
prepare slides. This helps in determining gitudinal section: the plant organ is cut
the distinguishing features of vascular tangentially. Sections are made by cutting
arrangement (Hyland, 1941). perpendicularly to the radius.
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 15
16 Chapter 3
Collection of desired plant materials ester wax is used for the tissue embedding
medium (Chayen and Gahan, 1959).
Fixation and storage
Fig. 3.1. Basic steps in microscopic observation 3.5.2 Preparation of whole mounts
of plant parts. and macerations
Objective piece
Media used for mounting vary between gly-
cerine 10%, glycerine jelly, lactophenol, eryth-
Slide
rosine or Canada balsam (or DPX mounting
Adjustment
medium), depending whether they are for
Stage
temporary or permanent preparations.
Mirror
3.7.2 Safranin
surfaces of solid specimens. The and in a natural condition. For this a simple
beam current is very low (10−10–10−12 technique can be used:
amp), so even delicate biological sur-
faces can be examined with a mini- 1. Take a small amount of xylene (C6H4
mum of heat damage. (CH3)2) in a glass Petri dish.
• Scanning transmission electron 2. Gradually dissolve thermocol (a polysty-
microscopy (STEM): it uses a fo- rene commonly available in shops) in the
cused incident probe across a speci- xylene and stir with a glass rod. Progres-
men that has been thinned to sively add more thermocol to the solution
facilitate detection of electrons. and stir with a glass rod until it turns into
• Reflection electron microscopy a honey-like solution (gummy).
(REM): an electron beam is incident 3. Apply the solution by finger once only
on the surface. The reflected beam on the lower and upper surfaces of the leaf
of elastically scattered electrons is at different places.
detected. 4. Allow the solution to semi-dry.
5. Put a transparent cellophane tape on the
semi-dried area of the leaf.
3.9 Maceration of Wood/Fibres 6. Remove the tape gently and paste on to
the labelled microscope slide.
Wood is a hard substance and the cells are 7. Observe the slide under low and high
closely attached to each other. For studying power for the epidermal tissue system.
the components of wood the cells must be free
from one another. For this a technique called
maceration is used to separate the cells. Place 3.11 Estimation of Pollen Viability
a piece of wood in a glass test tube. Add a
mixture of concentrated HNO3 (10%) and
Different methods for the estimation or
chromic acid (10%). Heat the material in a test
determination of pollen viability are avail-
tube on a burner for 10–15 min and then cool
able. Three generally used techniques are:
the test tube in running water at normal
(i) the iodine potassium iodide (IKI) tech-
temperature for 20 min. Remove the piece of
nique; (ii) tetrazolium; and (iii) the Evans
wood and wash with water and place the
blue method.
wood on a slide and press it to separate the
elements. Cover with a cover slip and observe
under the microscope. Different types of vessels,
parenchyma and fibres can then be examined. 3.11.1 Iodine potassium iodide technique
References
Chayen, J. and Gahan, P.B. (1959) An improved and rapid embedding method. Quarterly Journal of Microscopic
Science 100, 275–277.
Greenwood, M.S. and Berlyn, G.P. (1968) Feulgen cytophotometry of pine nuclei: effects on fixation role of
formalin. Stain Technology 43, 111–117.
Hyland, F. (1941) The preparation of stem sections of woody herbarium specimens. Stain Technology 16,
49–52.
4
General Anatomy of Crop Plants
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 21
22 Chapter 4
Ribosome
Vacuole
Nucleolus
Chloroplast
Cell
Plasma Protoplasm
membrane
Nucleus Cytoplasm
begins. Cells performing similar functions Various types of tissues are found in the
organize into groups. A group of such cells, plant body (Fig. 4.5). Based on the activity,
which are similar in origin, structure and all the tissues of plants are classified into
function, is described as tissue. In fact these mainly two types, meristematic tissues and
tissues are formed in response to the basic permanent tissues.
division of labour.
In complex organisms cells organize 1. Meristematic tissues are groups of imma-
into tissues, which in turn organize into tis- ture cells, which are primarily concerned
sue systems. Various tissue systems consti- with the production of new cells.
tute the organ and different organs form the 2. Permanent tissues are groups of differen-
organism. tiated cells, which have lost the power of
24 Chapter 4
Cell
Protoplasmic Non-protoplasmic
components components
Fig. 4.4. Detailed outline of plant cell structure and their components.
Tissues
Based on origin
(characteristics of initial cells) 4.7.1 The root
Primary meristem
Secondary meristem Introduction
Tissue systems
Intra stelar
Pericycle
Medulla
Medullary rays
Vascular bundles
absorption and serve only for conduction, This region helps in lateral conduction
support and storage. of water and salts. After secondary
growth the general cortex ruptures.
Anatomy of dicotyledonous root (ii) The endodermis is very distinctive in
roots. It is single layered, having com-
Dicot plants show a taproot system. In the
pactly arranged barrel-shaped cells.
root hair region it shows the primary struc-
Endodermal cells are characterized by
ture; behind the root hair region secondary
the presences of Casparian strips or
growth is initiated.
Casparian bands. Casparian bands are
The initial organization of the primary
made of suberin. In transverse section,
dicot root is simple when compared to that
Casparian bands appear as lens-shaped
of the stem. Transverse section of the root
thickening on the radial walls of the
shows three main parts: the epidermis, cor-
endodermis. Endodermis is also called
tex and stele (Figs 4.9, 4.10).
the starch sheet layer, as starch grains
1. The epidermis of the root is also called are present in the endodermal cells.
the rhizodermis, epiblema or siliferous layer, The root endodermis acts as a barrier
and is the outermost region of the root. It is between the cortex and the stele. Endo-
uniseriate, having a single layer of com- dermis ruptures due to secondary
pactly arranged thin-walled living cells. growth.
Both cuticle and stomata are absent. Epidermis 3. The stele consists of: (i) pericycle;
gives protection and absorption of water (ii) vascular strands; and (iii) pith.
and minerals. Epidermal cells produce uni- (i) The pericycle is the outermost region
cellular root hairs, which are useful in of the stele. It is uniseriate and paren-
increasing the surface area for absorption of chymatous. Cells of the pericycle
water and helps in absorbing water from retain meristematic activity. Lateral
soil. Generally root hairs are short lived. roots arise endogenously from the peri-
Epidermis ruptures after secondary growth. cycle and help in absorption of water.
2. The cortex is the middle region, lying be- During secondary growth, pericycle
tween the epidermis and the stele (Fig. 4.10). forms phellogen or cork cambium and
Cortex is well developed and is larger and a small amount of vascular cambium.
thicker than the stele. It is multiseriate and (ii) Vascular strands are composed of the
relatively homogeneous. It consists of general vascular tissues, xylem and phloem.
cortex and endodermis. Xylem conducts water and salts,
(i) The general cortex is extensively devel- whereas phloem transports food
oped in the root. It consists of loosely materials. Xylem and phloem are
arranged thin-walled parenchyma cells arranged in the form of separate
with prominent intercellular spaces. strands on different radii. They alter-
The cells are usually round or oval. The nate with each other. Hence the vas-
cells are colourless and store starch. cular strands in the root are described
General Anatomy of Crop Plants 27
Dicotyledonous root
(a) (b)
Epidermis
Cortex
Endodermis
Protoxylem
Phloem
Metaxylem
Fig. 4.10. (a) Transverse section of young dicotyledon (cotton root 40×) showing epidermis, multilayer
cortex and single layer endodermis. (b) Sector enlarged (100×) showing xylem strands (tetrarch condition).
Bark
Secondary xylem
Secondary phloem
Phloem
(a) (b)
Fig. 4.11. (a) Transverse section of matured dicotyledonous (cotton 40×) root showing secondary growth.
Secondary xylem vessels are larger in size. (b) Sector enlarged (100×) illustrating bark and the cortex
is compressed due to increase in secondary structures.
Monocotyledonous root
Epidermis
Ground tissue
Endodermis
Phloem
Xylem
Pith
Fig. 4.13. Transverse section of monocotyledonous (maize) root (100×) showing epidermis, multilayered
ground tissue (cortex), endodermis, phloem, xylem (polyarch) and central pith.
vascular strands in root are described (iv) The medulla is distinct and usually
as separate and radial. Xylem is exarch, parenchymatous. The medulla stores
having protoxylem towards the pericy- food materials.
cle and metaxylem towards the centre (v) Conjunctive tissue: the non-vascular
or medulla (centripetal development). tissue left between xylem and phloem
Xylem is polyarch, having more than eight strands is called conjunctive tissue. It
protoxylem points. Cambium is absent may be parenchymatous or sclerenchy-
and hence vascular bundles are descri- matous.
bed as closed type.
(iii) Pith or medulla: pith is absent. The
non-vascular tissue present between 4.7.2 The stem
xylem and phloem strands is called
conjunctive tissue. It may be totally Introduction
parenchymatous or a part of it may be
converted into sclerenchyma. During The part of the axis of the plant that is aerial
secondary growth the cambial cells in nature and consists of branches, leaves
that originate from the pericycle lying and reproductive structures is called the
against the group of protoxylem func- stem. The stem possesses nodes and inter-
tion as ray initials. nodes, which are absent in the root system
30 Chapter 4
Dicotyledonous stem
round or oval with prominent intercel- (ii) Vascular bundles comprise vascular
lular spaces. This region also stores tissues of xylem and phloem, in which
food materials temporarily and flavo- 15–20 vascular bundles are arranged
noids are deposited. in one ring called the eustele. Each
(iii) The endodermis is the innermost layer vascular bundle is wedge-shaped. It
of the cortex (Fig. 4.15). It is usually dis- consists of xylem towards the centre of
tinct in stems without Casparian bands. the axis and phloem towards the
In young stems the innermost layer of periphery. In the vascular bundle
the cortex contains abundant starch, xylem and phloem are present together
hence this layer is called the starch (conjoint) on the same radius (collat-
sheath. The starch sheath layer is eral), with a strip of cambium (fascicu-
homologous to endodermis, but mor- lar cambium) between xylem and
phologically unspecialized, hence such phloem (open). Hence the vascular
a layer may be called endodermoid. bundle is described as conjoint, collat-
After secondary growth the endodermis eral and open type. Xylem is endarch
ruptures. with protoxylem present towards the
3. The stele is the central region of the centre. The xylem consists of many
stem and is larger and thicker than the vessels. Xylem vessels are arranged in
cortex. It consists of pericycle, vascular rows.
bundles, and medulla and medullary (iii) Medulla and medullary rays: this reg-
rays. ion is distinct, present at the centre
(i) The pericycle (perivascular region) is of the stem and is usually parenchy-
the outermost non-vascular region of matous. The cells are round or oval with
the stele. It is multiseriate and scleren- intercellular spaces. Medulla stores food
chymatic. Sclerenchyma is mechanical materials. The extensions of parenchy-
tissue and gives strength and rigidity to matous pith between vascular bundles
the stem. are called primary medullary rays,
Cuticle
Epidermis
Parenchyma
Collenchyma
Endodermis
Phloem
Cambium
Xylem
vessels
Medulla
Fig. 4.15. Transverse section of dicotyledonous stem (Amaranthus 100×) showing uniseriate epidermis
followed by four to five layers of collenchyma and single layered endodermis. The stele is at the centre of
the stem, consisting of medulla and vascular bundles. Secondary growth has begun with cambium forming
secondary tissues.
32 Chapter 4
which help in lateral conduction and pericycle etc. are indistinguishable. The
may gives rise to secondary meristem whole region is parenchymatic and is
(inter-fascicular cambium). best described as ground tissue (or con-
junction tissue).
(ii) Vascular bundles are numerous and
Monocot stem are scattered irregularly in the ground
tissue (Fig. 4.17). Usually the periph-
Generally monocots lack secondary growth.
eral bundles are small and closely
The stem is usually circular in outline in
arranged, and the central bundles are
transverse section and is differentiated mainly
large and are widely arranged. Each
into epidermis, cortex and stele (Fig. 4.16).
vascular bundle is oval in shape and
1. The epidermis is the outermost region of is surrounded by fibrous bundle
the stem, and is meant for protection. It is sheath. It is many cells thick towards
uniseriate, having a single layer of compactly outer and inner regions and only a few
arranged barrel-shaped living cells. The outer cells thick at lateral regions. Hence
surface of the epidermis is covered by thick the vascular bundles are described as
cuticle, which checks transpiration. Few fibrovascular bundles. Vascular bun-
stomata are present in the epidermis for dles are conjoint, collateral (with
the exchange of gases and transpiration. All xylem and phloem present on the
the epidermal cells are colourless except the same radius) and closed (i.e. without
guard cells. cambium). The xylem is present
2. The cortex is highly reduced and usually towards the centre of the axis and con-
represented by a narrow hypodermal region. sists of a few vessels, which are
In mature stems, hypodermis is two- to arranged in the form of letter ‘Y’. The
three-layered sclerenchymatic tissue and protoxylem often disintegrates and
gives mechanical strength. In young stems, forms protoxylem lacuna. Phloem is
chlorenchymatic patches are present just present towards the outside, between
below the stomata region and perform the metaxylem vessels, and consists of
photosynthesis. sieve tubes, companion cells and a few
3. The stele is the central part of the stem and parenchymatous cells at the sides.
is larger and thicker than cortex. The stele in
the monocot stem, with numerous vascular
bundles scattered irregularly in the ground 4.7.3 The leaf and the petiole
tissue, is called an atactostele, and is the most
advanced type of stele. The stele consists of
Introduction
ground tissue and vascular bundles.
(i) Ground tissue: in monocot stems usually The leaf is a laterally growing organ on
there is no clear demarcation between the stem. It performs various metabolisms,
the cortex and pith since endodermis, which include photosynthesis, respiration
Monocotyledonous stem
Cuticle Xylem
Phloem
Epidermis
Sclerenchyma
Ground tissue
Phloem
Xylem
Vascular bundle
Fig. 4.17. Transverse section of monocot (maize) stem (100×) showing scattered vascular bundles in ground
tissue (parenchyma). Note the peripheral vascular bundles are smaller and inner (central) vascular bundles
are somewhat larger in size.
and transpiration. To effectively carry out distinct dorsal and ventral surfaces. In
these metabolisms, leaves have mesophyll transverse section the leaf shows three dis-
consisting of chlorophyll and vascular tissues. tinct regions, the epidermis, mesophyll and
Leaves show variation in anatomy vascular bundles (Fig. 4.18).
based on the habitat. Usually, leaves show
1. The epidermis is mainly for protection
two types of structures: dorsi-ventral leaves
and is present on both surfaces of the leaf
and isobilateral leaves.
blade. The epidermis covering the upper sur-
1. Dorsi-ventral leaves: in this types of leaf face of the leaf is called the upper epidermis
morpho-anatomical variations are present or ventral epidermis or adaxial epidermis.
in the dorsal and ventral surfaces. The dor- The epidermis present towards the lower
sal surface is dark green in colour and is side of the leaf is called the lower epidermis
directly exposed to sunlight and the ventral or dorsal epidermis or abaxial epidermis.
surface is light green in colour. Mesophytic Epidermis is single layered, having com-
leaves show dorsi-ventral symmetry. These pactly arranged barrel-shaped (tabular)
types of leaves are generally present in cells. The outer walls of the epidermal cells
dicotyledons, e.g. cotton and sunflower. are cutinized. Moreover, the epidermis on
2. Isobilateral leaves: do not show any var- its outer surface is covered by a continuous
iations either morphologically and ana- layer of cutin called cuticle. The cutinized
tomically between the dorsal and ventral outer walls and cuticle reduce the loss of
surfaces. The leaf blade is nearly erect and water due to transpiration. Anisocytic sto-
both surfaces usually receive direct and mata are present in both of the epidermal
equal amounts of sunlight. The internal layers, but more stomata are usually present
structure is more or less similar in both the in the lower epidermis. Stomata facilitate
upper and lower halves. This type of leaf is the exchange of gases between the leaf and
generally present in monocots and hydro- environment. All the epidermal cells except
phytes, e.g. leaves of grasses. guard cells are colourless. Epidermis con-
tains multicellular hairs.
Anatomy of dorsi-ventral leaf (dicot) 2. The ground tissue of the leaf, present
between the upper and the lower epider-
A typical mesophytic dicot leaf shows a dorsi- mal layers, is called mesophyll (Fig. 4.19).
ventral nature, i.e. the leaf blade consists of The mesophyll is chlorenchymatic and is
34 Chapter 4
Palisade tissue
Xylem
Phloem
Spongy tissue
Parenchyma
Lower epidermis
Fig. 4.19. Transverse section of dorsi-ventral leaf (cotton 100×) showing organization of tissues. Stellate
trichomes on upper and lower epidermis, palisade towards upper and spongy tissue towards lower
epidermis can be observed. Central vascular bundle can be observed, consisting of xylem towards the upper
epidermis and phloem towards the lower epidermis.
General Anatomy of Crop Plants 35
epidermis and phloem towards the lower stress, smaller cells lose water and as a
epidermis. The vascular bundle is surroun- result the leaf coils and rolls thereby pre-
ded by parenchymatous bundle sheath. The venting transpiration. The epidermis is uni-
cells of the bundle sheath are called border seriate and composed of more or less oval
parenchyma. The larger vascular bundles are cells (Fig. 4.21). The outer walls of the epi-
supported by hypodermal parenchymatous dermal cells are cutinized and also covered
strands. These strands are considered to be by cuticle. Cuticle reduces the loss of water
the sheath extension. due to transpiration. Stomata are covered
by two dumbbell-shaped guard cells. The
Anatomy of isobilateral Leaf graminaceous type of stomata is present in
both epidermal layers, but the stomata are
A typical monocot leaf shows isobilateral usually more concentrated in the lower epi-
nature. The monocot leaf shows three dis- dermis. The epidermis is rough in nature
tinct regions, the epidermis, mesophyll and due to the presence of silica crystals. Silica
vascular tissue (Fig. 4.20). crystals offer drought resistance by covering
1. Both sides (upper and lower) of the leaf some parts of the epidermis and form a
are covered by epidermis. The upper epi- physical barrier. Bulliform cells are present
dermis may be easily identified due to the in the upper epidermis.
presence of xylem and bulliform cells com- 2. As the leaf is isobilateral, the mesophyll
posed of large and small cells. Under water is not differentiated into palisade and
spongy tissues. It is composed of compactly
arranged, thin-walled, isodiametric chloro-
Isobilateral leaf phyllous cells, having well developed inter-
cellular spaces among them.
Epidermis Mesophyll Vascular bundle 3. The vascular tissue occurs in the form
Epidermal cells (Spongy tissue) Xylem of discrete bundles called veins. The vas-
Guard cells Phloem cular bundles are collateral and closed.
Most of the bundles are small in size but
Trichomes
fairly large bundles also occur at regular
Fig. 4.20. Outline of the transverse section of an intervals. The xylem is found towards the
isobilateral leaf. upper side and phloem towards the lower
Mesophyll
Lower epidermis
Vascular bundle Xylem Phloem
Fig. 4.21. Transverse section of isobilateral leaf showing different parts. Note: sclerenchyma patches
are more frequent in lower epidermis. Vascular bundles are situated just above the lower epidermis.
Parenchyma occupies at the centre of the leaf.
36 Chapter 4
Petiole
Trichome
Epidermis
Hypodermis
Ground tissue
Endodermis
Phloem
Xylem
Vascular bundle
Ground tissue
Fig. 4.23. Transverse section of cotton petiole (small portion 100×) showing different parts. New vascular
bundles are developing and show similar structure to the stem.
General Anatomy of Crop Plants 37
flexible, being composed of cellulose and are called floral parts or floral organs. The
pectin. The hypodermis gives considerable sepals and petals constitute the calyx and
strength, flexibility and elasticity to young corolla, respectively. The calyx and corolla
petioles. Having chloroplasts it may carry are the sterile parts. The stamens consist of
out photosynthesis. Due to the presence of an androecium, where free or united carpels
the thick layer of angular collenchyma, the compose the gynoecium (Fig. 4.24). The sta-
petiole is strong, stiff and gives strength. mens and the carpels are the reproductive
3. The ground tissue occurs just below the parts.
hypodermis, and consists of thin-walled The flower shows definite growth. In
parenchymatous cells having well defined the flower the apical meristem ceases to be
intercellular spaces among them. Vascular active after the formation of floral parts. In
bundles are arranged in half rings scattered more specialized flowers there is a shorter
in ground tissue. growth period and they produce a smaller
4. The vascular bundles are of various sizes and more definite number of floral parts
in the same petiole. Each vascular bundle is than the more primitive flowers. In still
wedge-shaped. In the petiole, the xylem is more advanced flowers there are special-
always found towards the upper side ized characters, such as whorled arrange-
whereas phloem occurs towards the lower ment of parts instead of parts within a
side (as in the leaf ). whorl, zygomorphy instead of the actino-
morphic condition, and epigynous instead
Features of special interest include:
of hypogynous condition.
• Generally a groove is present towards The sepals show similar structure to
the upper side of the petiole. leaves in their anatomy. Each sepal consists
• Typically the vascular bundles are dis- of ground parenchyma, a branched vascular
tributed in a semicircle in ground tissue.
• Generally the central bundle is largest
and remains encircled by the endoder- Stigma
mal sheath.
• The xylem is always present towards Anther
the upper side and phloem towards the
lower side.
Introduction Petal
system and an epidermis. The chloroplasts of both anther and filament. The vascular
are found in the green sepals but there is bundle is found throughout the filament and
usually no differentiation into palisade and culminates blindly in the connective tissue
spongy parenchyma. They may contain cells, situated in between the two anther-lobes.
laticifers, tannin cells and other idioblasts. The outermost wall layer of the anther
The epidermis of sepals may possess stomata is the epidermis. Just beneath the epidermis
and trichomes. there is endothecium, which usually con-
The petals also resemble leaves in their tains strips or ridges of secondary wall
internal structure. They contain ground material mainly on those walls that do not
parenchyma and more or less branched vas- remain in contact with the epidermis. The
cular system and an epidermis. They may innermost layer is composed of multinucle-
also contain crystal-containing cells, tannin ate cells; this is nutritive in function and
cells, laticifers and certain other idioblasts. known as tapetum. The wall layers that are
They contain pigments containing chro- located between the endothecium and tape-
moplasts. Very often, the epidermal cells of tum are often destroyed during the develop-
petals contain volatile oils, which emit the ment of pollen sacs. On the maturation of
characteristic feature fragrance of the flow- the pollen the tapetum disintegrates and
ers. In certain flowers the anticlinal epider- the outer wall of the pollen sac now consists
mal walls of the petals may be convex or of only the epidermis and endothecium. At
papillate. The epidermis may also contain the time of dehiscence of the anthers the pol-
stomata and trichomes. len are released out through the stomium.
Usually the stamen contains two lobed
four-loculed anthers. The anther is situated Structure of the pollen grain
on a slender filament that bears a single vas-
cular bundle. In certain primitive dicoty- Pollen grains are haploid (x), unicellular
ledonous families the stamens are leaf-like and uninucleate male spores, each of which
and possess three veins, whereas in advanced has two wall layers, an outer thick exine and
types they are single veined. inner thin intine. Exine may be smooth or
The structure of the filament is quite ornamented. In the exine, one or more un-
simple. The vascular bundle is amphicribral thickened slit-like or circular areas called
and remains surrounded by parenchyma. The germ pores are present. The position of germ
epidermis is cutinized and bears trichomes. pores is constant and has taxonomic value
Stomata may also be present on the epidermis (Fig. 4.25).
(a) (b)
Fig. 4.25. Structure of the pollen grain (400×): (a) okra: spherical in shape containing spine-like structures
and ornamentation; and (b) maize: smooth, isobilateral and without any ornamentation.
General Anatomy of Crop Plants 39
Each pollen grain divides to form two (2n), formed as a result of fertilization. The
unequal cells, a vegetative and a generative mature embryo consists of cotyledons,
cell. Pollination occurs at this two-celled hypocotyl (stem-like embryonic axis below
stage. Further development of male gameto- the cotyledons) and radicle (embryonic root).
phyte takes place on the stigma and style. Endosperm is formed due to triple fusion; it
consists of food storage tissue, triploid (3n)
Gynoecium as a result of double fertilization; two-thirds
of the genome is of maternal origin.
A single unit of a gynoecium is called a car-
pel. A flower may possess a solitary carpel Classification
or more than one. If two or more carpels are
present they may be united or free from one Based on the endosperm the seeds are clas-
another. When the carpels are united the gynoe- sified into two types.
cium is known as syncarpous; a gynoecium
1. Endospermic seeds: the endosperm is
with a single carpel is said to be apocarpous.
present in the mature seed and serves as a
The apocarpous gynoecium is termed a sim-
food storage organ. Testa and endosperm
ple pistil and the syncarpous gynoecium is
are the two covering layers of the embryo.
termed a compound pistil.
The amount of endosperm in mature seeds
is highly species-dependent and varies from
The vascular skeleton of the flower an abundant endosperm layer (Nicotiana
In structure, the petiole is a typical stem, tabaccum) to a single layer (Arabidopsis
herbaceous or woody, with a ring of vascu- thaliana).
lar bundles or with an unbroken cylinder of 2. Non-endospermic seeds: the cotyledons
vascular tissue. In the receptacle the stele is serve as the sole food storage organs as in
modified in shape, often expanding and the case of sunflower (Helianthus annuus).
becoming like an inverted or flattened cone During embryo development the cotyledons
or pyramid. From this receptacular stele absorb the food reserves from the endosperm.
depart the traces to the various floral organs, The endosperm is almost degraded in the
traces which in origin, structure, and behav- mature seed and the embryo is enclosed by
iour are similar to those of leaves. Gaps the testa.
accompany the exit of traces, and the crowd-
ing of the organs thus breaks the cylinder up Seed coat
into a network of strands. The traces pass
off successively to sepals, petals, stamens The seed coat is the outer protective layer of
and carpels, according to the manner of the seed, developed from the integuments
arrangement of these parts in the flower. of the ovule, diploid maternal tissue. The
seed coat covers the embryo as protection.
The seed coat consists of two layers, the
testa and the tegman.
4.7.5 Anatomy of seed
1. The testa is the outermost layer of the
Introduction seed coat, and is formed from the outer integ-
ument of the ovule. It is usually hard and
Seeds are the dispersal and propagation impermeable to water and sometimes con-
units of the Spermatophyta, which includes tain ornamentations. Testa contains macro-
gymnosperms (conifers and cycads and and microsclereids. Macrosclereids are
related species) and angiosperms (flower- arranged very compactly and are dumbbell-,
ing plants). Seed is an integumented ferti- boat-, bone-shaped (osteosclereids). Micro-
lized ovule consisting of embryo. The sclereids are small in size with a square
constituent parts of a seed include a seed shape and dentate edges. Hardness of the
coat, cotyledons, endosperm and embryo. seed coat depends on thickness of the testa
The embryo is a young sporophyte, diploid and the arrangement of sclereids. Rate of
40 Chapter 4
absorption of water (imbibition) depends on materials unused by the embryo are known
the thickness of the testa. Very hard and as perisperm. Usually perisperm consists of
thick testa causes seed dormancy in some large parenchyma cells with stored food
plants (Fig. 4.26). materials like oil granules and tannins. It is
2. The tegman is the inner layer of the seed observed only in some species, e.g. Coffea
coat. It is formed from the outer integument arabica and Beta vulgaris.
and is thin and soft. It covers the embryo.
growth of the integuments on the micro- Embryo development begins in the em-
pylar end. The caruncle helps to absorb bryo sac, following fertilization and
water during seed germination. divi sion of the zygote. In general, the
mature angiosperm embryo consists of
an embryonic axis, with either a single
cotyledon (monocotyledons) or a pair of
4.7.6 Anatomy of embryo and young cotyledons. There are apparent exceptions,
seedling e.g. in some species of Apiaceae and
Ranunculaceae the cotyledons are fused at
Anatomy of embryo maturity. In angiosperms the number of
cotyledons rarely exceeds two, though
The morphology and underlying anatomy
individuals of some species may develop
of the embryo in mature seed varies con-
three or more cotyledons (Magnolia gran-
siderably among species (Figs 4.27, 4.28).
diflora). Gymnosperm embryos frequently
have more than two cotyledons, sometimes
as many as 12 (polycotyledonous), e.g.
Pinus, Pinea, depending on species.
The embryonic axis bearing the coty-
Seed coat
ledons commonly shows a polarity from
the earliest stages of embryogenesis, with
Cotyledons the proximal end towards the micropyle
being the radicle containing the root meri-
stem, and at the distal end of the shoot the
meristem (plumule), sometimes with rec-
ognizable leaves. The radicle is usually
adjacent to the micropyle, through which
it emerges on completion of germination.
The first shoot segment above the cotyle-
dons is the epicotyl, and the region of the
Fig. 4.27. Transverse section of cotton seed (40×) axis between the radicle and the point of
illustrating dark seed coat and cotyledons showing insertion of the cotyledon is the hypocotyl.
black spots indicating gossypol glands. The cotyledons are often well developed
Coleoptile
(a) (b)
Corneous
endosperm
Floury
endosperm
Radicle
Fig. 4.28. Transverse section of maize (40×) seed showing coleoptile and radicle (a) and corneous
and floury endosperm (b).
42 Chapter 4
and serve as food storage organs in non- Table 4.1. Families showing number of traces.
endospermic seeds. Number of traces Family
The embryo contains thin-walled paren-
chyma cells. The cells are round or polyhe- 2 Most of the lower
dral in shape with intercellular spaces, monocotyledonous
and often packed with starch or aleurone. families
2 Liliaceae
Some isolated areas of embryo bear pro-
2 Zingiberaceae
cambium and a few mature protoxylem and
2 Amaryllidaceae
protophloem. 1, 2–3 Iridaceae
1 to several Araceae
Several Cannaceae
Anatomy of cotyledon 2 Ranales
References
Cereals are the carbohydrate rich food grains, of rice are recognized, indica, japonica and
which are consumed largely by humans and javanica. Indica rice is adapted to tropical
provide the daily need of nutrients for sur- and subtropical humid climates, while japo-
vival and livelihood. Examples of cereals nica is adapted to temperate regions. The
include rice (paddy), wheat, maize, sorghum, javanica rice is limited to tropical regions of
pearl millet etc. This book is mainly concen- South-east Asia.
trated on the anatomy (including general
botany) of the following field crops: rice,
maize, wheat, sorghum and pearl millet.
5.1.2 Origin, distribution and adaptation
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
44 (R. Maiti et al.)
Cereals 45
Today, the majority of all rice produced comes also be made into various types of noodles.
from China, India, Pakistan, Indonesia, Raw, wild, or brown rice may also be con-
Bangladesh, Vietnam, Thailand, Myanmar, sumed by raw-foodist or fruitarians if soaked
the Philippines and Japan. Asian farmers and sprouted.
still account for 92% of the world’s total
rice production. Rice is grown in all parts
of India, northern and central Pakistan.
Basmati rice cultivated in the northern plains 5.1.4 Morphological description
of the Punjab region is famous all over the
world for its smell and quality. Vegetative characters
Rice is well adapted to diverse environ-
mental conditions and climates from sea Rice is an herb having a plant height of
level up to high elevations. It is grown in 2–4 m. The plant generally takes 3–6 months
north-eastern China at 53°N of the equator to complete its life cycle. Most cultivated
and in New South Wales at 35°S in Australia species are hydrophytic and a few species
and is widely adapted below sea level at are mesophytic. Rice has a fibrous root sys-
Kuddanad (Kerala) to above 2000 m in tem, adopted for mostly hydrophytic condi-
Kashmir. Most of the rice areas lie between tions. Seminal roots help in the establishment
the equator and 40°N latitude and between of the seedling and are short lived. In peren-
70° and 140°E longitude. The highest yields nial rices, short underground shoots densely
are recorded between 30° and 45°N of the covered with sheathed leaves are present.
equator. Annual species have long but slightly branc-
O. sativa is the dominant rice species, hed adventitious roots, whereas a main root
cultivated in Asia, Africa, Europe, north- and subterranean shoots are completely
Central and South America and Oceania. absent. The stem is referred to as a culm or
O. glaberrima is another cultivated species stalk. The culm is composed of nodes and
limited to Africa. Indica rices grow widely internodes, and is enclosed within the
in tropical regions and are adapted to cooler sheath, but a small portion of the culm just
areas and largely grown in temperate cli- below the panicle is exposed after heading.
mates such as central and northern China, The number of elongated internodes ranges
Korea and Japan. A number of biological, from three to eight. Internode elongation is
environmental and socio-economic condi- associated with growth duration. Tillers
tions cause the existence of the large differ- develop from the leaf axils at each un-
ences in rice productivity around the world. elongated node of the main shoot or other
tillers during vegetative growth. Tillers
should not be lodging; tiller stems with strong
mechanical tissue (sclerenchyma) may impart
5.1.3 Utilization non-lodging. Tiller stems may be screened
microscopically for the variations in the inten-
As a cereal grain, it is the most important sity of sclerenchyma (Maiti, unpublished).
staple food for a large part of the world’s Rice leaf is composed of the leaf sheath, the
human population, especially in East, blade, the ligule and the auricles. The sheath
South, South-east Asia, the Middle East, is elongated, with a ribbon-shaped leaf base
Latin America and the West Indies. Rice is enrolling the culm. The blade is narrow, flat
cooked by boiling or steaming, and absorbs and longer than the sheath. At the base of
water during cooking. In some countries the leaf blade, a white band structure called
parboiled rice is popular. Raw rice may be the collar is present. A pair of hairy and
ground into flour for many uses, including sickle-shaped auricles is situated at the
making many kinds of beverages such as junction between the collar and the sheath.
amazake, horchata, rice milk and sake. Rice Leaf sheaths may possess cilia. Like stalks,
flour does not contain gluten and is suitable leaf sheaths also have a clearly marked or
for people on a gluten-free diet. Rice may ribbed venation and become keel-like by
46 Chapter 5
the protrusion of the mid-vein in the upper enclosed in the lemma and palea with awns.
part of the sheath beneath the leaf lamina. Flowers are sessile, bisexual, ligulate, zygo-
Leaf lamina consists of parallel venation. morphic, epigynous and trimerous. In every
The ligule is oblong or blunt and 1–5 mm flower, lemma, palea, two lodicules, pistil
long in most of the species. In O. latifolia, and stamens are present. There are three sta-
the ligule is short but breaks up at the apex mens, long anthers, dithecous, basally fixed;
into a row of hard, sometimes woolly hairs. the stamens have about 2–2.5 mm long,
linear-oblong anthers. The anthers dehisce
Floral characters through a longitudinal slit. The pistil is com-
posed of the stigmas, styles and ovary. The
The rice panicle consists of the base, axis, ovary is unilocular, unicarpellary, with a sin-
primary and secondary branches, pedicel and gle ovule on basal placentation, silky-like
glumes, and bears hundreds of spikelets, the style, and bifid feathery, plumose stigma.
characteristics of many cereals (Fig. 5.1). Two fleshy bodies called the lodicles are
The panicle axis is extended from the pani- situated at the base of the ovary. The lodi-
cle base to the apex and may contain from cles swell with water, thereby separating
eight to ten nodes. Panicles are developed the lemma and palea at flowering. Anthesis
from the nodes of primary and secondary involves a series of events. At the beginning
branches on the top of which spikelets are of anthesis, the tips of the lemma and palea
located. Articulation is present in all rice begin to open, the filaments elongate, and
species between lower and upper empty anthers begin to emerge from the lemma
glumes. This is fairly distinct in most spe- and palea. The filaments elongate more to
cies because the spikelets fall easily on rip- push the anthers out of the lemma and
ening of seeds. However, it is less distinct in palea. Anthesis lasts about 1–2.5 h. Anther
the cultivated species O. sativa, where dur- dehiscence take place just before the exten-
ing the ripening of fruits the spikelets do sion of the lemma and palea, with many
not drop off but remain on the branches. pollen grains falling on the sticky stigmas.
The spikelet is borne on the pedicel, a short Rice is a self-pollinated crop and the plants
stalk or secondary branch. Two rudimentary are pollinated by wind (anemophily). The
glumes are present on the upper part of the complete process from pollination to fertili-
pedicel. A pair of sterile lemma and the zation takes from 18 to 24 h. The fruit is a
rachilla are situated between the rudimen- caryopsis and generally called grain. Fruits
tary glumes and the spikelet. The flower is are small, compressed from the sides, 1–8
mm long, oblong, elliptical or orbicular,
depending on the shape and size of the
spikelet. Small fruits of rice are initially yel-
lowish, and then turn dark brown. Seeds are
endospermic.
Root anatomy
absorption of water from the soil. Root hairs The root system in rice is poorly developed
are generally short lived. The epidermis is compared to other cereal crops and root
present as a layer of sclerenchyma imparting hairs and root cap are absent, but in some
strength to the root system. genotypes there may be a root packet. A high
Cortex is parenchymatous and in the proportion of air chambers in the root is
central region, present between epidermis suitable for aeration of internal tissues under
and the stele. The cortex is poorly devel- anaerobic submerged conditions, which help
oped. Numerous air spaces form aerenchyma in gaseous exchange. In some genotypes there
in the cortical region for gaseous exchange may not be air chambers in the root. There is
(Fig. 5.2). a reduction in mechanical tissue and vascu-
The centre region is occupied by vas- lar tissue, but that reduction may be less or
cular tissue forming stele. The vascular more in some genotypes. The presence of
system is poorly developed because the sclerenchyma below the epidermis is desir-
rice plant is adapted for a hydrophytic able for imparting strength to the tissue sys-
habitat. Xylem and phloem are arranged in tem of roots.
the form of separate strands on different Traditional upland indica cultivars
radii. Xylem consists of very small xylem possess a larger diameter of root stele and
vessels. Sclerenchyma is almost absent; xylem vessels compared to modern upland
cambium and pith are absent. varieties. Japonica rice has a significantly
In hydroponically grown rice, roots are larger stele diameter relative to the root
observed to be bounded by an epidermis, diameter (Kondo et al., 2000).
exodermis, and a fibrous layer (Clark and Asian cultivated rice (O. sativa L.) has
Harris, 1981). The exodermis has a suberin genetic diversification of root characteris-
lamella along its inner tangential wall. tics, but this variation has not been eluci-
dated completely with reference to the
Significance of variations in root anatomy genetic background. To elucidate the varia-
tions in root anatomical and morphological
The structural anatomy of rice root is adapted traits among diverse varietal groups of culti-
more towards a hydrophobic environment. vated rice, Uga et al. (2009) analysed four
anatomical traits (stele transversal area, root
COR ARC END thickness, total transversal area and number
PC
HYP X of late metaxylem vessels) and two morpho-
EP PHL
logical traits (ratio of deep rooting and root
length index) in 59 accessions. An earlier
principal-coordinate analysis study using
data on 179 restriction fragment length poly-
morphisms categorized these accessions
into three varietal groups: 13 japonica, 21
indica-I, and 25 indica-II. Based on a prin-
cipal-components investigation of the six
traits, the japonica group had extensive var-
iation in root anatomy compared to the two
indica groups. In particular, japonica upland
MED rice was characterized by a larger stele and
xylem structures (Uga et al., 2009).
Epidermis is the outermost region of closely arranged, and the central bundles are
the stem and is meant for protection. It is large and are widely arranged. Each vascular
uniseriate, having a single layer of com- bundle is oval in shape and is surrounded by
pactly arranged barrel-shaped living cells. fibrous bundle sheath. It is many celled thick
The outer surface of the epidermis is cov- towards outer and inner regions and few
ered by a thin cuticle. A few stomata (grami- celled thick at lateral regions, hence the vas-
naceous type) are present in the epidermis cular bundles are described as fibrovascular
for the exchange of gases necessary for adap- bundles. Vascular bundles are conjoint, col-
tation to the hydrophytic system. lateral (with xylem and phloem present on
Cortex is highly reduced and is usu- the same radius) and closed (i.e. without
ally represented by a narrow hypodermal cambium). The xylem is present towards the
region. In mature stems hypodermis is scle- centre of the axis and consists of few vessels
renchymatic and gives mechanical strength. which are arranged in the form of letter ‘Y’.
In young stems chlorenchymatic patches The protoxylem often disintegrates and
are present just below the stomata regions forms protoxylem lacuna. Phloem is present
and perform photosynthesis. The cortex towards outside, between the metaxylem
shows a lot of air chambers (Fig. 5.3). vessels. It consists of sieve tubes, companion
Vascular bundles are present just below the cells and a few parenchymatous cells at the
epidermis. sides. The stele consists of numerous vascu-
Stele is the central part of the stem and lar bundles scattered irregularly in the ground
is larger than the cortex. The stele consists tissue, and is called an atactostele. It is the
of ground tissue and vascular bundles. most advanced type of stele.
There is no clear demarcation between the
cortex and pith since endodermis, pericycle
etc., are indistinguishable. The ground tis- Significance of variations
sue also shows air chambers. The central in stem anatomy
part of the stem is parenchymatic and is
best described as ground tissue (or conjunc- Rice stem is less defined in structure than in
tion tissue). sorghum or wheat stem. It adapts well to
Vascular bundles are numerous and are anaerobic conditions by formation of cham-
scattered irregularly in the ground tissue. bers (aerenchyma) in the cortex region for
Usually the peripheral bundles are small and the exchange of gases. Variation in scleren-
chyma structure arises from differences in
the thickening of the cell wall in the outer
cortical parenchyma and the number of
sclerenchyma cell layers, and contributes to
ARC both biotic and abiotic stress tolerance.
GT
VB Since japonica rice has higher thickening of
EP sclerenchyma than indica rice, it is more
tolerant to stem borers and desiccation. Rice
exhibits thinner cuticle and less wax depo-
sition on the epidermis compared to other
cereal crops, although genotypic variation
for cuticle thickness is observed among and
within subspecies.
Leaf anatomy
section shows epidermis, mesophyll and nature due to the presence of silica crystals.
vascular bundles (Fig. 5.4). The upper epidermis may be easily identi-
The epidermis is uniseriate and composed fied due to the presence of xylem and bulli-
of more or less oval cells. Bulliform cells are form cells. As the leaf is isobilateral, the
observed in the adaxial surface of the leaf mesophyll is not differentiated into palisade
blade, which helps in rolling the leaves and spongy tissues. The mesophyll tissue of
under low water levels. The outer walls of rice leaf is composed of lobed chlorenchyma
the epidermis are covered by a thin layer of with a length:breadth ratio of approximately
cuticle. Graminaceous type of stomata is 2:1. Large air chambers (aerenchyma) are
present in both epidermal layers. Stomata present in the mesophyll tissue, which is a
are covered by two dumb-bell shaped guard characteristic feature of a hydrophytic plant.
cells. Stomata are usually more frequent in Papilla and trichomes cover the entire leaf
the lower epidermis. Epidermis is rough in surface except on the adaxial surface of the
leaf sheath.
The vascular system is poorly devel-
ARC oped because the rice plant is adapted for a
SCL
hydrophytic habitat. The vascular bundles
MES are collateral and closed. Most of the bundles
UPEP are small in size but fairly large bundles also
occur at regular intervals. Xylem is found
towards the adaxial side and phloem towards
the abaxial side of the bundles (Fig. 5.5).
There are distinct anatomical differ-
ences between the early leaves (first to third)
TRC and leaves that appear later. The first leaf of
SCL
rice lacks a leaf blade and is very small in
VB
size compared to other leaves. The fifth and
LOEP PAR
MES BRC
later leaves have a strong midrib, which pro-
vides mechanical strength to the leaves. The
Fig. 5.4. Transverse section of leaf (100×) showing leaves also differ in the capability of photo-
upper epidermis (UPEP), lower epidermis (LOEP), synthetic activities; early leaves are photo-
sclerenchyma patches (SCL), vascular bundles (VB), synthetically less efficient than later leaves.
aerenchyma (ARC), parenchyma (PAR), mesophyll Development of leaf blade and leaf
(MES) and brachisclereid (BRC). sheath in rice is well coordinated. The leaf
Parenchyma MX
PRX
Aerenchyma
Fig. 5.5. (a) Transverse section of leaf (400×) showing the aerenchyma and vascular bundle; (b) vascular
bundle (400×) showing proto- and metaxylem regions.
50 Chapter 5
develops from the shoot apical meristem by developed when compared with xylem. These
rapid cell division. Leaf sheath elongation characters exhibit better adaptation to an
starts only after leaf blade elongation is aquatic nature.
completed. Air spaces known as lacuna are
formed in the leaf sheath and midrib of the Anatomy of rachilla
leaf blade.
Transverse section of the rachilla shows
ridges and grooves in outline (Fig. 5.6).
Significance of variations in leaf anatomy
Single-layered epidermis covers the internal
Rice and wheat leaves are distinct from tissues. There are four to five layers of scler-
other C3 monocot leaves in terms of higher enchyma present beneath the epidermis in
photosynthetic capacity, which has prima- the region of ridges and grooves. Mesophyll
rily resulted from the extensive selection tissue is encircled by sclerenchyma in the
and breeding for higher productivity. Rice region of grooves. Small vascular bundles
leaves differ from wheat in allocation of leaf are situated in the ridges. Parenchyma forms
nitrogen to rubisco (ribulose-1,5-bisphosphate ground tissue, leaving a central airspace.
carboxylase/oxygenase), the principal pho- Numerous large vascular bundles are dis-
tosynthetic enzyme, and rice also has a tributed throughout the ground tissue. Each
higher mesophyll cell area than wheat. In vascular bundle is collateral, closed, and
the nitrogen responsive cultivars, these areas covered by sclerenchyma tissue.
increase consistently along with increase in
intercellular air space favouring photosyn- Fruit anatomy
thetic activities. Air chambers are present in
rice mesophyll cells, which help in oxygen Rice fruit is called a caryopsis. Pericarp
supply as well as re-fixation of CO2 gener- tightly covers the grain (seed) and is gener-
ated from photorespiration. These air cham- ally known as husk. The grain is covered
bers are large, regular and are separated by by an aleurone layer consisting of proteins.
partitions of photosynthetic tissues. Higher Endosperm occupies the major portion in
adaptation of rice than wheat in hot and grain. The embryo (germ) is situated one
humid areas is attributed to better photosyn-
thetic activity in rice leaves due to better Sclerenchyma
spatial arrangements of chloroplasts in meso-
Collenchyma
phyll cells, higher mesophyll cell number
and availability of air spaces to re-capture
CO2 released through photorespiration
(Makino, 2011). The outer layer of chloren-
chyma cells in rice is covered by chloroplast
and stromules (stroma-filled appendages),
which is absent in wheat (Sage and Sage,
2009). The general features of rice leaves are
also conserved largely in the wild relatives, Aerenchyma Epidermis
which suggests that selection did not bring
much change in the leaf structure; rather, Ground
rice has been selected for having leaf struc- tissue
ture with high photosynthetic capacity.
The strength in rice leaves is provided Vascular bundle
by the large midrib, sclerenchyma strands
and often by the presence of sclereids. There Fig 5.6. Transverse section of rachis (100×) showing
is a great reduction in vascular tissue, small sclerenchyma, collenchyma in the region of ridges,
and big vascular bundles are arranged in an ground tissue, vascular bundles and central
alternate manner. In small vascular bundles, aerenchyma. Larger vascular bundles are distributed
xylem elements are lacking. Phloem is well in ground tissue.
Cereals 51
side of the bottom. The endosperm consists Several factors were involved in the diversi-
of protein starch-storing parenchyma cells fication of maize, including the existence of
called amyloplasts. primitive landraces, the influence of exotic
varieties from the northern part of the coun-
try, natural hybridization with other races
5.2 Maize as well as favourable geographical condi-
tions for growth of short-season maize.
5.2.1 Introduction Maize was considered as ‘mother corn’ by
the Native American Indian communities. It
Maize (Zea mays, 2n = 20) is the most was introduced in northern America around
important food and animal feed crop of the 700 years ago, where it was adapted within
western hemisphere, and is grown through- a very short time period. During the Spanish
out the world under a wide range of cli- discovery of the New World, maize was
mates. Since pre-Hispanic time, it has been brought to the soil of Europe, from where it
the basic food for the majority of people in spread rapidly to Asia and Africa, primarily
Mexico and Latin American countries. From through the establishment of Portuguese
about AD 900 to AD 1300 there was a shift colonies in Africa and Asian countries.
from intensive foraging with some agricul- Today it is a major food crop of almost all
ture to intensive farming with some forag- the continents.
ing. This was accompanied by more densely The origin of maize has been debated
settled population and greater cultural com- for a long time, but major research findings
plexity among people in the mid-western support that the most probable progenitor
hemisphere. The maize in the upper mid- of maize is teosinte (Zea mexicana) or
west around AD 600 was poorly adapted to ‘Balsas’ teosinte (Zea mays parviglumis), a
the short growing season and photoperiod weedy member of genus Zea (Beadle, 1939).
of the more northerly latitudes. After AD However, how the simple ear of teosinte
1100, the strains known as Northern Flint evolved into the large maize cob within a
Corns were developed with better photo- short evolutionary period of about 10,000
periodic response, yield, pest resistance years is a baffling question to maize research-
and hardiness. In scientific and formal ers. Several theories have been put forward
usage, ‘maize’ is normally used in a global for explaining the origin of maize. The
context. Equally, in bulk-trading contexts, teosinte theory supported by major research
‘corn’ is used most frequently. groups is contradicted by other theories, which
consider maize originated from Tripsacum
dactyloides (Mangelsdorf and Reeves, 1938),
Zea diploperennis, hybrids of Tripsacum
5.2.2 Origin, evolution and domestication and teosinte (tripartite theory), from trans-
mutation of teosinte male ear to maize
Maize is considered to have originated on female cob (Iltis, 1983) or from ancestral
the Pacific slopes of the Mesoamerica 7000– maize species through duplication of chro-
10,000 years ago (Wilkes, 2004). Specimens mosomes (Gaut and Doebly, 1997).
of primitive cobs of maize have been recov-
ered in caves in Oaxaca, Mexico, that date
back to 8000 BC. Mexico is one of the centres 5.2.3 Genetic resource
of origin of maize, where several native
landraces grow in abundance in diverse Since a population of maize is a mixture of
environments. From Mexico, it diffused to heterogeneous and heterozygous genotypes, it
South America through the Andes during is best described as a race (a group of indi-
6000–2000 BC. Studies on microsatellite var- viduals having enough common traits for
iation of maize genomic DNA indicates that grouping). The studies on numerical taxon-
maize arose from a single domestication event omy and molecular phylogeny of maize have
around 9000 years ago (Matsuoka et al., 2001). helped to identify a number of races and their
52 Chapter 5
geographical relationships. Early classifica- There are also other speciality types of maize
tions of maize races were based on morpho- such as blue corn, high lysine corn, high oil
logical, physiological, genetic and agronomic corn, baby corn or ornamental corn.
features and cytogenetic traits. It has been
observed that the Mesoamerican and South
American regions show greater genetic diver- 5.2.4 Utilization
sity and race differences than the northern
American regions. For example, more than 65
Maize and cornmeal (ground dried maize)
races of maize have been identified in Mexico,
constitutes a staple food in many regions of
while in the USA, only 14 races are observed.
the world. Maize meal is also used as a
The number of races of maize in South
replacement for wheat flour, to make corn-
American countries such as Bolivia (77) and
bread and other baked products. Popcorn
Peru (68) are higher than that of North
and corn flakes are a common breakfast
American countries (Hernández, 2009).
cereal found in many countries all over the
However, based on kernel (seed) char-
world. Sweetcorn, a genetic variety that is
acteristics, the following subtypes of maize
high in sugars and low in starch, is usually
are commonly recognized by growers and
consumed in the unripe state. When maize
consumers:
is ground into flour, maize yields more flour,
with much less bran, than wheat does.
1. Dent corn (Zea mays indentata) – upon
However, it lacks the protein gluten of wheat
maturity, the kernels become indented in
and, therefore, makes baked goods with poor
the crown of the seed. Dent corn is com-
rising capability and coherence. Certain
monly used as human food, for industrial
varieties of maize have been bred to produce
purposes and livestock feeds all over the
many additional developed ears. These are
world. The denting is caused by different
the source of the ‘baby corn’ used as a vege-
degrees of drying and shrinking of starch.
table in Asian cuisine.
2. Flint corn (Zea mays indurata) – hard,
Grain alcohol from maize is tradition-
smooth kernel of flint corn contains various
ally the source of bourbon whiskey. Starch
proportions of hard and soft starch. It is
from maize can also be made into plastics,
popular in Central and South America.
fabrics, adhesives and many other chemical
3. Sweetcorn (Zea mays var. saccharata/Z.
products. The corn steep liquor, a plentiful
mays var. rugosa) – immature kernels are
watery by-product of maize wet milling
sweet (contains ~10% sugar) and consumed
process, is widely used in the biochemical
raw. In mature kernels, sugar is converted to
industry and research as a culture medium
starch, so the sweetness is reduced. The
to grow many kinds of microorganisms.
sweetness is caused by a recessive gene sug-
Maize is increasingly used as a feedstock for
ary-1 (su-1), which retards conversion of
the production of ethanol fuel. Maize leaves
sugar to starch in the endosperm.
are used as forage in most countries and
4. Flour corn (Zea mays amylacea) – kernels
sometimes as mulching material.
are soft, smooth and filled, and mainly used
for production of white flour. Its use is lim-
ited in parts of the USA and South America
because it is used mostly as feed grain.
5. Popcorn (Zea mays everta) – a type of 5.2.5 Morphological description
flint corn with very hard exterior. The starchy
endosperm explodes and becomes fluffy upon Maize has been described in numerous
heating, giving the characteristic roundish botanical books and illustrations. The Aztec
shape of popcorn. and Mayan civilizations had their own god
6. Waxy corn (Zea mays) – a mutant type, named after maize, whose pictures revealed
having waxy endosperm, where all the amy- maize cobs. After the discovery of the New
lose of starch is replaced by amylopectin. It is World, the first detailed description and
caused by a recessive mutation ‘waxy’ (wx). illustration of maize can be found in ‘Historia
Cereals 53
Natural y General’ by Gonzalo Fernandez de short and narrow, arranged in two rows on
Oviedo (1535) of Spain, although a descrip- stem, with an expanded leaf base with leaf
tion of maize can be found in Columbus’s sheath, wavy margin and parallel venation.
chronicle as early as in 1511 (Finan, 1948).
Since then, various descriptions of the maize Floral characters
plant have been documented in several
books, monographs and articles. In this sec- Maize has a diclinous inflorescence, with
tion, a brief general morphology of modern male and female inflorescences arranged on
cultivated types of maize is outlined. the same plant. The male inflorescence is
known as tassel and is present at the apex
Vegetative characters (top) of the stem. The female inflorescence
is known as a cob, and is present on the tips
Maize is a mesophytic herb, less drought of lateral branches, which are developed
tolerant compared to sorghum and pearl from the axils.
millet; it grows up to 2–4 m in height. The The male inflorescence is modified into
root system is fibrous; stilt roots are formed a panicle (inflorescence of male flowers),
on the lower nodes of the stem and a ring of with spikelets in pairs. In each pair, one is
crown roots arise from the stem just above sessile. The spikelets are covered by glumes.
the ground level. The stem is aerial, erect, The female inflorescences are the ears,
greenish, hard and succulent, with nodes which are tightly covered over by several
and internodes reaching 20–30 cm; the apex layers of leaves, and so closed-in by them to
of the stem ends in the tassel (Fig. 5.7). the stem that they do not show themselves
Leaves are simple, exstipulate, alternate, easily until the emergence of the pale yel-
sessile, lower leaves are 50–100 cm long low silks from the leaf whorl at the end of
and 5–10 cm wide, upper leaves somewhat the ear. The silks are elongated stigmas that
look like tufts of hair, at first green and later
red or yellow. Spikelets are arranged in
pairs on the cob and are covered by cupules;
they are sessile, having a female flower.
Flowers are sessile, unisexual, zygo-
morphic, epigynous; in every flower there
is lemma, palea and two lodicules. The
androecium has three stamens, long anthers,
dithecous and basally fixed. The gynoecium
is unilocular, unicarpellary, single ovule in
basal placentation, with a silky-like style
and a feathery stigma.
When the tassel is mature and condi-
tions are suitably warm and dry, anthers on
the tassel dehisce and release pollen. Maize
pollen is anemophilous (dispersed by wind)
and because of its large settling velocity
most pollen falls within a few metres of the
tassel. Each silk may become pollinated to
produce one kernel of maize.
Fruit is generally referred as grain or
kernel and is botanically called a caryopsis.
The kernel of maize has a pericarp of the
fruit fused with the seed coat, typical of the
Fig. 5.7. Maize plant morphology: tassel occurs at grasses. The cob is close to a multiple fruit
the apex of stem and silk at the centre of the stem in structure, except that the individual fruits
from the ground level. (the kernels) never fuse into a single mass.
54 Chapter 5
The grains are about the size of peas, and vascular bundles that contain xylem and
adhere in regular rows round a white pithy phloem in alternate bands (Fig. 5.8).
substance, which forms the ear. An ear con- The epidermis consists of elliptical
tains from 200 to 400 kernels, and is from cells subtended by two layers of hypoder-
12–25 cm in length. They are of various col- mis. In the early stage of root development,
ours: blackish, bluish-grey, purple, green, the cortex is made up of ovoid parenchyma-
red, white and yellow. tous cells with considerable intercellular
Seeds are endospermic and filled with space.
carbohydrates; based on the nature of carbohy- As the root ages, the cortical cells elon-
drates they are of different shapes and sizes. gate to assume a plate-like appearance.
A single layer of ovoid to cubical cells of
cortical cells of cortical parenchyma (mul-
tilayered) encircles the endodermis. The
5.2.6 Anatomical description endodermal cells are barrel- to boat-shaped
and are thickened with suberin on the inner
Anatomical features of maize have been tangential wall. Crystals are present in the
well documented by different researchers. endodermal cells; their shape and size vary
However, a comprehensive collection of with cultivar and with age of the root. The
anatomical features describing essential pericycle below the endodermis is com-
anatomical structures of root, stem, leaf, flo- posed of one to several layers of thick-walled
ral parts and seed along with their biologi- sclerenchymatous cells, of which the outer-
cal significance is lacking. most layers are highly lignified.
Xylem and phloem tissues are present
Root anatomy on a diffident radius and are called radial
vascular bundles. Xylem parenchyma sur-
There are two distinct types of roots in the rounding the metaxylem may be thick-
maize root system, the embryonic roots that walled or lignified. The xylem and phloem
develop from young embryo and the postem- show a typical closed radial arrangement.
bryonic (adult) roots that develop during Protoxylem bundles are present on the exter-
different stages of crop growth. The general ior side of the metaxylem. The size of the
anatomical features of both types of root are metaxylem bundle varies according to cul-
similar and are typical of monocots. The tivar. The pith is solid with round inter-
maize root is composed of epidermis, ground mediate to compactly arranged parenchyma
tissue (cortex), endodermis surrounding the cells.
E X PX C
(a) (b)
XV
MX
ME
Fig. 5.8. (a) Transverse section of root (100×) showing different parts: cortex (C), endodermis (E), protoxylem
(PX), metaxylem (MX) and medulla (ME). (b). Sector enlarged (400×) showing xylem vessels (XV).
Cereals 55
Significance of variations in root anatomy soil for extraction of available moisture com-
pared to drought-susceptible lines (Hund
Cultivar differences are expressed in a num- et al., 2009).
ber of root characteristics in maize (Fig. 5.9a, b).
These are:
Stem anatomy
1. Presence or absence of sclerenchymatous
exodermis in the cortex imparting strength Stem anatomy of maize is typical of that of
and protection to internal tissue. the grasses. The ontogeny and anatomy of
2. Thickness of endodermal cell walls. vascular bundles in maize has been descri-
3. Size and shape of crystals in the endo- bed earlier by Esau (1943).
dermal cell cavity.
4. Intensity of lignification in the pericycle YOUNG STEM The epidermis consists of a
cell layers and in cells surrounding the vas- single layer of cells with thickened walls on
cular bundles. the outer side. The ground tissue consists
of thin-walled parenchyma. The vascular
The presence of exodermis, higher thick-
bundles, conjoint collateral in type, are scat-
ness and higher lignification provides better
tered in the ground tissue.
structural integrity to the root system. Thus
genotypes exhibiting these characteristics
are expected to have better crop stand and MATURE STEM In a transverse section, the
resistance to lodging. Heavy lignification of epidermis consists of cubical to boat-shaped
exodermis of maize root gives higher bend- epidermal cells containing elongated crys-
ing strength, thus providing better anchor- tals. Just below the epidermis, and opposite
age (Ennos et al., 1993). Thicker roots are the ridges of the stem, there are alternate
advantageous for higher root growth and bands of large and small thick-walled scler-
help in better water transport. Higher root enchymatous patches. Each sclerenchyma-
diameter is associated with higher xylem tous band alternates with a broad band of
diameter in maize, which helps in better chlorenchymatous tissue corresponding to
solute transport. A number of drought-tolerant the furrows of the stem outline. These hypo-
maize lines developed by the International dermal bands of sclerenchyma are in turn
Maize and Wheat Improvement Centre connected to the subtending broad cylinder
(CIMMYT) have thicker root systems; thus of sclerenchyma, which contains the ring of
they are able to penetrate deeper in the vascular bundles. There are four to five rows
(a) (b)
Fig. 5.9. Transverse section of root (400×) showing (a) Casparian thickening in endodermal cells
and (b) Casparian thickening in endodermal cells and crystals in the pericycle.
56 Chapter 5
(a) GR (b) GR
HYP VB
EP
VB
PH
Fig. 5.10. (a) Transverse section of stem (100×) showing atactostele. (b) Sector enlarged (400×) showing
epidermis (EP), hypodermis (HYP), scattered vascular bundles (VB) in ground tissue (GR), xylem (X) and
phloem (PH).
Cereals 57
(a) (b)
BSS
HPS
X
PH M
PR
Fig. 5.11. (a) Transverse section of stem (400×) showing lignified patches of hypodermal sclerenchyma
(HPS) and bundle sheath cells (BSS). (b) Transverse section of stem (400×) showing vascular bundles
with protoxylem cavity (PR).
Fig. 5.12. (a) Strong stereome (400×) showing large vascular bundles. (b) Intermediate stereome (400×)
showing medium size vascular bundles. (c) Weak stereome (400×) showing small bundles and smaller
number of sclerenchyma patches below the epidermis.
the transposon tagging approach, a gene Bk2 semi-lunar. Bulliform cells are absent in the
controlling cellulose deposition has been iden- upper epidermis of the midrib and vascular
tified, which shows high expression in the bundles of different sizes are present below
vascular bundle in the wild type (Bk2/bk2) and the upper epidermis. In some genotypes the
reduced expression in the mutant (bk2/bk2). midrib has three large vascular bundles, of
which one is in the centre and one is to each
Leaf anatomy side at the junction of leaf lamina midrib.
Chlorophyllous tissue is present between the
The anatomy of the maize leaf is characteris- vascular bundles, but confined to the lower
tic of that of a mesophytic grass. There is portion of the midrib. The sclerenchyma
some variation among the genotypes studied, forms a band below the upper epidermis. The
but in general, leaf anatomy does not vary sig- vascular bundles are of three types in the lam-
nificantly from genotype to genotype. The ina and midrib: (i) the large central vascular
epidermis is cuticularized, and composed of bundle corresponding to the main vein; (ii)
rectangular or oval epidermal cells. The lower medium size vascular bundles; and (iii) very
epidermis is entire in outline. Silica crystals, small vascular bundles. The first two types
prominent in some genotypes and not so in are generally fibro-vascular, containing
others, protrude from the adjoining cell walls patches of sclerenchyma on both sides (con-
of two adjacent epidermal cells towards the necting the lower or upper epidermis), or at
cuticle. Both the upper and lower epidermis one side only (connected to the lower epi-
contain unicellular long trichomes (Fig. 5.13a). dermis). The third type – the small vascular
Upper epidermal cells are wavy in outline bundles – are generally present in the lamina,
and interspersed with zones of bulliform and are without any fibrous bands. These
cells, the size of which differ in different geno- laminar bundles are again of two sizes, alter-
types. Two dumb-bell shaped guard cells nating with each other. Each vascular bun-
cover the stoma, known as a graminaceous dle remains surrounded by a bundle sheath
type of stomata (Fig. 5.13b). The chlorophyl- consisting of thin-walled parenchyma cells
lous tissue surrounding the vascular bundles (Fig. 5.14a,b). The cells of the bundle sheath
may be loose or compact. The surface structure generally contain large chloroplasts and form
of leaf shows variation in stomatal frequency, starch; grana are absent. This special struc-
its size, and trichome density. The shape of ture was first identified by Haberlandt in
the midrib in transverse section is almost 1882. The structure appears like a garland
(a) (b)
Fig. 5.13. (a) Leaf upper epidermal impression (40×) showing unicellular long trichomes. (b) Leaf upper
epidermal impression (400×) showing epidermal cells, stomata (graminaceous type) consisting of dumb-bell
shaped guard cells.
Cereals 59
Mesophyll
Bulliform cells
X
PH
Lower epidermis
CHL
Fig. 5.14. (a) Transverse section of leaf (40×) showing different features. (b) Sector enlarged (400×) with
fibro-vascular bundle showing chloroplasts (CHL), bundle sheath (BS), xylem (X) and phloem (PH).
(a) (b) T B
P
SCL
UE
2
LE
1 M VB
Fig. 5.15. (a) Transverse section of leaf at middle vein (100×) showing big (1), medium (2) and small (3) size
vascular bundles. (b) Transverse section of leaf showing upper epidermis (UE) and lower epidermis (LE),
bulliform cells (B) and trichomes (T) in UE. Sclerenchyma (SCL) patches are present below the UE,
parenchyma (P) present between the UE and mesophyll (M).
(a) Sclerenchyma (b) Cuticle apex with the formation of panicle meri-
Sclerenchyma stem (panicle initiation stage). The panicle
meristem assumes a bulbous shape with a
Big VB constriction at its base.
The panicle meristem shows a definite
cellular organization. There are prominent
tunica and corpus layers in the panicle axis,
with two to three layers of seriately arranged
tunica cells overlying the corpus. The cel-
lular organization in the corpus layer is not
regular. The protoplasm in the tunica cells
and the nuclei with prominent chromatic
material are distinct.
Small VB
Spikelet primordia
(a) (b)
Cuticle
Sclerenchyma
Sclerenchyma
Fig. 5.17. Transverse section of leaf (400×) showing variation in thickness of sclerenchyma patches: (a) large
amount and (b) smaller amount of sclerenchyma.
with a constriction on the sides, indicating and division ultimately lead to the formation
the boundary between the adjacent spikelet of four pollen grains. In mature anthers
primordia. This sequence of development approaching the time of dehiscence, some
of the spikelet primordia gradually pro- fringe-like outgrowths are observed all along
gresses towards the tip of the panicle. The the outer wall of the anther.
differentiating vascular traces in the panicle
axis gradually establish vascular connection Structure of the ovule
with the developing spikelets.
About 3 days after initiation, the panicle The single ovule is present in the cavity of
is much more elongated and the formation of the ovary and is attached laterally to the
the spikelet primordia has advanced towards ovarian wall by a short funicle. The ovarian
the tip. Each primordium divides to give wall is composed of a single layer of cubical
two spikelets (in pairs), which subsequently epidermal cells, subtended by several layers
develop floral parts. Each spikelet is enclosed of thin-walled cells.
by a pair of glumes (one lower glume and one
upper). Beginning at this stage, each individ-
Seed anatomy
ual spikelet primordium divides and extends
laterally to give rise to two floret primordia. The fruit is known as a caryopsis, and usually
Each floret primordium then develops floral referred to as grain (seed). The seed consists
parts. Although spikelet development takes of seed coat, endosperm, scutellum (cotyledon)
place acropetally, subsequent development and germ (embryo).
displays a basipetal gradient.
Seed coat
Microsporogenesis
In a transverse section of the mature cary-
Microsporogenesis in maize is as in other opsis the structure of the seed coat and
angiosperms. The young anther (8 days after endosperm is distinctly visible. Thick cuti-
panicle initiation) appears as a mass of tissue cle overlies the epidermis. The pericarp
in which the microsporangia are embedded. consists of epicarp, hypocarp and meso-
The microsporangia are present as four sepa- carp. The epicarp is made up of elongated
rate masses in the four lobes of the anther. epidermal cells (Fig. 5.18a, b). The hypo-
These cells divide along periclinal walls, carp cells are obliterated. Owing to compres-
leading an inner mass of primary sporoge- sion, the mesocarp cells, which are two- to
nous tissue and outer primary parietal cells. three-layered, have lost their cell outlines at
The ultimate cells of sporogenous tissue some places. Elongated cross cells are only
are pollen mother cells, which by reduction rarely visible and the tube cells which are
62 Chapter 5
(a) (b)
PC
PC
END
END
Fig. 5.18. (a) Longitudinal section of grain (caryopsis 100×) showing endosperm (END) and pericarp (PC).
(b) Sector enlarged (1000×).
distinct at an early stage are mostly obliter- from the latter by a single layer of epithe-
ated. Below the mesocarp there is a single lial cells. The embryo consists of a radicle
layer of large regularly oriented cubical and and a plumule (Fig. 5.19). The radicle is
rectangular cells called the aleurone layer. partially covered and protected by cole-
Staining bodies are present in the aleurone orhiza. The width of the pericarp at the
cell and starch grain variation among hilar region is much reduced. The radicle
genotypes. is partially covered and protected by cole-
orhiza. The plumule is partially covered
Endosperm and protected by the aleurone cells, which
are small in size, compact, and rectangular
Below the aleurone layer is the peripheral
in shape. The endosperm cells between
corneous endosperm, a single layer of well
the aleurone layer and the scutellum are
demarcated cells with compactly arranged
compact. The scutellar cells are elongated
starch and protein granules. The endosperm
and form a single layer surrounding the
occupies about two-thirds of the total seed
embryo. The embryo cells are highly com-
and is located in the broader part of the
pact, especially at the hilar region. The
maize grain. There is a continuous layer of
structure of the caryopsis at the black-layer
the large cubical cells immediately beneath
region is similar, cells outside the black
the hull called the aleurone layer. This aleu-
layer present at the hilar region are highly
rone layer contains protein granules. The
compact. The black layer shows a semi-
rest of the endosperm consists of starch-
lunar ring of vacuolated cells in a network
laden cells, which also contain some lipid.
pattern, which appears to cut off the vas-
The cells of this layer are cubic, elon-
cular connection from the pedicel to the
gated, small and tightly pressed against the
grain. Formation of this black layer coin-
aleurone layer. Below this peripheral layer,
cides with the termination of grain
the endosperm cells are loose, large-sized,
development.
and elongated and cubic to angular in
A longitudinal section of the maize
shape. These constitute the floury endo-
grain soaked for 2 or 3 days should be taken
sperm. Starch and protein granules become
along the endosperm–embryo axis and
increasingly less compact towards the cen-
stained with dilute iodine solution in
tre of the endosperm.
order to study its structure. The outermost
Embryo coat enclosing the entire grains formed of
inseparably fused fruit coat (pericarp) and
The embryo of the maize grain is located seed coat is called hull. The hull is not
beneath the endosperm. It is demarcated stained by iodine. In a longitudinal section
Cereals 63
Radicle
Fig. 5.19. (a) Longitudinal section of grain (caryopsis) showing plumule, floury endosperm, corneous
endosperm and (b) radicle.
Maize is considered as a model species for Wheat (Triticum spp.) is the most impor-
seed development and embryogenesis stud- tant cereal crop of the world in terms of
ies in grasses. Quite a large number of mutant production (World production 685.61 Mt
types for kernel characters have been identi- in 2009), closely followed by rice (685.24
fied in maize. Anatomical studies of seed Mt in 2009) (FAO, 2011). China, India, the
mutants in maize have helped to understand USA and the Russian Federation are
a number of basic and applied questions the principal wheat growers of the world.
related to seed development, maturation and The wheat crop refers to different species
embryogenesis. Anatomies of embryogenic of genus Triticum, including diploid,
(emb) mutants, which are defective in differ- tetraploid and hexaploid species. Bread
ent stages of embryogenesis, are commonly wheat (Triticum aestivum, 2n = 6x = 42)
used for understanding the process of embryo- is the major species cultivated world-
genesis in maize and related species (Elster wide, covering more than 85% of wheat
et al., 2000). The kernel mutants are also area. Other economically important spe-
used for programmed cell death (PCD), cies are macaroni or durum wheat
which leads to cellular differentiation, struc- (Triticum durum, which is used to make
turation of different organs such as scutel- pasta, spaghetti and macaroni), poulard
lum and suspensor, mobilization of stored wheat (Triticum turgidum, 2n = 4x = 28),
products, kernel denting and other processes club wheat (T. aestivum subsp. compac-
involved with endosperm and embryo tum, 2n = 6x = 42), macha wheat (T. aesti-
development. vum subsp. macha) etc. A few other wheat
Variation in distribution and amount of species were grown earlier, namely spelt
floury and corneous endosperm may deter- wheat (Triticum spelta), emmer or wild
mine the water uptake and nutritional qual- emmer wheat (Triticum diccoides) and
ity of grains (Maiti, unpublished). einkorn wheat (T. monococcum), but these
64 Chapter 5
5.3.3 Utilization
close to the ground, called tillers (primary), interception. The whole process is complete
and the tillers produce additional tillers within about 5 min. Pollination is anemo-
known as secondary and tertiary tillers hav- philous. The pollen grain, which has a
ing their own root system. lifespan of about 5 h, when settled on a
Wheat leaves are alternate, simple, ses- stigma, germinates in about 1.5 h to produce
sile, exstipulate, with parallel venation a pollen tube. At normal temperatures, the
arranged in two rows on the stem. Leaves pollen tube reaches the embryo sac in about
consist of two parts, the leaf sheath, which 40 min. The wheat fruit (usually called a
encircles the stem, and the blade that bends grain) is a caryopsis with a thin-walled peri-
away from the stem. The thickened junc- carp enclosing a single seed coat and testa
tion is called a collar and the membranous is fused with pericarp. The shape is oval
outgrowth is called a ligule. The claw-like with a smooth and rounded dorsal surface
appendages attached to the base of the leaf and grooved or furrowed ventral surface at
and closely clasping the sheath constitutes the centre. The furrow or groove is called a
the auricles. The foliage arrangement on the crease. The colour is either red, white or
culm is opposite. amber. The grain consists of grain coat,
nucellar epidermis, endosperm and embryo.
Floral character The seed is endospermic, oblong in shape
and constitutes about 82–86% of the grain
The panicle is called an ear or spike. This is a and consists of starch and gluten.
compound distichous spike, the axis bearing
two opposite rows of lateral spikelets and
a single terminal spikelet, but in case of
T. monococcum there is no terminal spikelet. 5.3.5 Anatomical description
The spikelets are sessile, have two glumes at
their base and are arranged alternately. Each Root anatomy
spikelet is attached to a rachis node and is
known as a floret. Flowers are called florets. The young seminal root consists of a root
Flowers are stipulate, ligulate, bisexual, zygo- cap. In the transverse section, the root shows
morphic, trimerous and epigynous. The flo- three distinct regions, epidermis, cortex and
ret is composed of three stamens and a pistil, vascular tissue (Fig. 5.21).
which consists of ovary. The flower is Epidermal cells have long unicellular
enclosed by a lemma (outer bract) and a palea root hairs in the root hair zone. These cells
(inner bract). The lemma tip is extended to are compactly arranged and single layered.
form awns. Sepals and petals are together The cortex with multilayered paren-
called tepals, and each floret consists of two chymatous cells surrounds an inner cylin-
tepals. There are three stamens, which are der of tissue called the stele. The inner layer
dithecous, versatile and yellow in colour. The of cortex that surrounds the stele shows
gynoecium is unilocular, unicarpellary, with a Casparian stripes of endodermis.
single ovule on basal placentation, silky-like The stele has alternate bands of xylem
style and bifid feathery stigma. Anthesis and phloem arranged around a central
occurs over about 3–10 days, depending on metaxylem vessel. In the root hair zone,
the environment, after the ear emerges from lateral branch roots arise from the stele,
the flag leaf sheath, when a number of closely adjacent to the phloem. They grow through
correlated events occur in a very short time. the cortex and ramify into the soil, their
As the filament grows, the anther dehisces, structure resembling that of the main root.
each chamber developing a longitudinal In the older regions of the root, the cortex
split, starting at the tip of the anther, through dies leaving only the stele so that the root
which pollen is released. The stigma lobes, appears to become thinner as it gets older.
which are pressed together before anthesis, In the deeper regions of the soil, the anat-
move apart, and the receptive branches are omy of the nodal roots is similar to that of
spread widely giving a large area for pollen a seminal root.
66 Chapter 5
Cortex
Endodermis
Xylem
Phloem
Ground tissue
Sclerenchyma
Epidermis Vascular bundle
Fig. 5.22. Transverse section of stem (small portion 400×) showing epidermis, sclerenchyma,
ground tissue and vascular bundle.
wheat (Zhu et al., 2004). Thickness of scle- single row of stomata between each rank of
renchyma, thickness of epidermis and bulliform cells and the vascular tissue.
density of epidermis are important factors Each stoma is made up of two characteris-
for digestibility of wheat straw as rumi- tically shaped guard cells and has two
nant feed (Travis et al., 1996). The culti- associated accessory cells. There are more
vars less susceptible to lodging contain on the upper surface and these are more
lower amount of neutral digestible fibre, densely distributed towards the tip. On
and thus have less nutritive value as ani- the other flank of the row of stomata, over
mal feed. the veins there are long cylindrical cells
characterized by thickened wavy walls.
Leaf anatomy The long cells are interspersed in a regular
manner by short cells of two types, cork
There are three main features of the anat- cells and silica cells. Short, unicellular
omy of the leaf, which includes epidermis, hairs occur mainly over the veins and on
mesophyll and vascular bundles (Fig. 5.23). either side of the row of stomata. The lower
The upper and lower epidermis of the epidermis has fewer cell types, mainly the
mature leaf encloses the mesophyll, which long cylindrical cells with wavy walls
is traversed at intervals by the vascular tis- interspersed by short cells. Stomata occur
sue. The upper epidermis is a complex tis- in the same position relative to the veins
sue with several cell types. The bulliform as in the adaxial epidermis and, although
cells are the largest cells lying between the hairs occur, they are less frequent than on
veins at the bottom of the furrows. Next to the upper epidermis. The epidermis on
the bulliform cells are long cylindrical both surfaces of the leaf has a cuticle with
cells with a smaller diameter than the bul- strongly developed epicuticular wax. The
liform cells, alternating in a regular man- form of the wax depends upon the sur-
ner with stomata. There is generally a face of the leaf and the position of the
68 Chapter 5
Bulliform cells
Upper epidermis
Mesophyll
Trichome
leaf on the stem. This occurs as lobed of cells radiate in a regular manner from the
plates, simple plates, flat ribbons and vascular bundles. Prominent sub-stomatal
tubes, the amount and form of the wax cavities occur, particularly beneath the sto-
depending on the position and surface of mata of the abaxial surface of the leaf. The
the leaf. The vascular tissue lies beneath vascular tissue and mesophyll are organ-
the ridges of the lamina and the associated ized in alternate strips of tissue running
thickening capping the vascular bundle of parallel with each other along the long axis
the midrib, and the major veins extend of the leaf.
from the ad- to the abaxial epidermis. The vascular bundle has the structure
The mesophyll cells are of a complex typical of a C3 plant. The phloem is abaxial
lobed shape, resembling armed palisade to the xylem and in the larger bundles con-
cells. When viewed in transverse section, sists of regularly arranged sieve tubes and
the sub-epidermal cells of the mesophyll companion cells. The xylem has two large,
are elongated similar to palisade cells. The prominent xylem vessels between which
cells in the middle layers of the leaf are not are smaller metaxylem vessels and fibres.
so elongated. Viewed in longitudinal sec- Adaxial to the metaxylem, there is an area
tion, the lobed nature of these cells is appar- of disrupted protoxylem. The conducting
ent. In the leaves at the base of the plant, the elements are surrounded by an inner (mestome)
degree of lobing is low and the dimensions sheath and an outer (parenchyma) sheath,
of the lobes are large. With ascending leaf though these are not as clearly defined as in
position up the stem, the degree of lobing some other C3 plants. The cells of the mestome
increases and the diameter of the lobes sheath are small and thick-walled and are
decreases. The effect of these changes is to without chloroplasts. Those of the outer bun-
increase the cell surface area per unit area of dle sheath are large and thin-walled and
leaf with ascending leaf position up the contain chloroplasts. In longitudinal sec-
stem. There is variation in the compactness tion, the cells of the bundle sheaths are
and arrangement of the mesophyll cells. elongated with blunt ends. The walls of the
Some cultivars have a relatively loose mestome sheath are lignified, and some-
arrangement of cells, while in others the times the wall adjacent to the conducting
cell arrangement is more compact and files elements is thicker than the other walls of
Cereals 69
the cell. The complex fine structure of the act as a mechanical barrier to fungal pene-
mestome sheath is important in regulating tration or insect-pest attack (Carver and
the transport of water and solutes. The small Thomas, 1990). Leaf thickness is thus con-
veins that interconnect the main longitudi- sidered to be a selection criterion for devel-
nal veins consist only of a single sieve tube opment of drought-tolerant wheat cultivars.
and xylem vessel and two files of parenchyma Wheat genotypes adapted to the very hot
cells. They pass through the mestome and climates of Baluchistan region of Pakistan
parenchyma sheaths and connect directly show increase in thickness of both epider-
with the metaxylem and metaphloem of the mis and hypodermis under xeric adapta-
main bundles. They do not have bundle tion (Hameed et al., 2002). These drought-
sheaths, and the vessel walls have a complex tolerant genotypes also exhibited lower
fine structure depending on the adjacent stomatal density on the adaxial surface than
mesophyll walls. The major bundles run par- the abaxial surface, a common mechanism
allel with each other the whole length of the observed for reducing transpiration in plants
leaf. The small transverse veins, which con- under drought stress. A lower ratio of xylem
stitute about 7% of the total length, occur to mesophyll tissues, increase in lignin depo-
every 2.5–3 mm, towards the tip of the leaf, sition and thickening of sclerenchyma have
the smaller longitudinal bundles terminate been suggested to be indicative of drought
in a Y-shape, the forks of the Y comprising tolerance in wheat (Ridley and Todd, 1966).
small transverse veins that link to the longi- The photosynthetic activity of wheat
tudinal veins at either side. At the pointed leaves is higher than other C3 crops except
tip of the leaf, the veins converge and con- rice, which is due to high number of meso-
nect with each other. The distance between phyll cells per unit area, arrangement of
the longitudinal veins varies from about 0.3 chloroplasts in cells to maximize solar radi-
mm in the first leaf to about 0.15 mm in a ation and the high concentration of rubisco
culm leaf. protein in the leaves.
as those found in the leaf, but are kidney- (Sorghum bicolor) is the fifth most impor-
shaped, similar to dicotyledon guard cells. tant cereal crop and is the dietary staple of
more than 500 million people in more than
Embryo development 30 countries. It is grown on 42 Mha in 98
countries of Africa, Asia and the Americas.
Division of the fertilized egg nucleus com- Nigeria, India, the USA, Mexico, Sudan,
mences later than that of the endosperm. China and Argentina are the major produc-
The early divisions produce a five-celled ers. Other sorghum-producing countries are
embryo with a basal cell, although variation Mauritania, Gambia, Mali, Burkina Faso,
in the pattern of development has been Ghana, Niger, Somalia and Yemen, Chad,
observed. Continuing cell division produces Sudan, Tanzania and Mozambique. Sorghum
at first a club-shaped structure, which ulti- is a self pollinating diploid (2n =2x = 20)
mately differentiates to form a mature with a genome (1C = 735 Mbp) about 25%
embryo in the ripe seed. the size of maize or sugarcane.
The growth of the embryo is supported
by nutrients derived from the antipodal cells
and from the hydrolysis of parenchyma cells 5.4.2 Origin and domestication
of the nucellus and neighbouring endosperm
cells. Later in development, transfer cells Nothing is known about when S. bicolor
appear in the nucellar endosperm epider- was first brought into cultivation, but
mis, near the base of the embryo and the pro- Murdock (1959) stated that, along with sev-
vascular tissue. eral West African crops, it was domesticated
some 7000 years ago.
Endosperm development The sorghum in India has been known
since ancient times, but the wild species,
Following the fusion of the sperm nucleus more or less adapted, is found throughout
and the polar nuclei, cell division is, for a the world. Engravings have been found in
time, synchronous, the number of endosperm ruins in Asia that represent sorghum in 700
cells doubling every 4–5 h. At first, the BC. It was probably introduced into the inte-
endosperm is coenocytic, but after about 3 rior of Asia at the beginning of the Christian
days cell walls are formed. Cell wall growth era, when sorghum was also known in the
commences at the edge of the embryo sac Mediterranean region and tropical Africa.
and furrows inwards to the central vacu- The domestication of sorghum was ini-
ole. The cells at the periphery of the endo- tiated by allelic changes in only two loci,
sperm divide, and eventually the entire resulting in different sections, followed by
embryo sac is cellular. After cell formation the innovation of the cropping techniques.
is complete, the sub-cellular structures, Wild sorghum disperses its seeds by the
which will synthesize the protein bodies rupture of inflorescence nodes with subse-
and the other cell components, are formed. quent spacing of the seeds. The essential
Amyloplast division ceases before cell divi- step adapted for the domestication was the
sion, and starch grains differ in growth rate harvest of the inflorescence, and the use of
in different cell layers of the endosperm. grain for sowing. The plants are cultivated
in warmer climates worldwide. Species are
native to tropical and subtropical regions of
5.4 Sorghum all continents in addition to the south-west
Pacific and Australasia.
5.4.1 Introduction
tropical areas sorghum grain is important as and seedling drought resistance. Source
food and as livestock feed. The stem and material for the glossy trait is maintained by
foliage are used as a green crop, hay silage the Genetic Resources Unit at ICRISAT.
and pasture. The stems are also used as fuel
and building materials. In temperate areas it Vegetative characters
is a major source of cattle feed, except in
China, where it is primarily used for food. Sorghum is a genus of numerous species of
Sorghum is used in the preparation of dif- grasses. It reaches to a height ranging from 2
ferent types of food, and unleavened bread to 4 m, having various stems with tillers,
is the most common food made from sor- each bearing one inflorescence in a panicle.
ghum flour. The dough is sometimes fer- One fertile sessile spike is accompanied by
mented before the bread is prepared and the two sterile pendant spikes, the characteris-
grain is boiled to make a porridge or gruel. It tics of the genus. It is well adapted to semi-
is also used in the preparation of biscuits. arid conditions; most varieties are drought
Beer is prepared from sorghum grain in tolerant and heat tolerant, and are especially
many parts of Africa. As well as these pro- important in arid regions where the grain is
ducts, popped and sweet sorghum, which the staple or one of the staples for poor and
are parched, are also eaten in some coun- rural people. It has a fibrous root system,
tries (House, 1980). Sweet sorghum is and forms stilt roots. The stem is aerial,
emerging as a feedstock for ethanol produc- erect, greenish and hard, succulent with
tion. Sorghum grain has high levels of iron nodes and internodes. Leaves are simple,
(>70 ppm) and zinc (>50 ppm) and is hence exstipulate, alternate, sessile and long,
being targeted as a means to reduce micro- arranged in two rows on stem, with an
nutrient malnutrition globally. expanded leaf base with leaf sheath, wavy
margin and parallel venation. Deposition of
silica crystals in the leaves is a characteris-
5.4.4 Morphological description tic feature of grasses.
Epidermis Endodermis
Hypodermis Pericycle
Cortex Phloem
Protoxylem
Metaxylem
Fig. 5.25. Transverse section of root (100×) showing various parts. Radial vascular bundles are visible in the
stele with prominent pith.
that high root endodermal silicification might fibro-vascular bundle with thick-walled
be related to drought tolerance. sclerenchyma, which is of smaller sizes, fol-
lowed by rings of bigger vascular bundles in
Significance of variations in root anatomy the cortex (Fig. 5.27).
In transverse section, the epidermis con-
Cultivar differences are expressed in a sists of cubical to boat-shaped epidermal cells
number of root characteristics in sorghum. with thickened outer wall with cutin. The
These are: ground tissue consists of thin-walled paren-
1. Presence or absence of sclerenchymatous chyma. The vascular bundles are pointed col-
exodermis in the cortex. laterally and scattered in the ground tissue.
2. Thickness of endodermal cell walls There is a continuous layer of sclerenchyma
(Fig. 5.26). just below the epidermis. There are alternate
3. Density, size and shape of crystals in the bands of large and small vascular bundles
endodermal cell cavity. below the hypodermal sclerenchyma band.
4. Intensity of lignification in the pericycle The vascular bundles towards the centre are
cell layers and in cells surrounding the vas- larger. The mechanical tissues in the outer
cular bundles. vascular bundles are extensive, particularly
in the peripheral region.
The presence of exodermis, higher thick- In the stem, different types of vascular
ness and higher lignification provides better bundles are found:
structural integrity to the root system. Thus
genotypes exhibiting these characteristics 1. Large bundle is sheathed by sclerenchy-
are expected to have better crop stand and matous mass on lateral and xylary polar
resistance to lodging. ends and a voluminous phloic cap of scler-
enchyma just beneath the epidermal layer.
Stem anatomy 2. Similar to above, but without epidermal
sclerenchyma sheath.
A transverse section of a stem shows cuticu- 3. Comparatively small with smaller xylary
larized epidermis followed by a sub-epidermal fibrous strands and the phloic cap equal in
74 Chapter 5
Epidermis
Hypodermis
Ground tissue
Phloem
Sclerenchyma
Vascular
bundle
Protoxylem
Metaxylem
Mesophyll
BS VB
Lower epidermis
Protoxylem Sclerenchyma
Metaxylem Phloem
Fig. 5.28. Transverse section of leaf (100×) showing various parts, including upper epidermis,
sclerenchyma, mesophyll and fibro-vascular bundle (VB), bundle sheath (BS), protoxylem and phloem.
cells) are present in both of the epidermal sorghum is a C4 plants it shows Kranz
layers, but the stomata are usually more anatomy (Fig. 5.30). Usually each bundle
frequent in the lower epidermis. All the epi- remains surrounded by a bundle sheath
dermal cells except the guard cells are consisting of thin-walled parenchyma cells.
colourless. Epidermis is rough in nature due The cells of the bundle sheath generally
to the presence of silica crystals (Fig. 5.29a). contain large chloroplasts devoid of photo-
Two types of hairs can be observed in sor- system I (without grana) and starch grains
ghum, non-glandular and glandular trichomes. in them.
Leaf trichome density (non-glandular pointed
trichomes) and glossiness intensity were Leaf wax – cuticle
considered reliable and very simple indica-
tors in the selection of sorghum genotypes Analysis of S. bicolor bloomless (bm)
tolerant to shootfly. The presence of bilobed mutants with altered epicuticular wax (EW)
glandular trichomes (Fig. 5.29d) induces structure uncovered a mutation affecting
susceptibility to this insect in sorghum. As both EW and cuticle deposition. The cuti-
the leaf is isobilateral, the mesophyll is not cle of the mutant line was about 60% thin-
differentiated into palisade and spongy tis- ner and approximately one-fifth the weight
sues, but is composed of compactly arranged of the wild-type parent cuticles. Reduced
thin walled, isodiametric chlorophyllous cuticle deposition was associated with
cells having well developed intercellular increased epidermal conductance to water
spaces among them. The leaf receives the vapour. The reduction in EW and cuticle
vascular supply from the stem. The vascular deposition increased susceptibility to the
tissue occurs in the form of discrete bundles fungal pathogen Exserohilum turcicum.
called veins. The vascular bundles are col- Evidence suggests that this recessive muta-
lateral and closed. Most of the bundles are tion occurs at a single locus with pleio-
small in size but fairly large bundles also tropic effects. Chemically induced mutants
occur at regular intervals; the xylem is had essentially identical EW structure,
found towards the upper side and phloem water loss and cuticle deposition (Jenks
towards the lower side of the bundles. As et al., 1994).
76 Chapter 5
(a) (b)
(c) (d)
Fig. 5.29. Leaf surface structures (SEM): (a) bilobed and trilobed silica crystals; (b) glossy line showing
a smooth wax layer and a trichome (SEM); (c) Genotype IS 4776 showing long wax strands (SEM);
(d) Genotype IS 844 showing glandular bicellular trichomes (Maiti, 1996).
Fig. 5.30. Scanning electron micrograph of the midrib showing thick-walled epidermis, trichome,
sclerenchyma sheath and Kranz anatomy (Maiti, 1996).
Trichome
pointed trichomes) and glossiness intensity
Sorghum leaf surfaces show the presence of were considered reliable and very simple
non-glandular pointed trichomes oriented indicators in the selection of sorghum geno-
at acute angles. types resistant to shootfly. The presence of
Two types of hairs can be observed in bilobed glandular trichomes induces suscep-
sorghum: are non-glandular and glandular tri- tibility to this insect in sorghum. The presence
chomes. Leaf trichome density (non-glandular of non-glandular trichomes acts as barriers
Cereals 77
to the laying of eggs and migration of larva 5. Reduction in stomatal density on lower
downwards through the collar. In addition, and upper epidermis minimizes the loss of
the glossy leaf surface offers difficulty egg- water from leaf.
laying, which may cause the eggs to slide 6. Presence of long size bulliform cells pre-
off. There is a large variation in the inten- vents evapo-transpiration.
sity of trichomes and it has been found
that shootfly tolerance increases with The seed
intensity of trichomes (Maiti et al., 1984;
Maiti, 1996). Two types of endosperm are found in the
Genetic study has been undertaken on seed, corneous and floury (Fig. 5.31). The
the inheritance of glossy leaves in sorghum. corneous endosperm is also called vitre-
In a study it was reported that the glossy ous endosperm and is present at the
leaf is a genetic trait that showed pleiotropic periphery, while the floury endosperm is
effects on disease and pest resistances. True- found in the inner core of the main bulk of
glossy (tg), glossy (g) and non-glossy (G) were the endosperm. Cells in the corneous
multiple alleles located in a glossy locus endosperm are oval or slightly elongated
(Tarumoto, 2006). with compactly arranged starch and pro-
tein granules, while the cells in the floury
part are much more elongated and broad
Significance of variations with loose arrangements of starch and
in leaf anatomy protein granules in the lateral and dorsi-
ventral sections of the endosperm. In lat-
The different sorghum genotypes show dif- eral section, the scutellar tissue, along
ferences in the following leaf anatomical with the embryo can be seen clearly near
features: the broad hilar region. The pointed end
1. Amount of cuticularization. of the embryo is inclined towards the lat-
2. Intensity of silica crystals in the upper eral wall of the seed. The corneous endos-
epidermal cells offering rigidity to the leaf perm encloses the inner mass of floury
as well as resistance to leaf insects. endosperm.
3. Size and shape of bulliform cells helps in The seeds can be classified according
leaf rolling, thereby avoiding loss of transpi- to the intensity of corneous over floury
ration under drought situations. endosperm (Fig. 5.32a–d):
4. Type and arrangement of vascular bundles
both in the lamina and midrib. 1. Almost the entire endosperm is corneous –
5. Characteristic size and shape of bundle only a very small floury portion may be
sheath chloroplasts. present in the centre; completely corneous
(IBPGR).
From observing these variations, the follow-
2. Over 50% of the endosperm is corneous –
ing inferences can be drawn on the signifi-
the endosperm is broad; almost corneous
cance of leaf anatomical features:
(IBPGR).
1. Presence of thick cuticle on the epider- 3. The endosperm is almost floury – a
mis of the leaf reduces transpirational loss thin layer of corneous endosperm may
of water. be present that should not exceed 10% of
2. High density of trichomes covers the sur- the entire mass that surrounds the broad
face and reduces the direct sunlight effect floury endosperm mass; almost starchy
and minimizes leaf temperature, thereby (IBPGR).
reducing transpirational loss of water. 4. More than 75% of the endosperm is
3. High density of trichomes offers insect floury – the endosperm layer is medium
resistance, especially sucking pest tolerance. broad; partly corneous (IBPGR).
4. More numerous and larger size vascular 5. The whole endosperm is floury – a thin
bundles help in efficient translocation of corneous field of the endosperm may be
water. present, full of starch (IBPGR).
78 Chapter 5
(a) (c)
(d)
(b)
Fig. 5.31. Scanning electron microscopic analysis of the corneous and floury endosperm of the sorghum
seed. (a) Longitudinal section of the corneous and floury endosperm. (b) Typical structure of the corneous
endosperm with a continuous face between the flour and the protein, resulting in a space without air
between the starch granules; the corneous endosperm appears extremely dense and vitreous. (c) Transition
section with corneous and floury endosperm cells. (d) The floury endosperm contains air spaces between
the starch granules with a clustered and loose appearance. SG, starch granules; PM, protein matrix;
PB, protein bodies; BG, broken granules (Courtesy L. Rooney, Texas A&M University).
Floury Endosperm
Endosperm
Totally Almost Partially
corneous corneous corneous
Coleoptile
Almost Totally
floury-corneous floury
Plumule Scutellum
Radicle Coleorrhiza
Fig. 5.32. Parts of the seed and texture of the endosperm (Maiti, 1996).
Cereals 79
MC HP CT
EP
PERICAR
P
CC
TC
Fig. 5.33. Transverse section of a mature caryopsis (400×) showing cuticle (CT), epicarp (EP), hypocarp
(HP), mesocarp (MC), cross cells (CC) and tube cells (TC) (Courtesy L. Rooney, Texas A&M University).
80 Chapter 5
Just beneath the corneous portion of the and drought stress. Therefore, there is a great
endosperm are located several layers of necessity to screen sorghum cultivars and
large elongated and vacuolated cells form- germplasm for these traits and to utilize these
ing the floury endosperm portion. The traits in sorghum crop improvement. There
starch granules are spherical and are not is also a necessity to select sorghum geno-
held together by a protein matrix. The pro- types with minimum sclerenchyma in the
tein bodies and matrix are present in the stem for increasing the acceptability as fod-
floury endosperm. der for cattle.
Embryo
5.5 Pearl Millet
The embryo contributes 10% of the total
dry weight of the seed. The scutellum, con- 5.5.1 Introduction
sisting of vacuolated parenchyma cells,
has a well-developed vascular system and
Pearl millet (Pennisetum glaucum (L.) R.
helps in the translocation of nutrient from
Br.) is an important grain crop as well as
the endosperm into the developing roots
food for semi-arid and arid regions of world.
and leaf tissues of the embryonic axis
In India it is known as bajra, and is prima-
during germination. The hilum helps in the
rily consumed in the states of Haryana,
translocation of nutrients from the vegeta-
Rajasthan, Gujarat and Madhya Pradesh. It
tive plant parts into the ovule during cary-
is used as hay and for pasture. It is one of
opsis development. Translocation of the
the most important food and forage cereals
nutrients into developing endosperm takes
in the semi-arid tropics of many African
place through specialized transfer cells in
countries, in India, Pakistan, Bangladesh,
the scutellum.
Burma, Sri Lanka, Argentina and the USA.
India is the largest producer of pearl millet.
Pearl millet is the staple food and fodder
Research trends on sorghum anatomy crop of farmers in the semi-arid areas of
Remarkable research advances have been north-west India.
undertaken with respect to anatomy of dif-
ferent parts of the plants with respect to
insect and drought tolerance. With respect to 5.5.2 Origin, domestication
general anatomy there exists a large variation and adaptation
in leaf, stem and root anatomical structures
among sorghum genotypes. There also exists Pearl millet has been present in the African
a large variation in leaf surface structures semi-arid tropics since 1100 BC. The origin
with respect to glossiness, non-glossy trait, of pearl millet has been traced to tropical
types of trichomes and intensity and silica Africa, with its centre of diversity for the
content, bulliform cells and Krantz tissue. crop in the Sahel zone of West Africa.
Similarly, variation exists in stem anatomy Cultivation subsequently spread to east and
with special reference to the intensity of southern Africa, and southern Asia. Records
mechanical tissue-stereome, which offers exist for cultivation of pearl millet in the
resistance to stem borers as well as drought. USA in the 1850s, and the crop was intro-
The presence and intensity of non-glandular duced into Brazil in the 1960s. The greatest
pointed trichomes and glossiness are inver- morphological diversity of pearl millet can
sely correlated with shootfly resistance. The be found today in the Sahel zone, located in
intensity of trichomes and glossiness on the western Africa, south of the Saharan desert
other hand may create a microclimate thereby and north of the forest zones. It has been
reducing radiation load and transpiration, concluded that pearl millet is the product of
emphasizing the great importance of these multiple domestications due to the high
traits in sorghum crop adaptation to both insect diversity present.
Cereals 81
Pearl millet is well adapted to produc- its base, usually fuzzy and sometimes hairy.
tion systems characterized by drought, low The fuzz is rough and rigid. Leaves are sim-
soil fertility and high temperature. It per- ple, exstipulate, alternate, sessile, long,
forms well in soils with high salinity or low arranged in two rows on stem, with an
pH. Because of its tolerance to difficult expanded leaf base with leaf sheath, wavy
growing conditions, it can be grown in areas margin, and parallel venation. The leaves
where other cereal crops, such as maize or are straight, 20–100 cm long and 0.5–5 cm
wheat, would not survive. wide, usually glabrous with spaced fuzz.
The base of the leaf is slightly auricular. The
ligule is narrow, membranous, with a strip
5.5.3 Utilization of fuzz that almost surrounds the stem. The
leaf sheath surrounds the stem completely
and is thicker than the blade; the external
Pearl millet grains are consumed as food in
surface of the sheath can be glabrous or
most of the African countries and some
covered with rough hairs in some fuzzy
states of India (Haryana, Rajasthan, Gujarat
varieties.
and Madhya Pradesh). It is also used as pas-
ture. Pearl millet is an important food across
the Sahel; it is often ground into flour, rolled
into large balls, parboiled, and then con- Floral characters
sumed as porridge with milk. Sometimes it
Bisexual flowers are formed on sessile
is prepared as a beverage.
spikelets and staminate or male flowers are
formed on pedicillate spikelets. This is a
cross-pollinated crop (80%). The terminal
5.5.4 Morphological description inflorescence is a cylindrical, bulbous pani-
cle. The peduncle is thin, cylindrical and
covered with soft fuzz under the spikelets.
Pearl millet is a genus of numerous species
The rachis is straight, cylindrical, solid and
of grasses. Pennisetum americanum is an
covered with soft and short fuzz. The small
annual erect plant with profuse tillering
rachis produces a series of florets placed in
habit that grows up to 2–4 m.
a spiral form on the rachia. The rachillas
are small and covered with short hairs,
Vegetative characters shorter in the apex of the panicle and den-
ser approximately in the middle portion.
Pearl millet is a herb, height 2–4 m, and is a Flowers are sessile, unisexual, zygomorphic
mesophyte, well adapted to semi-arid con- and epigynous; in every flower there is
ditions. The root system is somewhat large lemma, palea and two lodicules. Each spike-
and fibrous and therefore has a great capa- let consists of two sterile glumes and two
city for efficient use of the soil. It can also florets, the lower floret is generally male
utilize soil nutrients better than other crops and the one above hermaphrodite; glume
due to the deep penetration of its roots into 1 is small or 0, orbicular; glume 2 is small
the interior layers of the soil. The plant is or 0, variable. Lemma 1 is oblong to ovoid,
aerial, erect, greenish and hard, with succu- palliated; the palea is membranous and
lent nodes and internodes. The nodes are staminated; lodicules are absent. There are
slightly swollen, softly haired (pubescent), three stamens: typical pencilated, long
while the internodes are cylindrical and anthers, dithecous, basally fixed. The carpel
glabrous. The length of the internodes dec- is unilocular, unicarpellary, with a single
reases gradually from the base of the stem ovule on basal placentation, silky-like style
upwards. One light groove can be found and bifid feathery stigma. The gynoecium is
above each node containing the axillar bud. protogynous. The grain is grey, pearl-white,
The bud is slightly swollen and produces and rarely yellow and endospermic. It is a
the ring of adventitious root primordium in caryopsis (Fig. 5.34).
82 Chapter 5
Root anatomy
Stem anatomy
The anatomical structure of the root of
pearl millet is characteristic of monocots. It The anatomy of the young pearl millet stem
is composed of an outer epidermis, ground is characteristic of the grasses. The epider-
tissue, cortex below the epidermis, an mis consists of a single cell layer with
endodermis surrounding the vascular thickened walls on the outer side. Thick
bundles that consist of xylem and phloem cuticle covers the epidermal cells and
in alternating bands, and pith (Fig. 5.35). reduces the transpiration. The ground tis-
The epidermis consists of elliptical cells sue consists of thin-walled parenchyma
subtended by two layers of hypodermis. In (Fig. 5.37). Vascular bundles of the collat-
the early stages of growth, the cortex of the eral type are scattered in the ground tissue.
root epidermis consists of ovoid parenchy- A transverse section through a mature stem
matous cells with considerable intercellu- shows an epidermis consisting of cubical
lar space. As the root ages, the cortical to boat-shaped epidermal cells, containing
cells elongate and take a plate-like shape. elongated crystals. Just below the stem epi-
A single layer of ovoid or cubical paren- dermis, there are alternating bands of large
chyma cells surrounds the endodermis. The and small thick-walled sclerenchyma
endodermal cells are barrel- to boat-shaped, patches. Each sclerenchyma band alter-
and are thickened with suberin on the nates with a broad band of sclerenchyma-
inner tangential wall. Crystals are present tous tissue that corresponds to the furrows
in the endodermal cells, their shape and in the stem outline. These hypodermal
size varying with the cultivar and age of bands of the sclerenchyma are in turn con-
root (Fig. 5.36). The pericycle below the nected to the subtending broad scleren-
endodermis is composed of one to several chyma cylinder, which contains the ring of
layers of thick-walled sclerenchyma cells, vascular bundles. In a young stage plant,
of which the outer layer is highly ligni- hypodermis consists of collenchyma tis-
fied; the parenchyma that surrounds the sue offering great photosynthetic capacity
metaxylem may be thick-walled or ligni- (Fig. 5.38). There are four to five rows of
fied. The size of the metaxylem varies vascular bundles, the smaller ones found
among cultivars. toward the outside. Alternating small and
Cereals 83
END COR
EP
HP PH
Fig. 5.35. Transverse root section showing the pattern of cortical cells and arrangement of vascular bundles
(75×): EP, epidermis; HP, hypodermis; COR, cortex; END, endodermis; PH, phloem; and X, xylem
(Maiti and Wesche-Ebeling, 1997).
Epidermis
Hypodermis
Ground tissue
Vascular bundles
Fig. 5.37. Transverse section of stem (ground plan 40×) illustrating epidermis, hypodermis and scattered
vascular bundles in ground tissue. Note: the large amount of sclerenchymatic hypodermis offers great
strength to the stem.
system of the plant. Millet genotypes show entire in its outline. Silica crystals are
large variations in anatomical structure of prominent in some genotypes, but not in
stem with respect to the distribution and others. The upper epidermal cells are wavy
intensity of sclerenchyma, on the basis of in outline and interspersed with zones of
which we can classify the genotypes into bulliform (motor) cells, that show high var-
three groups: with a weak, intermediate or iation in size among genotypes. The scler-
strong stereome system. While breeding for enchyma surrounding the vascular bundles
new genotypes, anatomical studies thus may be loose or compact. In transverse sec-
may help in better selection of genotypes tion bulliform cells are present in the upper
according to the target environment and epidermis of the midrib and vascular bun-
breeding objectives. Some points to be dles of different sizes can be found. The
noted are: midrib has three vascular bundles in some
genotypes, one in the centre and the other
1. It is expected that the genotypes having a
two at the junction of the leaf lamina and
strong stereome will be resistant to lodging.
midrib. The vascular bundles in the lamina
A large amount of sclerenchyma gives
and midrib can be of three types: (i) a large
mechanical support and avoids the loss of
central vascular bundle present in the main
water from internal parenchyma tissue by
vein; (ii) medium sized vascular bundle;
evapo-transpiration.
and (iii) very small vascular bundle. The
2. It is expected that medium to large size
first two types are fibro-vascular and con-
and a high number of xylem vessels are
tain patches of sclerenchyma in both sides
required for efficient translocation of water
connecting the upper or lower epidermis.
under drought conditions.
Each vascular bundle remains surrounded
Leaf anatomy by a bundle sheath consisting of thin-walled
parenchyma cells (Fig. 5.39). The cells of the
The anatomy of pearl millet leaves is char- bundle sheath generally contain large chlo-
acteristic of a mesophytic grass. The anat- roplasts, grana are absent and large starch
omy of the leaves does not vary significantly grains occur in the chloroplasts. Mesophyll
from genotype to genotype. The cuticularized cells contain normal chloroplast. This special
epidermis is composed of oval or rectangular structure was first identified by Haberland;
epidermal cells. The lower epidermis is it appears like a garland (Kranz = wreath,
Bulliform cells
Upper epidermis
Mesophyll
Bundle
sheath
Lower epidermis
Vascular bundle
Fig. 5.39. Transverse section of leaf (100×) showing upper and lower epidermis, bulliform cells, mesophyll
and vascular bundles. Each vascular bundle is covered by bundle sheath cells. The large size vascular
bundle is at the central (mid-vein) region.
86 Chapter 5
garland), so it is known as Kranz anatomy. 2. High density of trichomes cover the surface
Kranz anatomy is the special characteristic and reduces the direct sunlight effect and
of C4 plants. Kranz anatomy is one of the minimizes leaf temperature, thereby reducing
adaptations for drought resistance in plants transpirational loss of water.
of semi-arid zones. 3. High density of trichomes offers insect
resistance, especially sucking pest tolerance.
4. A greater number and large size vascular
Significance of variations in leaf anatomy bundles help in efficient translocation of
water.
The different millet genotypes show differ-
5. Reduction in stomatal density on lower
ences in the following leaf anatomical
and upper epidermis minimizes the loss of
features:
water from the leaf.
1. Amount of cuticularization. 6. Presence of long size bulliform cells pre-
2. Intensity of silica crystals in the upper vents evapo-transpiration by rolling the leaves.
epidermal cells offering rigidness to the
leaf, thereby offering resistance to the leaf
Reproductive growth – panicle
insects.
development
3. Size and shape of bulliform cells helps in
leaf rolling, thereby avoiding loss of transpi- At the end of the vegetative stage approxi-
ration under drought situations. mately 16–28 days after emergence, the
4. Type and arrangement of vascular bun- shoot apex is transformed into a reproduc-
dles, both in the lamina and midrib. tive apex with the formation of a panicle
5. Characteristic size and shape of bundle meristem (stage of panicle initiation)
sheath chloroplasts. (Fig. 5.40). The panicle meristem assumes
From observing these variations, the follow- a bulbous shape with a constriction in
ing inferences can be drawn on the signifi- its base with definite cellular organization.
cance of leaf anatomical features: There are prominent tunica and corpus lay-
ers in the panicle axis, with two to three
1. Presence of thick cuticle on epidermis of layers of serially arranged tunica cells over-
leaf reduces transpirational loss of water. lying the corpus. The cellular organization
Fig. 5.40. Transverse section of panicle meristem, showing elongation of spikelet primordia with distinct
tunica and corpus (75×): SPP, spikelet primordia; T, tunica; C, corpus; VTR, vascular trace
(Maiti and Wesche-Ebeling, 1997).
Cereals 87
in the corpus layer is irregular. The pro- divides to give two spikelets, which sub-
toxylem in the tunica cells and the nuclei sequently develop flower parts. Each
with prominent chromatic material are spikelet is enclosed by a pair of glumes,
distinct. the upper and lower (Fig. 5.42). Subsequently
The spikelet primordial develops ini- each individual spikelet primordium
tially at the base and gradually progresses divides and extends laterally to give rise to
towards the tip of the panicle meristem. two floret primordia which develop into
Initiation of the spikelet primordial is marked flower parts.
by the condensation of protoplasm and by Two types of florets exist in each spike-
an increase in the volume of the flank meri- let in pearl millet, the upper is perfect and
stem. The spikelet primordium assumes a the lower is male. Each floret is enclosed
bulbous appearance (Fig. 5.41) with con- by two glumes. In a perfect flower (bisex-
striction at the sides, indicating the bound- ual), the flower primordium develops in
ary between the adjacent spikelet primordia the axil of the lateral primordial, which
(Maiti and Bidinger, 1981). form two glumes, upper and lower at the
About 3 days after initiation, the pani- base (Fig. 5.43). The floret primordium
cle is much more elongated and the gives rise to the primordial forming palea
formation of the spikelet primordial has and lemma and the central primordium,
advanced toward the tip. Each primordium which functions as the carpel primordium.
(SPP
VTR
Fig. 5.41. Longitudinal section of a young panicle, showing initiation of spikelet primordia at the base.
Note: spikelet primordia are developing at the base of the panicle; towards the tip, spikelet primordia are
not yet developed. SPP, spikelet primordia; VTR, vascular trace (Maiti and Wesche-Ebeling, 1997).
88 Chapter 5
SPI
SPII
GL
LM SPP
LGL LM
VTR ST
PL
Three primordial stamens develop from for the presence of the carpel. It has a
the central primordium afterwards. The lemma and lacks a palea. The central pri-
developmental pattern of a male flower is mordium gives rise to the three stamens
similar to that of a bisexual flower, except in the male flower.
Cereals 89
Eight days after panicle initiation a tissue A single ovule is present in the cavity of the
mass in which the microsporangia are ovary and is laterally attached to its wall by
embedded appears in the anther. Micro- a short funicle. The ovary wall is composed
sporangia are present as four separate of a single layer of cubical epidermal cells,
masses in the four lobes of the anther. These subtended by several layers of thin-walled
cells divide along periclinal walls, leading cells (Fig. 5.45).
towards an inner mass of primary sporang-
ial tissue and outer primary parietal cells. Fruit (caryopsis)
The ultimate cells of sporangial tissue are
the pollen mother cell, which by reduction In a transverse section of a mature cary-
division ultimately leads to the formation of opsis the structure of the seed testa and
four pollen grains (Fig. 5.44). In mature endosperm become visible. A thick
anthers approaching the time of dehiscence, cuticle covers the epidermis; the peri-
some fringe-like outgrowths are observed carp consists of an epicarp, hypocarp
all along the outer wall of the anther. and endocarp. The epicarp is composed
PG
ANS
Fig. 5.44. Transverse section through the anther, showing the pollen grains (75×). PG, pollen grains; ANS,
anther sac (Maiti and Wesche-Ebeling, 1997).
90 Chapter 5
FN
OVL
OV
Fig. 5.45. Longitudinal section through the ovary, showing the ovule (75×). OV, ovary; OVL, ovule; FN,
funicle (Maiti and Wesche-Ebeling, 1997).
of long epidermal cells, the hypocarp rone layer; they contain staining bodies
cells are thickened. The two to three lay- (Fig. 5.46).
ers of mesocarp cells have lost their out-
line due to the compression of outer cell Endosperm
layers. Elongated cross cells are only
rarely visible and tube cells, which are Below the aleurone layer is the peripheral
distinct at an early stage, are mostly corneous endosperm, made up of a single,
obliterated. Below the mesocarp exists a well-demarcated layer of cells, with closely
single layer of large cubic and rectangular arranged starch (Fig. 5.47) and protein gran-
cells, regularly oriented, called the aleu- ules. The large, long, cubic to rectangular
Cereals 91
ENP
EPC
MS
AL
Fig. 5.46. Transverse section through a developing seed, showing epicarp, mesocarp, aleurone layer
and endosperm (75×). EPC, epicarp; ENP, endosperm; MS, mesocarp; AL, aleurone layer
(Maiti and Wesche-Ebeling, 1997).
endosperm cells are loose below this periph- endosperm cells between the aleurone
eral layer and contain the floury endosperm. layer and the scutellum are compact. The
The starch and protein granules become scutellar cells are elongated and form a
progressively less compact towards the single layer that surrounds the embryo.
midpoint of the grain endosperms, from the The embryo cells are compact, especially
milky stage to maturity. in the hilar region. The cells outside the
black layer in the hilar region are compact.
Embryo The black layer shows a semi-lunar ring,
composed of vacuolated cells. The forma-
The aleurone cells in the embryo region tion of this layer coincides with the termi-
are small, compact and rectangular. The nation of grain development.
92 Chapter 5
Fig. 5.47. Scanning electron microscope micrograph of grain endosperm showing starch granules
(Maiti and Wesche-Ebeling, 1997).
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6
Pulses
Pulses are important sources of proteins mechanisms. In a previous study early matu-
and other nutrients. This chapter concen- rity chickpea lines were found, which can
trates only on a few important pulses, such escape terminal drought stress. The results
as chickpea, common bean, green gram, red showed that tolerant lines had a significant
gram, cowpea and pea. difference at 1% level of probability over sus-
ceptible lines for drought tolerance. These
lines produced a significantly higher yield
6.1 Chickpea than control drought tolerant lines. To pro-
duce the highest yield, lines require drought
6.1.1 Introduction tolerance with high adaptability, early maturity
and large seed size (Sabaghpour et al., 2006).
Chickpea (Cicer arietinum), the world’s third
most important food legume, is currently
grown on about 11 Mha, with 96% of cultiva- 6.1.2 Origin and distribution
tion in the developing countries. It is also called
Bengal gram, Garbanzo bean, Indian pea, ceci Chickpea is an ancient crop, first grown in
bean. Chickpea is a self-pollinating diploid Turkey about 7450 BC and in India about 4000 BC.
(2n = 2x = 16) with genome size 1C = 740 Mbp. It Chickpea originated in Mediterranean count-
is the second most important dry grain crop. ries and is cultivated in Asia, south Europe
Chickpea falls into two types: desi and and Mexico. The probable place of origin is
kabuli. The desi type plants are shorter with the region between Cuacaso and north-eastern
smaller leaves, white, pink or blue flowers Persia (van der Maesen, 1972), however,
and pigmented seeds, and are grown in the Ladizinsky (1975) proposes that the centre
semi-arid tropics. The kabuli types are taller of origin was south-eastern Turkey. The most
plants with larger leaves, white flowers and important importing countries are Spain,
longer, more rounded, cream-coloured seeds Algeria, Iran, Libya and the USA.
and are grown in temperate regions. The desi
types predominate in the Indian subcontinent
and kabuli types in other countries. Kabuli 6.1.3 Utilization
types dominate American production.
Chickpea plants mainly adapt to drought Chickpea is used as food in the following
environments either through escape, or tolerance ways. Chickpea is in great demand for its
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 95
96 Chapter 6
nutritive seeds, which have a high protein The stem is erect, branched, viscous, hairy,
content of 25.3–28.9% after dehulling. terete, herbaceous, green and solid. The
Sprouted seeds are eaten as a vegetable or branches are usually quadrangular, ribbed
salads. Young plants and green pods are and green. There are primary, secondary
eaten like spinach. Tinned chickpea is sold and tertiary branches. Thick, strong and
in Turkey and Latin America. Mature chick- woody primary branches (ranging from one
peas can be cooked and eaten cold in salads, to eight) arise from ground level as they
cooked in stews, ground into a flour called gram develop from the pumular shoot as well as
flour (also known as besan and used prima- lateral branches of the seedling. Secondary
rily in Indian cuisine), cooked and ground branches (from 2 to 12) develop at buds
into a paste or roasted, spiced and eaten as a located on the primary branches. They are
snack. Chickpeas are low in fat and most of less vigorous than the primary branches.
this is polyunsaturated. The nutrient profile The number of secondary branches deter-
of desichana (the smaller variety) is differ- mines the total number of leaves, and hence
ent, especially the fibre content, which is the total photosynthetic area. Tertiary bran-
much higher than the light coloured variety. ches arise from the secondary branches. The
Chickpea is also used as animal feed. primary branches form an angle with a ver-
Chickpea is also used for medicinal pur- tical axis, ranging from almost a right angle
poses. Chickpea can use as a hypocholes- to an acute angle. Usually stems are incurved
teraemic agent. Germinated chickpea was at the top, forming a spreading canopy.
effective in controlling cholesterol levels in Chickpea leaves are petiolate, compound
rats. Glandular secretions of the leaves, stems (Fig. 6.1) and uniparipinnate (pseudo-
and pods contain malic and oxalic acids, giv- imparipinnate). Some lines have simple
ing a sour taste. In India, these acids are col- leaves. The rachis is 3–7 cm long with
lected from the crop canopy by spreading grooves on its upper surface. Each rachis
thin muslin over the crop during the night. supports from 10 to 15 leaflets, each with a
This solution is a popular medicine used in small pedicel. The leaflets do not end at the
curing bronchitis, catarrh, cutamenia, cholera, true terminal position, but at the sub-
constipation, diarrhoea, dyspepsia, flatulence, terminal position. The leaflets are 8–17 mm
snake bite, sunstroke and warts. Raw chick- long and 5–14 mm wide, opposite or alter-
peas contain protease (enzyme that breaks nate with the terminal leaflet. Leaflets are
down proteins) inhibitors, which counter- serrated, the teeth covering about two-thirds
act the digestion enzymes in our body. of the foliar blade. Leaflets are obovate to
elliptical with the basal and top portions
cuneate and rounded. Leaves are pubescent.
6.1.4 Morphological description The stipules are ovate to triangular in shape
and serrated with from two to six teeth, 3–5
Vegetative characters mm long and 2–4 mm wide. The longest
margin is toothed and the smallest one
Chickpea is an herbaceous annual crop. The entire. The external surface of the chickpea
growth habit varies from prostate to erect. It plant, except the corolla, is densely covered
is grown in tropical, subtropical and tem- with glandular or non-glandular hairs. The
perate regions. It possesses a strong taproot hairs vary in dimensions: short stalked,
system with three or four rows of lateral multicellular stalked (both glandular and
roots. The parenchymatous tissues of the non-glandular) and unicellular, some geno-
root are rich in starch. All the peripheral types, however, do not possess any hair.
tissues disappear at plant maturity, and are
substituted by a layer of cork. The roots Floral characters
grow 1.5–2.0 m deep and bear Rhizobium
nodules. They are of the carotenoid type, The solitary flowers are born in an axillary
branched with laterally flattened ramifi- raceme. Sometimes there are two to three
cations, sometimes forming a fanlike lobe. flowers on the same nodes; such flowers
Pulses 97
Fig. 6.1. Chickpea plant morphology showing compound leaves and pods.
possess both a peduncle and pedicel. The mm long and about 4 mm wide with an auri-
racemose peduncle is 60–30 mm in length. culate base. The auricules are over the pedi-
At flowering, the floral and racemal portions cel and form a packet in the basal upper part,
of the peduncle form a straight line, giving which is covered by the vexillum. The keel is
the appearance that the flowers are placed on 6–8 mm long, rhomboid, with a pedicel 2–3
the leafy axil by a single peduncle. Chickpea mm long. Two-thirds of the frontal side of its
flowers are bisexual and have a papiliona- ventral face is adnate. The wings do not show
ceous corolla. They are white, pink, purple concrescence with the keel. There are ten sta-
or blue in colour. In coloured flowers, the mens in diadelphous (9)+1 condition. The
peduncles may be of different colours, the filaments of nine of the stamens are fused,
floral part purplish and the raceme green. forming an androecial sheath; the tenth sta-
The axillary inflorescence is shorter than the men is free. The stamina column is persist-
subtending leaf. The calyx is dorsally gibbous ent. The fused part of the filament is 4–5 mm
at the base. There are five sepals with deep long and the free part 2–3 mm, upturned and
lanceolate teeth. The teeth are longer (5–6 dilated at the top. The apex of the stem is
mm) than the tube (3–4 mm) and have promi- oblique. The stamens facing the petals are a
nent midribs. The five sepals are subequal. little longer than the others. The anthers of
The two dorsal (vexillar) sepals are closer to these stamens are bicelled, basifixed and
each other then they are to the two lateral round. The other anthers are dorsifixed,
ones in the ventral position. The fifth calyx ovate, and longer than the basifixed ones at
tooth is separate from the others. The pedun- flowering. The anthers burst longitudinally.
cles and the calyx are glabrous. The calyx The pollen grains are orange. The gynoecium
tube is oblique. Chickpea flowers have five is monocarpellary, unilocular and superior,
sepals, which are generally celeste and pur- with marginal placentation. It is ovate with a
plish red or light pink in colour. The petals pubescent surface. The ovary is 2–3 mm long
are polypetalous, i.e. consisting of vexillum, and 1–15 mm wide. There are 1–3 ovules,
wings and keel. The vexillum is obovate, rarely four. The style is 3–4 mm long, linear,
8–11 mm long, 7–10 mm wide, and either upturned, and glabrous except at the bottom.
glabrous, or pubescent with no glandular hair The stigma is globose and capitates. Some-
on its external surface. The wings are also times it may be as small as the style. Anther
obovate with short pedicles. They are 6–9 dehiscence takes place inside the bud 1 day
98 Chapter 6
before the opening of the flower. When pol- highly wrinkled seeds. The beak above the
len is first liberated, the stigma is still above micropyle is produced by the tip of the radi-
and quite free from the base of the anthers. cle. The shape of cotyledons varies from
The filament gradually elongates to carry semi-spherical to oviform. The length of the
the anthers above the stigma. This process seed ranges from 4 to 12 mm and its width
is completed before the flower opens, thus from 4 to 8 mm. The seed mass varies from
facilitating self-pollination. Anthesis in chick- 0.10 to 0.75 g per seed. The seed colour
pea is occurs through the day. ranges from whitish (even chalky) and cream
to deep black, although many other colours
such as red, orange, brown, green and yellow
Pod and seed development
may be found. The cotyledons are cream,
Pod formation begins 5–6 days after fertili- green, or orange coloured.
zation. The number of pods per plant varies Kabuli seeds contained fewer pectic
between 30 and 150 depending on the envi- polysaccharides and less protein, and had a
ronmental conditions and the genotype. thinner seed coat due to thinner palisade
The pod wall is 0.3 mm thick with three lay- and parenchyma layers. The outer palisade
ers: exocarp, mesocarp and endocarp. The layer thickness varied from one to two cells,
exocarp is hairy and glandular, the meso- leading to a textured and sometimes wrin-
carp has six to eight layers of parenchyma kled appearance of the seed surface. The desi
and the endocarp consists of three to four palisade layers were rigid and extensively
cell layers with fibres in its outermost region thickened. Hourglass cells were homogeneous
and five to six layers of parenchyma. Pod for both seed types, but not in an interspecific
size ranges from 15 to 30 mm in length, 7 to desi line (containing Cicer echinospermum
14 mm in thickness and 2 to 15 mm in parentage), which had heterogeneous cells.
width. The pod shape varies from rhom- The inner surface of the seed coat contained
boid, oblong to ovate. The number of seeds both pectic and proteinaceous materials
per pod ranges from one to two, with the (Wood et al., 2011).
maximum being three.
The number of trichomes and length of
trichomes plays an important role in pod 6.1.5 Anatomy
borer resistance. Pod trichomes’ length and
density showed negative correlation to pod Root anatomy
borer damage in chickpea, i.e. genotypes
having longer pod trichomes and or more A transverse section of the root shows three
pubescent pods received less pod borer dam- parts, epidermis, cortex and stele (Fig. 6.2).
age. The chickpea genotypes having a thicker The epidermis consists of uniseriate
pod wall received lower pod borer damage. compactly arranged barrel-shaped cells with-
Pod length, breadth and area of respective out intercellular spaces and has unicellular
genotypes showed a significant effect in root hairs.
resistance mechanism against pod borer The cortex is the middle region, lying
damage (Hossain et al., 2008). Therefore, between the epidermis and the stele, and is
there is a great necessity in screening peanut multiseriate, parenchymatous and relatively
cultivars and selecting cultivars for trichome homogeneous. Outer layers of the cortex
density on pods, which may then be incor- consist of loosely arranged thin-walled par-
porated into the breeding programme for pod enchyma cells with prominent intercellular
borer resistance. spaces and contain food deposits. The cells
are usually round or oval, colourless and
Seed store starch. This region helps in lateral con-
duction of water and salts. The inner layer of
The seeds are owl’s head shaped, and the the cortex consists of compactly arranged
surface may be smooth or wrinkled. The barrel-shaped cells with Casparian strips
two cotyledons are separated by a groove in or Casparian bands, called endodermis.
Pulses 99
Hypodermis Cortex
Epidermis
Pericycle
Xylem
Endodermis
Pith
Phloem
Fig. 6.2. Transverse section of root (100×). Note: secondary growth in the stele has started and the
endodermis and cortex are undergoing compression due to enlargement of the stele.
In transverse section, Casparian bands Chickpea, being a leguminous plant, has the
appear as a lens-shaped thickening on the capacity to fix atmospheric nitrogen with
radial walls of the endodermis. the help of rhizobium present in such root
Pericycle is the outer most region of the nodules. The symbiotic response of chick-
stele. It is uniseriate and parenchymatous. pea and rhizobium is dependent on genetic
Cells of the pericycle retain meristematic and physiological factors of both species.
activity. Lateral roots arise endogenously Rhizobium spp. (fast-growing species) and
from the pericycle. Stele consists of xylem Bradyrhizobium spp. (slow-growing species)
and phloem and is arranged in the form of are predominantly aerobic chemoorgano-
separate strands on different radii, alternat- trophic soil genera that form root nodules
ing with each other. Xylem is exarch having by symbiotic association with legumes and
protoxylem towards the pericycle and meta- non-legumes, but specially legumes.
xylem towards the centre. Generally four
(tetrarch) protoxylem regions are present ROOT INVASION AND NODULATION Rhizobia are
and form xylem tissue. Phloem occurs rhizosphere organisms multiplying on the
below the pericycle and consists of a large root surfaces and in the surroundings. They
amount of sclerenchyma. The centre of the move to the rhizosphere in response to the
stem is occupied by a small amount of pith nutrients and other substances containing
(Fig. 6.2). the root excretions. In this active soil vol-
ume, population growth is stimulated until
Root nodules the rhizobia occupy the whole chickpea root
surface and attach to epidermal cells, partic-
Members of the Leguminosae family usu- ularly to the root hairs (Fig. 6.3). Some bacte-
ally are found in symbiosis with a bacterium ria specifically interact with newly emerging
of the family Rhizobiaceae, which associ- root hairs, however, and initiate a pronounced
ates with the root cells of the plant forming curling of these growing hair cells. To initiate
specialized structures termed root nodules. this reaction rhizobia have evolved flavonoids.
100 Chapter 6
RHIZOMA IN
(a) RHIZOSPERE
(b) (e) PRIMARY ROOT
APICAL MERISTEM
NODULE
ENDODERMIS
XYLEM
Fig. 6.3. Sequential events in root invasion and nodulation up to differentiation of an active nodule.
(a) Rhizobia moves to the rhizosphere in response to the nutrients and other substances present in the root
secretions. (b) After detection of specific flavonoids released by root cells some rhizobia interact with newly
emerging root hairs and initiate a pronounced curling of hair cells. (c) Invasion of plant cells via induction
of an infection thread that penetrates the plant tissue and continues to grow and ramify in the root cortex.
(d) Infection thread eventually invades a focus of dividing plant cells and rhizobia are released into these
cells. The bacteria continue to grow and ultimately differentiate into bacteroids capable of fixing nitrogen.
(e) An active root nodule is formed (Maiti and Wesche-Ebeling, 2001).
These compounds are released by the plant of cells at different stages of development
into the root rhizosphere and act as positive toward the proximal point of attachment of
regulators of infection genes (Nod genes). the nodule to the parent root. The cytoplasm
Following the initiation of an infection pro- of infected cells in the nitrogen-fixing region
cess, rhizobia entrapped within curled root of the nodules is densely packed with sym-
hair cells began the invasion of these plant biosomes, most of which contain a single
cells. Invasion occurs via the induction of an bacteroid. In later stages of development
infection thread that penetrates the plant tis- infected cells become enlarged and highly
sue and continues to grow and ramify in the vacuolated, and eventually lose their con-
root cortex. The infection thread releases tents. Uninfected cells in the central region
rhizobia and these are embedded within a are smaller than infected cells and are also
plant membrane called the peribacteroidal highly vacuolated.
membrane. The bacteria continue to grow
and ultimately differentiate into bacteroids
Stem anatomy
capable of fixing nitrogen.
The outline of the stem in transverse section
NODULE GROWTH AND MORPHOLOGY Chickpeas is quadrangular in shape. It shows an outer
have a nodule type termed intermediate epidermis, central cortex and inner stele
(Fig. 6.4), which is characterized by the (Fig. 6.5).
presence of a defined meristem during nod- Epidermis contains uniseriate compactly
ular growth, an open vascular system con- arranged cells with unicellular long tricho-
necting the root vascular system with the mes. The epidermis is cover by thick cuticle.
nodule meristem, vacuolated infected cells, Below the epidermis, four to six layers of col-
aspargine/glutamine as the major transloca- lenchyma with some chlorenchyma patches
tion products of nitrogen fixing activity, and are present. Collenchyma gives considerable
enlarged simple bacteroids. strength, flexibility and elasticity to young
The features of developing and senesc- stems. Having chloroplasts, it may carry out
ing bacteroids include the presence of per- photosynthesis.
sistent meristematic tissue at the distal end Multilayered parenchymatous cortex is
of the multi-lobed nodules, and a gradient present between the collenchyma and stele.
Pulses 101
NODULE CORTEX
OPEN
VASCULAR
SYSTEM
INFECTED REGION
WITH ACTIVE
VASCULAR NITROGEN-FIXING
CYLINDER BACTEROIDS
Fig. 6.4. Schematic structure of an indeterminate root nodule. The open vascular system distinguishes
it from a determinative root nodule (Maiti and Wesche-Ebeling, 2001).
Trichome
Epidermis
Cortex
Collenchyma
Phloem
Xylem
Inter-fascicular cambium
Fascicular cambium
Medulla
Fig. 6.5. Transverse section of stem (100×) showing unicellular long trichomes, collenchyma, single layer of
epidermis and the medulla. Secondary growth is just initiated, so inter-fascicular cambium and fascicular
cambium are forming secondary vascular tissues.
Cells of the inner layer of the cortex, the bundles, medulla and medullary rays. Xylem
endodermis, are thickened by Casparian and phloem organize in the form of vascular
strips. It is uniseriate with stored food mate- bundles; 10–12 wedge-shaped vascular bun-
rial and covers the stele. dles are arranged in one ring, called eustele.
Stele is the central region of the stem and It contains xylem towards the centre of the
larger than the cortex. It consists of vascular axis and phloem towards the periphery.
102 Chapter 6
In the vascular bundle, xylem and phloem are specialized type of photosynthetic tissue.
present together (conjoint) on the same radius The upper surface of the leaf is dark green in
(collateral), with a strip of cambium (fascicu- colour due to palisade tissue. The lower part
lar cambium) between xylem and phloem. of the mesophyll present towards the lower
Hence the vascular bundle is described as epidermis is the spongy tissue. Cells of the
conjoint, collateral and open type. Xylem is spongy tissue are thin walled, irregular in
endarch with protoxylem present towards the shape, and arranged loosely with large con-
centre. Medulla and medullary rays are dis- tinuous intercellular spaces. Stomata open
tinct, present at the centre of the stem and into large intercellular spaces called sub-
usually parenchymatous. The cells are round stomatal chambers. The cells of spongy
or oval with intercellular spaces. Medulla parenchyma contain a smaller number of
occupies a large area compared to the other chloroplasts, hence the lower surface of the
parts of the stem, and stores food materials. leaf is light green in colour.
The extensions of parenchymatous pith The vascular tissue occurs in the form
between vascular bundles are called primary of discrete bundles called veins. Veins are
medullary rays, which help in lateral conduc- interconnected to form reticulate venation.
tion and may give rise to secondary meristem Vascular bundles (veins) occur in between
(inter-fascicular cambium). the palisade and spongy tissue (median
in position). They are collateral and closed.
Leaf anatomy In the vascular bundle, xylem is present
towards the upper epidermis and phloem
The leaf shows a dorsi-ventral nature, i.e. towards the lower epidermis. The vascular
the leaf blade consists of distinct dorsal and bundle is surrounded by parenchymatous
ventral surfaces. Leaf transverse section bundle sheath. The cells of the bundle
shows three distinct regions, the epidermis, sheath are called border parenchyma.
mesophyll and vascular system. A large number of trichomes are present
The epidermis is single layered with on the leaf surface of chickpea (Fig. 6.6).
compactly arranged barrel-shaped (tabular) During late vegetative growth, chickpea
cells. The outer surface of the epidermis is leaves and stem are covered with aqueous
covered by a thin continuous layer of cutin glandular droplets. If these droplets persist
called cuticle. Anisocytic type of stomata at low humidity for a long time, there may be
are present in both epidermal layers. The substantial loss of water via glandular tri-
stomatal index of upper epidermis is 28 to chomes. It is concluded that water loss via
33 and lower epidermis is 35 to 37 under trichomes could lower leaf temperature by
40× magnification. Stomata facilitate the several degrees within a wide range of atmos-
exchange of gases between the leaf and envi- pheric conditions. Seasonal variations were
ronment. All the epidermal cells except observed in hydrochloric acid, malic acid
guard cells are colourless. and Ca ions secreted by chickpea trichomes.
The ground tissue of the leaf, present It is suggested that malate, chloride ions and
between the upper and the lower epidermal Ca ions are linked to the amount of daily
layers, is called mesophyll. The mesophyll sunlight and to the day of the year, while the
is chlorenchymatic and concerned mainly pH of secretions is not directly related to
with photosynthesis. The mesophyll is dif- sunlight level (Lazzaro et al., 1995).
ferentiated into an upper palisade tissue and
lower spongy tissue. The palisade tissue is Role of glandular trichomes
present below the upper epidermis. The pali- in insect resistance
sade cells are thin walled, cylindrical and
contain numerous chloroplasts. The cells are Chickpea leaves bear glandular trichomes
compactly arranged in one layer and perpen- on the leaf surface. Little research has been
dicular to the upper epidermis. Narrow, inter- undertaken into the role of these trichomes
cellular spaces are present in the palisade in insect resistance.
tissue and allow the loss of water by transpi- Late vegetative growth chickpea leaves
ration. Palisade tissue is the most highly and stems can be covered with aqueous
Pulses 103
Upper epidermis
Palisade tissue
Spongy tissue
Trichome
Xylem
Phloem
Lower epidermis
Fig. 6.6. Transverse section of leaf (100×). Note: palisade tissues are long and compactly arranged,
and epidermis consists of bilobed trichomes. Xylem is towards the upper epidermis and phloem towards
the lower epidermis.
glandular droplets. In a study it was con- were investigated. However, there was a
cluded that water loss via the glandular significant negative correlation between
trichomes can be enough to lower leaf tem- pod damage and oxalic acid levels. Oxalic
perature by several degrees centigrade acid, which had been reported to have an
within a wide range of atmospheric condi- antibiotic effect on H. armigera larvae, has
tions. The exudate solutes were primarily an important role in resistance to this pest
malic, hydrochloric and oxalic acids. With- in chickpea (Yoshida et al., 1997).
out the strong acids a chickpea leaf, wet Chickpea trichome exudates contain
even on dry days, would be ripe for para- malic acid, oxalic acid and succinic acid.
sitic attack (Lauter and Munns, 1986). Oxalic acid has an antibiotic effect on
The role of exudates from trichomes the larvae of the pod borer, H. armigera,
were studied as follows: the plant charac- which results in reduced pod damage. A
ters responsible for the absence of egg dense mat of non-glandular trichomes
parasitoids and the feasibility of increasing in chickpea species prevents the small
parasitism levels on chickpea by mass- larvae from feeding on the pods. Gland-
releasing Trichogramma chilonis Ishii ular trichomes and their chemical exu-
were investigated. The residence time of dates may also be harmful to insect
female T. chilonis on chickpea leaves was predators and parasites (Peter and
affected by trichomes and the acidic Shanower, 1998).
trichome exudates secreted on all green The comparative assessment of density
parts of the plant. The parasitoids spent a of glandular hairs, population and size of
longer time on chickpea leaves where the stomatal apertures in chickpea cultivars
acidic trichome exudates washed off than were studied. In the resistant reaction group,
on unwashed leaves. When placed on there were higher number of glandular hairs
unwashed chickpea leaves, 6.8% of the on the dorsi-ventral sides and higher
parasitoids were trapped and killed by the number of leaf stomata as compared to the
exudates (Romeis et al., 1999). susceptible group. The resistant group had
Effects of malic acid and oxalic acid the smallest stomatal apertures (Randhawa
on oviposition of Helicoverpa armigera et al., 2009).
104 Chapter 6
reniform. The hilum is in the form of a half- acts as a barrier between cortex and stele.
moon shape. Raphe formed by two elevated The endodermis ruptures due to secondary
locules of clear shiny coffee colour. The growth.
micropyle is round. Cotyledons are large, Pericycle is the outermost region of the
hard and a cream colour. The seed surface stele, and is uniseriate and parenchyma-
is reticulate, lobular in appearance and of tous. Cells of the pericycle retain meristem-
variable colour, often coffee with dark atic activity. Lateral roots arise endogenously
spots. from the pericycle and help in absorption of
water. During secondary growth the pericy-
cle forms phellogen or cork cambium and a
6.2.5 Anatomical description small amount of vascular cambium.
In the vascular tissue, xylem and
Root anatomy phloem are arranged in the form of separate
strands on different radii, alternating with
The epidermis is uniseriate, having a single each other. Hence the vascular strands in
layer of compactly arranged thin-walled liv- root are described as separate and radial.
ing cells. Both cuticle and stomata are Xylem is exarch having protoxylem towards
absent. The epidermis gives protection and the pericycle and metaxylem towards the
absorption of water and minerals. Epidermal centre and scattered secondary xylem ves-
cells produce unicellular root hairs, which sels. Cambium is absent and hence vascular
are useful in increasing the surface area for bundles are described as closed type.
absorption of water and helps in obtaining
water from soil. The root hairs are generally Stem anatomy
short lived, and the epidermis ruptures after
secondary growth. A transverse section of primary stem shows
Cortex is the middle region, lying three regions, the epidermis, cortex and stele.
between the epidermis and the stele, and is Epidermis is the outermost region sur-
well developed and larger than the stele. It rounding the stem. It is uniseriate, having a
is multiseriate, relatively homogenous and single layer of compactly arranged barrel-
consists of general cortex and endodermis. shaped living cells with trichomes. The
Cortex is extensively developed in the root outer walls of the epidermal cells are highly
and consists of loosely arranged thin-walled cutinized, which reduces transpiration.
parenchyma cells with prominent intercel- Few stomata are present in the epidermis
lular spaces and flavonoid deposits. The for exchange of gases and transpiration. All
cells are colourless and store starch and are the epidermal cells are colourless except
usually round or oval. This region helps in the guard cells. Multicellular hairs are
the lateral conduction of water and salts. present on the epidermis.
Rhizobium bacteria lives as symbionts in The cortex is the middle region present
cortical cells and fix biological nitrogen. between the epidermis and stele, and is
After secondary growth the general cortex smaller than the stele. Hypodermis is multi-
ruptures. layered (from two to six layers) and col-
Endodermis is very distinctive in roots. lenchymatic; the collenchyma is living
It is single layered with compactly arranged mechanical tissue. The hypodermis pro-
barrel-shaped cells. Endodermal cells are vides considerable strength, flexibility and
characterized by the presence of Casparian elasticity to young stems. Having chloro-
strips or Casparian bands made up of plasts, it may carry on photosynthesis. The
suberin. In transverse section, Casparian endodermis is the innermost layer of the
bands appear as a lens-shaped thickening cortex, and is usually distinct in stems with-
on the radial walls of the endodermis. The out Casparian bands. The pericycle is multi-
endodermis is also called the starch sheet seriate and sclerenchymatic. Sclerenchyma
layer, as starch grains are present in the is mechanical tissue and gives strength and
endodermal cells. Endodermis of the root rigidity to the stem.
106 Chapter 6
Xylem and phloem form the vascular smaller, more irregularly shaped epidermal
tissues. They organize into wedge-shaped cells than the smooth leaves.
vascular bundles, with 15–20 vascular bun-
dles arranged in one ring called a eustele.
Leaf anatomy and adaptation to drought
The vascular bundle consists of xylem
and insects
towards the centre of the axis and phloem
towards the periphery. Both xylem are Leaf anatomical traits play important roles
phloem are present together (conjoint) on in adaptation to biotic and abiotic stress
the same radius (collateral), having a strip factors.
of cambium (fascicular cambium) between A study was conducted to find out
xylem and phloem (open), hence the vascu- genetic variability for trichome distribution
lar bundle is described as conjoint, collat- and density among three diverse dry bean
eral and open type. Xylem is endarch with (P. vulgaris L.) cultivars, and to characterize
protoxylem present towards the centre. The the types of trichomes present among the
xylem consists of many vessels arranged in cultivars. Different types of trichomes were
rows. The extensions of parenchymatous identified, on both the abaxial and adaxial
pith between vascular bundles are called leaf surfaces of the cultivars ‘Bill Z’, ‘Pom-
primary medullary rays, which help in lat- padour Checa’ and ‘Diacol Calima’. More
eral conduction and may give rise to second- straight trichomes are present on the abaxial
ary meristem (inter-fascicular cambium). leaf surface (Dahlin et al., 1992).
In common bean (P. vulgaris L.) leaf
Leaf anatomy epidermal characteristics may play a role
in plant defence against pathogens and/or
The leaf epidermal cell surface of a common drought. Crystalloids were present on
bean shows hook-shaped trichomes on the both surfaces of the leaves which were
abaxial and adaxial leaf surfaces. In a trans- devoid of wax. Trichome density and dis-
verse section, the leaf epidermal cells of tribution differed among the common bean
various sizes are covered by a thin cuticle. types. The abaxial leaf surface always pre-
This is followed by one layer of non- sented more trichomes than the adaxial
compact palisade parenchyma with abun- surface. Common bean leaves presented
dant chloroplasts. paracytic, anomocytic and anisocytic sto-
Leaflets of P. vulgaris contain calcium matal types. All common beans presented
oxalates in the adaxial bundle sheath exten- the adaxial epidermis of the leaves with
sions. A comparative study was made on a lower density of bigger stomata than on
the leaf epidermal cell structure of four bean the abaxial epidermis (Sebastian Stenglein
cultivars. Glandular trichomes were gener- et al., 2004).
ally present on the veins. It was found that Cuticle thickness plays a more impor-
there is an inverse relation between leaf tant role than calcification of cell walls in
growth and the number of stomata, epider- the resistance of older plants to R. solani
mal cells and trichomes at the lower and (Stockwell and Hanchey, 1983).
upper surface of the leaves. The decrease in
number of these anatomical variables was Seed anatomy
associated with the elongation and expan-
sion of the cells with growth. In a transverse section the cuticle visible
Leaf pubescence in P. vulgaris has been and macrosclereids are thin and elongated
associated with resistance to Uromyces in the form of a cylinder. There is a large
appendicularis. The density of trichomes number of starch granules distributed uni-
varied among cultivars and among leaves formly in the cotyledon cells. Protein gran-
from different nodes. There was an inverse ules are amorphous and distributed around
relationship between density of hairs and starch granules. The species showed large
hooked trichomes. In a study, wrinkled variations in form, size, position of hilum,
leaves had more trichomes and stomata and micropyle, and seed surface and colour.
Pulses 107
Fig. 6.7. Green gram plant morphology illustrating simple leaves with ovate shape and acute apex, yellow
flowers and slender pods.
Epidermis
Phloem
Xylem Collenchyma
Cortex
Groove
Ridge
Trichome
Medulla
Fig. 6.8. Outline of the transverse section of stem (100×) has ridges and grooves and collenchyma is present
below the ridges only. In the vascular bundle xylem is towards the periphery and phloem towards centre.
The central portion is occupied by the medulla.
below the ridges and small vascular bun- consists of parenchyma with large inter-
dles below the grooves. Xylem is endarch cellular spaces. Stomata open into large
with protoxylem present towards the cen- intercellular spaces called sub-stomatal
tre. The xylem consists of many vessels, chambers.
which are arranged in rows. Pith is prom- Vascular bundles (veins) occur between
inent and occupies a larger amount of the palisade and spongy tissue (median
space. in position). They are collateral and closed.
In the vascular bundle, xylem is present
towards the upper epidermis and phloem
Leaf anatomy towards the lower epidermis.
The presence of loose palisade cells leads
Leaf shows a dorsi-ventral nature, with uni- to a large amount of transpiration loss and
seriate compactly arranged epidermis present makes the plant drought susceptible.
on both surfaces of the leaf. Epidermis is
covered by a thin layer of cuticle. The aniso-
cytic type of stomata is present in both epi- Petiole anatomy
dermal layers. The stomatal index of the
upper epidermis is 25 and of the lower epi- A transverse section of the petiole shows
dermis is 28. three regions, the epidermis, ground tissue
The mesophyll is chlorenchymatic and vascular bundles (Fig. 6.10).
and differentiated into palisade and spongy The epidermis is uniseriate, having a
tissue. The palisade cells are thick wal- single layer of compactly arranged barrel-
led, cylindrical and loose and contain shaped living cells. Unicellular hairs are
numerous chloroplasts (Fig. 6.9). Mesophyll present on the epidermis.
110 Chapter 6
Spongy tissue
Xylem
Phloem
Lower epidermis
Collenchyma
Fig. 6.9. Transverse section of leaf (100×) showing arrangement of tissues. Palisade cells are not very
compactly arranged. At the centre of the leaf (midrib) two to three layers of collenchyma are present
above the lower epidermis.
Epidermis
Collenchyma
Ground tissue
Stele
Vascular trace
Trichome
Fig. 6.10. Transverse section of petiole (100×) showing epidermis with trichomes, collenchyma, a large
amount of ground tissue made up of parenchyma and central (stele) vascular tissue. Note: Vascular trace
is clearly visible in the stele making the stele appear like a ‘C’ shape.
6.4 Red Gram (Pigeonpea) early, small, few-seeded ‘tur’ type of Madhya
Pradesh, western and peninsular India.
6.4.1 Introduction
Cajanus cajan is one of the most drought- cm wide and pubescent, as is the stem.
tolerant legume crops, with a wide range of Lateral petioles are 2–3 mm long, with the
rainfall tolerance, but prefers more than 625 terminal petiole reaching 10–20 mm. Stipules
mm and in elevated areas exceeding 2000 m are linear and 2–3 mm long, whereas stipu-
cold nights and cloudy weather interfere lets are filiform and 1–2 mm long.
with fertilization of flowers. It flowers well in
regions where rainfall is 1500–2000 mm.
However, it will grow on deep, well-structured Floral characters
soils where the rainfall is only 250–375 mm.
Red gram is an annual, or more usually Flowers form in racemes, having from five to
short-term perennial shrub, that may reach ten flowers on top of an axillary, little-divided
4–5 m in height, but usually only 1–2 m, peduncle. Flowers are usually yellow but
woody at the base, with a variable habit but they may also be striated with purple streaks
usually erect. It has a taproot system, which or plain red. Flowers are pedicellate, bract-
is deep and quick growing. Root nodules eate, complete, bisexual, zygomorphic, pen-
are present on the lateral roots, with Rhizo- tamerous and perigynous. Usually two flowers
bium bacteria living as symbionts in the open at the same time on one inflorescence.
root nodules and performing the biological The flowers are self-pollinated, pollination
fixation of atmospheric nitrogen. Generally taking place before the flowers open. The
root nodules appear on the roots 3 weeks calyx is 10–12 mm long with five linear teeth.
after seeding, and the young nodules are the Pods are flat, with an acuminate tip, pubes-
most active in fixing nitrogen. The stem is cent and of variable colour, 5–9 cm long ×
green to brown in colour and thick. The 12–13 mm wide. There are five gamosepalous
angular stem results from three ribs starting sepals. The corolla is 20–25 mm long, with the
from the base of each petiole. flag 18–20 mm wide. There are five petals,
Leaves are trifoliate and alternate, set in red in colour, free, in a papilionaceous
a spiral along the stem (Fig. 6.11). Leaflets arrangement, and the posterior petal is large,
are oblong, lanceolate, 5–10 cm long × 2–4 called a vexillum. Wing petals are present
both sides of the vexillum; keel petals are
present on the anterior side. There are ten
stamens, diadelphous, dimorphic, dithecous,
introse, unicarpellary, unilocular. From one to
three ovules are present on marginal placen-
tation, simple hairy stigma. The fruit is legu-
minous, with green pods containing two to
nine seeds in shades of brown, red or black and
rounded. Husks bearing deep oblique furrows
underline the septa between the seeds.
Root anatomy
Stem anatomy
Medulla and medullary rays are dis- cylindrical and contain numerous chloro-
tinct, present at the centre of the stem and plasts. Spongy tissue consists of paren-
are usually parenchymatous. The cells are chyma with large intercellular spaces.
round or oval with intercellular spaces. The Vascular bundles (veins) occur in between
medulla stores food materials. The exten- the palisade and spongy tissue (median in
sions of parenchymatous pith between vas- position) and are collateral and closed. In
cular bundles are called primary medullary the vascular bundle, xylem is present
rays, which help in lateral conduction and towards the upper epidermis and phloem
may give rise to secondary meristem (inter- towards the lower. The leaf is bifacial and
fascicular cambium). covered with multicellular hairs. The
stoma were more dense in the lower epi-
Leaf anatomy dermis than the upper epidermis, and both
sides of the leaf had a corneous layer (Yin
The leaf shows a dorsi-ventral nature, and et al., 2004).
uniseriate compactly arranged epidermis is The presence of a thick cuticle, high
present on both surfaces of the leaf. The density of trichomes and compactly arran-
epidermis is covered by a thin layer of cuti- ged palisade tissues reduce drastically tran-
cle. There is a high density of unicellular spiration, thereby contributing to the drought
trichomes on the upper and lower epider- resistance of red gram.
mis (Fig. 6.13).
The mesophyll is chlorenchymatic and Role of trichomes in insect resistance
differentiated into palisade and spongy tis-
sue. The palisade cells are thick walled, Three types of glandular (types A, B, and E)
trichomes and two non-glandular (types C
and D) types were identified in red gram.
Types A, B, C and D were present on leaves,
Cuticle Upper epidermis
pods and calyxes of all three Cajanus spp.,
Palisade tissue except for Type A, which was not found on
pods and calyxes of most C. scarabaeoides
Spongy tissue
accessions examined. Because of their small
size and rarity, pods of C. scarabaeoides were
the most densely pubescent, followed by pods
of C. cajan and C. platycarpus.
Trichome density on pods varied sig-
nificantly among red gram genotypes and
different accessions of C. scarabaeoides.
The resistance of C. scarabaeoides pods to
Trichome Helicoverpa armigera (Hübner) larvae repor-
ted in an earlier study is due to the high
density of non-glandular trichomes. This
Lower epidermis wild species may thus be an important
source for developing insect-resistant red
Xylem Phloem
Collenchyma gram (Romeis et al., 1999).
Trichome length and density, sugars,
proteins and phenols were found to be asso-
Fig. 6.13. Transverse section of leaf (100×) showing ciated with resistance to M. vitrata in short-
high density of trichomes on the lower epidermis.
duration red gram genotypes. Trichome
Note: palisade cells are long and extremely
compactly arranged and the lower epidermis is fully
density on upper and lower surfaces of
covered by trichomes, there is a thick cuticle and the leaf, and length and trichome density
large xylem vessels. All these characters contribute and length on pods were found to be nega-
drought resistance to red gram compared with other tively correlated with the resistant genotype
legume crops. (Sunitha et al., 2008).
Pulses 115
Floral characters
6.5.3 Utilization
Flowers are in a terminal racemose inflo-
Cowpea is cultivated for the seeds (shelled rescence and are pedicellate, bracteate,
green or dried), the pods or leaves that are complete, bisexual, zygomorphic, penta-
consumed as green vegetables or for pas- merous and perigynous. There are five
ture, hay, silage and green manure. Tender gamosepalous sepals and five petals, free,
cowpea leaves and shoots contain 4% pro- in a papilionaceous arrangement; the pos-
tein, 4% carbohydrates and are rich in cal- terior petal is large, called the vaxillum.
cium, phosphorus and vitamin B. Wing petals are present on both sides of the
Dried seeds contain 22% protein and vexillum; keel petals are present on the
61% carbohydrates. According to the USDA anterior side. The male flower contains ten
food database, cowpea has the highest per- stamens, which arise in two whorls of five
centage of calories from protein among vege- each. The stamens of the outer whorl are
tarian foods. The leaves may be dried and opposite to the sepals and the inner whorl
stored for later use. Cowpea has the useful is opposite to the petals; hence it is called
ability to fix atmospheric nitrogen through its diplostemonous. Anthers are diadelphous,
root nodules, and it grows well in poor soils dithecous, basifixed and introrse. The female
with more than 85% sand and with less than flower is unicarpellary and unilocular, with
0.2% organic matter and low levels of phos- one to three ovules present on marginal pla-
phorus. In addition, it is shade tolerant and centation, with simple stigma and a hairy
therefore compatible as an intercrop with style. The fruit is a legume, long, slender
maize, millet, sorghum, sugarcane and cot- and green in colour.
ton. This makes cowpea an important com- There are 12–20 seeds per fruit, roun-
ponent of traditional intercropping systems, ded, and showing variation in colour among
especially in the complex and elegant sub- varieties.
sistence farming systems of the dry savan-
nahs in sub-Saharan Africa. In Africa, where 6.5.5 Anatomical description
cowpea is the preferred food legume, they are
consumed in two basic forms: (i) cooked Root anatomy
together with vegetables, spices and often
palm oil, to produce a thick bean soup, which A transverse section of the root shows three
accompanies the staple food (cassava, yams, regions, the epidermis, cortex and vascular
plantain); and (ii) decorticated and ground tissue.
into a flour and mixed with chopped onions Epidermal cells are compactly arranged
and spices and made into cakes. as thin-walled living cells. The cortex is
Pulses 117
Fig. 6.15. Cowpea plant morphology, showing simple leaves, yellow flowers and long slender pods.
the middle region, lying between the epi- elongated with a nucleus and more vacu-
dermis and the stele. oles. The bacteroids within cowpea cells
The cortex is well developed and is were arranged without any particular order
larger than the stele. It is multiseriate and with more space for host cellular material.
relatively homogeneous and consists of They were mostly present singly in peri-
parenchyma. The innermost layer of the bacteroidal membrane sacs, which sometimes
cortex is called the endodermis. This is fused to enclose more than one bacteroid. In
single-layered with compactly arranged the differentiation of nodular tissue, the hosts
barrel-shaped cells. The endodermis is also seem to play the dominant role in the mor-
known as the starch sheet layer, as starch phogenesis of bacteroids in symbiotic systems
grains are present in the endodermal cells. induced by the same strain of Rhizobium
The stele is covered by uniseriate paren- (Sen et al., 1986).
chymatous pericycle.
The stele (vascular strands) consists of Stem anatomy
xylem and phloem. Xylem conducts water
and salts and phloem transports food mater- The epidermis is uniseriate, having a single
ials. Vascular strands in the root are sepa- layer of compactly arranged barrel-shaped
rate and radial. Xylem is exarch, having living cells. The outer walls of the epider-
protoxylem towards the pericycle and mal cells are cutinized. The epidermis is
metaxylem towards the centre and scattered covered by a separate layer of cutin called
secondary xylem vessels. cuticle on its outer surface (cuticularized).
Few stomata are present in the epidermis
ROOT NODULES The role of nitrogen fixa- for the exchange of gases and transpiration.
tion is shown by the structure and organ- All the epidermal cells are colourless except
ization of nodular tissues and bacteroids the guard cells. Multicellular trichomes are
of cowpea induced by Rhizobium sp., 4 to 5 present on the epidermis.
weeks after inoculation, when nitrogenase The cortex is the middle region present
activity reaches a peak. All cell types in between the epidermis and stele, and is
cowpea nodules were larger. The inner cor- smaller than the stele. The hypodermis is
tex of cowpea had an ‘endodermis-like’ layer multilayered (four to six layers) and collen-
of cells. Many cells of cowpea remained free chymatic. Collenchyma is a living mechani-
of bacteroids. Cells in cowpea were mostly cal tissue. The endodermis is the innermost
118 Chapter 6
layer of the cortex without any Casparian significantly only in trichome number (sus-
strips. ceptible cultivated cowpeas have more) and
Xylem and phloem are organized into non-glandular trichome length. Trichome
wedge-shaped vascular bundles, with 15–20 length and angle to pod surface seemed to
bundles arranged in one ring called a be more important than density (Jackai and
eustele. It consists of xylem towards the Oghiakhe, 1989).
centre of the axis and phloem towards the Damage to cowpea (Vigna unguiculata)
periphery. In the vascular bundle xylem by the legume pod borer M. testulalis (Geyer)
and phloem are present together (conjoint) showed that trichome cover on individual
on the same radius (collateral) and have a cultivars varied in trichome length and
strip of cambium (fascicular cambium) density. Trichome density on different parts
between xylem and phloem (open); hence decreased with increasing plant age. Signi-
the vascular bundle is described as conjoint, ficant (P < 0.05) negative correlations were
collateral and open type. The xylem is endarch obtained between total trichome density on
with protoxylem present towards the cen- pods, pod infestation and damage severity.
tre. The xylem consists of many vessels Results suggest that trichome length may be
arranged in rows. less important than density in reducing pod
damage by M. testulalis in cultivated cow-
Leaf anatomy peas. It is concluded that breeding for a
higher level of trichomes in high yielding
Unicellular trichomes are present on both and agronomically desirable cowpea culti-
the epidermal layers and thick cuticle cov- vars will serve as an important component
ers the epidermis. Well developed meso- in the integrated management of M. testula-
phyll is distributed between the upper and lis (Oghiakhe et al., 1992).
lower epidermis. Studies revealed that there was a sig-
The mesophyll is highly differentiated nificant positive correlation between larval
into palisade and spongy tissue. The pali- penetration time on pods and length of non-
sade cells are cylindrical in shape, thick glandular trichomes. Weight loss, number
walled and consisting of numerous chloro- of feeding punctures and number of larvae
plasts. Sponge tissue consists of paren- on whole pods of wild varieties were lower
chyma with intercellular spaces and is for and significantly different from cultivated
storage of food materials. and semi-cultivated. This study showed
Vascular bundles are closed, collateral that it would be advantageous to use wild
and occur between the palisade and spongy varieties in a breeding programme to incor-
tissue (median in position). porate pubescence into high-yielding com-
mercial cultivars for resistance against M.
Role of trichomes to insect resistance testulalis and possibly other major pests
(Oghiakhe, 1995).
Cowpea possesses leaf trichomes. Few stud-
ies have been undertaken the role of tri- Disease resistance
chomes to various biotic stresses.
The role of pubescence was investi- The infection process of Colletotrichum
gated in the resistance of two wild cowpea destructivum, a hemibiotrophic anthracnose
varieties. Resistance to Maruca testulalis fungus, was studied in two cowpea (V. ungui-
(Geyer) was based on trichomes in the first culata) cultivars that differ in disease reaction
instance and phytochemicals, with previ- type. In the susceptible cultivar there was
ous reports indicating that phytochemicals formation of large, multilobed, intracellular
may be the principal factor in the resistance infection vesicles, followed by necrotrophic,
of TVNu 72 and TVNu 73 to this species. radiating, secondary hyphae produced in
Glandular and non-glandular trichomes were tissues. But in the resistant cultivar, the pro-
found to be present on both the cultivated and duction of appressoria and their melaniza-
wild cowpea. Two types of cowpea differed tion resulted in reduced penetration. Where
Pulses 119
Petiole anatomy
6.6 Pea
There is a close similarity between petiole
and stem with regard to the structure of 6.6.1 Introduction
epidermis. The ground parenchyma of the
petiole is like the stem cortex in arrange- A ‘pea’ is most commonly the small spheri-
ment of cells and in number of chloro- cal seed or the seed-pod of the legume Pisum
plasts. The supporting tissue is collenchyma sativum Linn. Each pod contains several
in relation to the arrangement of vascular peas. Peapods are botanically a fruit, since
tissues in the stem; the vascular bundles of they contain seeds developed from the ovary
the petiole are collateral. of a (pea) flower. However, peas are consid-
The epidermis is uniseriate, having a ered to be a vegetable in cooking. The name
single layer of compactly arranged barrel- is also used to describe other edible seeds
shaped living cells. The outer walls of the from the Fabaceae such as the pigeonpea
epidermal cells are cutinized, which reduces (C. cajan), the cowpea (V. unguiculata) and
transpiration. Unicellular hairs are present the seeds from several species of Lathyrus.
on the epidermis.
Multilayered (three to six layers) hypo-
dermis of angular collenchyma cells is
found immediately beneath the epidermis. 6.6.2 Origin, distribution and adaptation
Collenchyma is living mechanical tissue. The
hypodermis gives considerable strength, The wild pea is restricted to the Medi-
flexibility and elasticity to young petioles. terranean basin and the Near East. The ear-
Just beneath the hypodermis ground tissue liest archaeological finds of peas come
is found. This consists of thin-walled paren- from Neolithic Syria, Turkey and Jordan. In
chymatous cells with well-defined intercel- Egypt, early finds date from ca. 4800–4400
lular spaces among them. BC in the Nile delta area, and from ca.
Vascular bundles are arranged in the 3800–3600 BC in Upper Egypt. The pea was
form of a half-moon shape in ground tissue. also present in Georgia in the 5th millen-
The wedge-shaped vascular bundles are of nium BC. Farther east, the finds are more
various sizes in the same petiole. recent. Peas were present in Afghanistan ca.
2000 BC, in Harappa, Pakistan, and in north-
Research needs west India in 2250–1750 BC. In the second
half of the 2nd millennium BC this pulse
In view of the literature discussed above, it crop appeared in the Gangetic basin and
may be concluded sufficient research inputs southern India.
have been directed on the roles of glandular The pea is a green, pod-shaped veget-
trichomes in insect resistance in cowpea. able, widely grown as a cool-season veget-
The cuticle also has some role but studies able crop. The seeds may be planted as soon
are not available on the roles of these impor- as the soil temperature reaches 10°C (50°F),
tant traits on drought resistance. Several with the plants growing best at tempera-
field crops discussed in this volume show tures of 13–18°C (55–64°F). They do not
the important traits for drought resistance; thrive in the summer heat of warmer tem-
the presence of profuse trichomes and cuti- perate and lowland tropical climates but do
cle reduces transpiration loss. grow well in cooler high-altitude tropical
120 Chapter 6
areas. Many cultivars reach maturity about support and can climb to 1–2 m high. A tra-
60 days after planting. Generally, peas are ditional approach to supporting climbing
grown outdoors during the winter, not in peas is to thrust branches pruned from trees
greenhouses. Peas grow best in slightly or other woody plants upright into the soil,
acidic, well-drained soils. providing a lattice for the peas to climb.
Branches used in this fashion are sometimes
called ‘pea brush’. Metal fences, twine, or
6.6.3 Utilization netting supported by a frame are used for
the same purpose. In dense plantings, peas
give each other some measure of mutual
Pisum are used as vegetables. The peapod is
support. Pea plants can self-pollinate.
eaten flat, while in sugar/snap peas, the pod
becomes cylindrical but is eaten while still
crisp, before the seeds inside develop. The Vegetative characters
seeds have a high protein content.
Pisum is an annual herb, mesophytic, and
has a taproot system with bacterial root
nodules. The stem is weak and a tendril
6.6.4 Morphological description climber (Fig. 6.16), aerial, greenish, rounded
and pubescent. The leaf base is pulvinate
Pisum sativum is an annual plant, with a and the leaves are pinnately compound, alter-
life cycle of 1 year. It is a cool-season crop nate, ovate, with an acute apex and reticulate
grown in many parts of the world; planting venation. Terminal leaflets become tendrils;
can take place from winter through to early stipules are foliaceous.
summer depending on location. The aver-
age pea weighs between 0.1 and 0.36 g. Floral characters
The species is used as a vegetable, fresh,
frozen or canned, and is also grown to pro- Flowers are a terminal racemose type and are
duce dry peas such as the split pea. These pedicellate, bracteate, complete, bisexual,
varieties are typically called field peas. zygomorphic, pentamerous and perigynous.
The veining cultivars grow thin tendrils There are five gamosepalous sepals and five
from leaves that coil around any available free petals in a papilionaceous arrangement;
Fig. 6.16. Pea plant morphology. The pea has a weak stem and is a tendril climber, with pinnately
compound leaves, and a terminal racemose inflorescence.
Pulses 121
the posterior petal is large, called a ‘vexil- The cortex is the middle region, lying
lum’. Wing petals are present on both sides between the epidermis and the stele. Cortex
of the vexillum, with keel petals present on is well developed and is larger than the
the anterior side. There are ten stamens, stele (Fig. 6.17). It is multiseriate and rela-
which actually arise in two whorls of five tively homogeneous and consists of paren-
each. The stamens of the outer whorl are chyma. The innermost layer of the cortex is
opposite to the sepals and inner whorl of called the endodermis, which is single lay-
stamens is opposite to the petals, hence ered with compactly arranged barrel-
it is called diplostemonous. The gyno- shaped cells. The endodermis is also called
ecium is diadelphous, dithecous, basi- the starch sheet layer, as starch grains are
fixed, introse, unicarpellary and unilocular. present in the endodermal cells.
There are from one to three ovules present The stele is covered by uniseriate paren-
on marginal placentation, simple hairy chymatous pericycle. Stele (vascular strands)
stigma. The fruit is leguminous (pod), slen- consists of xylem and phloem. Xylem conducts
der and green coloured, with from five to water and salts and phloem transports food
ten seeds per fruit, rounded, and green in materials. The xylem is exarch, with proto-
colour. xylem towards the pericycle and metaxylem
towards the centre and scattered secondary
xylem vessels.
6.6.5 Anatomical description
Stem anatomy
Root anatomy
The stem is triangular in transverse sec-
Internal tissues of the root are covered by tion. A transverse section of the stem
uniseriate compactly arranged thin-walled primarily shows epidermis, cortex and
living cells called epidermis. vascular bundles.
Endodermis
Phloem
Pericycle Secondary xylem
Fig. 6.17. Transverse section of root (100×) showing organization of various tissues. The epidermis has root
hairs, and multilayered parenchymatous cortex occupies the major portion of the root. Note: secondary
growth in stele has started with secondary xylem vessels formed towards the centre and phloem towards the
outer side.
122 Chapter 6
The epidermis is uniseriate with a single xylem and phloem (open), hence the vascu-
layer of compactly arranged barrel-shaped lar bundle is described as conjoint, collat-
living cells. The epidermis is covered by a eral and open type. The xylem is endarch
thin cuticle. with protoxylem present towards the cen-
The cortex is multilayered and paren- tre. The xylem consists of many vessels
chymatic. Below the epidermis there are arranged in rows.
four to five layers of collenchyma
restricted to the ridges (Fig. 6.18). The Leaf anatomy
cells are round with prominent inter-
cellular spaces. The endodermis is inner- Epidermis covering the upper surface of the
most layer of the cortex. Due to leaf is called upper epidermis or ventral epi-
the secondary growth, vascular bundles dermis or adaxial epidermis and on the
are spread into the cortex. Pericycle is lower side of the leaf is called lower epider-
the outermost non-vascular region of the mis or dorsal epidermis or abaxial epider-
stele. It is multiseriate and sclerenchy- mis. The epidermis is single layered with
matic. Scleren chyma is mech anical tis- compactly arranged barrel-shaped (tabular)
sue and gives strength and rigidity to the cells. The outer walls of the epidermal cells
stem. are cutinized. Anisocytic stomata are pre-
Xylem and phloem form the vascular sent in both epidermal layers, but the sto-
tissues and are organized into wedge- mata are usually more frequent in lower
shaped vascular bundles. From 10–15 vas- epidermis.
cular bundles are arranged in one ring called The ground tissue of the leaf, present
a eustele. A weak stereome system can be between the upper and the lower epi-
observed. In the vascular bundle xylem and dermal layers, is called mesophyll. The
phloem are present together (conjoint) on mesophyll is not clearly differentiated
the same radius (collateral), having a strip between an upper palisade tissue and
of cambium (fascicular cambium) between lower spongy tissue (Fig. 6.19). Mesophyll
Epidermis Collenchyma
Cortex Cortical
vascular bundles
Phloem
Xylem
Medulla
Fig. 6.18. Transverse section of stem (100×) showing ridges and grooves; four to five layers of collenchyma
are present specially below the ridges. Cortical vascular bundles can be seen (special feature in some
legumes). Secondary growth has just initiated.
Pulses 123
Xylem
Lower epidermis
Phloem
Fig. 6.19. Transverse section of leaf (100×) illustrating the arrangement of various tissues.
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7
Oil Crops
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 125
126 Chapter 7
where it was readily adapted. Selection for sharing the same receptacle. As in other
high oil in Russian began in 1860 and was Asteraceae, it looks superficially like a sin-
largely responsible for increasing oil content gle flower. The sunflower’s ‘seed’ is actually
from 28% to 50%. The high producing lines an achene: a monocarpellate (formed from
from Russia were introduced into the USA one carpel) and indehiscent (they do not
after the Second World War, which rekindled open at maturity) fruit containing a single
interest in the crop. seed that nearly fills the pericarp, but does
Carbonized seeds of domesticated sun- not adhere to it. H. annuus is diploid with
flower (H. annuus var. macrocarpus Ckll.) 2n = 34 chromosomes.
recovered from the Hayes site in middle
Tennessee yielded an accelerator date of 4265 Vegetative characters
+/− 60 BP. This is the earliest date for domesti-
cated sunflower, extending the known age of The plant is a mesophytic herb, with a taproot
this eastern North American domesticate by system with highly branched lateral roots.
1400 years. The process of domestication has The stem is aerial, erect, greenish, rough, and
changed plants from being multi-stemmed hard, pubescent and monopodial in branch-
with a number of flower heads to ones that are ing and grows up to 3 m. Leaves are simple,
tall and unbranched, with a single large head. exstipulate, alternate, pubescent petiolate, ovate
shape, with a serrate margin, acute apex, and
rough, reticulate venation (Fig. 7.1).
7.1.3 Utilization
Floral characters
Every part of the plant may be utilized for
The inflorescence is a terminal, head inflo-
some economic purpose: the leaves can be
rescence or capitulum or anthodium, and the
used as cattle feed; the stems contain a fibre,
which may be used successfully in paper
making; and the flowers contain a yellow
dye.
The whole seed of sunflower is sold as
a snack and can be processed into a deli-
cious peanut butter alternative, sun butter.
It is also sold as food for birds and can be
used directly in cooking and salads. The
seed is rich in oil, which is said to approach
more nearly to olive oil than any other vege-
table oil known and is largely used as a sub-
stitute. Sunflower oil, extracted from the
seeds, is used for cooking, as carrier oil and
to produce biodiesel, for which it is less
expensive than the olive product.
head consists of a highly condensed disc- i.e. the pollen is spread by insects, normally
like peduncle on which numerous small honeybees. The sunflower’s ‘seed’ is actu-
sessile flowers called florets arise. The flo- ally an achene: a monocarpellate (formed
rets are surrounded and protected by involu- from one carpel) and indehiscent (they do
cres of bracts. The florets are of two types not open at maturity) fruit containing a sin-
(ray and disc florets), hence it is called a gle seed that nearly fills the pericarp, but
heterogamous head (Fig. 7.2). Ray florets does not adhere to it.
are present at the periphery of the head, and
are bracteate, sessile, incomplete, unisex-
ual, zygomorphic and epigynous. The disc 7.1.5 Anatomical description
florets are present at the centre of the head,
and are bracteate, sessile, complete, bisex- Root anatomy
ual, actinomorphic and epigynous. Sepals
are modified into a persistent pappus. There A transverse section of matured root shows a
are five gamopetalous petals. In disc florets thin layer of bark at the outer side of the root
the corolla (petals) is regular and tubular. In that is produced due to rupture of the epi-
ray florets the corolla is zygomorphic and dermis. Below the epidermis a two- to five-
legulate because the two posterior petals are layered cortex is present in compressed
reduced. The androecium is absent in the form (Fig. 7.3). Xylem is exarch with con-
ray florets; in disc florets there are four sta- tinuously formed secondary xylem vessels.
mens, epipetalous and syngenesious, the Pith is absent at the centre. Xylem vessels are
filaments are free and anthers connate later- endarch with prominent secondary xylem
ally to form a tube around the style. The vessels present at the centre of the root.
anthers are dithecous, basifixed and hooded.
The gynoecium is present in both ray and
Significance of variations in root anatomy
disc florets, and consists of a bicarpellary,
syncarpous, inferior, unilocular ovary hav- Cultivar differences are expressed in a num-
ing only one ovule with basal placentation. ber of root characteristics: (i) the presence
The style is terminal, simple and long, and or absence of sclerenchymatous exodermis
the stigma is forked and re-curved.
Sunflower is generally a cross-pollinated
crop. If cross pollination does not occur it can Epidermis Cortex
self-pollinate by the use of a safety mechanism.
Pollination occurs through entomophily,
in the cortex and the size of vascular bundle, The cortex is the middle region present
which may be related to drought resistance; between the epidermis and stele, and con-
and (ii) the thickness of endodermal cell tains hypodermis, middle cortex and endo-
walls. dermis. The hypodermis is multilayered
(with two to six layers) and collenchymatic
Stem anatomy (Fig. 7.4). Collenchyma may be in the form
of complete cylinder or in the form of dis-
Stem anatomy of sunflower is typical of crete strands. The hypodermis gives con-
that of the dicots. The mature stem shows siderable strength, flexibility and elasticity
secondary growth. The primary stem shows to the young stem. The endodermis is the
three regions, the epidermis, cortex and innermost layer of the cortex. In the young
stele. stem the innermost layer of the cortex con-
The epidermis is the outermost region tains abundant starch, hence this layer is
surrounding the stem, and is uniseriate; called the starch sheath.
having a single layer of compactly arranged Pericycle is the vascular region of the
barrel-shaped living cells. The outer walls stele. It is multiseriate and sclerenchy-
of the epidermal cells are cutinized. The matic. Sclerenchyma fibres may be in the
epidermis is also covered by a separate layer form of bundle caps. Wedge-shaped vascu-
of cutin called cuticle on its outer surface, lar bundles are arranged in one or two
which reduces transpiration. Stomata are rows. The vascular bundles are described
present in the epidermis for the exchange of as conjoint, collateral and open type. The
gases and transpiration. Multicellular hairs xylem consists of many vessels and is
are present on epidermis. endarch with protoxylem present towards
Endodermis
Trichome
Xylem
Medulla
Fig. 7.4. Transverse section of stem (100×) showing epidermis, multilayered collenchyma, cortex,
endodermis, xylem, phloem and medulla. Note: collenchyma layers are highly compactly arranged and long
trichomes are visible on the epidermis.
Oil Crops 129
the centre. The extensions of parenchyma- The size of the vascular bundles and the
tous pith between vascular bundles are number of vascular bundles reveals the
called primary medullary rays, which help capacity of conduction in sunflower culti-
in lateral conduction and may give rise to vars. It can be observed that the anatomy of
secondary meristem. young stem shows the presence of a thin
cuticle and multicellular trichome; this is
subtended by several layers of collenchyma,
Significance of variations in stem anatomy imparting flexibility of the sunflower stem.
Epidermal study reveals the following vari-
ability between genotypes: Leaf anatomy
• Density of trichomes may vary. The leaf shows a dorsi-ventral nature, i.e. the
• Cuticle thickness of the epidermis var- leaf blade consists of distinct dorsal and ven-
ies among genotypes. tral surfaces. The epidermis is single layered,
• Presence or absence of multicellular having compactly arranged barrel-shaped
trichomes; in some genotypes there (tabular) cells. The cell walls are cutinized
may be unicellular and multicellular and also covered by a thin cuticle. The cuti-
trichomes, which may be related to nized outer walls and cuticle reduce the loss
insect resistance. of water due to transpiration. Anisocytic
• Number of hypodermal layers may type of stomata are present in both the epi-
differ among genotypes, imparting flex- dermal layers, but the stomata are usually
ibility and varying the shape of hypo- more frequent in the lower epidermis; gener-
dermal cells (collenchyma cells). ally the stomatal index of upper epidermis is
• Distribution of collenchyma cells 33 and lower epidermis is 39. Stomata facili-
(Fig. 7.5). tate the exchange of gases between the leaf
• Number of pericycle layers and the and environment. The epidermis contains
shape of sclerenchyma cells differ. two types of trichomes: they are small and
• Sclerenchyma fibres may be in the unicellular and are either spine or hook
form of bundle caps in some genotypes, shaped. The mesophyll is chlorenchymatic
offering strength. and differentiated into an upper palisade
Collenchyma
Vascular bundle
Fig. 7.5. Transverse section of petiole (100×) showing the variation in collenchyma layers and size of the
vascular bundles: (a) five to six layers of collenchyma and medium size bundles and (b) two to three layers
and large size bundles.
130 Chapter 7
tissue and lower spongy tissue (Fig. 7.6). The decreased in relative volume, while inter-
palisade tissue is thick and compact in cellular space increased. There was a pro-
drought-resistant genotypes. Cells of the portional decrease in the amount of contact
spongy tissue are thin-walled, irregular in between mesophyll and vascular tissue,
shape, and arranged loosely with large con- while the surface area of mesophyll cells
tinuous intercellular spaces. Stomata open and vascular tissue in contact with inter-
into large intercellular spaces called sub- cellular space increased (Fagerberg and
stomatal chambers. The cells of spongy paren- Culpepper, 1984).
chyma contain a smaller number of chloroplasts,
hence the lower surface of the leaf is light Cuticle development
green in colour. Collateral and closed vascular
bundles (veins) occur between the palisade Cuticle development depends on moderate
and spongy tissue (median in position). water deficit (MWD). MWD generated from
Studies have been undertaken on the early to late anthesis had an effect on quanti-
development of leaf and cuticle. tative development of the cuticle, and qualita-
tive and quantitative development of the
Anatomical changes during cuticular waxes (CW) of the pericarp in two
leaf development hybrids grown under field conditions. At har-
vest maturity (HM), plants grown under MWD
Leaves of H. annuus L. were divided into showed higher CW content (31 to 47%) and
five parts: palisade, spongy, vascular and thicker cuticles (13%) in both experiments
epidermal tissue, and intercellular space. compared to controls. Epicuticular wax (ECW)
The percentage of whole-leaf volume that crystals showed a granular morphology. They
each compartment occupied and the relative determined a reduction of CW of 29% (mg
surface area of compartment cells in contact CW/g pericarp) during the development of
with intercellular space and adjacent cells the fruit, from reproductive stage (stage
for plants grown under low light were meas- when ray flowers have lost their turgidity) to
ured. During development all tissue parts HM (stage when water content of the fruit
was 11%). These results show how internal
Cuticle Upper epidermis mechanisms and external variables regulate
Palisade tissue pericarp wax content, and that fruit dehydra-
tion affects both the quantity and quality of
wax formation from the time of fertilization
to maturity (Franchini et al., 2010).
(a) (b)
VB
TR VB
TR
Col Col
Fig. 7.7. Transverse section of leaf (100×) illustrating variation in trichome (TR) density, distribution
of vascular bundles (VB) and collenchyma layers (Col) at the midrib; (a) high density of trichomes,
three narrow vascular bundles and more layers of collenchyma; (b) low density of trichomes,
smaller amount of collenchyma and five vascular bundles (widely spread).
Trichome
Xylem
Phloem
Vascular bundle
Fig. 7.8. Transverse section of petiole (100×) showing epidermis, cortex, xylem and medulla. Note the high
density of long and short trichomes covering the epidermis; latex cavities are distributed in the cortex; and
vascular bundles are large with six to eight xylem rows.
six layers) hypodermis of collenchyma cells among them. Vascular bundles are arranged
is found immediately beneath the epider- in a half ring and scattered in ground tissue.
mis and, having chloroplasts, it may carry The wedge-shaped vascular bundles are
out photosynthesis. Just beneath the hypo- of various sizes in the same petiole. In the
dermis ground tissue is found, which con- petiole, xylem is always found towards
sists of thin-walled parenchymatous cells the upper side whereas phloem is towards
having well-defined intercellular spaces the lower side.
132 Chapter 7
(a) (b)
Fig. 7.9. Transverse section of petiole (100×) showing variation in trichome density among the different
genotypes: (a) high trichome density; and (b) low trichome density.
(a) (b)
Collenchyma
Collenchyma
Vascular bundle
Vascular bundle
Fig. 7.10. Transverse section of petiole (100×) showing the variation between collenchyma layers and size
of the vascular bundles: (a) one to two layers of collenchyma and medium size bundles; and (b) three to four
layers and large size bundles.
Oil Crops 133
The density of trichomes has been impli- applied Si alleviates drought stress in
cated as a factor in this resistance. The sunflower cultivars by preventing mem-
important role of trichomes in the deter- brane damage (Gunes et al., 2008).
rence of feeding by Cylindrocopturus The salt-tolerant line was superior to
adspersus LeConte, a sunflower stem wee- the salt-sensitive one in growth perform-
vil (SSW), was examined. Trichomes were ance under drought conditions. Stomatal
randomly associated with feeding activ- conductance, transpiration and gas ex-
ity. The morphology and feeding behav- change in the salt-tolerant line were not
iour of this insect enables it to avoid affected by water deficit, whereas they were
sunflower trichome defences. The results affected in the salt sensitive line. The salt
indicate that selective sunflower breeding tolerant line had significantly greater val-
for increased trichome density would have ues for all these variables than the salt sen-
minimal benefit for increased resistance to sitive line. The lines did not differ in
C. adspersus feeding and oviposition water-use efficiency. From this study it is
(Barker, 1990). clear that salt tolerance and drought toler-
A great variability occurs among sun- ance of the two lines examined share
flower genotypes in trichome density (Maiti, osmotic effect (Ashraf and O’Leary, 1996).
unpublished). This reveals that there is a
great necessity to screen and identify sun-
flower genotypes with greater trichome Seed coat anatomy
density and utilize these in breeding for
insect and drought resistance. Integuments of fertilized ovule become the
seed coat. The seed coat covers the embryo
Drought and salt tolerance and provides protection. The seed coat
consists of two layers, the testa and the
The identification of sunflower genotypes tegman.
exhibiting tolerance to moisture stress as The testa is the outermost layer of the
well as those showing least percentage seed coat and is formed from the outer
reduction in their growth and yield attri- integument of the ovule. Generally it is
butes under stress is necessary for the hard and impermeable to water and some-
improvement of productivity under rainfed times shows ornamentations. The testa
environment. Several seedling, physiologi- contains macro- and microsclereids. Macro-
cal, morphological and biochemical traits sclereids are arranged very compactly and
have been shown to play a significant role are dumb-bell/boat/bone-shaped (osteo-
in crop adaptation to water stress among sclereids). Microsclereids are small in size
which cuticle thickness, compact palisade with a square shape and dentate edges.
cells, and a stout petiole with thick collen- The hardness of the seed coat depends on
chyma layers are important (Geetha et al., the thickness of the testa and the arrange-
2011; Maiti, unpublished). ment of sclereids. The rate of water absorp-
Application of silica prevents mem- tion (imbibition) also depends on the
brane damage (membrane permeability) thickness of testa. Very hard and thick testa
and increases leaf relative water content. causes seed dormancy in some plants.
Catalase activity was significantly decrea- The tegman is the inner layer of the seed
sed by drought stress, but supplemental coat and is formed from the outer integument.
Si increased it. In general, superoxide It is thin and soft and covers the embryo.
dismutase and ascorbate peroxidase The macro- and microsclereids are not
activities of the cultivars were increased very compactly arranged; therefore water
by drought and decreased by application enters easily into the seed and on imbibi-
of Si. Under drought stress the non- tion it ruptures easily. As a result of this the
enzymatic antioxidant activity of the culti- time of imbibition is less, therefore germi-
vars was significantly increased by Si. nation time is about 36 h at 25–30°C tem-
Based on this, it can be concluded that perature in laboratory conditions.
134 Chapter 7
taproot system with lateral branches con- three seeds per fruit. The seeds are rounded
taining root nodules formed by a symbi- and usually pink in colour.
otic association of Rhizobium bacteria,
which fixes atmospheric nitrogen by mean
of biological nitrogen fixation – nitrogenase 7.2.5 Anatomical description
enzyme. The stem is weak, aerial, erect,
greenish, rounded and pubescent. The leaves Root anatomy
are compound (Fig. 7.11), exstipulate and
alternate, with two to three pairs of leaflets The epidermis forms the outermost region
on a rachis. Each leaflet is ovate shape, of the root and is uniseriate; having a single
with an acute apex and unicostate reticulate layer of compactly arranged thin-walled liv-
venation. ing cells. Epidermal cells produce unicellu-
lar root hairs, which are useful in increasing
Floral characters the surface area for absorption of water and
helps in obtaining water from soil. The root
Flowers are born on a solitary, axillary inflo- epidermis is also called the rhizodermis,
rescence, pedicillate, stipulate, complete, epiblema and siliferous layer
bisexual, zygomorphic, pentamerous and The cortex is multiseriate and contains
perigynous. There are five sepals, green, relatively homogeneous parenchymatous
free and pubescent. The corolla consist of tissue between epidermis and stele. The
five yellow petals, free, in a papilionaceous symbiotic association with Rhizobium
arrangement, and the posterior petal is large, bacteroids occurs in the cortex region. The
called a vexillum. Wing petals are present endodermis forms the innermost layer of
on both sides of the vexillum; keel petals the cortex and has uniseriate barrel-shaped
are present on the anterior side. The androe- cells.
cium consists of ten stamens in a monadel- The pericycle is the outermost region of
phous condition, dimorphic, dithecous, the stele, and is uniseriate and parenchyma-
introse, and out of the ten stamens two are tous. Cells of the pericycle retain meristem-
staminode. The gynoecium consists of a atic activity. Lateral roots arise endogenously
unicarpellary, unilocular ovary with one to from the pericycle. The lateral roots help in
three ovules present on marginal placenta- absorption of water.
tion. The stigma is simple with hairy. The Xylem and phloem are arranged in the
leguminous fruit usually contain one to form of separate strands on different radii.
They alternate with each other.
Bundle cap length and number of xylem
vessels per vascular bundle showed a signi-
ficant positive correlation with peg strength.
Analysis revealed that bundle cap length
had a direct positive effect on peg strength
whereas number of xylem vessels per vascu-
lar bundle played only a minor role. Thus
bundle cap length emerged as a reliable ana-
tomical character, which can serve as a
selection criterion in breeding cultivars with
strong pegs to reduce harvesting losses in
groundnut (Tiwari et al., 1988).
Due to the proliferating cell divisions
derived from the pericycle (incomplete sec-
ond state of endodermal development), dis-
Fig. 7.11. Groundnut plant morphology showing tinct types of lateral roots can be recognized
compound alternate leaves and solitary, axillary in groundnut: long, first-order lateral roots,
inflorescence. forming the skeleton of the root system, and
136 Chapter 7
thin and short second- and higher-order lat- xylem towards the centre of the axis and
eral roots, ‘feeder roots’. The formation of phloem towards the periphery. In the vascu-
root nodules at the base of the lateral roots lar bundle xylem and phloem are present
was the result of proliferating cell divisions together (conjoint) on the same radius (col-
derived originally from the pericycle (Tajima lateral), with a strip of cambium (fascicular
et al., 2008). cambium) between the xylem and phloem
(open), hence the vascular bundle is descri-
Stem anatomy bed as conjoint, collateral and open type. The
xylem consists of many vessels and is endarch
The outline of the stem in transverse section with protoxylem present towards the centre.
shows ridges and grooves. A transverse sec- The presence of the thick cuticle and
tion of primary stem shows three regions, the collenchyma imparts strength and flexibility
epidermis, the cortex and the stele (Fig. 7.12). to the stem.
The epidermis is the outermost region
surrounding the stem. It is uniseriate, having Leaf anatomy
a single layer of compactly arranged barrel-
shaped living cells. The outer walls of the epi- The leaf shows a dorsi-ventral nature, i.e.
dermal cells are cutinized, and the epidermis the leaf blade consists of distinct dorsal and
is covered by cuticle on its outer surface. ventral surfaces. The epidermis is mainly
The cortex is smaller than the stele. The meant for protection and is present on both
hypodermis is multilayered (two to three) surfaces of the leaf blade. The epidermis is
and collenchymatic and gives considerable single layered with compactly arranged
strength, flexibility and elasticity to young barrel-shaped (tabular) cells. A thin cuticle
stems. The endodermis is innermost layer layer covers the epidermis and it reduces
of the cortex and consists of uniseriate com- the loss of water due to transpiration.
pactly arranged barrel-shaped cells. Anisocytic stomata are present in both epi-
There are from ten to 15 wedge-shaped dermal layers. The stomatal index of the
vascular bundles arranged in one ring called upper epidermis is 26 and of the lower
a eustele. Each vascular bundle consists of epidermis is 32 under 40× magnification.
(a) (b)
Epidermis
Collenchyma
Cortex
Vascular bundle
Endodermis
Phloem
Cambium
Xylem
Medulla
Vascular bundle
Fig. 7.12. (a) Transverse section of stem (40×) showing ridges and grooves in the outline (hexagonal),
vascular bundles arranged in the form of a ring (eustele) and the central hollow region. (b) Sector enlarged
illustrating organization of various parts. The vascular bundles are wedge shaped and the cambial ring
is formed by the union of inter- and intra-fascicular cambium (secondary growth has just initiated).
Oil Crops 137
The ground tissue of the leaf, present bet- water storage cells, cell sizes, and number
ween the upper and the lower epidermal of mesophyll cells per unit leaf surface
layers, is called mesophyll. Long palisade varied significantly between them (Ferreyra
tissue is present below the upper epider- et al., 2000).
mis, and is compactly arranged (Fig. 7.13). Few research activities have been under-
Palisade tissue is the most highly special- taken on the contribution of anatomical char-
ized type of photosynthetic tissue. The acters to insect and disease resistance, but
upper surface of the leaf is dark green in literature is rare on drought resistance.
colour due to palisade tissue. The lower
part of the mesophyll present towards the ROLE OF TRICHOME IN INSECT RESISTANCE The role
lower epidermis is the spongy tissue. Cells of long trichomes was studied in ground-
of the spongy tissue are thin walled, irre- nut. Groundnut (A. hypogaea L.) cultivars
gular in shape, and arranged loosely with of different susceptibility to the jassid,
large continuous intercellular spaces. Vascu- Empoasca kerri Pruthi, were studied to deter-
lar bundles (veins) occur between the pali- mine the inheritance of trichomes on the
sade and spongy tissue (median in position), adaxial surface of the leaf, leaf midrib and
and are collateral and closed. In the vascu- petiole, and their association with resistance
lar bundle, xylem is present towards the to E. kerri. Genotypes or crosses with long
upper epidermis and phloem towards the trichomes on the leaves and petioles showed
lower epidermis. a high level of resistance to jassids (leafhop-
The presence of the thick cuticle and pers) as evidenced by a very low percentage
compactly arranged palisade tissue reduce of yellowed foliage (hopper burn). The pres-
transpiration loss thereby imparting drought ence of long trichomes increases resistance
resistance, but there exists variability among to leafhoppers (Dwivedi et al., 1986).
genotypes.
In an anatomical study, two cultivars DISEASE RESISTANCELeaves from resistant and
showed different characteristics: stomatal susceptible groundnut genotypes (Arachis
density, leaf thickness, relative volume of hypogaea L.) were observed after inoculation
Cuticle
Upper epidermis
Palisade tissue
Spongy tissue
Collenchyma
Xylem Phloem Lower epidermis
Fig. 7.13. Transverse section of leaf (100×) showing the upper and lower epidermis and vascular bundle
at the centre (vein). Note: the thick cuticle on the epidermis and the long and very compactly arranged
palisade tissue helps in reducing the transpirational loss of water thereby offering drought resistance.
138 Chapter 7
with Cercospora arachidicola Hori and Cer- anatomical traits of stems, pegs, roots, or juve-
cosporidium personatum (Berk. and Curt.) nile plant leaflets and field pod-rot reaction
Deighton. Both the pathogens induced almost was not consistent among all genotypes.
similar anatomical responses in the inocu- Lignin distribution in pods, and an
lated leaves. The epidermal and mesophyll index representing µm palisade cells/mm of
cells were shrunken and there was more dam- leaf blade individually or in combination,
age to cell walls. This is due to depletion of might be used effectively to supplement field
polysaccharides, proteins, ascorbic acid and evaluations in screening breeding lines for
nucleic acids from the diseased host tissue at pod disease reaction (Godoy et al., 1985).
the site of contact with the pathogen. It was
not observed in healthy tissues of these gen- Petiole anatomy
otypes (Kaur and Dhellon, 1988).
Pod rotting-resistant cultivars (PEANUT) The petiole is covered by thick cuticle. Two-
showed the following anatomical charac- layered hypodermis of angular collenchyma
ters: the palisade mesophyll cells of 50-day cells is found immediately beneath the epi-
old plants were arranged more compactly in dermis. The hypodermis gives considerable
pod rot-resistant than in susceptible geno- strength, flexibility and elasticity to young
types. An index representing total width (µm) petioles and contains chloroplasts for per-
of palisade cells/mm leaf blade was more dis- forming photosynthesis. Just beneath the
criminative in distinguishing among geno- hypodermis ground tissue is found. It consists
types than average of either cell width or of thin-walled parenchymatous cells with
cell number alone. The distribution of lignin well defined intercellular spaces among them.
in groundnut shells was correlated with Wedge-shaped vascular bundles are of vari-
pod rot resistance. Resistant lines possessed ous sizes in the same petiole. In the petiole,
lignified cell walls in the epicarp and scler- the xylem is always found towards the
enchymatous mesocarp than in the suscep- upper side whereas phloem is found towards
tible genotypes. The relationship between the lower side (Fig. 7.14).
Collenchyma Epidermis
Groove
Xylem
Cambium
Ground tissue
Vascular
bundle
Phloem
Fig. 7.14. Transverse section of petiole (100×) showing epidermis, cortex, xylem and medulla. The upper
side of the petiole has a groove.
Oil Crops 139
dithecous and introse. The gynoecium is are usually round or oval, colourless and
bicarpellary, syncarpous, with many ovules store starch. The epidermis is single layered
present on parietal placentation due to a with compactly arranged barrel-shaped
false septum ovary having two chambers. cells. Endodermal cells are characterized by
The fruit is a siliqua, 1.25–6.25 cm in length. the presence of Casparian strips. Pericycle
Seeds are numerous, small, rounded, brown forms the outermost region of the stele and
in colour and arranged in rows. The crop is is uniseriate and parenchymatous. Cells of
of economic importance for the mustard oil the pericycle retain meristematic activity.
and rapeseed oil that can be extracted from The vascular strands are radial; the xylem is
the seeds of brassicas. Mustard seeds are exarch, of the closed type.
used as a condiment.
Stem anatomy
Epidermis Cortex
Secondary Phloem
xylem Xylem
Cambium
Medulla
Medullary ray
Fig. 7.17. Transverse section of stem (100×)
showing epidermis, three to four layers of
Fig. 7.16. Transverse section of root (100×) collenchyma and a smaller amount of cortex.
illustrating various features. The epidermis and Vascular bundles are arranged in the form of a ring.
endodermis have been ruptured due to secondary A large volume of medulla occupies the central
growth in the stele. The cortex is also compressed. portion of the stem.
Oil Crops 141
collateral, open and endarch; 15–20 vascu- (Fig. 7.19). Palisade cells are compactly
lar bundles are arranged in one ring called arranged and occupy more area when com-
eustele. The central part of the stem is occu- pared to spongy tissue, which is absent
pied by parenchymatous pith or medulla. below the veins. Vascular bundles occur
The cells are round or oval with intercellu- between the palisade and spongy tissue
lar spaces. Medulla stores food materials (median in position), and are collateral and
and aids in lateral conduction. closed. In the vascular bundle, xylem is
The presence of the medium-thick col- present towards the upper epidermis and
lenchyma layer contributes to the flexibility phloem towards the lower epidermis.
to the stem, offering resistance to breakage. The presence of a thin cuticle and loose
palisade associated with large stomata on
Leaf anatomy the leaf surface imparts susceptibility to
drought.
The epidermis is single layered with com-
pactly arranged barrel-shaped (tabular) cells Petiole anatomy
on both sides of the leaf. The outer walls of
the epidermal cells are cutinized. Anisocytic Epidermis covers the petiole and is the
stomata are present in both of the epidermal outer layer. It is uniseriate with a single
layers. The stomatal complex consists of layer of compactly arranged barrel-shaped
three subsidiary cells covering two kidney- living cells. Two layers of collenchyma are
shaped guard cells (Fig. 7.18). the stomatal present below the epidermis. Collenchyma
index of upper epidermis is 28 and lower is living mechanical tissue consisting of
epidermis is 31. The presence of epicuticu- chloroplasts (chlorenchyma) below the cor-
lar wax on both sides of the epidermis is an ners of the collenchyma cells. Just beneath
important character. The mesophyll is chlo- the hypodermis ground tissue is found
renchymatic and concerned mainly with (Fig. 7.20), consisting of thin-walled paren-
photosynthesis. The mesophyll is differen- chymatous cells with well-defined intercel-
tiated into palisade tissue and spongy tissue lular spaces among them. There are three to
four wedge-shaped vascular bundles arran-
ged in a ‘C’ shape in the ground tissue, and
these bundles can be various sizes in the
Stoma same petiole.
The presence of the thin cuticle and
medium-thin collenchyma contributes to
the fragile nature of the petiole.
Conclusion
EC
7.4 Sesame
GC
7.4.1 Introduction
Fig. 7.18. Leaf epidermal impression (400×)
showing stomatal complex. Two kidney-shaped
guard cells (GC) are covering stoma (pore), and Sesame (Sesamum indicum) is a flowering
three subsidiary cells (SC) are adjacent to the guard plant in the genus Sesamum. Numerous
cells – anisocytic stomata. wild relatives occur in Africa and a smaller
142 Chapter 7
Palisade tissue
Spongy tissue
Xylem
Phloem
Fig. 7.19. Transverse section of leaf (100×) showing palisade and spongy tissue, and a vascular bundle at
the centre of the leaf.
7.4.2 Origin
Xylem
Vascular
bundle Epidermis Despite the fact that the majority of the
Ground tissue Collenchyma wild species of the genus Sesamum are
Phloem
native to sub-Saharan Africa, sesame was
Fig. 7.20. Transverse section of petiole (100×) first domesticated in India. This is revealed
showing epidermis, cortex, xylem and ground by morphological and cytogenetic affinities
tissue. between domesticated sesame and the south
Indian native S. mulayanum Nair., as well
as archaeological evidence that it was culti-
number in India. It is widely naturalized in vated at Harappa in the Indus Valley between
tropical regions around the world and is 2250 and 1750 BC, and a more recent find of
cultivated for its edible seeds, which grow charred sesame seeds in Miri Qalat and Shahi
in pods. The flowers of the sesame seed Tump in the Makran region of Pakistan.
Oil Crops 143
Stem anatomy
Hypodermis Cortex
Epidermis
Secondary
xylem vessel
Medullary ray
Fig. 7.22. Transverse section of root (100×) showing secondary growth. Secondary xylem vessels
are larger in diameter and medullary rays are noticeable. The cortex is compressed due to secondary
growth.
Epidermis
Collenchyma
Cortex
Endodermis
Secondary phloem
Secondary
xylem
Cambium
Primary xylem
Medulla
Fig. 7.23. Transverse section of stem (100×) illustrating various parts. Secondary growth has started,
with secondary xylem larger in size and diameter than primary xylem.
Epidermis
Collenchyma
Ground tissue
Endodermis
Phloem
Xylem
Medulla
Fig. 7.26. Transverse section of petiole (100×) showing various parts. Three to four layers of collenchyma
and multilayered ground tissue can be observed.
Global castor seed production is around if the seed coat is drilled for stringing. Castor
1 Mt/year. Leading producing areas are oil in a processed form, called polyglycerol
India (with over 60% of the global yield), polyricinoleate or PGPR, is currently being
China and Brazil, and it is widely grown as used in chocolate bar manufacture as a less
a crop in Ethiopia. expensive substitute for cocoa butter.
sometimes with a reddish tinge, as they brownish mottling. Castor seeds have a
mature (Fig. 7.27). The leaves of some other warty appendage called the caruncle, which
varieties are green practically from the start, is a type of elaiosome. The caruncle pro-
whereas in yet others a pigment masks the motes the dispersal of the seed by ants
green colour of all the chlorophyll-bearing (myrmecochory).
parts, leaves, stems and young fruit, so that
they remain a dramatic purple to reddish-
brown throughout the life of the plant. 7.5.4 Anatomical description
Plants with the dark leaves can be found
growing next to those with green leaves, so Root anatomy
there probably is only a single gene control-
ling the production of the pigment in some Epidermis forms the outermost region of the
varieties at least. The stems (and the spheri- root. It is a uniseriate, having a single layer of
cal, spiny seed capsules) also vary in pigmen- compactly arranged thin-walled living cells.
tation. The fruit capsules of some varieties The epidermis ruptures after secondary
are more attractive than the flowers. growth (Fig. 7.29). Cortex forms the middle
region, lying between the epidermis and the
Floral character stele. It is multiseriate and relatively homo-
geneous and consists of loosely arranged
The flowers are borne in terminal panicle- thin-walled parenchyma cells with promi-
like inflorescences of green or, in some vari- nent intercellular spaces. The cortex rup-
eties, shades of red monoecious flowers tures or is compressed due to secondary
without petals. The male flowers are yel- growth. The endodermis is very distinctive
lowish green with prominent creamy sta- in roots and is single layered with compactly
mens and are carried in ovoid spikes up to arranged barrel-shaped cells. Endodermal
15 cm long; the female flowers, borne at cells are characterized by the presences
the tips of the spikes, have prominent red of Casparian strips or Casparian bands.
stigmas (Fig. 7.28). Endodermis is also called a starch sheet
The fruit is a spiny, greenish (to reddish layer, as starch grains are present in the endo-
purple) capsule containing large, oval, shiny, dermal cells. The pericycle is uniseriate and
bean-like, highly poisonous seeds with variable parenchymatous. Xylem and phloem are
Fig. 7.27. Castor plant morphology showing glossy leaves with long-stalked, alternate and palmate leaves
with 5–12 deep lobes with coarsely toothed segments and panicle-like inflorescence.
Oil Crops 149
arranged in the form of separate strands on pericycle and metaxylem towards the centre
different radii and alternate with each other, and uniformly distributed secondary xylem
hence the vascular strands in the root are vessels. Cambium is absent and hence vascu-
described as separate and radial. The xylem lar bundles are described as closed type. The
is exarch, having protoxylem towards the central part of the root is occupied by a small
amount of parenchymatous pith.
Anatomy of stem
Epidermis
Secondary
xylem
vessel
Secondary
phloem
Medulla
Medullary
ray
Fig. 7.29. Transverse section of mature root (100×) illustrating secondary growth. The epidermis, secondary
phloem, secondary xylem, medullary rays and medulla are clearly visible.
150 Chapter 7
Xylem vessels
(a) (b)
Xylem vessels
Fig. 7.30. Transverse section of mature root (100×) showing variability in size of the secondary xylem
vessels: (a) secondary xylem is smaller in size; (b) secondary xylem is larger in size.
Medulla
Phloem
Cambium
Medulla
Meta-xylem
(a) Protoxylem (b)
Medullary ray
Fig. 7.31. (a) Transverse section of stem showing different parts (100×). (b) Sector enlarged 400×: resin-filled
lysogenous cavities, and metaxylem showing larger vessels.
the epidermis. The presence of two layers of the centre of the axis and phloem towards
chlorenchyma indicates the photosynthetic the periphery. The xylem consists of many
activity of the stem. The endodermis is uni- vessels arranged in rows, and is endarch
seriate, and the compactly arranged layer with protoxylem present towards the cen-
covers the stele. Xylem and phloem form tre. Medulla and medullary rays are distinc-
the vascular tissues and are organized into tive, present at the centre of the stem and
vascular bundles with eight to ten wedge- usually parenchymatous. The cells are round
shaped vascular bundles arranged in one or oval with intercellular spaces. The medulla
ring. Each bundle consists of xylem towards stores food materials.
Oil Crops 151
(a) (b)
Fig. 7.32. Transverse section of stem showing variability in different parts: (a) large xylem vessels and
collenchyma layers; (b) small xylem vessels.
152 Chapter 7
Lower
epidermis
Collenchyma Parenchyma
tissue is present below the upper epider- bundle sheath, the cells of which are termed
mis. Palisade cells are thin walled, cylin- border parenchyma.
drical and contain numerous chloroplasts.
The cells are compactly arranged in one Significance of variations
layer and perpendicular to the upper epi- in leaf anatomy
dermis. Narrow intercellular spaces are
present in the palisade tissue. Palisade tis- The arrangement of palisade tissue is dif-
sue is the most highly specialized type of ferent in different genotypes (Fig. 7.34) as
photosynthetic tissue. The upper surface is the number of stomata, trichome den-
of the leaf is dark green in colour due to sity and types of trichomes. The type and
palisade tissue. Cells of the spongy tissue arrangement of vascular bundles both in
are thin walled, irregular in shape and the lamina and midrib (Fig. 7.35) contains
arranged loosely with large continuous inter- variability. The characteristic size and shape
cellular spaces. Stomata open into large of bundle sheath chloroplasts also varies
intercellular spaces called sub-stomatal cham- among genotypes.
bers. The cells of spongy parenchyma contain Drought-resistant genotypes possess
a smaller number of chloroplasts, hence thick, compactly arranged long palisade cells
the lower surface of the leaf is light green and a thick cuticle, which reduces transpi-
in colour. ration loss compared to the drought suscep-
Vascular bundles (veins) occur between tible varieties.
the palisade and spongy tissue (median in
position). They are collateral and closed. Anatomy of petiole
In the vascular bundle, xylem is present
towards the upper epidermis and phloem A transverse section of petiole shows four
towards the lower epidermis. Vascular bund- regions, the epidermis, hypodermis, ground
les are surrounded by parenchymatous tissue and vascular bundles (Fig. 7.36).
Oil Crops 153
(a) (b)
Palisade
Palisade
tissue
tissue
Fig. 7.34. Transverse section of leaf (400×) showing variability in palisade tissue: (a) short and loosely
arranged palisade cells; (b) long and compactly arranged palisade cells.
P
P
COL COL
Fig. 7.35. Transverse section of leaf (40×) showing variability in collenchyma layers (COL)
and the distribution of xylem (X) and phloem (P) tissue.
(a) (b)
Fig. 7.37. Transverse section of petiole showing variation in collenchyma layers: (a) five layers; (b) seven
layers.
References
Ashraf, M. and O’Leary, J.W. (1996) Effect of drought stress on growth, water relations, and gas exchange of
two lines of sunflower differing in degree of salt tolerance. International Journal of Plant Sciences 157(6),
729–732.
Barker, J.F. (1990) Sunflower trichome defenses avoided by a sunflower stem weevil, Cylindrocopturus
adspersus LeConte (Coleoptera: Curculionidae). Journal of the Kansas Entomological Society 63(4),
638–641.
Blamey, F.P.C., Joyce, D.C., Edwards, D.G. and Asher, C.J. (1986) Role of trichomes in sunflower tolerance to
manganese toxicity. Plant and Soil 91(2), 171–180.
Dwivedi, S.L., Amin, P.W., Rasheedunisa, Nigam, S.N., Nagabhushanam, G.V.S., Rao, V.R. and Gibbon, R.W.
(1986) Genetic analysis of trichome characters associated with resistance to jassid (Empoasca kerri
pruthi) in peanut. Peanut Science 13(1), 15–18.
Fagerberg, W.R. and Culpepper, G. (1984) A morphometric study of anatomical changes during sunflower leaf
development under low light. Botanical Gazette 145(3), 346–350.
Ferreyra, R.A., Pachepsky, L.B., Collino, D. and Acock, B. (2000) Modeling peanut leaf gas exchange for the cali-
bration of crop models for different cultivars. Ecological Modelling 131(2–3), 285–298.
Franchini, M.C., Hernández, L.F. and Lindstrom, L.I. (2010) Cuticle and cuticular wax development in the
sunflower (Helianthus annuus L.) pericarp grown at the field under a moderate water deficit. International
Journal of Experimental Botany 79, 153–161.
Geetha, A., Saidaiah, P., Suresh, J. and Siva Sankar, A. (2011) Morpho-physiological and biochemical basis of
drought tolerance in sunflower – a review. Agricultural Reviews. College of Agriculture, Acharya N.G.
Ranga Agricultural University, Rajendranagar, Hyderabad, India.
Godoy, R., Smith, O.D., Taber, R.A. and Pettit, R.E. (1985) Anatomical traits associated with pod rot resistance
in peanut. Peanut Science 12(2), 77–82.
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Gunes, A., Pilbeam, D.J., Inala, A. and Cobana, S. (2008) Influence of silicon on sunflower cultivars under
drought stress, i: growth, antioxidant mechanisms, and lipid peroxidation. Communications in Soil
Science and Plant Analysis 39, 13–14.
Kaur, J. and Dhellon, M. (1988) Histological and histochemical changes in groundnut leaves inoculated with
Cercospora arachidicola and Cercosporidium personatum. Phytoparasitica 16(4), 327–335.
Kim, K.S., Park, S.H. and Jenks, M.A. (2007) Changes in leaf cuticular waxes of sesame (Sesamum indicum L.)
plants exposed to water deficit. Journal of Plant Physiology 164(9), 1134–1143.
Maiti, R.K. and Wesche-Ebeling, P. (2002) Advances in Peanut Science. Science Publishers, USA,
376 pp.
Maiti, R.K., Singh, V.P., Purohit, S.S. and Vidyasagar, P. (2007) Research Advances in Sunflower (Helianthus
annus L.). Agrobios (International), ISBN 81-904309-2-0, 512 pp.
Tajima, R., Abe, J., New Lee, O., Morita, S. and Lux, A. (2008) Developmental changes in peanut root struc-
ture during root growth and root-structure modification by nodulation. Annals of Botany 101(4),
491–499.
Tiwari, S.P., Murthy, T.G.K., Johnson, G.K. and Varmoda, D.L. (1988) Path-coefficient analysis of anatomical
characters affecting peg strength in groundnut. Euphytica 39(2).
8
Fibre Crops
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
156 (R. Maiti et al.)
Fibre Crops 157
8.1.4 Utilization
helps in absorbing water and minerals. Trans- A high density of vascular tissue (xylem)
verse section of a matured root shows a thin is essential under drought condition for effi-
layer of bark at the outer side of the root, which cient translocation of water to the plant parts.
is produced by rupture of the epidermis. Below Large xylem vessels help in efficient conduc-
this, the two- to five-layered cortex (Fig. 8.2) tion of water and minerals to the shoot sys-
band is compressed. Xylem is exarch with con- tem (Fig. 8.3).
tinuously formed secondary xylem vessels in
mature root and pith is absent at the centre. Root anatomy in applied field (biotic
Endarch xylem vessels with prominent sec- and abiotic stress)
ondary xylem vessels are present at the centre
PATHOGEN BARRIER Catechin (3,5,7,3′,4′-
of the root.
flavanpentol) and gallocatechin (3,5,7,3′,4′,
5′-flavanhexanol) were isolated from methanol
Significance of variations in root anatomy extracts of roots of 1-week-old ‘Acala 4-42’ cot-
Cultivar differences are expressed in a number ton seedlings and identified by infrared spec-
of root characteristics: (i) presence or absence trophotometry. Root chemical composition has
of sclerenchymatous exodermis in the cortex; a direct impact on pathogen penetration. The
and (ii) thickness of endodermal cell walls. localization of the catechins is secreted by the
(a) (b)
Co
MX Co
PrX
En
Fig. 8.2. Transverse section of young root. (a) Ground plan (40×) showing exarch, tetrarch, closed radial vascular
bundles. (b) Sector enlarged (100×) showing endodermis (En), cortex (Co), protoxylem (PrX), and metaxylem (MX).
(a) (b)
B
SP
SX
C
PX
MR
Fig. 8.3. Transverse section of mature root showing secondary growth: (a) medullary rays (MR) and bark (B)
are visible; (b) secondary xylem vessels (SX) are clearly visible in the picture, along with bark (B), secondary
phloem (SP), cambium (C) and secondary xylem (SX).
Fibre Crops 159
(a) ST (b) XP
SXV
PhP MR
Fig. 8.7. Transverse section of stem showing secondary phloem; (a) phloem parenchyma (PhP) and sieve
tubes (ST) can be observed in the picture; (b) xylem parenchyma (XP), secondary xylem vessels (SXV)
and medullary rays (MR).
(a) (b) B
FB
(c) (d)
Fig. 8.8. Transverse section of stem (after secondary growth) showing variations: (a) small size xylem
vessels with uniform distribution; (b) thick bark (B), compressed primary phloem, uniformly spread
small to medium size secondary xylem vessels and fibre bundles (FB, phloem fibres); (c) medium
thick bark and medium size secondary xylem vessels; (d) large size secondary xylem vessels and
compressed primary xylem vessels at the centre. Medulla is prominent at the centre of the stem.
162 Chapter 8
shape of sclerenchyma cells, number of sec- lower epidermis is 33. Stomata facilitate the
ondary xylem vessels and number and size of exchange of gases between the leaf and envi-
vascular bundles. Variability in the density of ronment. The epidermis contains two types
mucilaginous canals in the cortex region is of trichomes: small unicellular spine-like and
also noticed (Fig. 8.9). A large volume of scle- hook-like trichomes.
renchyma gives mechanical support and The mesophyll is chlorenchymatic and
avoids the loss of water from internal paren- differentiated into an upper palisade tissue
chyma tissue by evapo-transpiration. It is and lower spongy tissue. The palisade tis-
expected that medium to large size and a high sue is thick and compact in drought-resistant
number of xylem vessels are required for effi- genotypes, while in susceptible ones these
cient translocation of water under drought are loose leading to greater loss of water by
conditions (Fig. 8.8c). The presence of large transpiration. Cells of the spongy tissue are
vessels associated with dense lignified scler- thin-walled, irregular in shape and arranged
enchyma offers strength to the stem, as well as loosely with large continuous intercellular
drought resistance facilitating efficient trans- spaces. Stomata open into large intercellular
location of water and photosynthates through spaces called sub-stomatal chambers. The cells
phloem to the growing bolls. of spongy parenchyma contain lower num-
bers of chloroplasts, hence the lower surface
Leaf anatomy of the leaf is light green in colour.
Collateral and closed vascular bundles
The leaf shows a dorsi-ventral nature, i.e. the (veins) occur between the palisade and
leaf blade consists of distinct dorsal and ven- spongy tissue (median in position). The
tral surfaces. The epidermis is single layered, vascular bundle is surrounded by paren-
having compactly arranged barrel-shaped chymatous bundle sheath.
(tabular) cells. The cell walls are cutinized
and also covered by a thin cuticle, which Significance of variations in leaf anatomy
reduce the loss of water due to transpiration.
Anisocytic stomata are present on both the Ample genotypic variability is observed for
epidermal layers (Fig. 8.10). The stomata are the leaf anatomical structures under different
usually more frequent in the lower epidermis adaptive environments. Variations in the
(Fig. 8.11). The stomatal density varies among arrangement of palisade tissues (Figs 8.12,
cotton cultivars, but on average the stomatal 8.13), number of stomata, leaf trichome den-
index of the upper epidermis is 25 and the sity (Fig. 8.14), density of gossypol glands
(a) B (b)
MC
Fig. 8.9. (a) Transverse section of stem showing very thick bark, and a smaller number of mucilage cavities (MC)
in the cortex. (b) Transverse section of stem showing medium thick bark, and a greater number of mucilage
cavities in the cortex.
Fibre Crops 163
(Fig. 8.15), type and arrangement of vascular of water from internal parenchyma tissue by
bundles both in the lamina and midrib and evapo-transpiration.
size and shape of bundle sheath chloroplasts The presence of a thick cuticle on the
are found in cotton germplasm. A number of epidermis of the leaf reduces transpirational
mechanisms providing resistance to dehy- loss of water. Thick, compactly arranged
dration are observed in genotypes adapted to palisade tissue also helps reduced stomatal
water-limited environments. Sclerenchyma density to minimize water loss. Some addi-
gives mechanical support and avoids the loss tional mechanisms such as high trichome
density reduce the direct sunlight effect and
minimize leaf temperature, thus finally
reducing transpirational loss. The high den-
sity of trichomes also offers resistance to
sucking insect pests. Insect pest resistance
SP is also enhanced by the presence of a higher
GC number of glands producing gossypol.
There exists a large variation in the com-
SC
pactness of palisade cells among cotton hybrids.
SC Compactly arranged palisade cells in the leaf
contribute to reducing transpiration loss and in
SC drought resistance; on the other hand, loosely
arranged palisade cells leads to a higher rate
of transpiration.
Cotton hybrids show a large variation in
EC the types and intensity of trichomes, which
are related to insect pest tolerance. Three
types of trichome are present on cotton leaf
surface: (i) stellate; (ii) unicellular simple; and
(iii) bifurcate types varying among genotypes
(Fig. 8.16).
Fig. 8.10. Stomatal complex of the leaf epidermis
(400×) showing two kidney-shaped guard cells (GC)
The leaf surface of cotton shows the pres-
covering the stomatal pore (SP), with three unequal ence and intensity of gossypol glands, which
subsidiary cells (SC) surrounding the guard cells. are related to insect tolerance. There exists a
This type is called an anisocytic stomata. Epidermal large variation in dermal morphology among
cells (EC) are irregular in shape and unequal in size. cotton hybrids.
(a) (b)
Stomata
Fig. 8.11. Upper epidermis (a) of the leaf (400×) showing a lower frequency of stomata compared to lower
epidermis (b).
164 Chapter 8
(a) (b)
P
Col T
Fig. 8.12. Transverse section of leaf showing: (a) high density of trichomes (T) on both the epidermis, thick
compactly arranged palisade (P) tissue, thick cuticle, and three to four layers of collenchyma (Col) tissue above
the lower epidermis; and (b) very low density of trichomes on both the epidermis, thin, loosely arranged
palisade tissue, thin cuticle, and two to three layers of collenchyma tissue above the lower epidermis.
(a) (b)
Fig. 8.13 Transverse section of leaf showing portion of palisade tissue: (a) thick, long compactly arranged
and (b) thin, loosely arranged palisade cells.
(c)
Fig. 8.14. Leaf upper epidermal impression showing variability in density of trichomes: (a) high, (b) medium
and (c) low density.
leaf pubescence should be rated using the by the affected cells may contribute to
youngest, fully expanded main-stem leaves. increased sample variability and possibly
Within each field trial, leaf pubescence to errors in osmotic potential (ψ) measure-
ratings varied significantly among cultivars ments (Wullschleger and Oosterhuis, 1987).
and were consistent for cultivars at differ-
ent locations and across years. These data ANATOMICAL CHANGES UNDER INCREASED ULTRAVIOLET
indicate that the rating system has a mor- RADIATION Enhanced UV-B radiation increa-
phological basis and can be used to charac- sed epicuticular wax content on adaxial leaf
terize the leaf pubescence of cotton cultivars surfaces, and stomatal index on both adaxial
(Bourland et al., 2003). and abaxial leaf surfaces. Leaf thickness was
reduced following exposure to UV-B owing
CUTICLE ABRASION IN RELATION TO WATER POTENTIAL to a decrease in thickness of both the palisade
MEASUREMENTS Transverse sections viewed and mesophyll tissue, while the epidermal
with a transmission electron microscope thickness remained unchanged. The reproduc-
indicated substantial direct damage to the tive parameters were reduced at both ambi-
cuticle with large sections of cell wall devoid ent and high UV-B levels. The study shows
of a cuticular layer. Although the exposed that cotton plants are sensitive to UV-B at
cell walls were intact, the lateral cell walls both the whole plant and anatomical level
were physically compressed and distorted (Kakani et al., 2003).
during abrasion. In addition, the cytoplas-
mic and vacuolar membranes of the epider- GOSSYPOL GLANDS The intensity and the vol-
mal cells were also frequently ruptured. ume of the gossypol amount is an important
Evaluation of the damage following abra- part of defence against plant insects and dis-
sion indicated that the release of turgor eases. The number of gossypol glands on the
166 Chapter 8
(c)
Fig. 8.15. Lower epidermis of the leaf (10×) showing variation in density of gossypol glands (Gos): (a) high,
(b) medium and (c) low density.
stem, boll walls (carpels), leaves and seeds Trichome density on leaf surface to jas-
were visually counted and ranged from 0 to 142/ sid tolerance was observed. The existence
cm2 for stem, 0 to 135/cm2 for leaf, 0 to 85/cm2 for of a positive relationship between trichome
boll wall and 0 to 16/mm2 for embryo (kernel). density and jassid tolerance was also con-
The number of gossypol glands varied for the firmed through artificial screening (Kannan
genotypes and years but this variance was too et al., 2006).
low in some genotypes. Measured parameters Trichome density on lower leaf surface is
were positively and significantly correlated found to be related to sucking pest resistance
to each other. Correlations for number of gos- (Vibha Seeds).
sypol glands between stem and leaf (r = 0.77**), The intensity of gossypol glands influ-
boll wall (r = 0.65**) and seed (r = 0.32**) were ences insect and disease resistance.
significant (Bolek et al., 2010).
Petiole anatomy
THE ROLE OF TRICHOME INTENSITY ON INSECT RESIS-
TANCE This study revealed that pubescence There is a close similarity between petiole
provides a mechanism of resistance to move- and stem with regard to the structure of epi-
ment of newly hatched larvae. The study obser- dermis. The ground parenchyma of petiole
ved movements of newly hatched tobacco is similar to that of the stem cortex in the
budworm larvae, Heliothis virescens (F.), on arrangement of cells and in the number of
upper and lower leaf surfaces and petioles of chloroplasts. The supporting tissue is col-
four cotton (G. hirsutum L.) strains (Ramalho lenchyma. In relation to the arrangement of
et al., 1984). vascular tissues in the stem, the vascular
Fibre Crops 167
(a) (b)
Sim
St
(c) (d)
SBif
LBif
Fig. 8.16. Transverse section of petiole (portion of epidermis) showing different types of trichomes:
(a) stellate (St), (b) simple unicellular (Sim), (c) short bifurcated (SBif) and (d) long bifurcated type (LBif).
bundles of the petiole are collateral. The epi- an important role in hormonal regulation of
dermis contains unicellular trichomes. Multi- leaf abscission. Cotton petioles with a higher
layered (two to six layers) hypodermis of number of vascular bundles will thus help
collenchyma cells is found immediately increasing the longevity of leaves, thus incre-
beneath the epidermis and, having chloro- asing photosynthesis and biomass. Drought-
plasts, it may carry out photosynthesis. Just resistant cotton cultivars possess a thick
beneath the hypodermis ground tissue is cuticle (Fig. 8.17), high density of trichomes
found. This consists of thin-walled parenchy- (Fig. 8.18), high number of collenchyma layers
matous cells having well defined intercellular (Fig. 8.19) and larger sized vascular bundles,
spaces among them. Vascular bundles are thereby giving flexibility and strength to the
arranged in a half ring and scattered in the petiole. Drought-resistant cultivars also have
ground tissue. The wedge-shaped vascular a long and stout petiole. The petiole is also
bundles are of various sizes in the same pet- commonly used for nitrogen status analysis in
iole. In the petiole, xylem is always found cotton, as the nitrate taken up by root moves to
towards the upper side whereas phloem is the leaves through the petiole. Hence, the
found towards the lower side. higher number of vascular bundles and pres-
ence of thick collenchyma for stiffness and
Significant variation on petiole anatomy flexibility of the petiole will be helpful for
efficient translocation of photosynthates, par-
Plant water deficits induce both leaf and boll ticularly under water stress conditions. The
abscission from cotton under field conditions. presence of cuticle thickness subtended by
Petiole, being near the abscission zone, plays several layers of collenchyma contributes to
168 Chapter 8
Cuticle
(c)
Fig. 8.17. Transverse section of petiole showing variation in cuticle thickness: (a) thin layer of cuticle (Col)
on the epidermis; (b) moderately thick; and (c) very thick layer of cuticle.
Fig. 8.18. (a) Transverse section of petiole showing high density of trichome and medium size vascular bundles
(CVB) and small amount of centre ground tissue. (b) Low density of trichomes and high amount of centre ground tissue.
drought resistance. It has been observed that omy with special reference to the intensity of
the variation in leaf epidermal and leaf ana- trichomes and gossypol glands contributes
tomical traits may be used in specific charac- greatly to the tolerance or susceptibility of
teristics of a cotton cultivar. cotton cultivars to some insect resistance.
There exists a large variability in the intensity
Research trends of the leaf anatomical components, thereby
giving enormous opportunity in the selection
In the context of the above mentioned litera- of genotypes for tolerance to jassids and suck-
ture it can be assessed that leaf surface anat- ing pests. On the other hand, the cuticular
Fibre Crops 169
(a) (b)
Col Col
Fig. 8.19. Transverse section of petiole showing variability in collenchyma and distribution of vascular bundles:
(a) four to five layers of collenchyma tissue (Col) and irregularly distributed xylem vessels in the vascular bundle;
(b) six to seven layers of collenchyma tissue, uniformly distributed xylem vessels in vascular bundle.
Pedicel anatomy
Anther
Filament
Style
Filament
Ovary
Ovule
Axis
Thalamus
Pedicle
Fig. 8.21. (a) Longitudinal sections of flowers at different stages of development. (b) Flower showing
arrangement of floral parts on thalamus. Floral parts (sepals, petals and stamens) develop below the level
of the gynoecium (hypogynous flower).
Androecium
there is endothecium, which usually pos-
The anthers contain two lobes (Fig. 8.22), and sesses strips or ridges of secondary wall
are found to be situated on a slender fila- material mainly on those walls that do not
ment that bears a single vascular bundle. remain in contact with the epidermis. The
The structure of the filament is quite simple. innermost layer is composed of multinucle-
The vascular bundle is amphicribral and ate cells; this is nutritive in function and
remains surrounded by parenchyma. The known as tapetum. The wall layers that are
epidermis is cutinized and bears trichomes. located between the endothecium and tape-
Stomata may also be present on the epider- tum are often destroyed during the develop-
mis of both anther and filament. The vascular ment of pollen sacs. On the maturation of
bundle is found throughout the filament and the pollen the tapetum disintegrates and the
culminates blindly in the connective tissue outer wall of the pollen sac now consists of
situated in between the two anther-lobes. only the epidermis and endothecium. At the
The outermost wall layer of the anther time of dehiscence of the anthers the pollen
is the epidermis. Just beneath the epidermis is released out through the stomium.
Fibre Crops 171
L
H
AX
SEP
OV
(a) AX (b) F P
S
AX
Fig. 8.25. (a) Longitudinal sections of boll (fruit) at different stages of development, showing axis (AX)
and developing seeds. (b) Fully developed boll showing pericarp (P), axis (AX), seeds (S) and fibres (F).
(a) (b)
(c) (d)
Fig. 8.26. Developmental stages of fibre cell (1000×): (a) initiation, (b) elongation, (c) secondary thickening
and (d) maturation. The arrow points to the specific fibre cell being described.
exhibit tip synthesis do not exhibit increases and the water content decreases drastically
in cell diameter (Steer and Steer, 1989). (Fig. 8.26d). Increases in fibre dry weight are
the result of increases in the synthesis of cel-
EXPANSION Cell expansion is also regulated lulose (Meinert and Delmer, 1977). The cell
by the extensibility of the cell wall. Regardless wall that develops during this phase has been
of whether cell expansion occurs via tip syn- termed secondary cell wall and is composed
thesis or diffuse growth, the wall in the region almost exclusively of high DP (degree of poly-
of expansion must yield to turgor pressure if merization) cellulose (Marx-Figini, 1982).
the cell is to increase in size (Fig. 8.26c). For Unlike secondary cell walls of other plant cells,
this reason, cell expansion most commonly the cotton fibre secondary cell wall contains
occurs in cells that have only a primary cell little non-cellulosic component and no lignin.
wall (Cosgrove, 1997). Primary cell walls con- The secondary cell wall must be plastic
tain low levels of cellulose. Production of the enough to allow for the observed increases in
more rigid secondary cell wall usually signals fibre diameter. As lignin is one of the wall
the cessation of cell expansion. Secondary cell components thought to strengthen and make
wall formation is often indicated by the devel- the wall rigid, the lack of lignin is consistent
opment of wall birefringence. Fibre diameter with the observed increase in fibre length and
significantly increases as fibres grow and diameter (Fig. 8.26).
develop secondary cell walls. Fibre cells show There exists a large variability in mor-
increases in diameter; however, the specific phology of fibre cells, in length, breadth and
rates of change differed; fibres continue to wall thickness, which may be related to fibre
increase in diameter during the secondary wall quality.
synthesis stage of development, indicating that
the synthesis of secondary cell wall does not
Significance of variations in fibre anatomy
coincide with the cessation of cell expansion.
Cotton cultivars shows variation in fibre char-
MATURATION During the maturation stage the acters, which includes fibre length (staple
fibres are thickened, i.e. diameter is increased length), diameter, wall thickness (Fig. 8.27),
174 Chapter 8
(a) (b)
CW
BR
(c)
Fig. 8.27. Variability in fibre cell morphology in relation to wall thickness and breadth. (a) Coarse fibre
showing high cell wall (CW) thickness and high breadth (BR). (b) Medium-fine fibre showing medium
cell wall thickness and breadth. (c) Fine fibre showing less cell wall thickness and less breadth.
length/breadth (L/B) ratio and percentage of before the flower opens, and if fertilization
uniformity. These characters determine the occurs, continuing up to the 28th day after
quality of the fibre. flowering. The fibres at the base of the seed
It is generally observed that genotypes often develop first, and are longer than those
having highest fibre length, less fibre cell that arise at the tip of the seed. The elonga-
diameter, less wall thickness and high L/B tion of the fibre occurs during 25–30 days
ratio give finer and stronger fibre. The rate of after flowering. Towards the end of this
fibre cell elongation is an important parameter period there is a deposition of the secondary
in determining the longest fibre, with the higher wall, which may continue up to the 78th day
rate of fibre elongation giving fine quality fibre. after flowering. The growth in length and the
Fine and long fibres have the higher rate of cell deposition of the secondary wall depend on
elongation per day. Fibre with the highest uni- the variety of the cotton plant and the envi-
formity gives good spinning quality. Fibres ronmental conditions (Flint, 1950).
having higher fibre cell diameter and high
wall thickness produce coarse fibres. Seed anatomy
In the above parameters, the rate of fibre
cell elongation is very important for obtain- The cotton seed is composed of embryo
ing long staple fibres, which have commer- (Fig. 8.28), endosperm, perisperm, inner
cial demand. pigment layer, palisade (Malpighian) layer,
Differentiation of lint and fuzz is believed colourless layer, outer pigment layer and epi-
to be a continuous process, beginning just dermis, including lint hairs; traces of starch,
Fibre Crops 175
in addition to oil and protein, occasionally endosperm and embryo. Epidermal outgrowths
occur in the cells of both young and mature are in the form of cellulosic fibres, called lint
embryos. The palisade layer is a part of the (Reeves and Valle, 1932).
inner rather than of the outer integument. Integuments of fertilized ovule become
The ovules are campylotropous. Endosperm is the seed coat, which covers the embryo and
present and usually is fairly abundant. The gives it protection. The seed coat consists of
embryo and endosperm contain an abundance two layers, the testa and tegman. Testa is the
of oil and protein, but very little starch at matu- outermost layer of the seed coat and is for-
rity. In developing ovules, however, starch is med from the outer integument of the ovule.
commonly found in the integuments, nucellus, Generally it is hard and impermeable to water
and sometimes contains ornamentations.
The testa contains macro- and microsclereids
(Fig. 8.29). Macrosclereids are arranged very
compactly. Macrosclereids are dumb-bell/boat/
bone shaped (osteosclereids). Microsclereids
are small in size with a square shape and den-
tate edges (Fig. 8.29). The hardness of the
seed coat depends on the thickness of testa
and the arrangement of sclereids. The rate of
absorption of water (imbibition) depends on
the thickness of testa. Two types of fibres are
produced from epidermal layers: short fibres
called ‘fuzz’ and longer fibres known as ‘lint’.
The tegman is the inner layer of the seed
coat and is formed from the outer integument.
SC It is thin and soft and covers the embryo. This
thin seed coat and the fact that the macro-
and microsclereids are not very compactly
COT
Gos arranged, means that water enters easily into
the seed and on imbibition it ruptures easily.
Fig. 8.28. Transverse section of cotton seed The time of imbibition is less so germination
showing seed coat (SC), cotyledons (COT) time is about 36–40 h at 25–30°C under labo-
and gossypol glands (Gos). ratory conditions. The outer integument is
Testa
Microsclereids
Tegman
Fig. 8.29. (a) Transverse section of seed coat illustrating testa and tegman (100×); (b) the testa shows macro-
and microsclereids (400×).
176 Chapter 8
thin, usually being only two or three cells in Fibre crops have played a very important
thickness. The inner integument is much role in the world since primitive times to
thicker, but is partially reabsorbed during meet daily needs, and in modern civilization
development of the embryo. to meet industrial needs. The studies on fibre
In seed of tetraploid cotton (G. hirsutum crops deal with distribution, origin, botany,
and G. barbadense), approximately 30% of ecological conditions, agronomy, harvesting
the seed coat epidermal cells develop into method, quality and processing of different
cellulose-enriched fibres, while the embryos species of fibres. In addition, the different
synthesize oils and proteins. Hence, both the aspects of fibre plants grown in wild condi-
maternal and filial tissues of the cotton seed tions, the methods of extraction, processing
are of significant economic value. After initia- and uses of different species vary widely.
tion from the ovule epidermis at or just before This book provides a critical analysis of pros-
anthesis, the single-celled fibres elongate to pects and problems of fibre crops. It also dis-
2.5–6.0 cm long in the tetraploid species cusses the technological aspects and market
before they switch to intensive secondary cell potentials of several fibres.
wall cellulose synthesis. A number of candidate Although fibre science does not attract
genes and cellular processes that potentially the special interest of many scientists, it
regulate various aspects of fibre development deserves special attention for the mysteries
have been identified (Ruan, 2005). of its origin, its structure, its productivity
The presence of a thick cuticle and thick and its great economic importance in mod-
macrosclereids delays the time of initiation ern civilization.
of germination in cotton seeds.
The formation of a meshy network is the among crop species. It has been reported that
main characteristics of these bast fibres. The longer fibre cells contribute to greater stren-
ultimate fibre cells derived from fusiform ini- gth. In this respect C. olitorius (tossa jute) pro-
tials, which form the building skeleton of the duces good quality fibre for a desirable fibre
fibre filament, are pointed at the apex and bundle structure compared to C. capsularis
have a lumen, the morphology of which var- (white jute), which has an undesirable fibre
ies among fibre crop species. The general bundle structure.
characteristics of fibre bundle structure and In C. olitorius there exists large variations
ultimate fibre cells are depicted in the follow- in fibre bundle intensity and fibre bundle struc-
ing figures. A large variation is observed in ture, thereby offering scope for selection of
fibre bundle structure among crop species and varieties with high yield potential and high
among varieties of the same species with res- fibre quality. There is enough scope for
pect to shape of fibre wedge, number of layers, selection of genotype for better qualities,
number of fibre bundles and their structures. such as fineness and bundle with uniform
The fibre wedge is in general tapering to surface, and use them for genetic improve-
blunt with gradual reduction of fibre bundles ment of quality. The same is true in the case of
towards the tip. The interconnection of fibre C. capsularis. The main defects of this jute are
bundles forms the meshy structure. For exam- the irregular fibre bundle and more meshy
ple, the structure of fibre bundles and their structure compared to that of C. olitorius. There-
cross-sectional area varies among the varie- fore, emphasis should be given to improve
ties of capsularis jute. fibre quality using anatomical traits. A greater
A breeder has to select a plant on the number and more layers of fibre bundles per
basis of its morphological structure, as fibre fibre wedge may also be considered for yield
retting will destroy the crop without provid- improvement (Maiti, 1973).
ing seed for progeny population. Moreover, The fibre filaments with highly meshy
during the retting process microbial organ- structure offer obstacles during the carding
isms degenerate the parenchymatous tis- process, while fibre filaments with minimal
sues without affecting ligno-cellulosic fibre meshy structure are easier to handle during
filaments, thereby liberating fibre strands the carding process. A good example of bet-
after completing retting process and wash- ter fibre type is Malachra capitata, which
ing with water. Over-retting or under-retting produces ideal fibre bundles for the spin-
reduces fibre quality, therefore it is difficult ning process. The fibre bundles of this plant
to assess the fibre quality if the fibre is not are more or less rectangular with uniform
retted properly. In this respect anatomical surface structure and are scarcely meshy,
traits can give correct prediction of the revealing good fibre quality. The fibre filament
potential quality of the genotype before tenacity is also high. Hibiscus cannabinus
retting, which may be used in the breeding produce better quality fibre compared to that
process. Considering these limitations, ana- of H. sabdariffa. Different species of minor
tomy of the fibre filament depicts clearly Hibiscus spp. produce poor quality fibres
the genetic potential of the fibre quality of with irregular fibre bundles and highly
crop plants as discussed herein. meshy fibre strands such as H. panduraeformis.
Large variability occurs among bast On the other hand, H. suratensis produces
fibre crop species in the anatomical struc- uniform fibre bundles with less meshy struc-
ture of fibre bundles and their orientation ture. Similarly, Sida rhombifolia produce
and intensity in the fibre wedge. In the fol- strong and good quality fibre. In a study
lowing section, the variability of fibre bun- (Maiti, 1969), it was reported that Hibiscus
dle anatomy of a few fibre crop species and vitifolius possesses uniform rectangular fibre
their varieties is depicted. bundles, producing good quality fibre of high
The irregular contour of fibre bundles fibre strength. The fibre bundles of Abutilon
produces fibre filaments with irregular surface indicum are united laterally producing flat
of poor fibre quality. The morphology of ulti- ribbon-like fibre filaments after retting. There-
mate fibre cells, and tip and size vary widely fore, the anatomical structures contribute
178 Chapter 8
meristematic tissue of primary or secondary reliable selection criteria for genetic improve-
origin depending on the species. Fibres of ment of both fibre yield and quality.
ramie and flax are mostly cellulose. In the context of the above discussions
Apart from seed fibres (cotton), yield and it is observed that great variability was
quality determinations of vegetable fibres are observed in anatomical traits of fibre bun-
difficult to assess unless the fibre is extracted dles of different long fibres such as jute,
by the retting process. This involves a series kenaf and other bast fibres among species
of biological and physical changes in fibre and among varieties of the same species. It
structure induced by microorganisms. Once is suggested that the following parameters
the fibre retting is complete, its quality is may be considered as selection criteria for
determined by several parameters including yield and quality improvement.
fibre tenacity, fineness, surface contour, rigid-
ity etc., assessed by various testing machines.
During the retting process the fibre quality is 8.2.5 Simple technique for screening
affected severely by poor- or over-retting. varieties of main bast fibre crops for yield
Plant breeders use indirect methods to esti- and quality potentials
mate potential fibre yields. In general in the
case of stem fibres, selection criteria for This technique is suitable for main bast fibre
genetic improvement of fibre yield are plant crops such as jute, kenaf, ramie and flax.
height and basal diameter. It is expected that A small quadrant (piece) of bark is cut at the
the greater plant height and basal diameter base of the stem of the respective species at
contribute to higher fibre yield, but a greater flowering stage. Then transverse section of
basal diameter combined with a lower num- the bark is cut with a sharp razor blade and
ber of fibre bundles usually produces poor viewed under a microscope by adding a
yields. On the contrary, a greater basal dia- drop of safranin in the case of jute and
meter with a greater number of fibre bundles kenaf for distinguishing lignocellulosic
increases fibre yield. Therefore, the selection fibre strands and by adding bismark brown
by greater basal diameter may be misleading or light green in the case of ramie or flax for
in few cases. For example, for soft stem observing cellulose fibre cells in transverse
mutants, the stem diameter is large, but with section. By using this method, a large
scanty fibre bundles. There are large varia- number of plants can be screened and
tions among crop species and within the selected within a short time for desirable
same species in the intensity of fibre bundles yield-contributing and quality traits. This
and their structure. It is evident that the anat- technique gives an opportunity to utilize the
omy of bast fibre plays a great role in deter- selected plants in the crossing programme
mining fibre yield (Maiti, 1971a). for possible genetic improvement of yield
It may be stated that until now there have and quality. The genetics of these traits need
been no direct selection criteria for fibre yield to be investigated for understanding the
and quality, which is only possible after inheritance of these traits.
extraction of fibre by the retting process. The Taking a small portion of bark at the base
yield and fibre quality are highly affected by of stem will not affect the plant growth but
retting methods and condition. At present, give an opportunity to the breeder to utilize a
plant height and basal diameter of stem are selected plant in crossing with another desir-
indirect selection criteria, but these selection able selection. The selection procedure will
criteria are influenced by the amount of fibre continue in F1, F2 to F7 to obtain homozygous
bundles or cells in the stem bark and the plants for the combination of desirable yield
quality is affected highly by poor retting con- and quality parameters. For example, there is
ditions. Specific examples may be given a necessity to improve the quality of white jute,
in case of ‘Soft Stem Mutant’ and ‘Halome- C. capsularis, owing to the irregular fibre bun-
hara’, which possess scant amounts of fibre dle and highly meshy structure. As mentioned
bundles. earlier there exists large variability in fibre bun-
Therefore, it may be hypothesized that dle structure among capsularis varieties. The
anatomical traits of fibre bundles may be jute variety ‘Sudan’ is finer in bundle, which
180 Chapter 8
Screening of germplasm/varieties
Basal diameter
Improved variety
Fig. 8.30. Schematic hypothetical diagram for possible genetic improvement of yield and quality.
Fibre Crops 181
varying from 3 to 7 cm. The stipules are of two In C. capsularis, leaves have a bitter fla-
types, conical and crossed. The angle of leaves vour and vary in morphology among geno-
is highly variable, from sharp to open. types; the pod is small, globular, rough and
corrugated, and seeds are pyramidal; the
Floral characters fibre is highly reticulated.
It is reported that the varieties of C. cap-
Flowers are present in the axils in solitary sularis show wide variations in morphologi-
form or in groups of three to four (panicle). cal characters, varying in the nature of
Flowers are pedicillate, with a small pedicle, pigmentation and branching habit of stem,
bisexual, actinomorphic, pentamerous and pigmentation, shape and size of leaf lamina
hypogynous. The calyx consists of five sepals, and petiole. This indicates that there is a
united, green in colour, and with volvate aes- wide range of diversity in morphological
tivation. The corolla has five yellow petals, free, characters, thereby offering great scope to the
4.5 mm in length, with imbricate aestivation. breeders for the selection of desirable varie-
The androecium consists of numerous ties for a certain growth parameter. For exam-
stamens, free, short filaments, dithecous, ple, higher leaf area, leaf angle (45°C) and
and introrse gynoecium with five carpels, longer petiole length may lead to greater pho-
syncarpus, five locules and a superior ovary. tosynthetic capacity.
Numerous ovules are present in each loc- Tossa jute (C. olitorius) yields more fibre
ule, with axile placentation, a short style per unit area compared to white jute (C. cap-
and the stigma is flat. sularis). The fibre is finer, softer, more lustrous
The fruit is a capsule (dehiscent capsule) and less rooty than C. capsularis. The leaves
measuring 5–6 cm in length, 2 cm in diameter are sweet whereas capsularis leaves are bitter
with a corrugated appearance. At maturity in taste. The capsules in C. olitorius are cylin-
the capsule opens in five parts without show- drical, but are globose in C. capsularis.
ing partitions between the seeds. The seeds
are numerous, small, 2–3 mm, of dark colour
and pyramidal form. 8.3.6 Anatomy
Meristem
8.3.5 Variations between C. capsularis
The bud initials are laid in the usual method
and C. olitorius
in the primordial meristem in the case of
branching plants of C. capsularis. The non-
The two cultivated species of jute are distinct branching character of some varieties is due
in their morphology, but similar in their gen- to the different structural organization of
eral characteristics, with the exception of the the shoot apex. In the non-branching plants
leaves and the fruit. They differ in leaf shape the absence of bud initials is associated with
and size, pod shape, growth habit and fibre early vacuolation of the meristematic cells.
quality. Variability in morphological and Cases occur of sporadic development of a few
yield-contributing traits is of great impor- axillary and extra-axillary buds (Kundu and
tance in the genetic improvement of a crop. Rao, 1954).
In this respect, varieties and germplasm of
C. capsularis vary widely in pigmentation Nodal anatomy
and branching nature of the stem, the shape
and size of leaf lamina and petiole. The cotyledonary node is unilacunar single
In C. olitorius, leaves possess a sweet trace or two traces, and the first leaf node is
flavour, the stem is cylindrical and smooth unilacunar single trace or trilacunar depend-
in surface; the capsules are long, slender ing upon the species. The approximation of
and cylindrical with a ribbed surface; the two traces from the cotyledonary node and
seeds are smooth, the fibre has a less meshy three traces from the first leaf node forms a
structure, and is very little adhered to the single trace in the lamina region (Thanki
bark. The fibres are golden brown in colour. et al., 2000).
Fibre Crops 183
Fibre anatomy and development of fibre development of the fibre cells they become
cemented at the ends by cementing materials
The fibre is an anatomical structure present of hemicelluloses and pectins to form the
in the stem and derived mainly from the cam- fibre strands. The fibre strands make their
bial activity and the growth in length by the way through the stem in a zigzag pattern and
apical meristem. An assessment of fibre is become connected here and there, forming a
only possible after the extraction of the fibres meshy structure of jute and other bast fibres.
by the retting process. Therefore, the knowl- The fusiform initials not only cut off fibre ini-
edge of morphogenesis of the fibres could tials, but also at times undergo transverse
enlighten us on the fibre-producing capacity divisions leading to the formation of soft
of jute and other bast fibres. parenchyma lying intermingled between the
In jute, 90% of the total fibre of the plant fibre strands, thus making possible their sep-
is developed in the secondary phloem by the aration and isolation. This process leads to
activity of the cambium, and the remaining the formation of networks in bast fibres.
10% is formed in the protophloem region. The ray initials in turn cut off deriva-
The vegetative phase represents the period of tives to the formation of the ray cells,
greatest cambial activity. Cambium maintains which make their way at right angles to the
a definite ratio in its division with approxi- main axis of the stem. Thus, the deriva-
mately two xylem initials produced for each tives of cambial initials have a bidirec-
phloem initial, due to which the secondary tional momentum of force, one (fibre
wood developed by the cambium forms a initial) along the longitudinal direction
much thicker zone and occupies the greater and another (ray initial) along the trans-
part of the radial growth. The fibre cells verse direction. This bidirectional force is
developed from cambium undergo rapid associated with a vertical twisting growth
change in length and breadth. The secondary by apical meristem leading to the peculiar
wall of the young fibre cells becomes thicker mode of arrangement in the stem. The for-
by the gradual deposit of secondary layers of mation of a meshy network (Fig. 8.31) is a
cellulose materials. main characteristic of jute and other bast
It is reported that the outermost layers of fibre, depending on the activity of cam-
the fibre bundles in the fibre wedge are pro- bium. The degree of meshiness is associ-
tophloeic in origin. The sieve tubes and com- ated with tangential expansion and radial
panion cells are obliterated by the expansion growth of the stem. The meshes are loose
and elongation of the surrounding cells. The towards the top and compact towards the
procambium in the protophloeic region under- basal region. At the basal region, the
goes cell division and modification leading to meshes are loose towards the periphery
the protophloeic fibres. This process contin- and more compact towards the cambium.
ues towards the inner side, thereby crushing The meshiness is a drawback because it
and obliterating all the sieve tubes and the hinders the carding process. Coarse meshi-
companion cells, and the entire protophloem ness reduces the quality of fibres and
is covered by the solid fibre patches. The weakens the fibre strands during splitting
remaining bulk of the fibres are derived from and combing operations in the spinning
meristematic activity of cambium, which process.
contains two types of fibre initials, the fusi- Fibre filaments as they appear in com-
form initials and ray initials. Fusiform initials merce are developed and oriented in a given
are elongated cells that turn lengthwise along manner peculiar to each variety of species.
the axis of the stem and give rise to new fibres Fibre cells derived from the meristematic
by repeated longitudinal divisions. Ray ini- activity of cambium are assembled together
tials are short cells in rows that divide to give in a definite pattern. The differential cambial
rise to ray cells. The fibre initials after being activity that governs the yield and quality of
cut off from fusiform initials make their way fibres is characteristic of the genotype of the
longitudinally through the tissue by an apical different species or varieties. However, the
intrusive and symplastic growth. After full mode of arrangement of the fibre bundles in
184 Chapter 8
(a)
(b)
(c)
(d)
Fig. 8.31. Pattern of anatomical structure of fibre bundles in jute stem: (a) transverse section; (b) single fibre
wedge; (c) view of meshy structure of fibre filaments in longitudinal section; and (d) meshy structure
(Maiti, 1997).
the secondary phloem fibre differs from one towards the periphery with alternate patches
species to another. Generally it is a ribbed of fibre bundles and thin-walled tissue in
meshy structure broad at the base and grad- between. Fibre bundles are arranged radially
ually tapering upwards into free filiform in superimposed arcs along radii, tangential
strands, mostly composed of primary cells. In in the axis of the stem, and the bundles are
a transverse section these fibre strands appear traversed by phloem rays and show great var-
as an extra-cylinder ring of pyramidal wedges iability in the fibre bundle structure among
(see Fig. 8.31), having a centripetal broad base C. olitorius varieties. The ultimate fibre cells,
and a narrow conical and radially directed which form the building skeleton of the fibre
Fibre Crops 185
filament, are pointed at the apex and have a of C. olitorius indicated that growth rate of
lumen at the centre, although they vary between the meristem (intercalary and apical) for the
two species. The ideal criterion for the selec- formation of fibre and plant height showed
tion of jute for high yield is a combination of alternate peaks, but the period of activity for
three factors: plant height, basal diameter both components was different in other vari-
and fibre content (number of fibre bundles), eties (Fig. 8.32). The period of optimum cam-
which is influenced by the cambial activity bial activity among the varieties was similar
during the vegetative phase. at 40 days. However, the period of optimum
The secondary cambium forms the activity of the apical meristem (42 days) was
phloeic fibre in all peripheral directions, and very different from the activity of cambium
wood in a centripetal direction towards the (Maiti et al., 1974).
stem pith. The contribution of apical meri- Shaikh et al. (1980) applied several ana-
stem leads to the elongation of stem, and its tomical methods in order to select mutants of
direct contribution to quantity of fibre is very C. capsularis with good fibre yield. In this
low since the greater quantity of fibre corres- study, two morphological characteristics,
ponds to the secondary activity of the cam- plant height and basal diameter, were consid-
bium. Diverse anatomical studies show that ered and anatomical components were stud-
the activity and function of the apical meri- ied in transverse sections. The coefficients of
stem finishes when the flowering begins, but environmental variability and genetics and
the activity of the secondary cambium could their correlations were computed in order to
continue even after flowering. This study in confirm the effects of other agronomic and
the quantitative assessment of the cambial anatomical characteristics on fibre yield. Some
activity in relation to the production of fibre genotypes with good yield have very thick
bundles in some cultivars of C. capsularis bark and greater area of fibre bundles in each
indicated that the varieties differed widely in section. The number of cells per fibre bundle
the optimum growth responses. In another in each strip showed a negative influence on
study, the cambial activity with respect to the the yield of fibre. The same fibre cell area has
formation of fibre and wood in four varieties high variability in the genetic coefficient.
Fig. 8.32. Variability in fibre bundle structure among C. olitorius genotypes: 1, fibre wedge; 2–9, fibre
bundle morphology (Maiti, 1997).
186 Chapter 8
This indicates an opportunity for selecting 11 indigenous genotypes. Highest mean fibre
these characters. Shaikh et al. consider these cell length and thinnest fibre cell occurred in
characteristics the most effective for the deter- the upper part of the stem.
mination of a good yield potential, better than The genetic relationship between ana-
the agronomic parameters of plant height and tomical characters and fibre yield and quality
basal diameter. These observations support in varieties of C. capsularis was reported by
the findings of Maiti (1980), who emphasizes Chen et al. (1990) and by Chen (1991). The
the importance of anatomical characters in number of fibre bundles and layers was
determining yield as well as quality. highly correlated with plant height and stalk
The ultimate fibre cell (Fig. 8.33) is wide diameter. Secondary fibre cells began differ-
in the centre and gradually tapering toward entiation 21 days after emergence, but the
the ends; the cell presents lumen that on duration of development varied significantly
occasion gets obliterated. In C. olitorius, fre- among the varieties, especially in the number
quently the lumen is wider and uniform in of fibre cells and fibre bundles, mainly at the
thickness, and sometimes has a constriction middle and late growth stages. In mature fibre
on the cell wall in C. capsularis. The lumen cells, the cell cross-sectional diameter showed
is irregular with frequent constrictions. The significant positive correlation with cell wall
maceration of fibre filaments to individual thickness. Fibre cells took 30 days from dif-
fibre cells could be carried out in a mixture of ferentiation to ripening, the growth and ripen-
10% chromic acid and 10% nitric acid in a ing rate differing with variety, plant growth stage
proportion of 1:1, in an incubator at 60°C and temperature.
approximately for 1 h. The number of fibre bundles and number
In a study by Hao et al. (1993) in Fujian, of cells in the bundle are indicators of fibre
China, correlation coefficients among fibre yield as well as the irregularity and regularity
counts, fibre strength and single fibre cells were of the bundle surface, the length of the ultimate
determined in 16 accessions of C. capsularis. fibre, the length/breadth ratio and the area of
It is suggested that advanced generation for section of bundle. The improvement of fibre
selection of single fibre cell length, small cell quality is imperative where there is com-
cavity and thin cell wall may be effective in petition with synthetic fibres. Several studies
improving the quality of the fibre. This has have suggested that anatomical traits associ-
also been suggested by Maiti (1980). ated with morphological characters could be
Ali (1989) advocated the use of ultimate considered as selection criteria for the genetic
fibre cells for selection in C. olitorius. He improvement of jute for yield and quality.
determined the length, thickness, and length Some structural parameters of fibre bun-
to thickness ratio of the ultimate fibre cells dles, as revealed by the microscopic study of
from the top, middle and base of the stems of the transverse section of jute plant stems,
have been found to have high positive cor-
relations with filament fineness of extracted
fibres. Equations have been derived by lin-
ear regression analysis to predict the fineness
of extracted fibres. The study would not only
be very helpful in selecting the fine fibre-
producing plants in breeding programmes,
but will also offer some information on the
structure–quality relationship of jute fibre,
essential for genetic upgrading of fibre quality
(Majumdar, 2002a).
stem. This true-breeding mutant produces varieties of capsularis jute indicate that these
only primary phloeic fibres. The secondary show much structural variabilities, and it is
phloeic fibres, which are the derivatives of predicted that some of the anatomical charac-
vascular cambium, are found lacking, though ters are helpful in the selection of a variety in
there is formation of secondary phloem pyra- quality breeding. As fibres are in the stem, the
mids from the cambial tissue. Inheritance of fibre yield may be reflected by the attainment
fibre (bast) in jute (C. capsularis) is mono- of plant height and basal diameter, but the
genic and recessive (Mitra, 1984). yield may be less if the fibre content is less.
In a study the mutant grew more slowly, Fibre cells are broad and narrow-
had shorter internodes and yielded much lumened type; tips are straight and pointed,
less fibre after retting. The fibre of the mutant notches are present in denticulate manner in
contained 50% less lignin but comparatively some, the lumen is broader than wall thick-
more cellulose than that of the normal type. ness, constricted and blocked; and the cell
The lower and earliest developed part of the surface is generally regular.
mutant stem had no lignified fibre cells. This The tenacity values of bundles of jute
developmental deficiency in lignification of fibres belonging to different varieties have been
fibre cells was correlated to a similar deficiency predicted from the structure of their fibre bun-
in phenylalanine ammonia lyase activity, but dles as seen in the transverse section of jute
not peroxidase activity, in the bark tissue stems and the length of the constituent ulti-
along the stem axis. This mutant may be uti- mate cells by multivariate regression analysis.
lized to engineer low-lignin jute fibre strains Jute plant stems having fibre bundles compris-
(Sengupta and Palit, 2004). ing a greater number of compactly arranged
long and fine ultimate cells give fibres of higher
Yield on the basis of anatomy strength when retted under standard condi-
tions (Majumdar, 2002b).
Correlation studies showed that bark thick- There is a simple procedure for identify-
ness, number of phloem pyramids, number ing the textile bast fibres from flax, nettle/
of fibre bundles in phloem pyramids, num- ramie, hemp and jute. The procedure is based
ber of fibre bundle arcs in phloem pyramids on measuring the fibrillar orientation with
and area of phloem pyramids were posi- polarised light microscopy and detecting the
tively and significantly correlated with fibre presence of calcium oxalate crystals (CaC2O4)
yield in the genotypes. Genotypes showed in association with the fibres (Bergfjord and
strong association of the component charac- Holst, 2010).
ters with fibre yield except number of fibre
bundle arcs in phloem pyramids. Bark thick- Fibre quality parameters
ness showed highly significant correlation
with phloem pyramid areas in almost all the The quality of the jute fibre is considered in
genotypes (Ali, 1993). accordance with five major criteria: force,
fineness, lustre, colour and length. The ideal
fibres for spinning have high lustre, good col-
Fibre quality on the basis of anatomy
our and good length. For commercial pur-
Plant height and basal diameter are used as the poses, the most important quality parameters
conventional method of selection for improve- are fibre strength, fibre fineness, nature and
ment of yield of jute and allied fibres, but to extent of defects, root content of the fibre, col-
the breeders there are no authentic breeding our of the fibre and fibre density. Besides the
criteria for quality improvement. Breeders length of the fibre cell, the diameter and dis-
require some suitable and rapid techniques for tribution of fibre cells are also important
assessing quality on a single-plant basis. They quality parameters of jute fibre.
cannot depend on physical methods for esti-
mation of quality. Anatomists have attempted FIBRE STRENGTH Fibre strength is defined as
to correlate some of the anatomical compo- the amount of force required to break a fibre
nents with quality characters of fibres. Different of unit length and weight. The unit length is
188 Chapter 8
generally fixed at 5 cm or 10 cm and a mech- from three different varieties of yarns in a hot
anical jerk is given to break the fibre. The press and then studied for the bending pro-
fibre strength is expressed as g/tex. The fibres perties on a DCS-500 tester using a three-
have high initial modulus and low elasticity; point loading system. It was observed that the
therefore, they have small extension capabil- bending properties of composites, especially
ity before breaking. the transverse bending properties, are affected
by the mixing degree of reinforcing fibre/
FIBRE FINENESS Fineness is described as the matrix fibres and bulkiness of reinforcing
width of fibre filaments when fibre fila- fibres bundle (Li et al., 2002).
ments are separated. It is also expressed as
the width of fibre strands when separated Pulp quality
mechanically or manually. The fibre fine-
ness ultimately depends on the fibre cell Jute fibre pulp showed better papermaking
diameter. Jute fibre cells are not circular, properties than jute cuttings and caddis. The
rather elliptical shaped with many irregu- tear index of these raw materials was similar
larities. This makes direct estimation of to softwood. The bleachability of jute fibre
fineness difficult from fibre cell width. pulp was also better than that of cuttings and
caddis. Pulp yield and bleachability was
ROOT CONTENT The root content in the higher and kappa number lower for jute fibre
fibre describes the hard bark regions at the than jute cuttings and caddis, but α-cellulose,
lower end of stem. As the region is hard, S10 and S18 values and viscosity were almost
retting is not often complete in this region. similar in these three raw materials (Jahan
It is expressed as the percentage of rooty et al., 2007).
fibre weight to total fibre weight. When
fibre is exported, generally the rooty ends
are chopped out and sold separately, and 8.4 Kenaf
are called ‘cuttings’.
8.4.1 Introduction
FIBRE COLOUR AND LUSTRE Although fibre col-
our does not affect the end use of jute fibre as Kenaf (Hibiscus spp.) is a bast fibre and is
hessian, sacking or clothing, it is a primary next to the jute fibre in importance and a val-
determinant of jute fibre price in the market, uable substitute of jute for sack and rag pro-
especially for tossa jute. The capsularis jute duction. The main kenaf fibre-producing
is dull white in colour, while olitorius jute species of economic importance are H. can-
turns golden yellow after retting. The colour nabinus and H. sabdariffa (H. sabdariffa var.
is also indicative of extent of retting of fibre, altissima) and often the fibres from these two
thereby serving as a selection criterion for the species are difficult to distinguish in the mar-
buyer. The lustre of the fibre also indicates ket. They belong to the family Malvaceae.
the extent of retting as over-retted fibres will The major producers of kenaf fibre are
lose lustre and will become dull. India, China, Thailand, Vietnam, Cambodia,
Indonesia, Brazil and Cuba. China was the
FIBRE CELL LENGTH The individual length of largest producer of kenaf in the first decade of
jute fibre ranges from less than 1 mm to the 21st century.
greater than 5 mm, with an average around In India, these two species are also known
2.8 mm. Longer fibre cells indicate better as ‘roselle’ (H. sabdariffa), mesta pat or mesta
quality fibre and are more suitable for blend- (H. cannabinus and H. sabdariffa), Deccan
ing in clothes and paper pulp industries. jute and ambari. H. cannabinus is also known
as Java jute and siami jute in South-east Asia,
Yarn quality stokroos in South Africa and Guinea hemp in
West Africa. In northern Africa, particularly
The unidirectional thermoplastic commin- in Egypt, kenaf is known as teal or teel. In the
gled yarn composites have been fabricated European countries, kenaf was introduced
Fibre Crops 189
later and is generally named after the country objects, and also in combination with other
from which it was introduced, such as ‘cáñamo synthetic fibres or vegetable fibres such as
de la India’ in Spain. jute. In some countries the fruit is eaten. The
quality of the kenaf fibre is good except for its
8.4.2 Origin and distribution fineness and semi-meshy structure. Fibre
strands are more irregular than jute.
Kenaf seeds contain 16–22% oil and
H. cannabinus originates from Africa, where
32% protein, ideally suiting them for use in
it is found in its wild form, and also in the
cooking; they also contain unsaturated fatty
subtropical regions of Asia. There are some
acid in a low proportion, which is used in the
controversies with regard to the origin of
elaboration of margarine. After extraction of
H. sabdariffa var. altissima: some authors
oil, the residue contains approximately 35%
indicate that it originated in Africa and India;
protein and is used as livestock feed. The
however, it has been reported that H. sabdar-
seedlings, leaves and fruits are used to make
iffa var. altissima has been introduced in
sauces, jellies and wines.
India through single seed mixture with
Calapogonium muconoides.
H. cannabinus and H. sabdariffa are
widely distributed in the USA, Latin American 8.4.5 Morphological description
countries, Russia, Sudan, Egypt, India,
Australia, the Philippines, Java, Iran, Nigeria, Vegetative characters
Senegal, Thailand, China and Brazil. It has
Both species are herbaceous and present
been introduced into Cuba, Guatemala, El
two sexes in the flowers present in the leaf
Salvador, Colombia, Mexico, Costa Rica and
axils on the stem. They are annual crops.
Haiti as a source of fibre and into Tanzania,
The taproot is pivotal, and secondary and
Kenya and Australia for the production of
tertiary roots are derived to give support to
paper. It was introduced into Europe early in
the crop and absorb nutrients. The stem
20th century.
could be flat and thorny and reach a height
The crop was introduced into China
of 2–4 m depending on the environmental
from India through Taiwan and from Russia
conditions and varieties. Roselle (H. sab-
in the early 20th century. A cultivar called
dariffa) is an annual plant. The stem shows
‘Madras Red’ was introduced in Zhejiang
several growth patterns and is generally
province, China. In Russia, kenaf was culti-
purple. The leaves are polymorphic, pal-
vated on a large scale in the 1930s.
mate, extremely lobulate with three or five
lobes, and alternate on the stem with very
8.4.3 Adaptation deep venation (Fig. 8.34).
Kenaf is more adaptive than jute under Floral characters (H. sabdariffa)
diverse conditions of climate and soil, and is
very resistant to drought. Kenaf is a traditional The flowers of H. sabdariffa are smaller than
fibre crop with adaptability to both tropical in H. cannabinus, and solitary in the axiles of
and temperate environments, although a trop- the upper leaves on a short peduncle. The
ical environment is more favourable. It is peduncles are shorter than the petioles; the
considered as a fibre crop of marginal farm- bracts and the calyx show upward growth,
ing communities under drought-prone rain- the sepals are triangular in form and acumi-
fed conditions. nate, with more than half an intense purple
colour. The flowers are generally cream or
yellow, with scarlet to purple in the interior
8.4.4 Utilization part of the neck. The capsule (fruit) is ovoi-
dal, pointed, hairy, and shorter than the axis;
Kenaf fibres are used mainly in the fabrication the seeds are reniform, coffee-coloured and
of paper, cordage, fabric, yarn and decorative smaller than those of H. cannabinus.
190 Chapter 8
Fig. 8.34. Kenaf plant morphology: leaves are polymorphic, palmate, extremely lobulate with three or five
lobes; the flowers are cream or yellow in colour.
8.5.3 Utilization
differences for diverse anatomical character- transverse partition and nodes in an X form
istics in certain varieties of flax, such as the (Fig. 8.39). The individual fibre cell in cross-
thickness of the fibre patch, the preparation section is polygonal or circular. The fibre cell
of wood to the region of fibre, the area of has a length of 27 mm with a range of 9 to 70
cross-section of the fibre cells, the thickness mm and an average diameter of 23 µm.
of the fibre cell wall, and the number of lamella Cellulose is the main constituent of flax
in the cell wall. fibre. Cellulose deposition in the fibres of flax
On the basis of the distribution of the hypocotyls were observed by Goubet et al.
fibre cells, the overall varieties are classified (1993) using 14CO2. The glucose was incorpo-
into three groups: rated into storage polysaccharides (probably
starch) and used when needed for the secon-
1. Orientation of the fibre into a compact group.
dary deposition of cellulose.
2. Fibre cells in few groups or in isolated form.
3. Fibre cells isolated and distributed.
The fibre cells are cylindrical with a flat sur- Lignification and other deposition
face and pointed termination; they have a in the fibre cell
very narrow lumen and a very thick cell wall.
The process of lignification in flax fibre has
The lumen is not distinguishable near the tip
been studied in detail by a number of work-
of the fibre cell. One of the important charac-
ers. With the aid of spectroscopic and micro-
teristics of the flax fibre cell is the presence of
scopic observations, a definite relationship of
lignin deposition and fibre maturation has
been identified. Salnikov et al. (1993) studied
the ultrastructure of flax development using
transmission electron microscopy of ultra-thin
section of plant dry material and cytochemical
staining of the polysaccharides. Their findings
suggest that Golgi apparatus, endoplasmic
reticulum, microsatellites and plasmodesmata
take an active part in the synthesis of the
polysaccharide wall matrix. The orientation
of cortical microtubules and microfibrils of
secondary cell walls was annular. The two
layers of the secondary wall differed in the
distribution and content of polysaccharides.
It was also observed that the inner electron-
dense layer of the fibre cell wall became thinner
at fibre maturation.
Enzymes such as peroxidase were acti-
Fig. 8.38. A transverse section of flax stem showing vely involved in the process of fibre cell lig-
orientation of fibre cells in the cortex (Maiti, 1997). nifications in flax fibre. Cell wall peroxidase
(a) (b)
Fig. 8.39. Ultimate fibre cell of flax showing X-shaped cross-section (Maiti, 1997).
Fibre Crops 195
reached an optimum level around the time of Chemical composition of flax fibre
maximum xylem differentiation. There was
an alteration in the type of peroxidase en- Chemical analysis reveals that several monosac-
zymes at the time of active lignification of charides can be obtained from flax fibre on
fibre cells. In a recent study from dissolved decomposition. The fibre carbohydrate con-
fibre bearing tissue from the stem of flax, it tains primarily glucose (72–75%), galactose
was observed that the activities of cell wall (3–4%), mannanose (3–4%) and traces of ara-
peroxidases were correlated with the fibre lig- binose and xylose. Acid insoluble lignin con-
nification. The onset of fibre lignification was tent varies from 3 to 4%.
associated with an increase in the levels of
peroxidases both ionic and covalently bound 8.5.6 Effect of herbicide
to wall, but the rise was greater in the levels of
covalently bound wall peroxidases. From In flax, a range of glyphosate treatments was
PAGE analysis a number of cationic and ani- applied at three stages of flowering. During
onic enzymes were identified; these enzymes normal maturation, increase in fibre cell wall
may have specific roles in the lignification of thickness, lignification of the fibres and dif-
flax fibres (McDougall et al., 1993). ferentiation of the secondary xylem contin-
Love et al. (1994) suggested that the ued for 3–5 weeks after the beginning of
amounts of phenolic substances reported earl- flowering. This differentiation was halted by
ier are overestimated. They estimated deposi- the application of glyphosate. In some sec-
tion of bound phenolic substances in flax tions epidermal and cortical cells showed the
fibres by solid state NMR spectroscopy at low most damage, this being consistent with her-
field (25 MH2) under conditions to give quan- bicide uptake at the stem surface. In other
titative responses from the aromatic carbon sections phloem and associated parenchyma
atoms. This method gave a better estimate for cells showed the most damage. This resem-
determination of phenolic structures in fibres bled the release of fibre bundles that results
in comparison with single-pulse excitation from conventional post-harvest retting (Fraser
used earlier. et al., 1982).
Fig. 8.41. Transverse sections of ramie stem showing the distribution of fibre cells in the cortex (Maiti 1997;
World Fibre Crops).
cells are united at the ends with gums, wax researchers have studied the structure of the
and pectins. fibre of ramie by means of X-rays and elec-
A great deal of information is available tron micrograph.
on the helical structures, microfibrillar orien-
tation and stratifications in the cell wall of Technique for studying secondary wall
the fibre. For example, there are three stratifi-
cations (S1, S2, and S3) forming two helical A technique was developed by Maiti and
layers in the cell wall of the fibre, the external Ghosh (1974) and Maiti (1980) in India to study
layer in the form of the S and the internal one the lamellar orientation of the secondary wall
in the form of a Z, with the possibility of the of the ramie fibre. Two or three drops of H2SO4
existence of a third cellular covering. Several at 70% are added to a transverse section of
Fibre Crops 199
stem in a Petri dish at 40°C for 5 min. Sub- photosynthetic rate and decreasing electro-
sequently, the solution is removed and a few lyte ex-osmosis rate (Liu et al., 2001).
drops of chlorozinc-iodine are added to the
sample, and the sample is covered with a
cover slip for observation under a micro- 8.6.8 Drought – anatomical changes
scope. With this treatment, the lamellar struc-
tures of ramie fibre cells are visible. On the Drought-tolerant cultivars of ramie had more
basis of this study, it was concluded that fine hairs on abaxial leaf surfaces, produced
there are differences among the varieties of thicker leaf cuticles, better preserved leaf shape
ramie with respect to the lamellar orientation and erectness, and lost fewer leaves. They also
in the secondary wall, which is probably generated longer roots with larger root masses
related to the strength, rigidity and elasticity. and more storage organs, preserved higher
Also, the area of cross-section of the ramie root-to-shoot ratios, produced larger diameter
fibre cell indicates the fineness of the fibre in stem vessels, and better conserved cell integ-
the same study. These authors found that rity than drought-sensitive cultivars of ramie
there are large differences in the size of cross- when plants were grown under drought stress.
section of fibres among varieties of ramie. Fibre yield was better in drought-tolerant cul-
The fibre cells of ramie show a spiral struc- tivars of ramie, because these cultivars had
ture in the cell wall (Maiti, 1980). adapted root systems, leaf responses, cellular
responses and biochemical activities to allow
plants to continue higher levels of photosyn-
thesis and carbon deposition under more
8.6.6 Chemical properties of fibre stressful environments than the less drought-
tolerant cultivars (Liu et al., 2005).
Chemically, the fibre consists of cellulose and Under drought stress conditions there was
hemicellulose with very little amount of lignin. a decrease in yield, biomass and agronomic
Most of the cellulosic material of the fibre is characteristics, i.e. plant height, stem diameter
holocellulose. Harvested fibre has a very high and ribbon thickness. Under osmosis and
gum content, which comprises about 20% of drought stress conditions, the ramie genotypes
the fibre. resistant to drought had a lower withering
rate compared to that of genotypes of lower
drought resistance. Under drought stress
8.6.7 Effect of growth regulators conditions, drought resistant genotypes had a
greater leaf thickness and volume of root sys-
The effects of two mixtures of four plant tem, and a higher dry matter weight of under-
growth regulators (choline chloride, gibberel- ground parts (Cheng, 2000).
lin (GA3), benzyladenine (6-BA) and NaHSO3)
at 20:9:5:800 mg/kg (H1) and 20:42:43:2350
mg/kg (H3) (active ingredients), respectively, 8.7 Hemp
were investigated on yield and fibre quality
in ramie (B. nivea (L.) Gaud.). The mixtures 8.7.1 Introduction
were sprayed over the canopy at two growth
stages (10 and 20 days after the previous cut) Hemp (Cannabinus sativa) is used mainly for
of field-grown ramie. The treatments increa- fibre production from the stem in temperate
sed raw fibre yield by 13–18%, and improved countries, although it is also used for medi-
fibre fineness by 57–349 m/g, increased the cinal purposes. It has been used since the
number of leaves per plant and also improved first Chinese civilization. At present, hemp
all yield components. Treatment H1 resulted is exploited in temperate climates for fibre
in a denser distribution, smaller diameters and seed oil. As a medicinal plant, it pro-
and greater quantity of fibre cells in stem cross- portionates resin with high stimulating
section. Physiological responses included power or drugs known by the name of ‘has-
improving leaf water status, increasing net sis’, whose main hallucinogenic narcotic
200 Chapter 8
has an average breadth of 22 mm and length in the outer zone initiated a secretory cavity
of nearly 25 mm. The lumen is broader than among fibrous wall materials. Hyaline areas
in flax and is frequently straight and lineal; at accumulated conspicuous electron-dense
the apex, it is very narrow. The fibre cells are contents that were released into the secretory
rounded and present projections; the cell cavity, thereby forming rounded secretory
wall is very thick. Also, the fibre cell presents vesicles. Fibrous wall material secreted
some nodulations and sub-terminal ramifica- from the surface of the disc cells became dis-
tions, which do not occur in flax. tributed throughout the secretory cavities
In Cairo, comparative morphological among the numerous secretory vesicles.
and anatomical studies of male and female
plants of C. sativa were undertaken at dif-
ferent stages in their life cycle. No signifi- 8.7.6 Fibre quality
cant difference was observed in cannabinoid
content between sexes; the highest concen-
The fibre colour is white or creamy to coffee
trations were found in young stems and
colour or brown. However, when it is extrac-
leaves, and the concentrations decreased
ted by machinery, it is grey in colour. The
gradually with age.
fibre length varies from 1.2 to 2.1 m on aver-
The dermal sheath of glandular tri-
age and can be used for spinning and fabrics.
chomes of C. sativa was studied by trans-
Fibre quality is measured in terms of force,
mission microscopy. Prior to the secretory
fineness, uniformity, colour, lustre, length
activity, cuticle thickened selectively on the
and strength.
outer wall of disc cells of each trichome,
The quality of hemp stems as a raw
whereas thickening was less evident on the
material for paper was assessed at Wagenin-
dermal cells of the bract. Membraned secre-
gen, the Netherlands. Bark content decreased
tory vesicles that differ in size and appear-
during the growing season, and at harvest in
ance were the source of precursor for cuticle
September it ranged from 30 to 35% depend-
synthesis. Vesicle contents were released
ing on cultivar and plant density. The pro-
following the degeneration of the vesicle
portion of secondary bast fibre in the bast
membrane upon contact with the subcutic-
fibre fraction increased with stem weight
ular wall and finally contributed to both
from 10 to 45%. Differences in chemical
structural and amorphous phases of cuticle
composition within the sets of bark or core
development. The structural phase was evi-
samples were small compared with the dif-
dent by deposit and thickening of cuticle at
ference between bark and core. The bark of
the subcuticular wall–cuticle interface to
a French cultivar contained less cellulose
form a thickened cuticle. In the amorphous
than that of the Hungarian cultivars. Bark
phase precursors permeated the cuticle in a
quality for paper improved during the grow-
liquid phase, as evidenced by the fusion of
ing season, because the cellulose content in
cuticle and wax layers between contiguous
the bark increased while the content of
glands.
lignin and extractives decreased.
The development of the secretory activ-
ity and the formation of the subcuticular
wall of glandular trichomes in C. sativa
were examined by transmission micros- 8.8 Sunnhemp
copy. The secretory cavity originated at the
wall–cuticle interface in the peripheral wall 8.8.1 Introduction
of the discoid secretory cells. During the
pre-secretory phase in the development of Sunnhemp (Crotalaria juncea) is a fibre
the glandular trichome, the peripheral wall crop of Indian and Asiatic origin. It is next
of the disc cells became laminated into a to cotton, jute and kenaf in importance.
dense inner zone adjacent to the plasma The demand for sunnhemp is increasing
membrane and a less dense outer one adja- in the manufacture of specialized tissue
cent to the cuticle. Loosening of wall matrix paper and currency notes. The fibre has
202 Chapter 8
more tensile strength and durability under seed as well as in the extent of ramification
exposure to humidity than jute. Sunnhemp of the stems.
is a promising raw material for pulp pro-
duction in the USA. Little or no nitrogen Vegetative characters
fertilizer is required to be applied, it is
drought resistant and grows on poor soil. The plant is annual, erect, with cylindrical
The stalks dry out faster than other species stems and a height of 1.2 to 3 m. The tap-
after the killing frost. It has a lower yield root is long and presents great ramification
than kenaf, and the stems are fragile. It is a with enough nodules for the fixation of N
short-day plant, which restricts the period through bacteria. The stem is extremely
during which it can be grown. It is cross- green. The leaves are simple, narrow, ses-
pollinated, and seed production is depend- sile and lanceolate and covered with tiny
ent on bees. Sunnhemp belongs to the pubescences.
family Fabaceae or bean or pea family.
Floral characters
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Jasdanwala, R.T., Singh, Y.D. and Chinoy, J.J. (1977) Auxin metabolism in developing cotton hairs. Journal of
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Kakani, V.G., Reddy, K.R., Zhao, D. and Mohammed, A.R. (2003) Effects of ultraviolet-b radiation on cotton
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9
Vegetable Crops
Vegetables form our daily dietary supplements herb widely cultivated for its edible fruit.
as they are a rich source of vitamins, minerals Savoury in flavour; the fruit of most varie-
and fibres. Most vegetables are used for nutri- ties ripens to a distinctive red colour. It
tion but some even have therapeutic proper- belongs to family Solanaceae. Diploid chro-
ties used in medicine. There are diverse forms mosome number is 24 (2n = 2x = 24).
of vegetables in nature, some commonly found
in all geographical regions but a few restric-
ted to specific regions. Such species, termed 9.1.2 Origin
endemic species, have a small range of adapt-
ability, hence are found growing in small pock- The tomato is native to South America.
ets of climatic regions. Thus vegetable crops Genetic evidence shows the progenitors of
develop different adaptations, either morpho- tomatoes were herbaceous green plants with
logically and anatomically in order to survive small green fruit and a centre of diversity in
under different habitat conditions. Each crop the highlands of Peru. One species, Solanum
variety has its own distinct range of adapta- lycopersicum, was transported to Mexico
tions that give it a characteristic identity. where it was grown and consumed by Meso-
Large variability exists among vegetable american civilizations. The exact date of
crop species in their characters. Morphological domestication is not known. The first domes-
and anatomical characters differ from one ticated tomato may have been a little yellow
species to another. We can identify the species fruit, similar in size to a cherry tomato, grown
based on these characters. Anatomy gives the by the Aztecs of central Mexico. Aztec writ-
details of internal structure and functions of ings mention tomatoes were prepared with
different plant parts and how they are adapted peppers, corn and salt. The word ‘tomato’
to different environmental conditions. comes from the Aztec (Nahuatl) tomatl, liter-
ally ‘the swelling fruit’.
9.1 Tomato
9.1.3 Utilization
9.1.1 Introduction
Tomato is frequently used as a vegetable for
The tomato (Lycopersicon esculentum) is an curries all over the world. Tomatoes are
herbaceous and profusely branched annual now eaten freely throughout the world, and
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
206 (R. Maiti et al.)
Vegetable Crops 207
Stem anatomy
Spongy tissue
Xylem
Phloem
Parenchyma
Trichome
Collateral and closed vascular bundles (veins) It has been reported that the foliar tetra-
occur between the palisade and spongy tis- cellular glandular trichomes (tetrads) of the
sue (median in position). In the vascular tomato plant, Lycopersicon esculentum Mill.,
bundle, xylem is present towards the upper contribute significantly to the antibiotic effect
epidermis and phloem towards the lower of the leaf against the fruitworm Heliothis zea
epidermis. The vascular bundle is surroun- (Boddie), as measured by reduction in larval
ded by parenchymatous bundle sheath. growth. This effect is attributable to pheno-
lic compounds localized within the tetrads
Role of leaf trichome in insect resistance (Duffey and Isman, 1980).
Removal of the glandular trichome
Intensive research has been undertaken on exudate from leaflets of the wild tomato
the role of glandular trichomes in insect L. hirsutum by swabbing with ethanol resul-
resistance. ted in loss of resistance to larvae of the tomato
Tomato varieties used for commercial fruitworm (H. zea). H. zea larvae were killed
production in Dutch glasshouses possess a by fumes from the surface extract and from
high density of glandular trichomes on the pure 2-tridecanone. The air surrounding leaf-
stem, but a very low density on the leaves. lets was found to be rich in 2-tridecanone
The two-spotted spider mite Tetranychus vapours (Dimock and Kennedy, 1983).
urticae Koch, and the predatory mite, Phyto- The active compounds in the L. pennel-
seiulus persimilis Athias-Henriot disperse lii rinsates were identified as 2,3,4-tri-O-
from leaf to leaf via the stem and face a high acylglucoses bearing short to medium chain
risk of death caused by exudates of the length fatty acids. These compounds are
glandular trichomes. These risks have been localized in the glandular exudate of the type
quantified on the tomato cv. Turbo and an IV trichomes and may accumulate to levels
accession of Lycopersicon peruvianum that in excess of 400 g/cm2 (Goffreda et al., 1989).
is almost free of glandular trichomes (Van 2-Tridecanone/glandular trichome-
Haren et al., 1987). mediated resistance to the tobacco horn-
The tomato plant glandular trichome worm Manduca sexta (L.) and the Colorado
breaks to release a yellow chemical sub- potato beetle Leptinotarsa decemlineata
stance that produces the characteristic (Say) in a wild tomato, L. hirsutum f. glabra-
tomato plant smell. Current pest manage- tum C. H. Mull, PI 134417, adversely affects
ment techniques depend on pesticides but several species of parasitoids and predators
trichome-based host-plant resistance may of the tomato fruitworm H. zea (Boddie)
reduce pesticide use. Most studies are (Kennedy et al., 1991).
focused on Lepidoptera and Hemiptera, L. hirsutum proved to be the least affec-
although a few studies have been under- ted by the leaf curl virus because it did not
taken on Coleoptera, Diptera and Acarina. support whitefly infestation to any extent.
Both antibiotic and anti-xenotic effects L. hirsutum differed from other species by
have been demonstrated. It is emphasized the presence of glandular trichomes desig-
that trichome-based host-plant resistance nated type VIc. Whiteflies became entrapped
could be utilized as a pest management in the exudate of type VIc trichome glands
tool; trichomes of wild species need to be before they could transmit the virus. Thus,
introgressed into the cultivated tomato. it may be possible to control tomato leaf
Hybrids between the cultivated tomato and curl virus transmission by breeding plants
the wild species Lycopersicon hirsutum with certain types of trichomes, especially
f. glabratum, Lycopersicon pennellii and trichome type VIc (Channarayappa et al.,
Lycopersicon cheesmanii f. minor have 1992).
been produced, which demonstrate useful
levels of resistance to Acarina, Diptera and Cuticle
Hemiptera pests, although these effects may
be tempered by effects on natural enemies Fruit anatomy and ultra-structure associated
(Simmons and Gurr, 2005). with cuticle cracking (CC) were compared
Vegetable Crops 211
cells are cutinized. Cuticle reduces the tran- Just beneath the hypodermis ground
spiration. Multicellular hairs are present on tissue is found, consisting of thin-walled
the epidermis. parenchymatous cells with well defined inter-
Multilayered (two to six layers) hypo- cellular spaces among them. Vascular bun-
dermis of collenchyma cells is found imme- dles are arranged in a half ring scattered in
diately beneath the epidermis. Collenchyma ground tissue.
is living mechanical tissue. The hypodermis The vascular bundles are of various sizes
gives considerable strength, flexibility and in the same petiole. Each vascular bundle
elasticity to young petioles. As it contains is wedge-shaped. In the petiole, the xylem
chloroplasts, it may carry out photosynthesis. is always found towards the upper side
whereas phloem occurs towards the lower
side (as in the leaf).
Hypodermis Epidermis
Vascular bundles
Seed anatomy
Ground tissue
In full-grown tomato seeds the embryo is
surrounded by a large quantity of endosperm
cells and by the testa or seed coat. The
embryos are bent and flattened. The seeds
are discoid in nature and a micropylar cap-
like organization having endosperm and
testa covers the radicle tip. This micropylar
cap is the place of radicle protrusion, but
there is no observable distinction between
Trichome testa rupture and endosperm rupture. Pro-
fuse hairs (Fig. 9.6) are present on the thin
testa, which help in absorption of water,
hold the water and make the water easily
Fig. 9.5. Transverse section of petiole showing accessible for the embryo. Macrosclereids
epidermis, trichomes and vascular system. Note: are round in shape and not very com-
vascular bundles arranged in the form of a ‘C’ shape. pactly arran ged, so water easily enters into
Testa
Hairs
Endosperm
Fig. 9.6. Transverse section of seed (100×) showing abundant hairs, testa and plentiful endosperm.
Vegetable Crops 213
the embryo, reducing the germination time anticancerous and many other properties.
of the seed; this is 36 h under laboratory Extensive research activities have been direc-
conditions (Maiti, unpublished). ted for improvement of yield and quality as
well as for controlling biotic and abiotic fac-
Research needs tors affecting the crop growth and produc-
tivity. Diploid chromosome number is 24
In the context of the literature review (2n = 2x = 24).
mentioned earlier it is assessed that ana-
tomical traits play an important role in
improving tomato for tolerance to various 9.2.2 Origin
insects, such as the presence and abundance
of glandular trichomes which exude toxic
Chilli peppers originated in the Americas.
materials inhibiting the attack of insects
After the Columbian Exchange, many culti-
and some disease resistance specially related
vars of chilli pepper spread across the
to cuticle thickness. Tomato production is
world, used in both food and medicine.
highly affected by these biotic stresses,
which need to be addressed effectively. There
exists a large variability among tomato culti-
vars in the intensity of these traits, thereby 9.2.3 Utilization
offering good scope for selection of culti-
vars for tolerance to these biotic stresses. Chilli peppers are used around the world to
Therefore, concerted research activities need make a countless variety of sauces, known
to be directed in this direction for genetic as hot sauce, chili sauce, or pepper sauce.
improvement of tomato for biotic stress The fruit is eaten raw or cooked for its fiery
factors, including virus that is transmitted hot flavour, concentrated along the top of
by whitefly. the pod. The stem end of the pod has most
It has been reported that the presence of the glands that produce the capsaicin.
of thick cuticle and compact palisade is The white flesh surrounding the seeds con-
related to drought resistance. In addition, tains the highest concentration of capsaicin,
the presence of thick cuticle and thick col- therefore removing the inner membranes is
lenchyma contributes to stiffness of the effective at reducing the heat of a pod.
petiole as well as drought resistance in Chilli is sold fresh, dried and powdered.
tomato cultivars. More research emphasis Keitel et al. (2001) recommended the
needs to be directed towards these aspects, usefulness of Capsicum pain plaster in chronic
and there is a requirement to select geno- non-specific low back pain. In patients with
types possessing these traits. osteoarthritis or fibromyalgia, it was obser-
ved that a Capsicum plaster preparation can
also be used to advantage in chronic non-
specific back pain.
9.2 Chilli Takashi et al. (2001) investigated the can-
cer chemopreventive activity of carotenoids
9.2.1 Introduction in the fruits of red paprika C. annuum L. and
observed the presence of capsanthin and
Capsicum peppers are widespread and the related carotenoids.
fruit contribute to burning and stinging of Gupta et al. (2002) reported a hypo-
hands, mouth and eyes in conjunction with cholesterolaemic effect of the oleoresin of
food preparation and ingestion (Fett, 2003). C. annuum L. in gerbils (Meriones hurrianae
Capsicum pepper, popularly known as chilli Jerdon). It prevented the accumulation of
(Capsicum annuum L.), has a great demand cholesterol and triglycerides in the liver and
throughout the world being a very important aorta. The faecal excretion of cholesterol and
condiment of high commercial value. It also has triglycerides were significantly increased in
medicinal values and contains antioxidants, oleoresin-fed gerbils.
214 Chapter 9
The plant is a mesophytic perennial herb The root anatomy in transverse section con-
with a taproot system, and fine lateral roots sists of distinct epidermis, cortex and vas-
spreading superficially (not deep penetrat- cular tissue (Fig. 9.8).
ing). The stem is aerial, erect, branched, The epidermis is uniseriate with a sin-
quadrangular and greenish (Fig. 9.7). The gle layer of compactly arranged thin-walled
leaves are simple, opposite and pubescent, living cells. The epidermal cells produce
obviate, with a wavy margin, acute apex, root hairs, which are useful for absorption
stipulate, petiolate (small), with reticulate of water.
venation. The cortex is multiseriate and relatively
homogeneous. It consists of general cortex
Floral characters and endodermis.
The general cortex consists of loosely
The flowers are small and white, bracte- arranged thin-walled parenchyma cells with
ate, pedicellate, complete, bisexual, pen- prominent intercellular spaces. The cells
tamerous and hypogynous. There are five are colourless and store starch and are usu-
sepals, which are gamosepalous and per- ally round or oval.
sistent, and there are five gamopetalous The endodermis is single layered with
petals. There are five stamens, epipetal- compactly arranged barrel-shaped cells. Endo-
ous, and the anthers are dithecous and dermal cells are characterized by the pres-
basifixed. The gynoecium is a bicarpel- ence of Casparian strips.
lary syncarpus superior ovary with exile The stele forms the central part of the
placentation. The fruit is a berry with a root and consists of pericycle, vascular
marcescent calyx. Seeds contain capsai- strands and conjunctive tissue. Pericycle is
cin, and are rounded in shape, flat, with a the outermost region of the stele and is
brown colour, smooth surface, and sized uniseriate and parenchymatous. Cells of
1.5–2.3 mm. the pericycle retain meristematic activity.
Fig. 9.7. Chilli plant morphology: quadrangular stem, leaves are simple, obviate, with a wavy margin
and acute apex, and flowers are small and white.
Vegetable Crops 215
Epidermis
Endodermis
Cortex
Secondary xylem
vessel Medullary ray
Fig. 9.8. Transverse section of mature root (100×) showing secondary growth. Ruptured epidermis,
compressed cortex, secondary phloem, secondary xylem, medullary rays and medulla are clearly visible.
Later roots arise endogenously from the The cortex consists of hypodermis,
pericycle. Vascular strands are radial, the middle cortex and endodermis. Hypodermis
xylem is exarch and closed type. The sec- is outermost region of the cortex and is mul-
ondary xylem of chilli root is composed of tilayered (two to six layers) and collenchy-
mostly isolated xylem vessels accompanied matic. Chlorenchyma is of the angular type
with medium thick-walled sclerenchyma and is living mechanical tissue imparting
giving strength to the root system. Conjunctive flexibility to the stem. The hypodermis
tissue is the non-vascular tissue present gives considerable strength, flexibility and
between the xylem and phloem strands. It elasticity to young stems, and, having chlo-
consists of medullary rays or pith rays, roplasts, it may carry out photosynthesis.
which help in the lateral conduction of The presence of a thick layer of collenchyma
water and store food material. offers strength, rigidity and drought resist-
ance. The middle cortex is multilayered and
Stem anatomy parenchymatic. The cells may be round or
oval with prominent intercellular spaces.
A transverse section of stem shows three This region also stores food materials tem-
regions, the epidermis, cortex and stele porarily. Endodermis forms the innermost
(Fig. 9.9). layer of the cortex, and is usually distinct
The epidermis is uniseriate consisting and without Casparian bands.
of compactly arranged barrel-shaped living The stele is the central region of the
cells. The outer walls of the epidermal cells stem and is thicker than the cortex. It consists
are cutinized; the cuticle reduces the tran- of pericycle, vascular bundles and medulla.
spiration. Few stomata are present in the Medullary rays are prominent structures in
epidermis for the exchange of gases and mature stems. Pericycle is the outermost non-
transpiration. All the epidermal cells are vascular region of the stele and is multiseriate
colourless except the guard cells. Trichomes and sclerenchymatic. The eustele is formed
are absent on the epidermis. from 15–20 vascular bundles arranged in
216 Chapter 9
Endodermis Epidermis
Collenchyma
Xylem Cortex
Phloem
one ring. Each vascular bundle is wedge- grown under 660 or 660/735 LED arrays.
shaped, conjoint, collateral, open and The effects of spectral quality on anatomical
endarch. Medulla is distinct, present at the changes in stem and leaf tissues of peppers
centre of the stem and is usually parenchyma- generally correlate to the amount of blue
tous. The cells are round or oval with inter- light present in the primary light source
cellular spaces. The medulla stores food (Schuerger et al., 1997).
materials and assists in lateral conduction.
It can be observed that in the secondary Leaf anatomy
xylem vessels are arranged in radial rings
interspersed with medullary rays and medium Leaf anatomy in transverse section shows
thick-walled sclerenchyma, offering strength epidermis, mesophyll and vascular tissue
to the stem. (Fig. 9.10).
It was reported that cross-sectional areas Epidermis is present on both surfaces of
of pepper stems, thickness of secondary the leaf blade, i.e. upper epidermis and lower
xylem, numbers of intra-xylary phloem bun- epidermis. The epidermis is single layered
dles in the periphery of stem pith tissues, with compactly arranged barrel-shaped (tab-
leaf thickness, numbers of chloroplasts per ular) cells. The outer walls of the epidermal
palisade mesophyll cell, and thickness of cells are cutinized. The outer surface of the
palisade and spongy mesophyll tissues were epidermis is covered by a continuous layer
greatest in peppers grown under MH (metal of cutin called cuticle. The cutinized outer
halide) lamps, intermediate in plants grown walls and cuticle reduce the loss of water due
under the 660/blue LED array, and lowest in to transpiration. Anisocytic type of stomata
peppers grown under the 660 or 660/735 LED are present in both the epidermal layers,
arrays. Most anatomical features of pepper but the stomata are usually more frequent
stems and leaves were similar among plants in the lower epidermis. Stomata facilitate
Vegetable Crops 217
Phloem
Lower epidermis
Spongy tissue
Collenchyma Parenchyma
the exchange of gases between the leaf and palisade and spongy tissue (median in posi-
environment. The stomatal index of upper tion), and are collateral and closed. In the
epidermis ranges from 11 to 13 and lower vascular bundle, xylem is present towards the
epidermis from 26 to 29. All the epidermal upper epidermis and phloem towards the lower
cells except guard cells are colourless. epidermis. The vascular bundle is surroun-
The ground tissue of the leaf, present ded by parenchymatous bundle sheath.
between the upper and the lower epidermal
layers, is called mesophyll. The mesophyll Trichome in relation to leaf curl virus
is chlorenchymatic and is concerned mainly
Studies revealed that negative association
with photosynthesis. The mesophyll is dif-
of trichome density and length with thrips
ferentiated into an upper palisade tissue and
and mites scored at flowering and maturity
lower spongy tissue. Palisade tissue is
and corresponding positive association with
present below the upper epidermis. The
dry chilli yield in all the populations, indi-
palisade cells are thin walled, cylindrical and
cating the role of trichomes in conferring
contain numerous chloroplasts. The cells
resistance to the leaf curl virus complex
are arranged compactly in one layer, per-
(Yadwad et al., 2008).
pendicular to the upper epidermis. Narrow
intercellular spaces are present in the pali- Petiole anatomy
sade tissue. Palisade tissue is the most
highly specialized type of photosynthetic The chief internal tissues of the petiole can
tissue. The upper surface of the leaf is dark be seen as four main divisions: the epider-
green in colour due to palisade tissue. Spongy mis, hypodermis, ground tissue and vascu-
tissue forms the lower part of the mesophyll lar bundles (Fig. 9.11).
present towards the lower epidermis. Cells The epidermis is uniseriate with a single
of the spongy tissue are thin-walled, irregu- layer of compactly arranged barrel-shaped
lar in shape, arranged loosely with large living cells. The outer walls of the epider-
continuous intercellular spaces. Stomata mal cells are cutinized. Cuticle reduces the
open into large intercellular spaces called transpiration.
sub-stomatal chambers. The cells of spongy Multilayered (two to six layers) hypodermis
parenchyma contain a lower number of of collenchyma cells is found immediately
chloroplasts, hence the lower surface of the beneath the epidermis. Collenchyma is living
leaf is light green in colour. mechanical tissue. It is annular collenchyma.
The vascular tissue occurs in the form Hypodermis gives considerable strength,
of discrete bundles called veins. Veins are flexibility and elasticity to young petioles
interconnected to form reticulate venation. and, having chloroplasts, it may carry out
Vascular bundles (veins) occur between the photosynthesis.
218 Chapter 9
Xylem
Epidermis
Phloem
Collenchyma
Trichomes
Ground tissue
Just beneath the hypodermis ground which delays the imbibition of water thereby
tissue is found. This consists of thin-walled delaying time of germination.
parenchymatous cells with well-defined inter-
cellular spaces among them. Significance of variations in anatomy
Vascular bundles are arranged in a half
ring scattered in ground tissue. The vascu- Future research may be directed in the fol-
lar bundles are wedge-shaped and of vari- lowing directions. The possession of a
ous sizes in the same petiole. In the petiole, higher number of trichomes gives disease
the xylem is always found towards the resistance to a plant. The presence of thick
upper side whereas phloem is found towards epicarp will help in long storage and fruit
the lower side (as in the leaf). borer resistance. A high amount of capsai-
cin at the placental region has more com-
mercial value in the market. Insignificant
Seed coat anatomy research has been directed on the use of
anatomical characters for stress resistance
The seed coat of chilli has thick testa in chilli, unlike in other vegetable crops.
(Fig. 9.12). The testa and tegman are not
clearly differentiated; the rate of imbibition
is slow due to the thick seed coat so time
taken for germination is 4–5 days under 9.3 Aubergine, Eggplant or Brinjal
30°C temperature in laboratory conditions.
It can be observed from the seed coat anat- 9.3.1 Introduction
omy of chilli that there is a thick cuticle
giving a glistening appearance to the seed Aubergine (also called eggplant or brinjal)
surface and the presence of a highly com- (Solanum melongena L.) belongs to the fam-
pact layer of rectangular macrosclereids, ily Solanaceae and is extensively distributed
Vegetable Crops 219
Testa
Tegman
Macrosclereids
Microsclereids
Endosperm
throughout the temperate and tropical century, Arabic traders introduced this veg-
regions. The fruit is a berry and the seeds etable in the West. China currently leads the
have a large endosperm and are grown world in aubergine production, followed by
mainly for food and medicinal purposes. India, Japan, Turkey and Egypt.
The aubergine is usually called brinjal in
India, and is an edible vegetable fruit of
high commercial value and demand in vari- 9.3.3 Utilization
ous countries and well adapted. Aubergine
(S. melongena) is consumed extensively in Fruits of aubergine are used as vegetables
Brazil and on the Indian subcontinent. It has for cooking different dishes all over the
various medicinal values, but aubergine pol- world. Several studies report the reduction
len is reported to cause allergic reactions of cholesterol with the extract or powdered
(Gill et al., 2002). The diploid chromosome infusion of aubergine fruit. There are also
number is 24 (2n = 2x = 24). reports on its effects on cholesterol metabo-
lism and its possible hypocholesterolaemic
effect. Flavonoids extracted from the fruits
9.3.2 Origin of S. melongena orally administered at a
dose of 1 mg/100 g BW/day showed signifi-
Aubergine is native to India and Sri Lanka cant hypolipidaemic action in normal and
and has been widely cultivated in all tem- cholesterol fed rats. HMG CoA reductase
perate regions of the world. They have also activity was found to be enhanced, while
grown in southern and eastern Asian coun- activities of glucose-6-phosphate dehydro-
tries since time immemorial. In the 16th genase and malate dehydrogenase reduced
220 Chapter 9
significantly. Silva et al. (1999) studied five gamopetalous petals. The flower has
the effect of aubergine extract on serum five epipetalous stamens, the anthers are
and hepatic cholesterol and triglyceride dithecous (four-chambered) and basifixed,
levels in adult rats. The results indicated and show porous dehiscence (anthers open
that the aubergine extract increased serum by means of apical pores). The gynoecium
but decreased hepatic cholesterol and had is bicarpellary syncarpous superior ovary,
little or no effect on both serum and hepatic with axile placentation. The fruit is a berry,
triglycerides. i.e. fleshy with marcescent calyx (no further
growth of calyx after fertilization). Seed
is rounded in shape, brown, with a smooth
and shining surface.
9.3.4 Morphological description
Fig. 9.13. Aubergine morphology: leaves are simple, ovate, stipulate, with a wavy margin and acute apex.
The inflorescence is auxiliary with a scorpioid cyme. The flowers are mostly pink in colour.
Vegetable Crops 221
Secondary xylem
Cortex
Cambium
Fig. 9.14. Transverse section of root showing secondary xylem and compressed endodermis and cortex due
to secondary growth achieved by the division of cambium in the stele.
Cortex forms the middle region, lying retain meristematic activity. Lateral roots
between the epidermis and the stele. Cortex arise endogenously from the pericycle,
is well developed and is thicker than the which help in absorption of water. During
stele. It is multiseriate and relatively homo- secondary growth, pericycle forms phello-
geneous and is differentiated into general gen or cork cambium and a small amount of
cortex and endodermis. General cortex is vascular cambium. Xylem and phloem are
extensively developed in the root and con- the vascular tissues. These are arranged in the
sists of loosely arranged thin-walled paren- form of separate strands on different radii,
chyma cells with prominent intercellular alternating with each other; hence the vas-
spaces and flavonoid deposits. The cells are cular strands are described as separate and
usually round or oval, are colourless and radial. Xylem is exarch, having protoxylem
store starch. This region helps in lateral towards the pericycle and metaxylem
conduction of water and salts. After second- towards the centre and scattered secondary
ary growth the general cortex compresses. xylem vessels. Cambium is absent and
The endodermis is a very distinct single hence vascular bundles are described as
layer with compactly arranged barrel-shaped closed type.
cells. Endodermal cells are characterized by
the presences of Casparian strips or Caspa- Secondary growth in root
rian bands made of suberin. In transverse
section, the Casparian bands appear as lens- During secondary growth, pericycle becomes
shaped thickening on the radial walls of meristematic and divides, giving rise to
the endodermis. Endodermis is also called cork cambium or phellogen. It produces a
the starch sheet layer, as starch grains are few brownish layers of cork cells or phel-
present in the endodermal cells. The endo- lem towards outside, and the phelloderm
dermis of the root acts as a barrier between on the inside. Secondary xylem and second-
the cortex and stele, and ruptures due to ary phloem are formed due to the activity of
secondary growth. the cambium ring (Fig. 9.14). After second-
The stele consist of pericycle, vascular ary growth the secondary vascular tissues
strands and pith. Pericycle forms the outer- form a continuous cylinder and usually the
most region of the stele and is uniseriate primary xylem becomes embedded in it. Pri-
and parenchymatous. Cells of the pericycle mary xylem located at the centre and primary
222 Chapter 9
phloem elements are generally seen in a sheath. The starch sheath layer is homologous
crushed condition. It is observed that the to endodermis, but morphologically unspe-
secondary xylem of root contains isolated cialized, hence such layer may be called
scattered larger xylem vessels. Pith is absent. endodermoid. After secondary growth the
The non-vascular tissue present between endodermis ruptures due to pressure cre-
xylem and phloem strands is called con- ated by inner secondary tissues.
junctive tissue, and it may be totally paren- The stele is the central region of the stem
chymatous or a part of it may be converted and is thicker than the cortex. It consists of
into sclerenchyma. During secondary growth pericycle, vascular bundles and medulla
the cambial cells that originate from the peri- and medullary rays. Pericycle (perivascular
cycle lying against the group of protoxylem region) forms the outermost non-vascular
function as ray initials and produce medul- region of the stele; it is multiseriate and
lary rays. These rays are transversed in the sclerenchymatic. Sclerenchyma is mechani-
xylem and phloem through cambium. cal tissue and gives strength and rigidity to
the stem. Xylem and phloem are organized
Stem anatomy into wedge-shaped vascular bundles and
from 10 to 20 vascular bundles are arranged
Transverse section of the primary stem in one ring consisting of xylem towards the
shows epidermis, cortex and stele. centre of the axis and phloem towards the
The epidermis is the outermost region periphery. In the vascular bundle xylem
surrounding the stem. It is uniseriate with a and phloem are present together (conjoint)
single layer of compactly arranged barrel- on the same radius (collateral), with a strip
shaped living cells. The outer walls of the of cambium (fascicular cambium) between
epidermal cells are cutinized and the epi- xylem and phloem (open), hence the vascu-
dermis is covered by a separate layer of lar bundle is described as conjoint, collateral
cutin called cuticle on its outer surface and open type. Xylem is endarch with pro-
(cuticularized), which reduces transpira- toxylem present towards the centre. Medulla
tion. Few stomata are present in the epider- and medullary rays are distinct, present at
mis for exchange of gases and transpiration. the centre of the stem and are usually
All the epidermal cells are colourless except parenchymatous. The cells are round or
the guard cells. Multicellular hairs are pre- oval with intercellular spaces. The exten-
sent on the epidermis. The epidermis ruptures sions of parenchymatous pith between vas-
after secondary growth and bark occupies the cular bundles are called primary medullary
place of epidermis. rays, which help in lateral conduction and
Cortex is the middle region present may give rise to secondary meristem (inter-
between the epidermis and stele, and is thin- fascicular cambium). The medulla stores
ner than the stele. It shows hypodermis, mid- food materials.
dle cortex and endodermis. The hypodermis
is the outermost region of the cortex and is Secondary growth in stem
multilayered (two to six layers) and collen-
chymatic. Hypodermis gives considerable The ever increasing shoot system requires
strength, flexibility and elasticity to young the supply of plentiful water and mineral salts
stems. The middle cortex is also multilay- and the root system requires much food
ered and parenchymatic. The cells may be material. Primary vascular tissues are inad-
round or oval with prominent intercellular equate to supply them. The vascular cambium
spaces. This region also stores food materials becomes active and produces secondary
temporarily and flavonoids are deposited. The vascular tissue (Fig. 9.15). During the sec-
endodermis forms the innermost layer of the ondary growth cambial strips are formed
cortex and is usually distinct in stems with- from medullary rays; these are called inter-
out Casparian bands. In young stems the inner- fascicular cambium. Fascicular cambium
most layer of the cortex contains abundant and inter-fascicular cambium combines and
starch, hence this layer is called the starch forms a cambial ring. The meristematic cells
Vegetable Crops 223
Epidermis
Primary phloem
Cortex
Cambium
Pith
Secondary xylem
Fig. 9.15. Transverse section of stem (100×) showing different parts. Secondary growth has been initiated
and cambium is forming secondary xylem.
of the cambial ring form secondary xylem cells and small pith were the main charac-
towards the inner side and a smaller amount ters of tolerant varieties. On the other hand,
of secondary phloem towards the outside. a thinner cuticle and collenchymatous area
Phellogen or cork cambium is formed from (hypodermis), loose parenchyma cells in
the cells of hypodermis or middle cortex. Cork the cortical region, larger spaces between
cambium forms secondary cortex towards vascular bundles, a smaller number of tri-
the inner side and cork tissue towards the chomes, soft parenchymatous cells in the inter-
outer side. A greater amount of cork tissue fascicular region, might be responsible for
is produced than the secondary cortex. the susceptibility to brinjal shoot and fruit
It can be observed from the transverse borer (Hossain et al., 2002).
section of stem that the presence of thick
cuticle on the stem surface and several lay- Leaf anatomy
ers of collenchyma give strength to the stem.
The secondary xylem contains radial rows Epidermis is mainly intended for protection
of vessels interspersed with medullary rays and is present on both surfaces of the leaf
and sclerenchyma. blade. Epidermis covering the upper surface
of the leaf is called upper epidermis or ven-
Shoot and fruit borer tral epidermis or adaxial epidermis. The epi-
dermis present towards the lower side of the
The presence of a thick cuticle, broad and leaf is called lower epidermis or dorsal epi-
thick collenchymatous area (hypodermis), dermis or abaxial epidermis. The epidermis
compact parenchyma cells in the cortical is single layered with compactly arranged
tissue, small area in the cortical tissues, barrel-shaped (tabular) cells. The outer walls
more vascular bundles with narrower spaces of the epidermal cells are cutinized and the
in the inter-fascicular region, and compact epidermis on its outer surface is covered by
arrangement of vascular tissue with lignified a continuous layer of cutin called cuticle.
224 Chapter 9
The cutinized outer walls and cuticle reduce resistance. The lower part of the mesophyll
the loss of water due to transpiration. present towards the lower epidermis is the
Anisocytic type of stomata are present in spongy tissue. These cells are thin-walled,
both of the epidermal layers, but stomata are irregular in shape, and arranged loosely with
usually more frequent in the lower epider- large continuous intercellular spaces. Stomata
mis. The stomatal index of the upper epider- open into large intercellular spaces called
mis is 21.3 and the lower epidermis is 29.6. sub-stomatal chambers. The cells of spongy
Stomata facilitate the exchange of gases bet- parenchyma contain a smaller number of
ween the leaf and environment. All the epi- chloroplasts, hence the lower surface of the
dermal cells except guard cells are colourless. leaf is light green in colour.
Epidermis contains multicellular hairs. The leaf receives the vascular supply
The ground tissue of the leaf, present from the stem. The vascular tissue occurs
between the upper and the lower epidermal in the form of discrete bundles called veins,
layers, is called mesophyll. The mesophyll which are interconnected to form reticu-
is chlorenchymatic and is concerned mainly late venation. Vascular bundles (veins)
with photosynthesis. The mesophyll is dif- occur between the palisade and spongy
ferentiated into an upper palisade tissue and tissue (median in position). They are col-
lower spongy tissue. The palisade tissue is lateral and closed. In the vascular bundle,
present below the upper epidermis (Fig. 9.16) xylem is present towards the upper epi-
and the cells are thin walled, cylindrical and dermis and phloem towards the lower epi-
contain numerous chloroplasts. The cells dermis. The vascular bundle is surrounded
are arranged compactly in one layer, per- by parenchymatous bundle sheath, the cells
pendicular to the upper epidermis. Narrow of which are called border parenchyma. The
intercellular spaces are present in the pali- larger vascular bundles are supported by
sade tissue. Palisade tissue is the most highly hypodermal parenchymatous strands. These
specialized type of photosynthetic tissue. strands are considered to be the sheath
The upper surface of the leaf is dark green in extension.
colour due to palisade tissue. Compactly The presence of thick cuticle and com-
arranged palisade tissue reduces the high pactly arranged palisade cells definitely
transpiration loss of water and offers drought reduces transpiration loss, which needs to be
assessed in aubergine in relation to drought-
resistant cultivars. No studies are available
Upper epidermis Palisade tissue in the literature.
The leaf area, leaf thickness, trichome
Collenchyma density and chlorophyll content were cor-
related with leafhopper and whitefly popu-
Xylem lation. The data indicate that leaf area, leaf
Phloem Spongy tissue thickness and chlorophyll content exerted
no effect on leafhopper population, while
trichome density had a negative correlation
(Naqvi et al., 2009).
Lower epidermis
Effect of chromium on leaf anatomy
Testa
Tegman
Macrosclereids
Microsclereids
Endosperm
Fig. 9.17. Transverse section of seed (400×) showing different parts. Note: microsclereids are rectangular
in shape.
pods are usually harvested at 2–4 day inter- eastward. The lack of a word for okra in
vals. The pods are harvested at certain the ancient languages of India suggests
intervals depending on marketable size and that it arrived there in the Common Era.
consumer’s preference. In general the pods The plant was introduced to the Americas
should be harvested when the pods are by ships plying the Atlantic slave trade by
4.5–7 cm in length. 1658, when its presence was recorded in
Brazil. It was further documented in Suriname
in 1686.
Okra may have been introduced to
9.4.2 Origin and Distribution
south-eastern North America in the early
18th century. It was being grown as far north
The species apparently originated in the as Philadelphia by 1748.
Ethiopian highlands, though the manner of
distribution from there is undocumented.
The Egyptians and Moors of the 12th and
13th centuries used the Arab word for the 9.4.3 Utilization
plant, suggesting that it had come from the
east. The plant may thus have been taken Okra is a popular and important food world-
across the Red Sea or the Bab-el-Mandeb wide. Its tender fruits are used in making
strait to the Arabian Peninsula, rather than many dishes. Okra gum is used industrially.
north across the Sahara. One of the earliest The yield in oil, the quality of its proteins
accounts is by a Spanish Moor who visited and the use of the stem in paper-making
Egypt in 1216, who described the plant reveal that okra has economic potential for
under cultivation by the locals who ate the cultivation. Okra fruit rhamno-galacturon-
tender, young pods with meals. ans increased cell proliferation.
From Arabia, the plant spread around In Asian medicine the fruit of the okra
the shores of the Mediterranean Sea and plant A. esculentus (L.) Moench is used as a
Vegetable Crops 227
mucilaginous food additive against gastric sepals) sepals, which are bowl shaped
irritative and inflammatory diseases. with trichiferous nectar present at the base
of calyx. There are five petals, polypetal-
ous, large and attractive. The stamens are
9.4.4 Morphological description numerous and monodelphous and the
kidney-shaped anthers are monothecous.
Vegetative characters The gynoecium is multicarpellary, syncar-
pus, with a superior ovary and axile pla-
The plant is a mesophytic herb with a tap- centation. The number of stigma is equal to
root system with inclined deep lateral roots. number of locules in the ovule. The fruit is
The stem is aerial, erect, branched, and stel- a loculicidal capsule and seeds are round,
late hairs are present. The leaves are alter- grey-ash colour with a rough surface, and
nate, pubescent, simple, heart shaped, with the seed size is from 2.5 mm to 3 mm.
a dentate margin and obtuse apex, stipulate,
petiolate and have palmately reticulate
venation (Fig. 9.18).
9.4.5 Anatomical description
Floral characters
Root anatomy
Flowers are born on solitary inflorescence
The epidermis is uniseriate with a single
and are bracteate, pedicellate, with the pre-
layer of compactly arranged thin-walled liv-
sence of an epicalyx, complete, bisexual,
ing cells. Epidermal cells produce root hairs,
pentamerous and hypogynous, actinomor-
which are useful for absorption of water.
phic. There are five gamosepalous (united
The cortex is multiseriate, relatively
homogeneous and consists of general cortex
and endodermis. The endodermis is single
layered with compactly arranged barrel-
shaped cells. Endodermal cells are charac-
terized by the presences of Casparian strips.
Stele is the central part of the root and
consists of pericycle, vascular strands and
conjunctive tissue. The pericycle is the out-
ermost region of the stele and is uniseriate
and parenchymatous. Cells of the pericycle
retain meristematic activity. Later roots
arise endogenously from the pericycle. The
vascular strands are radial and the xylem
is exarch, closed type. The non-vascular
tissue present between xylem and phloem
strands is called conjunctive tissue. Pith is
absent. Secondary tissues are formed by the
activity of cambium causing the primary
structure of the root to lose its initial appear-
ance in transverse section (Fig. 9.19).
Stem anatomy
Cortex
Epidermis
Fig. 9.19. Transverse section of root (100×) showing secondary growth and various parts including
epidermis, cortex, secondary xylem and medullary rays.
reduces the transpiration. Few stomata are Medulla stores food materials and helps in
present in the epidermis for exchange of lateral conduction.
gases and transpiration. All the epidermal During the secondary growth primary
cells are colourless except the guard cells. phloem, endodermis and cortex are crushed
Stellate, tufted trichomes are present on the due to pressure created by inner secondary
epidermis, which give insect resistance. tissues (Fig. 9.20).
The hypodermis is the outermost region
of the cortex. It is multilayered (two to six Leaf anatomy
layers) and collenchymatic. Collenchyma is
living mechanical tissue. The hypodermis The epidermis is present on both surfaces of
give considerable strength, flexibility and the leaf blade, i.e. upper epidermis and
elasticity to young stems. As it contains lower epidermis. Epidermis is mainly meant
chloroplasts, it may carry out photosynthe- as a protective layer. The epidermis is single
sis. The endodermis is innermost layer of layered with compactly arranged barrel-
the cortex. It is usually distinct in stems shaped (tabular) cells. The outer walls of the
without Casparian bands. epidermal cells are cutinized. The cutinized
Pericycle is the outermost non-vascular outer walls and cuticle reduce the loss of
region of the stele, and is multiseriate and water due to transpiration. The stomatal
sclerenchymatic. Each vascular bundle is index of the upper epidermis is 25.9 and of
wedge-shaped, conjoint, collateral, open and the lower epidermis is 31.3. Anisocytic sto-
endarch. Medulla and medullary rays are mata (Fig. 9.21) are present in both of the
distinct, present at the centre of the stem, epidermal layers. Stellate, tuft trichomes
and are usually parenchymatous. The cells are present on the epidermis. All the epider-
are round or oval with intercellular spaces. mal cells except guard cells are colourless.
Vegetable Crops 229
Epidermis
Primary phloem
Cortex
Secondary phloem
Cambium
Secondary xylem
Fig. 9.20. Transverse section of stem (100×) showing secondary growth. The primary phloem
and endodermis is crushed due to pressure created by inner secondary tissues.
Upper epidermis
Cuticle
Palisade tissue
Xylem
Spongy tissue
Phloem
Collenchyma
Parenchyma
Lower epidermis
Cortex Epidermis
Collenchyma
Phloem
Endodermis
Xylem
Fig. 9.23. Transverse section of petiole (100×) showing epidermis, collenchyma and vascular bundles.
9.5.3 Utilization
that of the lower epidermis is 28.98. The bundles occur between the palisade and
foot cell of the trichome and subsidiary cell spongy tissue (median in position). They are
surrounding the foot cell contains cystolith. collateral and closed. In the vascular bundle,
Trichomes are of the glandular type. xylem is present towards the upper epider-
The mesophyll is differentiated into an mis and phloem towards the lower epider-
upper palisade tissue and lower spongy tis- mis. The vascular bundle is surrounded by a
sue. The palisade tissue is present below the parenchymatous bundle sheath.
upper epidermis. The palisade cells are thin
walled, cylindrical and contain numerous Petiole anatomy
chloroplasts. The cells are compactly arranged
in one layer, perpendicular to the upper epi- The epidermis is uniseriate, with a single
dermis. Narrow intercellular spaces are layer of compactly arranged barrel-shaped
present in the palisade tissue. Palisade tissue living cells. The outer walls of the epider-
is the most highly specialized type of photo- mal cells are cutinized. Multicellular hairs
synthetic tissue. The upper surface of the leaf are present on the epidermis. A multilayered
is dark green in colour due to palisade tissue. (two to six layers) hypodermis of angular
Spongy tissue forms the lower part of the collenchyma cells is found immediately
mesophyll present towards the lower epi- beneath the epidermis (Fig. 9.27). Collen-
dermis. Cells of the spongy tissue are thin- chyma is living mechanical tissue. The
walled, irregular in shape and arranged loosely hypodermis gives considerable strength,
with large continuous intercellular spaces. flexibility and elasticity to young petioles.
The cells of spongy parenchyma contain fewer As it contains chloroplasts, it may carry out
chloroplasts, hence the lower surface of the photosynthesis.
leaf is light green in colour. Just beneath the hypodermis ground
The vascular tissue occurs in the form of tissue is found. This consists of thin-walled
discrete bundles called veins. Veins are inter- parenchymatous cells with well-defined
connected to form reticulate venation. Vascular intercellular spaces among them. Vascular
Epidermal hair
Epidermis
Hypodermis
Cortex
Phloem
Xylem
Pith
bundles are arranged in a half ring scattered parthenocarpic ovules but was lost from the
in ground tissue. Bicollateral vascular bun- integuments of the unpollinated ovules by
dles are arranged in two rows. day 6. Pollinated and parthenocarpic ovules
The central part of petiole is hollow, contributed increasingly to fruit dry weight
which is an important character of Cucur- over the 9-day period.
bitaceae members. The ovule tissues, in particular the nucel-
lus and integument, may exert control over
Anatomical changes in stigma after pollination early development in both pollinated and
parthenocarpic fruits (Sedgley et al., 1977).
The following changes were observed on
stigma. The wall thickenings of the papilla
Anatomical changes under flooding stress
transfer cells contained callose and their
presence prior to pollination was confirmed There are morpho-anatomical changes in
using EM-autoradiography, freeze-fracture flooding lines comparing with control. The
and fixation. No further callose thickenings obtained results revealed that growing
were produced following pollination. Pol- watermelon on land flooded with a 2-week
lination resulted in a rapid increase in aque- period was the best. Parameters of plants in
ous stigma secretion and localized disruption this treatment exceeded those of unflooded
of the cuticle, which appeared to remain on plants in plant length (47.3%), plant stem
the surface of the secretion. The reaction diameter (26.9%), leaf area (43.7%), leaf
was localized to the papilla cells adjacent to area index (82.8%), fruit number per plant
the pollen tube only. Both pollen-grain wall (30%), yield (48.8%), number of stomata/
and stigma secretion contained proteins, mm2 (18.3%), number of cells/mm2 (24.3%)
carbohydrates, acidic polysaccharides, lip- and width of second trichome cell (64.4%),
ids and phenolics (Sedgley, 1982). narrowest vessel diameter (28.75%). More-
over, this treatment revealed superiority
Anatomical changes during fruit development over a 1-week flooding period in stem dry
matter percentage (26.9%), narrowest width
Parthenocarpic fruits were larger and had of vessel (27.84%), highest vessel width
higher fresh weight and percentage water (12.12%). A 2-week flooding period also
than pollinated fruits at day 1 but the posi- overwhelmed that of a 3-week period in fruit
tions were reversed by day 9. Unpollinated number per plant (26.2%), width of narrow-
fruits did not increase in size after day 3. est vessel (12.84%) and width of hair second
Pericarp cells were small, of regular shape cell (43.7%)(Abdel and Bamerni, 2011).
and showed no obvious change with either
time or treatment. Cell number increased in Anatomy of embryo sac
the pollinated and parthenocarpic but not
in the unpollinated fruits. Cells divided in In this study embryo sacs from watermelon
the flesh of the parthenocarpic but not of were observed over a 13-day period follow-
the pollinated fruits, which increased in size ing flowering with: (i) normal pollination;
by cell enlargement only. Starch, present in (ii) non-pollination; and (iii) induction of
the cells of the flesh and placenta at day 0, parthenocarpic fruit development with naph-
was absent from the unpollinated fruits at day 6. thalene acetic acid. With normal pollination,
Ovules grew in both pollinated and parthe- approximately 2 days after fertilization the
nocarpic fruits largely due to cell division embryo sacs completed development and
in the nucellus and integuments; the polli- consisted of two synergids with prominent
nated ovules were larger than the partheno- filiform apparatus, an egg cell, a central cell
carpic throughout. Embryo and endosperm with two polar nuclei and three antipodal
development occurred in the pollinated but cells. Sperm nuclei were observed within
not in the parthenocarpic ovules. Starch the embryo sac at 2 days and by 4 days the endo-
was present throughout the 9-day period sperm was proliferating. In the non-pollination
in the integuments of the pollinated and treatment the embryo sac was still intact
236 Chapter 9
after 4 days although the antipodal nuclei dried and cored thick outer skin has tradi-
were becoming hard to distinguish. By 7 tionally been used to make musical instru-
days only the two synergids and the egg cell ments like the tanpura. The baul singers of
were still well defined, the polar nuclei Bengal (India) and Bangladesh have their
appeared in some preparations to be fused, musical instruments made out of it. The
and the antipodals had degenerated. By 10 practice is also common among Buddhist
days the embryo sac was a structure-less and Jain sages. In Vietnam, it is used in a
watery mass. In parthenocarpic fruit the variety of dishes: boiled, stir-fried, soup
fate of the embryo sac was similar to that in dishes and as a medicine. The shoots, ten-
non-pollinated fruit except that final break- drils and leaves of the plant may also be
down was delayed past 10 days (Buttrose eaten as greens. The juice of bottle gourd is
and Sedgley, 1979). considered to have many medicinal proper-
ties and is very good for health.
Significance variations in anatomy
Stem anatomy
Epidermis
Cortex
Primary xylem
Secondary
Medullary ray xylem
Fig. 9.29. Transverse section of root (100×) showing epidermis, cortex, primary xylem, secondary xylem
and medullary rays.
Epidermis
Glandular trichome
Collenchyma
Outer phloem
Outer cambium
Xylem
Inner cambium
Endodermis
Inner phloem Cortex
Fig. 9.30. Transverse section of stem (100×) showing epidermis, glandular trichomes, collenchyma, cortex
and bicollateral vascular bundles.
Vegetable Crops 239
cells. The outer walls of the epidermal cells The vascular tissue occurs in the form
are cutinized and covered by a continuous of discrete bundles called veins, which are
layer of cuticle. Anamocytic stomata are interconnected to form reticulate venation.
present in both of the epidermal layers. The Vascular bundles (veins) occur between
stomatal index of the upper epidermis is the palisade and spongy tissue (median in
11.76 and of the lower epidermis is 19.39. position) and are collateral and closed. In
The foot cell of the trichome and the sub- the vascular bundle, xylem is present
sidiary cell surrounding it contain cysto- towards the upper epidermis and phloem
lith. Trichomes are of the glandular type towards the lower epidermis. The vascular
(Fig. 9.31). bundle is surrounded by parenchymatous
The mesophyll is differentiated into an bundle sheath.
upper palisade tissue and lower spongy tis-
sue. The palisade tissue is present below the Petiole anatomy
upper epidermis; the cells are thin walled,
cylindrical and contain numerous chloro- The epidermis is uniseriate with a single
plasts. The cells are arranged compactly in layer of compactly arranged barrel-shaped
one layer, perpendicular to the upper epi- living cells. The outer walls of the epider-
dermis. Narrow intercellular spaces are mal cells are cutinized. Cuticle reduces
present in the palisade tissue. Palisade tis- transpiration. Multicellular hairs are present
sue is the most highly specialized type of on the epidermis.
photosynthetic tissue. The upper surface of Multilayered (from two to six layers)
the leaf is dark green in colour due to pali- hypodermis of collenchyma cells is
sade tissue. The lower part of the mesophyll found immediately beneath the epidermis
present towards the lower epidermis is the (Fig. 9.32). Collenchyma is living mech-
spongy tissue. Cells of the spongy tissue are anical tissue and is of the angular type.
thin walled, irregular in shape, and arranged Hypodermis gives considerable strength,
loosely with large continuous intercellular flexibility and elasticity to young petioles
spaces. The cells of spongy parenchyma and, having chloroplasts, it may carry out
contain fewer chloroplasts. photosynthesis.
Upper epidermis
Glandular trichome
Cuticle
Palisade tissue
Lower epidermis
Xylem
Phloem
Spongy tissue
Collenchyma Parenchyma
Fig. 9.31. Transverse section of leaf (100×) showing epidermis, glandular trichomes, palisade and spongy
tissue and vascular bundles.
240 Chapter 9
Collenchyma
Glandular trichome
Ground tissue
Outer phloem
Xylem
Outer cambium
Inner phloem Inner cambium
Fig. 9.32. Transverse section of petiole showing glandular trichomes on epidermis, collenchyma
and bicollateral vascular bundles.
cucurbitaceous vegetable in spite of its consi- treat malaria and diabetes. The leaves are
derable nutritional and medicinal properties. allowed to steep in hot water before being
Monoecious bitter gourd (M. charantia strained thoroughly so that only the remain-
L. var. minima and maxima Williams & Ng), ing liquid is used for the tea.
a cucurbit of major economic importance,
is widely cultivated in India, China, Africa
and South America. The morphology (i.e. 9.7.4 Morphological description
growth habit and fruit shape, size, colour
and surface texture) of Indian bitter gourd
Vegetative characters
is diverse where gynoecious sexual forms
exist. The plant is a mesophytic herb, climbing,
Momordica charantia is a tropical and (with tendrils) highly pubescent and suc-
subtropical species belonging to the fam- culent. It has a taproot system with super-
ily Cucurbitaceae, and is widely grown for ficially spreading and not profuse roots.
its edible fruit, which is among the most The stem is aerial, weak, succulent, angu-
bitter of all fruits. Various names exist for lar, branched, highly pubescent and green-
the plant and its fruit, including bitter ish, with tendrils developed from the leaf
melon, bitter gourd, goya from the Japanese axial. The leaves are alternate, petiolate,
or Karela/Karella, ampalayá from Tagalog, (pubescent), stipulate, simple, palmately
and cerasee (Caribbean and South America; lobed (seven lobes) and heart shaped, with
also spelled cerasse). a serrate margin, acuminate apex and reti-
culate venation (Fig. 9.34).
9.7.2 Origin
9.7.3 Utilization
Epidermis Cortex
Primary xylem
Medullary ray
Secondary xylem
Fig. 9.35. Transverse section of root (100×) showing epidermis, cortex, primary xylem, secondary xylem
and medullary ray.
Vegetable Crops 243
Epidermis
Collenchyma
Cortex
Outer phloem
Outer cambium
Xylem
Inner cambium
Inner phloem
Fig. 9.36. Transverse section of stem (100×) showing epidermis, collenchyma, cortex and bicollateral
vascular bundles.
244 Chapter 9
Xylem
Spongy tissue
Lower epidermis
Palisade tissue is the most highly specialized layers) hypodermis of angular collenchyma
type of photosynthetic tissue. The upper cells is found immediately beneath the epi-
surface of the leaf is dark green in colour dermis. Collenchyma is living mechanical
due to palisade tissue. The lower part of tissue. The hypodermis gives considerable
the mesophyll present towards the lower strength, flexibility and elasticity to young
epidermis is the spongy tissue. Cells of the petioles, and, having chloroplasts, it may
spongy tissue are thin walled, irregular in carry out photosynthesis.
shape, arranged loosely with large continu- Just beneath the hypodermis ground
ous intercellular spaces. Stomata open into tissue is found (Fig. 9.38). It consists of thin
large intercellular spaces called sub-stomatal walled parenchymatous cells with well-
chambers. The cells of spongy parenchyma defined intercellular spaces among them.
contain a lower number of chloroplasts, Vascular bundles are arranged in a half ring
hence the lower surface of the leaf is light scattered in ground tissue.
green in colour. Bicollateral vascular bundles are arran-
The vascular tissue occurs in the form ged in two rows. The centre part of petiole
of discrete bundles called veins. Veins are is hollow, which is an important character
interconnected to form reticulate venation. of Cucurbitaceae family members.
Vascular bundles (veins) occur between
the palisade and spongy tissue (median in Seed coat anatomy
position) and are collateral and closed. In
the vascular bundle, xylem is present The seed coat is thick and consists of hairs.
towards the upper epidermis and phloem Macrosclereids are rectangular in shape and
towards the lower epidermis. The vascular very compactly arranged.
bundle is surrounded by parenchymatous
bundle sheath.
9.7.6 Fruit development
Petiole anatomy
Fruit development in bitter gourd depends
The epidermis is uniseriate with a single on temperature, and early development is
layer of compactly arranged barrel-shaped observed in summer compared to winter. It
living cells. The outer walls of the epider- took 12–16 days for harvest, with fruits turn-
mal cells are cutinized. Cuticle reduces ing yellow in 16 days in Taiwan, whereas it
transpiration. Multicellular hairs are present took 20–22 days for harvest under lower
on the epidermis. Multilayered (two to six temperatures in the cold growth season of
Vegetable Crops 245
Epidermis Collenchyma
Outer phloem
Outer cambium Ground tissue
Xylem
Inner cambium
Inner phloem
Fig. 9.39. Cucumber plant morphology: plant is a climbing (with tendrils) herb and pubescent. The stem
is aerial, weak, succulent, angular and branched. The leaves are simple, heart shaped, with a serrate margin.
Auxiliary inflorescence with the male and female flowers arranged in racemose.
The flower is mostly yellow in colour, ebrac- general cortex consists of loosely arranged
teate, unisexual, pentamerous and hypogy- thin-walled parenchyma cells with promi-
nous. There are five gamosepalous sepals; in nent intercellular spaces (Fig. 9.40), which
the male flower these are bell shaped, in the is compressed due to secondary growth. The
female flower the calyx tube is connate with cells are usually round or oval, colourless and
the ovary and pubescent. There are five free, store starch. This region helps in the lateral
yellow and pubescent petals. The male flower conduction of water and salts. Endodermis
has five syandrous stamens and the anthers are forms the innermost layer of the cortex and is
dithecous (four-chambered); the presence of a very distinct in roots. It is single layered, with
pistillode is notable. The female flower has compactly arranged barrel-shaped cells. Endo-
three to five staminodes present on the calyx dermal cells are characterized by the presence
edges. The gynoecium is unilacular, tricarpel- of Casparian strips or Casparian bands. After
lary syncarpous and with parietal placentation. secondary growth the general endodermis rup-
The fruit is a pepo, green to light yellow in tures. Endodermis of the root acts as barrier
colour and oblong to round. between the cortex and stele.
The stele consists of pericycle, vascular
strands and pith. Xylem and phloem form the
9.8.4 Anatomical description vascular tissues. Xylem is meant for conduc-
tion of water and salts and phloem for the
Root anatomy transport of food materials. The xylem is
exarch, with protoxylem towards the pericy-
Epidermis is the outermost region of the cle and metaxylem towards the centre and
root and is uniseriate, with a single layer of scattered secondary xylem vessels. Cambium
compactly arranged thin-walled living cells is absent and hence vascular bundles are
consisting of root hairs. The epidermis rup- described as closed type. Pith is absent. The
tures due to secondary growth. non-vascular tissue present between xylem
The cortex is the middle region, lying and phloem strands is called conjunctive
between the epidermis and the stele. Cortex tissue and may be totally parenchymatous
is well developed, multiseriate and relatively or a part of it may be converted into scleren-
homogeneous and is thicker than the stele. It con- chyma. During secondary growth the cam-
sists of general cortex and endodermis. The bial cells that originate from the pericycle
Vegetable Crops 247
Epidermis Cortex
Primary
xylem
lying against the group of protoxylem func- having chloroplasts, it may carry out photo-
tion as ray initials and produce medullary synthesis. The middle cortex is also multi-
rays. These rays are transversed in the xylem layered and parenchymatic. The cells may
and phloem through cambium; this is a be round or oval with prominent intercel-
characteristic feature of the roots. lular spaces. This region also stores food
materials temporarily. Endodermis is inner-
most layer of the cortex and is usually dis-
Stem anatomy
tinct in stems without Casparian bands.
The epidermis is the outermost region The stele is the central region of the
surrounding the stem and is uniseriate, with stem and is thicker than the cortex. It con-
a single layer of compactly arranged barrel- sists of pericycle, vascular bundles, and
shaped living cells. The outer walls of the medulla and medullary rays. Pericycle is the
epidermal cells are cutinized. Cuticle reduces outermost non-vascular region of the stele
the transpiration. Few stomata are present and is multiseriate and sclerenchymatic.
in the epidermis for the exchange of gases Bicollateral vascular bundles are arranged
and transpiration. All the epidermal cells are in two rows. Each vascular bundle is wedge-
colourless except the guard cells. Multi- shaped, conjoint, open and endarch. The
cellular hairs are present on the epidermis. central part of the stem contains a hollow
The cortex is the middle region present region, which is an important character of
between the epidermis and stele and is thin- Cucurbitaceae family members.
ner than the stele. In section, it shows hypo-
dermis, middle cortex and endodermis. Leaf anatomy
Below the ridges from three to four layers of
angular collenchyma are present (Fig. 9.41). Epidermis is present on both surfaces of the
Collenchyma is living mechanical tissue. leaf blade, i.e. upper epidermis and lower
The hypodermis gives considerable strength, epidermis. The epidermis is single layered, with
flexibility and elasticity to young stems and, compactly arranged barrel-shaped (tabular)
248 Chapter 9
Epidermis
Collenchyma
Cortex
Outer phloem
Outer cambium
Xylem
Inner cambium
Inner phloem
Fig. 9.41. Transverse section of stem (100×) showing epidermis, collenchyma, cortex and bicollateral
vascular bundles.
cells. The outer walls of the epidermal cells Glandular trichome Cuticle Palisade tissue
are cutinized and covered by a continuous Upper epidermis
layer of cuticle. Anamocytic stomata are
present in both the epidermal layers. The
foot cell of the trichome and the subsidiary
Spongy
cell that surrounds the foot cell of the tri-
tissue
chome contains a cystolith. Trichomes are of
Xylem
the glandular type. The stomatal index of the Lower
upper epidermis ranges from 18 to 23 and Phloem epidermis
that of the lower epidermis from 26 to 28.
The ground tissue of the leaf, present
between the upper and the lower epidermal
Parenchyma
layers, is called mesophyll. The mesophyll Collenchyma
is chlorenchymatic and is concerned mainly
with photosynthesis. The mesophyll is dif-
ferentiated into upper palisade tissue and
lower spongy tissue. Fig. 9.42. Transverse section of leaf (100×) showing
The palisade tissue is present below the epidermis, palisade and spongy tissue and vascular
upper epidermis. The palisade cells are thin bundle.
walled, cylindrical, contain numerous chlo-
roplasts and are compactly arranged in one The lower part of the mesophyll pre-
layer, perpendicular to the upper epidermis. sent towards the lower epidermis is the
Narrow intercellular spaces are present in spongy tissue (Fig. 9.42). Cells of the spongy
the palisade tissue. Palisade tissue is the most tissue are thin-walled, irregular in shape
highly specialized type of photosynthetic and arranged loosely with large continuous
tissue. The upper surface of the leaf is dark intercellular spaces. Stomata open into large
green in colour due to palisade tissue. intercellular spaces called sub-stomatal
Vegetable Crops 249
chambers. The cells of spongy parenchyma that a high density of glandular trichomes
contain fewer chloroplasts, hence the lower and chemicals secreted by them deter A.
surface of the leaf is light green in colour. gossypii and disturb aphid settling on aphid-
The vascular tissue occurs in the form of resistant varieties (Sarria et al., 2010).
discrete bundles called veins. Veins are
interconnected to form reticulate venation. Leaf senescence
Vascular bundles (veins) occur between the
palisade and spongy tissue (median in posi- Detached cucumber (Cucumis sativa L.)
tion). They are collateral and closed. In the leaves show senescence and rejuvenation
vascular bundle, xylem is present towards after cultivation in nutrient solution for 4
the upper epidermis and phloem towards the weeks. Rooting of the petiole elicited a
lower epidermis. The vascular bundle is sur- combination of different morphological,
rounded by parenchymatous bundle sheath. anatomical and physiological changes in
the lamina. Extensive growth in area and
Glandular trichomes – insect resistance thickness, changes of chloroplast structure
and activity, as well as the pattern of
The leaf epidermis and their effects on beha- Chl-protein complexes were observed and
viour of Aphis gossypii Glover were evalu- compared either to the corresponding
ated in two Cucumis melo L. genotypes parameters of young detached leaves. The
(aphid susceptible, aphid resistant). No dif- hormones showed mutuality in their
ferential effects of epicuticular waxes on effects, benzyladenine being responsible
aphid behaviour were observed. The type, for the growth of cells, while indolylacetic
distribution and number of trichomes on acid and kinetin promoted an increase in
melon leaves were also studied. Pubescence chlorophyll content (Kovács et al., 2007).
in melon, measured as the number of non-
glandular trichomes per square centimetre, Petiole anatomy
was not sufficient to prevent aphid settling.
There was a high density of type I glandu- The epidermis is uniseriate, with a single
lar trichomes on leaves of the aphid- layer of compactly arranged barrel-shaped
resistant genotype. These results indicate living cells (Fig. 9.43). The outer walls of
Epidermis
Collenchyma
Ground tissue
Vascular bundle
Fig. 9.43. Transverse section of petiole (100×) showing epidermis, collenchyma, ground tissue
and bicollateral vascular bundles.
250 Chapter 9
the epidermal cells are cutinized. Cuticle bundle to fork. A large schizo-lysigenous
reduces the transpiration. Multicellular cavity develops in the perianth tube,
hairs are present on the epidermis. extending into the connective and sta-
Multilayered (two to five layers) men prolongations. Stomata are found on
hypodermis of angular collenchyma cells the outer surfaces of the perianth tube
is found immediately beneath the epider- and the calyx lobes. The corolla seems
mis. Collenchyma is living mechanical tis- regularly free from stomata except at the
sue. The hypodermis (collenchyma) gives extreme tips of the lobes. The mesophyll
considerable strength, flexibility and elas- of the corolla is undifferentiated
ticity to young petioles and, having chlo- (Heimlich, 1927).
roplasts, it may carry out photosynthesis.
Just beneath the hypodermis ground tis- Seed coat anatomy
sue is found, which consists of thin-
walled parenchymatous cells having The seed coat is very thick; the testa is thick
well-defined intercellular spaces among but the macrosclereids are not very com-
them. Vascular bundles are arranged in a pactly arranged, so water enters into the
half ring scattered in ground tissue. seeds resulting in a low germination time,
Bicollateral vascular bundles are arranged i.e. 24 h (Fig. 9.44).
in two rows. The centre part of the petiole
contains a hollow region, which is an Significance variations in anatomy
important character of Cucurbitaceae
members. Future research may be directed in the fol-
lowing directions. The possession of a
higher number of multicellular trichomes
Development and anatomy gives disease resistance to insects. The epi-
of the staminate flower carp and pericarp is thick, so is helpful in
All organs arise in a low spiral arrange- long storage and fruit borer resistance. The
ment; the staminate flower produces presence of silica may provide resistance to
three stamens. The morphological nature powdery mildew.
of these is uncertain. Heimlich (1927)
inclines to the view that two of them are
normal, complete stamens, and the odd Testa
one a stamen in which one theca fails to
Macrosclereids
develop. One large bundle enters the base
of each complete stamen. This is a dou-
Microsclereids
ble bundle in the sense that an original
bundle coming from the vascular plate in
Tegman
the receptacle forks and then very soon
reunites its two branches. The large bun-
dle sends off a right and a left branch in
the broad connective. Each branch forks
Embryo
sharply to give off one ascending and one
descending branch. The main bundle
continues above the right and left
branches and ultimately forks again,
sending a branch into each of the two sta-
men prolongations. The odd stamen is
similar to the complete stamens except
that the whole vascular distribution is Fig. 9.44. Transverse section of cucumber seed
exactly one-half that of the complete showing inner embryo covered by outer testa and
stamen. This difference is apparently due inner tegman. Macrosclereids are almost rectangular
to the failure of the respective original in shape, microsclereids are small dot-like structures.
Vegetable Crops 251
Epidermis
Macrosclereids
Microsclereids
Vascular bundle
Tegman
Fig. 9.47. Transverse section of petiole (100×)
showing epidermis, collenchyma, ground tissue Fig. 9.48. Transverse section of seed coat showing
and vascular bundles. micro- and macrosclereids.
254 Chapter 9
Stem
Epidermis
Secondary
Cortex xylem
Primary
xylem
Medullary ray
Fig. 9.50. Transverse section of root (100×) showing epidermis, cortex, primary and secondary xylem
and medullary rays.
Epidermis
Cortex Collenchyma
Vascular
bundle
Fig. 9.51. Transverse section of stem (100×) showing epidermis, collenchyma and vascular bundles.
256 Chapter 9
The middle cortex is also multilayered cells. The outer walls of the epidermal cells
and parenchymatic. The cells may be round are cutinized and covered by a continuous
or oval with prominent intercellular spaces. layer of cuticle. The stomatal index of upper
This region also stores food materials tem- epidermis is 10.52 and that of the lower
porarily. Endodermis is innermost layer of epidermis is 14.6. Anamocytic stomata are
the cortex and is usually distinct in stems present in both of the epidermal layers. The
without Casparian bands. foot cell of the trichome and the subsidiary
Stele is the central region of the stem cell surrounding it contain cystoliths. The
and is thicker than the cortex. It consists of trichomes are of the glandular type.
pericycle, vascular bundles and medulla The ground tissue of the leaf, present
and medullary rays. Pericycle is the outer- between the upper and the lower epidermal
most non-vascular region of the stele, and is layers, is called mesophyll. The mesophyll
multiseriate and sclerenchymatic. Bicolla- is chlorenchymatic and is concerned mainly
teral vascular bundles are arranged in two with photosynthesis. The mesophyll is dif-
rows (Fig. 9.51). Each vascular bundle is ferentiated into an upper palisade tissue
wedge-shaped, conjoint, open and endarch. and lower spongy tissue (Fig. 9.52).
Medulla and medullary rays are absent, and The palisade tissue is present below the
the central part of stem contains a hollow upper epidermis. The palisade cells are thin
region, which is an important character of walled, cylindrical and contain numerous
Cucurbitaceae members. chloroplasts, and are compactly arranged in
one layer, perpendicular to the upper epi-
Leaf anatomy dermis. Narrow intercellular spaces are
present in the palisade tissue. Palisade tis-
Epidermis is present on both surfaces of the sue is the most highly specialized type of
leaf blade, i.e. upper epidermis and lower epi- photosynthetic tissue. The upper surface of
dermis. The epidermis is single layered, with the leaf is dark green in colour due to pali-
compactly arranged barrel-shaped (tabular) sade tissue.
Upper epidermis
Cuticle
Spongy tissue
Glandular trichome
Palisade tissue
Xylem
Phloem
Lower epidermis
Parenchyma
Collenchyma
Fig. 9.52. Transverse section of leaf (100×) showing epidermis, collenchyma, ground tissue and vascular
bundles.
Vegetable Crops 257
The lower part of the mesophyll present mal cells are cutinized. Cuticle reduces the
towards the lower epidermis is the spongy transpiration. Multicellular hairs are present
tissue. Cells of the spongy tissue are thin on the epidermis.
walled, irregular in shape, and arranged Multilayered (from two to six layers)
loosely with large continuous intercellular hypodermis of angular collenchyma cells is
spaces. Stomata open into large intercellular found immediately beneath the epidermis
spaces called sub-stomatal chambers. The (Fig. 9.53). Collenchyma is living mechani-
cells of spongy parenchyma contain fewer cal tissue. The hypodermis gives consider-
chloroplasts, hence the lower surface of the able strength, flexibility and elasticity to
leaf is light green in colour. young petioles and, having chloroplasts, it
The vascular tissue occurs in the form may carry out photosynthesis.
of discrete bundles called veins. Veins are Just beneath the hypodermis, ground
interconnected to form reticulate venation. tissue is found. It consists of thin-walled
Vascular bundles (veins) occur between parenchymatous cells having well-defined
the palisade and spongy tissue (median in intercellular spaces among them. Vascular
position) and are collateral and closed. In bundles are arranged in a half ring scattered
the vascular bundle, xylem is present in ground tissue.
towards the upper epidermis and phloem Bicollateral vascular bundles are arran-
towards the lower epidermis. The vascular ged in two rows. The central part of the petiole
bundle is surrounded by parenchymatous contain a hollow region, which is an impor-
bundle sheath. tant character of Cucurbitaceae members.
Xylem
Inner cambium
Inner phloem
Fig. 9.53. Transverse section of petiole (100×) showing epidermis, collenchyma, ground tissue and vascular
bundles.
258 Chapter 9
Testa
Tegman
Fig. 9.54. Transverse section of seed (100×) showing testa and tegman.
the partially unfolded outer leaves. Cabbage and waxy coating, exstipulate and sessile,
is used in a variety of dishes for its naturally and food material is preserved in the leaves
spicy flavour. The so-called ‘cabbage head’ (vegetative bud) (Fig. 9.55).
is widely consumed raw, cooked, or pre-
served in a great variety of dishes. It is the Floral characters
principal ingredient in coleslaw.
Cabbage is an excellent source of vita- The inflorescence is auxiliary, racemose
min C. It also contains significant amounts of type. The flower is ebracteate, pedicellate,
glutamine, an amino acid that has anti- complete, bisexual, tetramerous and hypo-
inflammatory properties. Cabbage can also gynous. There are four polysepalous sepals
be included in dieting programmes, as it is a and four petals, which are polypetalous, in a
low calorie food. In European folk medicine, cruciform arrangement. There are six tetra-
cabbage leaves are used to treat acute inflam- dynamous stamens and the anthers are
mation: a paste of raw cabbage may be placed dithecous and basifixed. The gynoecium has
in a cabbage leaf and wrapped around the a bicarpellary syncarpous superior ovary,
affected area to reduce discomfort. Some with parietal placentation. The fruit is a sili-
claim that it is effective in relieving painfully qua and the seed is round (approximately
engorged breasts in breastfeeding women. 1 mm), pink to black with a smooth surface.
Fresh cabbage juice has been shown to pro-
mote rapid healing of peptic ulcers.
9.11.5 Anatomical description
(a) (b)
Fig. 9.55. Cabbage plant morphology: leaves are thick, opposite, sessile, exstipulate, simple, ovate, with
a wavy margin and rounded apex, reticulate venation and waxy coating, and food material preserved
in leaves (vegetative bud).
260 Chapter 9
Epidermis
Cortex
Vascular cylinder
Fig. 9.56. Transverse section of root (100×) showing epidermis, cortex and central vascular cylinder.
The endodermis is single layered, with Cortex is thinner than the stele and shows
compactly arranged barrel-shaped cells. hypodermis, middle cortex and endoder-
Endodermal cells are characterized by the mis. Hypodermis is the outermost region of
presences of Casparian strips. the cortex. It is multilayered (from two to
Pericycle is the outermost region of the six layers) and collenchymatic. Collenchyma
stele and is uniseriate and parenchymatous. is living mechanical tissue. The hypoder-
Cells of the pericycle retain meristematic mis gives considerable strength, flexibility
activity. Later roots arise endogenously from and elasticity to young stems and, having
the pericycle. Vascular strands are radial, chloroplasts, it may carry out photosynthe-
xylem is exarch, closed type. Pith is absent. sis. The middle cortex is also multilayered
The non-vascular tissue present between and parenchymatic. The cells may be round
xylem and phloem strands is called con- or oval with prominent intercellular spaces.
junctive tissue. This region also stores food materials tem-
porarily. The endodermis forms the inner-
Stem anatomy most layer of the cortex. It is usually distinct
in stems without Casparian bands.
Epidermis is the outermost region surround- Stele is the central region of the stem
ing the stem and is uniseriate, with a single and is thicker than the cortex. It consists of
layer of compactly arranged barrel-shaped pericycle, vascular bundles and medulla and
living cells. The outer walls of the epider- medullary rays. Pericycle is the outermost
mal cells are cutinized. Cuticle reduces the non-vascular region of the stele and is mul-
transpiration. Few stomata are present in tiseriate and sclerenchymatic. Xylem and
the epidermis for exchange of gases and phloem are organized into vascular bundles.
transpiration. All the epidermal cells are Each vascular bundle is wedge-shaped, con-
colourless except the guard cells. Hairs are joint, collateral, open and endarch. Medulla
absent on the epidermis. and medullary rays are distinct, present at
Cortex is the middle region, present the centre of the stem and usually parenchy-
between the epidermis and stele (Fig. 9.57). matous. The cells are round or oval with
Vegetable Crops 261
Cortex
Epidermis
Secondary phloem
Cambium
Pith
Secondary xylem
Fig. 9.57. Transverse section of stem (100×) showing epidermis, cortex, secondary phloem, secondary
xylem and pith.
intercellular spaces. The medulla stores food tion. The epidermis is single layered, with
materials and helps in lateral conduction. compactly arranged barrel shaped (tabular)
cells. The outer walls of the epidermal cells
Anatomic changes during are cutinized. The epidermis is also covered
shoot development on its outer surface by a continuous layer of
Zhu et al. analysed shoot apical anatomic cutin called cuticle. The cutinized outer walls
changes during the development of Brassica and cuticle reduce the loss of water due to
plants Chinese cabbage (Brassica campes- transpiration. Anisocytic stomata are present
tris spp. pekinensis) and cabbage (Brassica in both of the epidermal layers, but the sto-
oleracea L.). It was shown that all of their mata are usually more frequent in the lower
apical meristems changed from the original epidermis. The stomatal index of the upper
tunica-corpus structure to the intergrade sub- epidermis is 19.1 and of the lower epider-
area, to typical five-subarea structure and to mis is 26.8. The stomata facilitate the
four-subarea structure when they went into exchange of gases between the leaf and envi-
reproductive development. The bilateral ronment. All the epidermal cells except guard
cells of sub-tunica of Chinese cabbage and cells are colourless. Hairs are absent on the
cabbage divided into apical leafy primordial, epidermis. The presence of epicuticular wax
in which lateral inflorescence meristem arose on both sides of the epidermis is an impor-
(Zhu et al., 2006). tant character.
The ground tissue of the leaf, present
between the upper and the lower epidermal
Leaf anatomy layers, is called mesophyll. The mesophyll
is chlorenchymatic and is concerned mainly
Epidermis is present on both surfaces of the with photosynthesis. The mesophyll is not
leaf blade, i.e. upper epidermis and lower differentiated into palisade tissue and spongy
epidermis. Epidermis is mainly for protec- tissue (Fig. 9.58).
262 Chapter 9
Epicuticular wax
Upper epidermis
Xylem
Phloem
Mesophyll
Lower epidermis
Collenchyma
Fig. 9.58. Transverse section of leaf (100×) showing epidermis, collenchyma, spongy tissue and vascular
bundles. Note: thick epicuticular wax and the mesophyll is not differentiated into palisade and spongy
tissues – these are anatomical features of cabbage.
The vascular tissue occurs in the form plants. In Ni5 plants the number of chloro-
of discrete bundles called veins. Veins are plasts in mesophyll cells was higher than in
interconnected to form reticulate venation. the Ni0 control (Molas, 1997).
Vascular bundles (veins) occur between the
palisade and spongy tissue (median in posi- Cuticle
tion) and are collateral and closed. In the vas-
cular bundle, xylem is present towards the A scanning electron microscope was used
upper epidermis and phloem towards the to observe variations in wax formations on
lower epidermis. The vascular bundle is sur- leaves of cabbage. The size and shape of
rounded by parenchymatous bundle sheath. the wax formations on adaxial and abaxial
surfaces of the same leaf differed in all gen-
Effect of nickel on leaf anatomy era. Individual wax crystals on the first-
formed leaves were two to three times
There are morpho-anatomical changes in cab- larger than those on the sixth-formed leaves
bage when grown on nickel (different concen- (Davis, 1971).
trations of 0, 5, 10 and 20 g/m3 (Ni0, Ni5, Ni10, Flea beetles (Phyllotreta spp.), cabbage
Ni20)) medium agar. Reduction of leaf blade stink bugs (Eurydema ventrale) and onion
area, of succulence and of leaf density and thrips (Thrips tabaci) cause significant eco-
growth of specific leaf area were noticed in nomic problems to cabbage growers in
plants treated with all concentrations of Ni. In Slovenia. The aim of this study was to assess
Ni-treated plants the total number of stomata the potential effect of the epicuticular wax
and open stomata decreased, and the number on leaves as defence mechanism against
of defective stomata in both adaxial and abax- these three cabbage pests. These insect pests
ial sides of leaves was higher. In all Ni-treated showed weak preference on cabbage heads
samples the volume of spongy and palisade with high epicuticular wax. There was a
mesophyll cells was smaller in comparison to strong negative relationship between epicu-
control. In comparison to control, the inter- ticular wax content and the level of plants
cellular spaces of mesophyll tissue decreased infested. High epicuticular wax was resist-
in Ni10 and Ni20 plants and increased in Ni5 ant to insects (Žnidarčič et al., 2008).
Vegetable Crops 263
Fig. 9.59. Coriander plant morphology: decompound leaves, spathulate, with a serrate margin.
The inflorescence has a compound umbel, and involucres and involucel are present.
Epidermis Cortex
Stele
Fig. 9.60. Transverse section of root (100×) showing epidermis, cortex and stele (vascular cylinder).
intercellular spaces are present in the pali- scattered in ground tissue. The vascular
sade tissue. Palisade tissue is the most bundles are of various sizes in the same pet-
highly specialized type of photosynthetic iole. Each vascular bundle is wedge-shaped.
tissue. The upper surface of the leaf is dark In the petiole, the xylem is always found
green in colour due to palisade tissue. The towards upper side whereas phloem is
lower part of the mesophyll present towards found towards the lower side (as in the
the lower epidermis is the spongy tissue. leaf).
Cells of the spongy tissue are thin walled,
irregular in shape, and arranged loosely
Seed coat anatomy
with large continuous intercellular spaces.
The cells of spongy parenchyma contain a The seed contain deposits of aromatic vola-
smaller number of chloroplasts, hence the tile compounds as storage food material.
lower surface of the leaf is light green in The testa is thin and there is no clear differ-
colour. ence between testa and tegman (Fig. 9.62).
The vascular tissue occurs in the form Rate of imbibition is slow.
of discrete bundles called veins. Veins are
interconnected to form reticulate venation.
Vascular bundles (veins) occur between Significance variations in anatomy
the palisade and spongy tissue (median in Future research may be directed in the fol-
position) and are collateral and closed. lowing directions. Possessing a greater
In the vascular bundle, xylem is present number of multicellular trichomes gives
towards the upper epidermis and phloem disease resistance to insects. The presence
towards the lower epidermis. The vascular of volatile compounds is helpful in fungal
bundle is surrounded by parenchymatous resistance.
bundle sheath.
Petiole anatomy
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10
Anatomical Adaptation to Defence
Against Biotic Stresses
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 269
270 Chapter 10
10.2 Structural Modifications Wax and lipids serve three major pur-
for Defence Against Insect Pests poses in defence against insect pests and
and Pathogens diseases.
1. Wax and lipids form a barrier layer, which
Insect pests damage the crop by phytophagy, inhibits penetration of fugal hypha, bacterial
through chewing the tissues or sucking saps spores or sucking insects. The deposition of
using specialized structures. In the battle cuticular waxes increases with plant age. In
against insect pests, plants rely on two many cases, increased cuticle thickness
mechanisms – structural barriers that pro- increases the resistance towards the patho-
hibit or inhibit insects from phytophagy gen, as observed in the case of charcoal rot
and chemical warfare through producing (Rhizoctonia solani) in beans, groundnut,
chemicals to kill, trap or stop insects from cotton and radish, powdery mildew (Sphaero-
feeding on the host plant. In both mecha- theca pannosa) of roses or grey mould (Botrytis
nisms, the structural anatomy of plants cinerea) of grapes (Mence and Hildebrandt,
plays a very important role in mechanical 1966; Gabler et al., 2003). However, for some
defence or delivery of the chemicals through pathogen such as Uncinula necator causing
pores or glands. powdery mildew of grape, the cuticle does
not seem to be problematic for penetration,
although spread of disease is reduced by
preformed barrier at the cuticular region
10.2.1 Modifications of outer (Ficke et al., 2004).
protective layers 2. The waxes are hydrophobic and repel
water, so pathogenic spores do not get water
Cuticle formation for germination on the plant surface. The
spore structures of rust fungi do not germi-
The epidermis, which constitutes the out-
nate properly on wax-coated leaves of wheat
ermost layer of plant tissues, is the first
and barley.
line of defence against insect pest or patho-
3. The wax is considered unpalatable by
gen attack (Riederer, 2006). It serves as a
many insect pests, so wax deposition reduces
protective tissue system in leaves, floral
insect attack.
parts, fruits, seeds, stems and roots of
plants. The epidermal cells of aerial plant The chemical composition of plant cuticu-
parts are often covered in a waxy cuticle lar lipids influences the choice of the insect
that not only prevents water loss from the pests. Wild groundnut species exhibit dif-
plant, but also prevents microbial patho- ferent cuticular lipid composition than the
gens from coming into direct contact with cultivated groundnut, which provides a
epidermal cells and thereby limits higher degree of resistance to fall armyworm
infection. The cuticle can be relatively thin and thrips (Yang et al., 1993). Studies have
as in aquatic plants, or extremely thick as shown that when corn earworm (Helicoverpa
in cacti. zeae) larvae are grown on diets having
Cuticles are formed from the outer epi- cuticular lipids extracted from silk of maize,
dermal cell layers by deposition of long the growth of the larvae was reduced. This
chain lipids on the surface of epidermal suggests that the presence of cuticular lip-
cells. These lipids are of two types – cutin ids negatively affects the growth and feed-
(C16 and C18 fatty acids and glycerol) and ing of earworms and provides resistance to
waxes (comprising of very long chain fatty the crop (Yang et al., 1992).
acids). These fatty acids are synthesized in Apart from serving as a barrier, plant
the plastids and endoplasmic reticulum of cuticle has also been proposed to play a cru-
epidermal cells by fatty acid elongases and cial role in systemic acquired resistance
are mobilized to the exterior of the epider- (SAR). SAR is activated in plants through
mal cells by lipid transporters (Kunst and signalling processes, which sense insect or
Samuels, 2009). pathogen attack in an attack site and induce
Defence Against Biotic Stresses 271
defence response in distal tissues in the 2003). Due to the accumulation in the epi-
host. Arabidopsis mutants with defects in dermis of leaves, soluble silicon activates
cuticle formation fail to transmit the SAR genes involved in the production of second-
response, indicating that cuticles play an ary compounds of metabolism, such as
important role in SAR (Xia et al., 2009). polyphenols, and enzymes related to plants’
defence mechanisms.
Silicification
Lignification and suberization
The role of silica in disease resistance in
cereal crops has been established for quite a Lignin is a complex phenolic substance result-
long time (Germar, 1935). Silicon uptake ing from the polymerization of p-coumaryl,
through roots increases insoluble silicon coniferyl and sinapyl alcohols in the plant
deposition, which acts as a physical barrier cell wall, which act as a physical barrier for
to the pathogen’s entrance (Zeyen et al., pathogen infection (Ride, 1983). The accu-
1983). The amorphous silica located in the mulation of lignin and lignin-like materials
cell wall has a marked effect on the cellular in plant parts for resisting pathogen invasion
wall physical properties. It induces forma- is a very common structural defence phe-
tion of papilla in the inner surface of epider- nomenon. Lignin is highly resistant to attack
mal cell walls and the deposition of callose by microorganisms, and lignified cell walls
(Bélanger et al., 2003). The papilla inhibits are an effective barrier to pathogen entrance
penetration of fungal hyphae and callose and spread. Lignification toughens the cell
encases the fungal haustoria, resulting in wall, which exhibits higher resistance to
resistance reaction. The system of silicon- mechanical pressure applied during pene-
mediated physical resistance works well in tration by fungal appressoria. Due to this
many host–pathogen interaction systems water-resistant barrier, other components of
including rice–rice blast, wheat–powdery the cell wall are less accessible to cell wall-
mildew (Blumeria graminis f. sp. tritici), degrading enzymes.
barley–powdery mildew (Erysiphe graminis There are many examples of pathogen
f. sp. hordei), oat–powdery mildew, coffee– inhibition in crop plants by lignification, and
Cercospora and sorghum–Rhizoctonia. probably in the majority of host–pathogen
Insoluble deposition of silica layers in the interaction it serves as a major component
penetration sites increases the resistance of host defence system. In wheat, lignification
response to powdery mildew pathogens. seems to be a specific response mechanism
Among the grass crops, rice shows the for protection against filamentous fungi, but
highest silicon uptake, which goes up to not for other physical stresses (Ride et al.,
10–15% of total dry weight. Silicon is 1989). It has been observed that silencing
deposited in all rice plant parts, the maxi- of monolignol synthesizing genes makes
mum being in stem and leaves. In leaves it wheat highly susceptible to powdery mildew
is deposited as a glass-like coating in the (Blumeria graminis f. sp. tritici) in suscepti-
epidermis, middle lamella, intercellular ble as well as in resistant lines by allowing
spaces, sclerenchyma, vascular bundle and penetration of fungal appressoria (Bhuiyan
bundle sheath, but not in cytoplasm (Kim et al., 2009).
et al., 2002). The formation of such an Panama wilt is a serious disease of
impervious layer with cuticular deposition banana caused by Fusarium oxysporum f. sp.
serves as a physical barrier for both insects cubense, infecting roots of the crop and
and pathogens, including blast (Magna- causing characteristic wilting symptoms.
porthe grisea), brown spot (Bipolaris oryzae) Comparison of root anatomy of resistant and
and sheath blight (Rhizoctonia solani). susceptible cultivars shows that higher ligni-
Besides the physical barrier, soluble fication and callose deposition in the roots
silicon also creates a physiological barrier provides a considerable level of resistance
through upregulation of plant defence sys- against this disease (De Ascensao and Dubery,
tems (Koga et al., 1988; Bélanger et al., 2000). Rapid building up of mechanical
272 Chapter 10
barrier in the root epidermal cells through found to bear a smaller number of pores and
cell wall esterification and lignification lenticels compared to susceptible plants
serves as the main defence system of banana (Eibach, 1994; Gabler et al., 2003).
against this disease. Stomata are yet other natural openings
Suberin deposition creates a barrier in the plant that are essential for physiologi-
layer that prohibits progress of fungal patho- cal activities but serve as entry points for
gens during infection and the release and pathogen attack. It has been observed that
spread of fungal enzymes, and also creates rust fungi sense the relative distance of the
toxicity to microbes due to the high propor- stomatal guard cells and accordingly pene-
tion of phenolic compounds (Kolattukudy trate the stomata at a desirable height (Hoch
and Espelie, 1989). It provides structural et al., 1987). When the fungus hyphae
resistance in many crops against pathogens, encounter a lip of the proper height, they
such as wilt of tomato and potato (Vertcillium undergo a developmental programme result-
dahliae) and root rot of soybean (Phytoph- ing in the formation of invasive structures
thora sojae). Suberin formation is common that enter the stomata and begin coloniza-
when disease progresses in the vascular bun- tion of the leaf interior. In mulberry plants,
dle. The vascular bundle develops a resistance susceptibility to powdery mildew is related
response by heavily impregnating surround- to higher stomatal density (Chattopadhyay
ing cells of the infection zone so that the et al., 2011); therefore low stomatal density
progress of disease is halted. Suberization is is a promising character for selection of
a common feature in the barks of tree crops resistant cultivars.
as a structural defence mechanism.
entrap and immobilize insects or contain of endogenous genes involved in these sec-
toxic or deterrent substances. Trichome ondary metabolic pathways provides
glands produce and secrete various types opportunity for development of genotypes
of exudates affecting insects, microbes and with higher resistance to insect pests. In
herbivores. Glandular trichomes are com- tobacco, expression of a P450 hydroxylase
mon in the nightshade family (Solanaceae). gene specific to the trichome gland leads to
Selection for presence of trichomes has formation of cembratriene-diol (CBT-diol)
been helpful in developing new varieties from its precursor cembratriene-ol (CBT-ol)
of potato and tomato that resist insect (Wang et al., 2001). CBT-diol, the main
pests because of glandular hairs on their exudate, is less effective in aphidicidal
leaves and stems. Other crop plants in activity than CBT-ol. Suppression of this
which glandular trichomes are being used gene increases concentration of CBT-ol
to breed for imparting pest resistance with a remarkable increase in resistance to
include lucerne, strawberry, sunflower aphids.
and tobacco. Both glandular and non-glandular tri-
Several groups of chemicals including chomes can coexist in the same genotype. In
phenols, quinones, resins and volatiles are pigeonpea, three glandular (Types A, B, and
released from the glandular trichomes. E) and two non-glandular (Types C and D)
These chemicals act in various ways to trichome types have been detected that are
reduce insect damage such as by reducing present on leaves, pods and calyxes of both
insect population growth, creating traps to the cultivated and wild species (Romeis
reduce mobilization of insects, making the et al., 1999). Presence of high density of
host tissues unpalatable, or by repelling non-glandular trichomes in pigeonpea
insects from coming near to the host. Many helps in preventing attack of pod borer
crop plants including potato and tomato (Helicoverpa armigera), which is the most
release phenolic compounds through their serious pest of this crop. The wild species
trichome glands, producing sticky substances having higher trichome density is more
to trap small insects such as aphids. Type A resistant to H. armigera infestation, which
glandular trichomes of wild potato Solanum establishes the importance of trichomes in
berthaultii produce phenolic exudates insect-pest resistance.
(glucose ester of p-hydroxyphenylpropionic
acid), which are oxidized by polyphenol Studies on inheritance of trichomes
oxidase (PPO), producing a cement-like
substance resulting in entrapment and death Studies have been undertaken on the inher-
of the aphids (Kowalski et al., 1993). High itance of trichomes. Segregation ratios in
PPO activity has also been correlated with successive generations of five single-cross
resistance to potato tuber moth (Phthori- matings between trichomed and trichome-
maea operculella). The glandular trichomes less parents indicated that the presence of
from trichomes of S. berthaultii also pro- leaf trichomes is controlled by a single
vide resistance to Colorado potato beetle recessive gene. Inheritance of this character
(Leptinotarsa decemlineata) by exudate in three crosses among four trichomed par-
deposition on larvae and also providing a ents involved the same locus, designated tr.
barrier to feeding (Neal et al., 1991). A sec- Trichome density on the abaxial leaf surface
ond type (type B) of glandular trichome has varied among trichomed lines from single
been identified in other wild potatoes, crosses. Heritability of trichome density
Solanum sparsipilum and Solanum pinna- between the F3 and F4, estimated in the
tisectum, which exhibits a strong repulsive cross IS1054 × B CK60, was 0.75 (Gibson
effect on oviposition of tuber moth (Musmeci and Maiti, 1983).
et al., 1997). Trichomed, segregating and trichome-
Many of the exudates are secreted less F3 and F4 lines from four trichomed ×
under the influence of trichome-specific trichomeless crosses and their parents
gene expression. Alteration or suppression were studied in the field in Patancheru.
Defence Against Biotic Stresses 275
Compared with trichomeless lines, tri- trichomes in non-glossy lines may also
chomed lines had significantly lower per- stimulate shoot fly attack.
centages of plants with Atherigona soccata
eggs 18 days after emergence and of dead Do all trichomes contribute
hearts at both 18 and 23 or more days after to plant defence?
emergence. The ratio of the difference
between the means of trichomed and tri- Not all trichomes are suitable for resistance
chomeless lines for the percentage of dead to insect pests. The ability of trichomes to
hearts to the corresponding difference impart resistance to insect pests depends on
between the parents ranged from 0.16 to the structure and the properties of the tri-
0.92 and exceeded 0.32 in seven of nine chomes. It has been observed that there is
comparisons, indicating that trichomes great variation in the structure of trichomes,
were a major factor in resistance. Means including length of trichome, body shape,
of parents and progenies were regressed tip shape and size, gland shape and size and
on four possible genetic models and number of cells in base and head. In tomato,
results indicated that, besides trichomes, seven types of trichomes have been
at least two loci that interact with each described by Luckwill (1943). Some of these
other were involved in resistance (Maiti trichomes (for example type VI, which has a
and Gibson, 1983). glandular head) are associated with higher
In a separate study, leaf surfaces of seven mortality of larvae of potato tuber moth than
genotypes of Sorghum bicolor, two of maize, other trichome types. Similarly, presence of
Zea mays, and two of pearl millet, Pennise- dense type I trichomes, which is longer in
tum americanum, were examined by scanning size than other trichomes, is inversely
electron microscopy for possible morpho- related to feeding ability of the potato tuber
logical differences. Glossiness was deter- moth larvae (Simmons et al., 2006).
mined by spraying water, which adhered to Presence of glandular trichomes that do
the glossy leaves. Cuticular transpiration of not exhibit insect resistance properties may
detached third and fifth leaves was estimated increase susceptibility to insect pests. In sor-
from the rate of water loss after abscisic acid ghum, genotypes having non-glandular tri-
induced stomatal closure. Glossy character chomes are more tolerant to shoot fly than
was correlated with the reduction or absence the genotypes with glandular trichomes
of wax deposits on the leaf surfaces, while (Maiti and Gibson, 1983). They proposed
hairiness might occur in either glossy or non- that the glandular trichomes of sorghum
glossy genotypes. Unlike sorghum and maize, may produce some chemicals that favour
in which all leaves after the fifth or seventh insect growth. In pigeonpea, glandular tri-
were glossy, pearl millet showed no glossi- chomes (type A) on the calyxes and pods
ness through the ninth leaf. Measurements contribute to susceptibility to pod borer
showed that cuticular transpiration of glossy (H. armigera), while the non-glandular tri-
leaves was often more than double that of chomes (trichome type C and D) are associ-
non-glossy leaves. Comparisons among sor- ated with resistance (Sharma et al., 2009).
ghums showed that non-glossy lines had Although most of the evidence on glandular
higher stomatal densities than glossy lines trichome research suggests negative effects
(Traore et al., 1989). of exudates on insects, recently Weinhold
Further study may reveal that the den- and Baldwin (2011) found that o-acyl sug-
sity of non-glandular pointed trichomes ori- ars, the most abundant metabolite of glandu-
ented at an acute angle on leaf surfaces lar trichomes in wild tobacco Nicotiana
offers barriers to the movement of maggots/ attenuata, is readily consumed by the larvae
larvae to reach the shoot apex through the of Lepidopteran insect pests Spodoptera lit-
collar. When associated with non-glandular toralis, Spodoptera exigua and Manduca
trichomes, waxy coatings impart glossiness sexta. These findings suggest that trichome
on the leaf surface, which prevents insects exudates may also attract insect pests thereby
from laying eggs. The presence of bicellular increasing the susceptibility of a genotype.
276 Chapter 10
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11
Anatomical Adaptation for Drought
and Waterlogging Stress Tolerance
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
278 (R. Maiti et al.)
Drought and Waterlogging Stress Tolerance 279
plants for drought-prone agriculture has led transpiration takes place through cuticle.
to accumulation of both these mechanisms Thus the ability to conserve water under
for increasing water use efficiency under drought stress depends on the thickness
drought conditions. and deposition of cuticle. If cuticle is thin
with no deposition, stomatal resistance
becomes ineffective under drought stress.
11.2.1 Mechanisms of avoidance Cuticle is a comparatively thin layer
of drought through anatomical (0.1–10 µm), so the resistance to water loss
modifications is primarily determined not by the thick-
ness of cuticle, but by the wax deposition
(see Chapter 9, this volume) on the cuticle
Reduction of transpiration by modulating
(epicuticular wax). Mechanical removal of
stomatal activity and density
epicuticular wax results in higher transpi-
Stomatal activity is more influenced by ration. The wax deposition benefits the
drought stress than photosynthetic activities plants under drought stress in two ways: by
in plants. Opening and closing of stomata is reducing transpiration and also by increas-
controlled by the guard cells surrounding ing reflectance of the leaf (Maiti, 1996).
the stomatal pore. Under water sufficiency Studies in maize show that cuticular depo-
conditions, water influx from surrounding sition is inversely related to epidermal
cells to guard cells causes the guard cells to water loss. However, some studies show
swell and rise, leading to opening of sto- that the amount of cuticular wax has no
mata. This is regulated by a cellular network correlation with transpirational loss (Ristic
involving ion channels and transporter pro- and Jenks, 2002). Thus rather than the
teins in the plasma membrane, and is influ- amount, the chemical composition may be
enced by many factors such as concentrations more pertinent to water loss through the
of K+, Ca2+, sugars and CO2 and also on the epidermis.
spectrum of light. The stomata can sense
drought conditions through the increase in Anatomical modification
abscisic acid (ABA) concentration, which in root characters
causes efflux of solute from guard cells
resulting in closure of stomata. It can sense Drought stress increases root growth com-
the rise in ABA concentration in nearer leaf pared to shoot growth as plants try to har-
tissues as well as in distant regions such as vest higher soil moisture. The root–shoot
roots through xylem-mediated ABA trans- ratio in almost all crop plants increases
port. Rapid stomatal sensitivity under under drought stress, indicating root growth
drought stress is a desirable mechanism to is a common adaptation across species
close the stomata and reduce water loss. under water-limited environments. This
Mutations affecting stomatal sensitivity growth is activated by and associated with a
readily reduce water availability, resulting number of signaling processes, including
in rapid wilting of plants. ABA accumulation, concentration of reserve
Stomatal density is directly correlated carbohydrates in the apical zone of root,
with stomatal conductance and thus crop accumulation of proline, cytokinins and
productivity, by helping to maintain the pho- expansion proteins. A characteristic symp-
tosynthetic activities in leaves. Sustenance tom of drought stress is activation of pri-
of stomatal conductance during the crucial mary root growth and cessation of secondary
drought cycles helps to increase productiv- root growth, which is driven by cytokinins-
ity under drought stress. mediated blocking of auxin accumulation.
Primary root growth enables the plant to
Cuticular resistance penetrate deeper in the soil to extract soil
moisture. In most of the landraces of crop
Along with stomata, plant cuticle is involved plants adapted to arid region, the deep root
in transpiration. When stomata are closed, system has been observed to be the major
280 Chapter 11
to better grain filling in cereals during post- (Kabuli type), mungbean (V. radiata), lentil
anthesis period, since the greener leaves (L. culinaris) and soybean (G. max). Only a
produce more photosynthates during this few legume species such as faba bean (Vicia
period and translocate it to the endosperm faba) are comparatively more susceptible to
of developing seeds. The stay green geno- drought.
types have developed internal structural The anatomical features contributing
modifications for better assimilation of water to drought tolerance in arid legumes are
and mineral nutrients. similar to general features of drought toler-
At metabolite level, increase in proline ance or avoidance (see Chapter 6, this volume),
and soluble concentration is a common which attempt to maximize water conser-
phenomenon among the grass crops. Proline vation by modifying stomatal conductance
provides tolerance to abiotic stress via and cuticular structures, maintenance of
osmo-protection as well as by increasing the photosynthetic activities, economization
antioxidant enzyme activities. The soluble of water distribution, efficient mobiliza-
sugars also contribute to stress tolerance as tion of storage reserve and enhanced water
signalling agents, osmo-protectants and as a uptake through development of a deep pri-
pool for metabolic activities. Studies have mary root system (White and Castillo, 1989;
shown that accumulation of these metabo- Ramirez-Vallejo and Kelly, 1998). Although
lites is much higher in drought-tolerant cul- epicuticular wax deposition is not com-
tivars of rice, wheat and sorghum than the monly found in legumes, black gram geno-
susceptible cultivars. In the case of rice, the types deposit epicuticular wax under
soluble sugars are accumulated more in leaf drought stress to reduce water loss. Simil-
sheaths than other plant parts, as sheaths arly, osmotic adjustment, which plays a
are more exposed to drought stress com- major role in drought tolerance in cereals,
pared to other plant parts (Cabuslay et al., seems to be of less importance in the drought
2002). Leaf sheaths also provide mechanical tolerance mechanism of legumes (Likoswe
protection to actively dividing meristematic and Lawn, 2008).
tissues. The drought-tolerant rice cultivars Pigeonpea is one of the most drought-
adapt to the water-limited environment by tolerant tropical legume crops. The ability
accumulating more soluble sugars and pro- to withstand high drought in this crop spe-
line for higher osmo-protection in the leaf cies is primarily attributed to the deep root
sheaths than leaf blades. system, better osmotic adjustment, higher
photosynthetic ability and multiple flow-
ering flushes (Odeny, 2007). However,
there is considerable within and between
11.2.3 Adaptations in arid legumes species variability in utilizing these mech-
anisms for drought tolerance among arid
The semi-arid and arid regions of the world legumes. For example, stomatal conduct-
are characterized by low agricultural pro- ance contributes to drought tolerance in
ductivity and higher percentage of popula- some common bean genotypes, but it does
tion under extreme poverty with nutritional not appear as a primary mechanism for
deficiency. The legume crops are important drought tolerance in other common bean
diets for poor people in this region; hence genotypes or drought-tolerant genotypes of
they are grown in intermittent or terminal faba bean (Ludlow and Muchow, 1990;
drought conditions mostly utilizing resid- Ehleringer et al., 1991; Grzesiak et al.,
ual soil moisture. The arid and semi-arid 1999). In some drought-tolerant genotypes,
legumes such as pigeonpea (C. cajan), cow- stomatal closure is faster in response to
pea (V. unguiculata), chickpea (C. arieti- drought. Common bean cultivars that are
num, desi type) and black gram (V. mungo) susceptible to drought exhibit reduction in
are drought-tolerant species. Some other thickness of epidermis, protoderm and area
legumes that have moderate drought toler- of parenchymatic cells along with plasmo-
ance are common bean (P. vulgaris), chickpea lysis and death of root cells (Peña-Valdivia
282 Chapter 11
et al., 2010). Stratification of root epider- zone. This symptom is very common in
mis and protoderm starts under drought rice fields of Asian countries. It has
stress. The cortex and xylem cell wall been long recognized as ‘bronzing’ of rice
thickens in response to drought stress, plants.
resulting in reduction in diameter of xylem The rice plant under submerged condi-
vessels, which limits water movement. tions exhibits two major anatomical adap-
These responses are also observed in non- tations: (i) the underground root zone by
legume crops under stress, such as maize. formation of aerenchyma; and (ii) enhanced
Studies on maize shows that in the drought- growth and shoot elongation under deep
tolerant cultivars the protoderm thickness water. Both these mechanisms are very
is less affected than in susceptible culti- unique among crop species, as no other crop
vars (Peña-Valdivia et al., 2005). Wild rela- plant grown on soil shows such extreme
tives of common bean show higher drought anatomical adaptations under submerged
tolerance and anatomical features better condition.
adapted to drought stress, which suggests
that during domestication, some traits con-
tributing to drought tolerance were lost,
which makes the present day common 11.3.1 Aerenchyma formation in rice
bean genotypes less adapted to arid regions as a response to waterlogging conditions
compared to their wild relatives (Peña-
Valdivia et al., 2010). Although some crop species can survive in
Optimal resource allocation is a critical waterlogged condition for a few days, rice
issue in arid legumes. Plants with large veg- is a remarkable crop whose life cycle is
etative biomass and larger leaf area lose completed mostly in the waterlogged anaer-
more water through transpiration, which is obic environment. It also grows well under
the main hindrance for the development of aerobic conditions (upland rice), which
a drought-tolerant mungbean cultivar with makes it a very good candidate for studying
high productivity. The mungbean leaves are various anatomical, physiological and cel-
large and distributed horizontally, resulting lular adaptations under aerobic as well as
in higher transpirational loss. Moreover, anaerobic environmental conditions. Rice
under drought stress root–shoot ratio is a semi-aquatic plant with a very high
increases considerably in all legumes, since adaptability to different environmental
root growth is less affected than shoot conditions. The most striking feature of rice
growth under drought. anatomical adaptation for waterlogging
stress tolerance is the formation of aeren-
chyma. Aerenchyma are formed in many
other plants under waterlogging condition,
11.3 Tolerance to Waterlogging including wheat, barley, maize, jute, tomato
(Submergence) and many forage grasses. However, while in
other plants, development of aerenchyma
Waterlogging and soil flooding results in is induced only in response to waterlogging
submergence of below-ground parts of the stress, rice has a special genetic mechanism
plant. This not only reduces the oxygen for constitutive aerenchyma formation even
availability in the soil, creating hypoxia or in aerobic condition (Kawai et al., 1998).
anoxia, it also changes the soil chemical However, flooding enhances aerenchyma
properties resulting in difficulties for formation in rice.
absorbance of chemical nutrients from the Aerenchyma is a cellular space filled
soil. A prominent outcome of flooding in with air, which provides oxygen to the
soil is the iron toxicity, which results in respiring cells under anoxic and hypoxic
the conversion of available iron in the soil condition by accumulating oxygen from
into non-available forms, creating deposi- the environment by diffusion (Colmer,
tion of iron oxides and sulfides in the root 2003). Aerenchyma create a diffusion
Drought and Waterlogging Stress Tolerance 283
path for transport of oxygen from aerial (System Rice Intensification or SRI)
parts to the waterlogged tissues as well as minimize the anaerobic environment by
helping in diffusion of other volatile com- maintaining non-flooded soil conditions.
pounds including CO2 and ethylene, both This management practice has led to
of which are crucial for survival of the increased rice productivity by increasing
rice plant under waterlogged conditions tiller number and better plant growth as
(Vartapetian and Jackson, 1997). Ethylene well as reducing the cost of cultivation
serves as a signalling agent for aeren- (Sato and Uphoff, 2007).
chyma formation in maize; however, it
plays little role in development of aeren-
chyma in rice. Underwater CO2 concen-
tration reduces in the daytime due to 11.3.2 Shoot elongation
photosynthesis while it increases in the in deep water rice
night time by respiration. Inadequate sup-
ply of CO2 during photosynthesis results Certain rice genotypes adapted to com-
in reduced submergence tolerance (Setter plete submergence (deep water rice) have
et al., 1989). the unique capability to shoot elongation
However, formation of aerenchyma and float over the water level for growth
leads to weakening of root structure, thus and development. The ability to elongate
its penetration ability and mechanical sup- under such an environment depends on
port to the plant is reduced. The rice plant several plant and environmental factors,
maintains a balance between aerenchyma such as genotype, age of plant, root and
formation and functional ability of root to shoot anatomical characters, light percep-
provide these mechanical supports. Rice is tion through water, temperature of the
commonly grown in environments in alter- water etc.
nate aerobic and anaerobic environments. Shoot elongation is a primary response
Maintaining stagnant water in the field for in rice and in many wetland plants under
days or weeks is a general practice in farm- submerged condition. The internodes of
ers’ fields, which increases the rate of for- shoots elongate vertically rapidly through
mation of aerenchyma in the roots, so that the water until they contact the atmos-
more oxygen can be harvested under phere (Jackson, 1985). As in roots, aeren-
anaerobic condition. Due to less rainfall or chyma plays a crucial role in elongating
withholding irrigation, water is often shoots by providing oxygen and other gas-
depleted from the soil, resulting in hard- eous elements to the shoot tissues. More
ening of soil. This exerts high mechanical internode air spaces develop under sub-
pressure on the rice roots, resulting in col- merged condition in rice. Development of
lapse of the aerenchyma structure. The internode air spaces is higher in submer-
rice plant then again shifts to the forma- gence-tolerant cultivars than susceptible
tion of vascular tissues in place of aeren- cultivars (Datta and Banerji, 1974). The
chyma. Anatomical studies reveal that air spaces also reduce the culm weight
rice roots develop a smaller amount of and help to maintain buoyancy in the
aerenchyma under irrigated cultivation water.
conditions (alternate wetting and drying) In deep water rice, the air layers formed
than complete submerged conditions in the leaves serve two purposes: (i) to accu-
(Mostajeran and Rahimi-Eichi, 2008). The mulate oxygen in the air chambers through
inherent plasticity of rice root to develop diffusion, which is used by the leaf for
aerenchyma under submerged conditions respiration (Matsukura et al., 2000); and
and degrade it under aerobic conditions is (ii) the air chambers increase floating capa-
a key anatomical adaptation for its semi- bility of the leaves above water, so that part
aquatic nature. The cost of maintaining of the leaves may directly come in contact
plasticity obviously affects the productiv- with the atmosphere for photosynthesis and
ity of rice. New rice cultivation techniques respiration.
284 Chapter 11
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12
Anatomical Adaptation in Crop Plants
to Harvest Higher Energy
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 285
286 Chapter 12
for genotypes that can harvest higher solar due to a lack in other important desirable
energy and convert this into chemical characters, such as higher harvestable
energy efficiently. The second issue is to product and usability as food, fuel or fod-
channel the storage compounds to storage der. A second example of structural change
organs efficiently (from source to sink). The and adaptation is the ability to develop a
structural anatomy of crop plants has thus symbiotic relationship with microorgan-
been modified accordingly to harvest higher isms for fixation of nitrogen. The specially
energy, conversion of energy to storage developed root nodules in legumes are fac-
product, minimization of vegetative growth tories for converting nitrogen into nitroge-
activities during reproductive maturity nous compounds, which are supplied to
phase, better assimilation of mineral nutri- the plant for growth and development.
ents from the environment (soil and air) Conservation of energy for increasing the
and reduction in unnecessary losses of amount of storage reserves and efficient
water from the plant. We should remember channelling of reserve foods toward stor-
that human selection has not brought all age tissues have also been accomplished
the changes in plants to be required as crop. by a number of structural modifications in
During the domestication phases of crop crop species.
plants, the human population has employed
opportunistic selection for crop species
from natural variability according to their
choice. The process of selection was prima- 12.2 Higher Photosynthesis
rily based on domestication-related traits
including higher productivity, suitability of Over 90% of the total biomass of the crop is
consumption and feasibility of cultivation derived from photosynthetic activities
as a crop. Per unit productivity was not (Zelitch, 1982). Thus adaptation for higher
very important to early agrarian popula- photosynthesis is a major driving force for
tions due to smaller consumption require- selection of crop plants. During the process
ment and availability of land for cultivation. of domestication, selection led to improve-
Crop yield has become the primary and ment of the photosynthetic capacity of crop
sometimes the sole concern with the plants. Comparison of photosynthetic abil-
increase in human population size and ity of modern crop plants with that of their
competition for resources and is the main wild relatives reveals that the crop plants
concern for modern day plant breeding. exhibit higher photosynthesis (Bhagsari and
The changes in structural anatomy of crop Brown, 1976; Saitoh et al., 2004). Rice and
plants are clear indications of this evolu- wheat, two principal C3 crops, exhibit
tionary process, which is being changed higher photosynthesis than other C3 crops,
continuously depending on the adaptation which contributes to the high yield of these
conditions and demand for higher two crops (Murata, 1981).
productivity. However, until recently, net photosyn-
During evolution, several anatomical thesis was considered not to be related to
changes in the plant kingdom have been crop yield (Makino, 2011). Researchers
found to be beneficial for humans, making found no correlation between net CO2
these plants suitable as a crop. A leading exchange per unit leaf area and yield per
example is the evolution of C4 plants from unit area, which is considered paradoxical
C3 plants. The higher photosynthetic abil- because the reserve food expressed as yield
ity with desirable features of higher har- would not have been produced if there was
vestable produce for consumption have no photosynthesis (Evans, 1975; Zelitch,
made a good number of C4 species crop 1982). But experiments with elevated CO2
plants, such as maize, sorghum, sugarcane concentration proved that yield is increased
and pineapple. However, several other when the rate of photosynthesis was higher,
grass species having the C4 photosynthesis provided the other factors are unaltered
mechanism were not selected as a crop, (Long et al., 2006).
Anatomical Adaptation to Harvest Higher Energy 287
part of crop species has certain utility, pro- place in this crop through increase in
ductivity is mainly concerned with the prin- biomass (Hay, 1995).
cipal economic product. In the case of grain
crops, seeds are the principal product, where
Partitioning of biomass through
the stored material in seed is used either as
anatomical adaptations
food (cereals and pulses) or for extraction of
oil (rapeseed, mustard, soybean, groundnut). According to Evans (1975), sink capacity
In these crops, the productivity is deter- and yield tend to increase in parallel until
mined by harvest index (ratio of grains pro- they reach a particular limit set by the pho-
duced and total biomass). The cereal crops tosynthetic capacity. The photosynthetic
have occupied the bulk of the agricultural and storage capacities of improved geno-
area not only for the supply of essential car- types of major crops are likely to maintain a
bohydrates, but also for their high harvest close balance. Thus improving one or the
index (HI) compared to other crops. The HI other will have a coordinated effect on pro-
of intensively cultivated varieties of major ductivity. In certain crops, the sink capacity
crops falls in the range of 0.4–0.6 (Hay, is limited by structural boundaries, such as
1995). grain size in maize. Thus even with a higher
During the early selection periods, rate of photosynthesis as a C4 plant, maize
productivity enhancement was mainly productivity is not higher than rice, a C3
achieved by increase in biomass (Kawano, plant, which has a lower rate of photosyn-
1990). This led to the development of locally thesis but higher sink capacity. Hybrid
adapted genotypes, which have a higher maize has increased the productivity
vegetative growth under the low input con- through higher vegetative growth as well as
ditions. Since early agriculture was not increase in size of cobs, but the HI did not
input intensive, genotypes responsive for increase significantly.
high input condition having higher harvest Structural limitations of source and
index were not paid much attention; pro- sink are primary bottlenecks of productivity
ductivity was enhanced primarily through enhancement. Zelitch (1982) proposed that
higher crop growth, resulting in both vege- the limitation of sink capacity in crop plants
tative and reproductive yield. However, cer- is the major limiting factor for realizing high
tain historical evidence suggests that the HI yield despite adequate photosynthesis. Crop
of crops grown by the ancient civilization ideotypes (described in Chapter 15) have
was not very low (Nagato et al., 1988; Amir been proposed by many workers that can
and Sinclair, 1994). HI of major cereal crops break the barrier of source or sink limita-
decreased in the middle ages with the tion. Remarkable achievements have been
increased alternative use of vegetative parts made in rice improvement using the ideo-
of cereal crops as animal feed. type concept.
The concept of HI in modern plant
breeding came in the early 20th century,
when Beaven proposed the term ‘migra-
tion coefficient’ in 1914 to describe the 12.3 Biological Nitrogen Fixation
ratio of grain weight to total plant weight
(Donald and Hamblin, 1976). The concept The capability of fixing atmospheric nitro-
gained importance in plant breeding only gen in symbiosis with microbial fauna is a
in the 1960s when Donald (1962) provided remarkable feature of plant species to meet
mathematical expressions for HI determi- the nitrogen demand. Higher nitrogen con-
nation. In the second half of the 20th cen- centration is a characteristic feature of
tury, modern plant breeding has led to a plants that exhibit biological nitrogen fixa-
substantial increase in the HI of grain crops tion. The additional nitrogen is required to
(Sinclair, 1998). However, in certain crops meet high nitrogen demand of the tissues
such as maize, the HI was already high, as well as to develop a number of nitroge-
and major genetic improvement has taken nous secondary metabolites that provide
Anatomical Adaptation to Harvest Higher Energy 289
protection from predation and pathogene- Table 12.1. Legume crop-rhizobial species/strain
sis. For examples, the leguminous nitro- specificity.
gen-fixing species produce a variety of
Legume crop host Rhizobial symbiont
polyphenols, which protect them from pest
and disease attack. Pea Rhizobium leguminosarum
The leguminous plants add substantial bv. viciae
amounts of nitrogen to the soil. It has been Soybean Bradyrhizobium japonicum
observed that the amount of nitrogen fixed Sinorhizobium fredii
by leguminous crops is about 40 Mt/year Bean Rhizobium leguminosarum
bv. phaseoli
(Vitousek et al., 1997). Although the ability
Rhizobium etli
of fixing nitrogen by different legume spe-
Clover Rhizobium leguminosarum
cies varies considerably (soybean fixes more bv. trifolii
nitrogen than chickpea), on an average, the
major legume crops fix about 9–245 kg
nitrogen/ha (Unkovich and Pate, 2000). and fix nitrogen in legumes, such as Blasto-
bacter, Devosia, Methylobacterium, Agro-
bacterium, Phyllobacterium, Ochrobactrium,
Cupriavidus, Herbaspirillum and Burkhol-
12.3.1 Plant-bacteria specificity
deria (Balachandar et al., 2007). This group
for N-fixation
is known as beta-rhizobia to differentiate it
from the rhizobial bacteria. These species in
Nodule formation in plants is induced by association with Rhizobium, Mesorhizobium
two groups of soil bacteria, rhizobia and and Bradyrhizobium add substantial amounts
actinobacteria. A new group of beta-rhizobia of nitrogen to the soil through leaf senescence.
has recently been identified, which can Nitrogen fixation by symbiosis is also observed
induce nodule formation in legumes. Rhizo- in other non-leguminous crops (sugarcane,
bial bacteria comprise the genera Rhizo- sweet potato), where rhizobia exist as endo-
bium, Sinorhizobium, Bradyrhizobium, phytes in plants and fix nitrogen, although
Azorhizobium, Allorhizobium and Meso- nodules are not formed (Dong et al., 1994;
rhizobium. Rhizobial nodulation is limited Terakado-Tonooka et al., 2008). These recent
to the plant families Leguminosae and Ulma- discoveries clearly suggest that the impact of
ceae. The interactions of rhizobia and the biological nitrogen fixation by crop plants in
plant species are specific, and each rhizobial symbiosis with microorganisms is much more
species has a particular host range. The than anticipated one or two decades ago.
actinobacteria (hyphae-forming bacteria)
are the members of genus Frankia, which
induce nodulation in many plant families
including Casurianaceae, Myricaceae, Rham- 12.3.2 Mechanism of nodulation
naceae, Rosaceae, Betulaceae, Datiscaceae,
Eleagnaceae and Coriariaceae. However, for What is the mechanism behind the specifi-
crop productivity, rhizobial-leguminous crop city of host–rhizobium interaction? The leg-
species interaction is most important. The umes release a group of isoflavonoid and
specificity of legume crops and rhizobia is betaine compounds through their root,
described in Table 12.1. which serve as chemo-attractants to these
Apart from the food crop species of bacterial species (Cooper, 2007). The recog-
Leguminosae, a large number of tree species nition of a particular host by the rhizobia is
of agroforestry systems (Leucaena leuco- specified by the structure of the flavonoid
cephala, Calliandra calothyrsus, Gliricidia compounds released. A group of genes in
sepium and Sesbania sesban) are capable of the bacterial genome, known as nod (nodu-
biological nitrogen fixation. lation) genes are activated, releasing nod
Recent discoveries show that several non- factors. Three of these genes are common to
rhizobial microbes can induce nodulation all rhizobial species (nodA, nodB and nodC,
290 Chapter 12
develop the basic structure of Nod factors), almost throughout the period of nodule
a few provide host specificity (nodE, nodF, activity; as a consequence, they are oval in
nodP, nodQ etc.), and one acts as regulatory shape. An apical meristem region is present
gene (nodD). The flavonoid compounds in the indeterminate types, which leads to
released by the host plant are recognized by continuous growth. In both the nodules, the
the NodD protein, inducing transcription of cortex region is surrounded by a periderm.
other nod genes. The Nod factors are lipochi- The characteristic pink colour is due to
tin oligosaccharides, a group of signal mole- presence of high concentration of leghaemo-
cules. The host root, sensing the presence of globin, which plays a pivotal role in nitro-
nodulating rhizobia, bends in a characteris- gen fixation.
tic fashion to enclose the rhizobial popula-
tion, known as ‘Shepherd’s crook’. The
rhizobial bacteria move to the root hair 12.3.4 Nitrogen fixation
region or in lateral root regions, colonize
and induce nodule formation. Structural Nitrogen fixation is performed by the fol-
alteration in the root hair cells and the epi- lowing biochemical reaction catalysed by
dermal region lead to curling of the roots the bacterial nitrogenase enzyme complex.
and development of an infection thread.
A small group of cells of root cortex oppo- N2 + 8e− + 8H+ + 16ATP ® 2NH3 + H2
site to the protoxylem region, known as +16ADP + 16Pi
nodule primordium, starts dividing rapidly The fixed ammonia is rapidly transferred to
under the influence of auxin influx and dif- surrounding cells and is converted to
ferentiates into a nodule. amides (asparagines, glutamine) or ureides
(allantoin, allantoic acid, citrulline) and is
transported to shoots via xylem. The energy
for nitrogen fixation comes from oxidative
12.3.3 Nodule anatomy
phosphorylation, which requires high
amount of oxygen influx in the nodules.
A nodule is a modified root, which attains However, the reaction itself requires a
spherical or ovoid shape due to cell divi- reducing environment. The exclusion of
sion of root cortex cells. In some cases peri- oxygen from the zones of nitrogen fixation
cycle cells are also involved in nodule is performed by leghaemoglobin, which has
formation (actinorrhizal nodule, indetermi- a very high affinity for oxygen (ten times
nate legume nodule). The anatomy of the more than blood haemoglobin). In many
nodule is typical of the root with a vascular legumes the hydrogen produced is oxidized
system. However, the vascularization is by hydrogenase enzyme. In other legumes
peripheral as in the stem, with a central the hydrogen produced diffuses from the
void which is a reserve for the rhizobial nodules into the soil, where it is taken up by
population. The rhizobia penetrates the microorganisms (Peoples et al., 2008).
nodule by forming an infection thread,
which enters the nodule primordium
through root epidermal transfer cells (Lin
et al., 2008). It lives there as a bacteroid, 12.4 Higher Nutrient Assimilation
where it is encircled by plant membrane
called peribacteroid membrane. Depending 12.4.1 Anatomical modifications
on the growth characteristics, the nodules for nutrient assimilation
are grouped into determinate nodules
(groundnut, soybean) and indeterminate Nutrient is essential for optimum growth
nodules (pea). In determinate nodules, of a crop plant through efficient xylem
growth is controlled and the shape of the for assimilation of minerals through root
nodule is spherical. In the case of indeter- and its translocation to the leaves and
minate nodules, the cell division continues stems, associated with the translocation of
Anatomical Adaptation to Harvest Higher Energy 291
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13
Anatomical Adaptation for Better
Reproduction Efficiency
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
(R. Maiti et al.) 293
294 Chapter 13
colourful to attract insects, and often con- are used for oviposition and gall formation
tain nectary glands. for hatching, while others are left for seed
The size of the insect pollinator corres- production. In dioecious fig, different syco-
ponding to the size of the flower largely niums are used for seed production and gall
influences the extent of pollen attached to formation (Moe et al., 2011).
the insect body and the amount of the pol-
len shedding on stigma. This character Flower anatomy favouring
along with the frequency of visits made by pollination by other agents
the pollinator charts the success of cross-
pollination. In cotton, the bumble bee is Wind pollination is the next most important
considered to be a more efficient pollinator pollination mechanism after insect pollina-
than the honeybee. Because of the relatively tion (Table 13.3). The mechanism of wind
larger size of the bumblebee corresponding pollination evolved much earlier than insect
to honeybee and the small size of the cotton pollination, which is the primary mode of
flower, the bumble bee cannot collect honey pollination of gymnosperms. The transition
without touching the anthers, thereby col- from wind pollination towards insect polli-
lecting pollen and depositing it on the nation is considered a major landmark in
stigma of self-flower or another flower. The evolution of angiosperms (Hu et al., 2008).
honeybees on the other hand are interested This change has been intrinsically associated
in extrafloral nectary and seldom enter the with the evolution of flower anatomy, where
cotton flower. Thus the cotton genotypes several features have been added, such as a
having extrafloral nectary are preferred large flower structure and organs to produce
more by the honeybee for pollination. In sticky and sugary substances. However, dur-
contrast, the rape flower anatomy fits well ing recent evolution within angiosperms, a
to the honeybee for the collection of pollen number of species have changed their polli-
and nectar. The positioning of anther and nation mechanism from animal to wind pol-
stigma helps the stigma to be in touch with lination, suggesting some common, recurrent
the surface of the bee carrying foreign pol- mutations may be involved in such transi-
len, thereby aiding pollination. The anat- tion. The evolution of insect pollination from
omy of florets of sunflower also is well wind pollination is more prominent in prim-
suited for honeybees, where it is the princi- itive families of angiosperms, such as
pal pollinator. However, in this crop, the Cyperaceae, Caryophyllaceae and Moraceae
numbers of flower units are quite high com- (Wragg and Johnson, 2011).
pared to the average bee population. Thus The anatomy of the inflorescence of
the seed yield of a sunflower crop largely wind-pollinated crops is characterized by
depends on the size of the bee population smaller size of flowers with increase in
visiting the crop. flower number. It is observed primarily in
An interesting flower anatomy and pol- monoecious (maize) and dioecious (coconut)
lination mechanism is observed in fig, a trop- crop plants. The wind-pollinated flowers
ical fruit tree (Ramirez, 1969). The pollination
is assisted by a group of wasps belonging to
family Agaonidae. The wasp completes its Table 13.3. Crops pollinated by wind and other
life cycle within the fig inflorescence (syco- mechanisms.
nium); males never emerge from the inflores- Crop Pollination mechanism
cence, but females after emerging from one
inflorescence invade another receptive inflo- Maize Wind
rescence through a small hole called an osti- Coconut Wind, insects
ole. It carries pollen from one syconium to Date palm Manual by hand pollination,
another and during the process of oviposi- wind, insect
Beet Wind
tion helps in pollination of female flowers
Vanilla orchid Hand pollination
within the syconium. In monoecious fig,
(Vanilla planifotus)
some female flowers within the syconium
Better Reproduction Efficiency 297
generally do not produce any nectar and are meiotic events, including chromosome
of unattractive colour or shape. doubling and distribution in egg cells and
companion reproductive cells (synergids and
Position and timing of pollen landing antipodal cells in the ovule). The metabolic
activities for these events draw the required
The position of landing of pollen on stig- energy from assimilates developed in the
matic surface and the timing of pollen dis- vegetative tissues. Consequently, deposition
persal largely determine the success of of food reserve in the endosperm sink is
pollination and fertilization. For example, reduced. Plant breeders often experience
cotton stigma remains receptive from morn- negative correlation between seed number
ing to noon, but receptivity drops sharply and seed size. Simultaneous improvement in
after noon. The cotton ovule contains about both the components is a challenging task,
50 seeds, thus at least 50 pollen grains must which requires high photosynthetic activity
germinate, reach to the ovary and release as well as balanced distribution of assimilates
male gametophyte for successful fertiliza- between metabolic activity spent for repro-
tion. The position of pollen adhesion to the duction and accumulation of food reserves in
stigmatic surface is crucial for pollen hydra- the endosperm.
tion and germination. The pollen that land
on the base of the stigma germinate poorly
and receive more mechanical hindrance for
germination and pollen tube growth. 13.3.2 Reduction in accessory cost
Abundance of pollen throughout the stig-
matic surface results in better seed set and Accessory costs of reproduction include
fibre yield. This is the reason for yield energy expense towards development and
improvement in cotton fields where bee vis- maturity of seed, excluding the direct energy
its are frequent, which helps in better pol- used for embryo and endosperm develop-
len dispersal and landing during the peak ment (Thompson and Stewart, 1981). Such
receptivity period of stigma. costs include energy utilized for develop-
ment of support structures for pollination,
fertilization and seed maturity as well as
energy spent towards aborted ovules. It is fur-
13.3 Reduction in Cost ther classified in two broad categories, pre-
of Seed Production pollination cost and post-pollination cost
(Haig and Westoby, 1991). The pre-pollination
13.3.1 Reduction in direct accessory costs include investment made in
cost towards seed development development of flower structure and pedun-
cles, which is required for pollen capture
Biologically, the direct cost for seed produc- and fertilization. Development of protective
tion is defined by the amount of energy spent structures of the seed, cost of ovule packag-
for development of the seed. However, in ing and structures needed for dispersal of
broader agronomic terms, this cost also seed (wings, tufts, hairs etc.) are included
includes the cost for whole fruit development in the post-pollination accessory costs.
where fruit is the economic product (vegeta- Investigations on 14 diclinous non-crop
bles, fruit crops). In most of the grain crops, plant species revealed that the accessory
productivity improvement has been achieved costs comprise 33–96% of the total repro-
through the increase in seed size and number ductive cost (Lord and Westoby, 2006). The
of seeds per plant. For increasing the number economics of seed development largely
of seeds per plant energy has to be distributed depends on efficient management in
towards a higher number of fertilization balancing these costs, since a larger amount
events, which is ensured by more ovule and of energy is consumed for development of
pollen production. Both of these events these structures compared to the direct cost
require expenditure of more energy towards involved in seed development. The grain
298 Chapter 13
crops, adapted for better translocation of food protection to the seed. In many cereal crops,
reserve towards endosperm or cotyledon, are the seed is encased by a number of pro-
expected to have reduced accessory costs. tective structures, such as lemma, palea,
However, studies on accessory cost determi- glumes and awn. Construction of these
nation in crops are limited. More information structures requires considerable energy.
is required for understanding the role of However, these structures not only provide
domestication and selection in economizing protection and hold the grain, but also
floral anatomy in crop species. contribute significantly towards photosyn-
thesis and assimilation of food reserve in
Economization of pollen capture grains. Thus selection in the natural envi-
ronment as well as under cultivated envi-
The structural anatomy of the flower deter- ronments has been directed towards
mines the cost required for development of balancing the floral anatomy so that the
floral organs. Reduction in cost of pollen accessory costs for seed production are
capture has been achieved by many plant optimized.
species by enclosing the stamens with petal In wheat, the photosynthetic contribu-
structures for ensuring self-pollination. The tion made by these components of ear can
legume flower structure is a prominent go up to 45% (Kriedemann, 1966; Wang
example of closing the flower so that pollen et al., 2001). The awn of wheat is particu-
of self-flower is not wasted outside and larly significant as a protective structure
entry of foreign pollen is restricted. In many prohibiting entry of pests and pathogens
cross-pollinated species, pollen capture and as a light-capturing photosynthetic
is economized by grouping the pollen- organ contributing significantly particularly
receiving flowers. Maize is a prominent at the grain-filling stages (Li et al., 2006).
example where female flowers are grouped Domestication of wheat has resulted in
together in a cob so that pollen capture modification of awn structure and number.
efficiency can be increased. However, being In wild wheat, two prominent awns are
a wind-pollinated crop, the cost spent observed, which act additionally as a bal-
towards pollen development is higher (2–5 ancing organ during seed dispersal as well
million pollen grains, equivalent to 0.5– as provide motility for dispersion in air and
1.25 million meiotic divisions) compared soil (Elbaum et al., 2007). Since self-
to self-pollinated cereal crops. dispersal is not a requirement in the culti-
vated wheat species, domestication has led
Reduction in cost of packaging, to the reduction and sometimes the elimina-
protection and dispersal of seed tion of awns (awnless varieties). The reduc-
tion in awn number is a good example of
A large proportion of accessory cost is spent economization of floral anatomy of wheat,
towards packaging of ovules and providing reducing the cost of awn development.
References
Chacoff, N.P., Morales, C.L., Garibaldi, L.A., Ashworth, L. and Aizen, M.A. (2011) The Americas Journal of
Plant Science and Biotechnology 3, 106–111.
Elbaum, R., Zaltzman, L., Burgert, I. and Fratzl, P. (2007) The role of wheat awns in the seed dispersal unit.
Science 316, 884–886.
Emberlin, J. (1999) A Report on the Dispersal of Maize Pollen. Available at: http://www.mindfully.org/GE/
Dispersal-Maize-Pollen-UK.htm (accessed 18 July 2011).
Haig, D. and Westoby, M. (1991) Seed size, pollination costs and angiosperm success. Evolutionary Ecology
5, 231–247.
Hu, S.S., Dilcher, D.L., Jarzen, D.M. and Taylor, D.W. (2008) Early steps of angiosperm–pollinator coevolution.
Proceedings of the National Academy of Sciences USA 105, 240–245.
Better Reproduction Efficiency 299
Klein, M.A., Vaissière, E.B., Cane, H.J., Steffan-Dewenter, I., Cunningham, S.A., Kremen, C. and Tscharntke,
T. (2007) Importance of pollinators in changing landscapes for world crops. Proceedings of Royal Society
of London (B) 274, 303–313.
Kriedemann, P. (1966) The photosynthetic activity of the wheat ear. Annals of Botany 30, 349–363.
Li, X., Wang, H., Li, H., Zhang, L., Teng, N., Lin, Q., Wang, J., Kuang, T., Li, Z., Li, B., Zhang, A. and Lin, J.
(2006) Awns play a dominant role in carbohydrate production during the grain-filling stages in wheat
(Triticum aestivum). Physiologia Plantarum 127, 701–709.
Lord, J.M. and Westoby, M. (2006) Accessory costs of seed production. Oecologia 150, 310–317.
McGregor, S.E. (1976) Insect Pollination of Cultivated Crop Plants. Available at: afrsweb.usda.gov/SP2UserFiles/
Place/…/OnlinePollinationHandbook.pdf (accessed 13 May 2011).
Miller, P.D. (1985) Maize pollen: collection and enzymology. In: Sheridan, W.F. (ed.) Maize for Biological
Research. A Special Publication of the Plant Molecular Biology Association, USA, pp. 279–282.
Moe, A.M., Rossi, D.R. and Weiblen, G.D. (2011) Pollinator sharing in dioecious figs (Ficus: Moraceae).
Biological Journal of the Linnean Society 103, 546–558.
Ramirez, W.B. (1969) Fig wasps: mechanism of pollen transfer. Science 163, 580–581.
Stebbins, G.L. (1970) Adaptive radiation of reproductive characteristics in Angiosperms I: pollination mecha-
nisms. Annual Review of Ecology, Evolution and Systematics 1, 307–326.
Thompson, K. and Stewart, A.J.A. (1981) The measurement and meaning of reproductive effort in plants.
American Naturalist 117, 205–211.
Wang, Z.M., Wei, A.L. and Zheng, D.M. (2001) Photosynthetic characteristics of non-leaf organs of winter
wheat cultivars differing in ear type and their relationship with grain mass per ear. Photosynthetica 39,
239–244.
Wragg, P.D. and Johnson, S.D. (2011) Transition from wind pollination to insect pollination in sedges:
experimental evidence and functional traits. New Phytologist, doi: 10.1111/j.1469-8137.2011.03762.x.
14
Anatomical Basis of Crop Ideotype
for Higher Productivity
©R. Maiti, P. Satya, D. Rajkumar and A. Ramaswamy 2012. Crop Plant Anatomy
300 (R. Maiti et al.)
Crop Ideotype for Higher Productivity 301
the case of food crops. The higher the HI, in parts of China and Japan. However, it
the higher is the channelling of the food cannot withstand chilling temperature as
reserve from non-harvestable to harvestable much as another grain crop, wheat.
produce. Where the economic product is Nevertheless, it can tolerate acidic soil (pH
seed, HI is the ratio of total seed yield to <4.0) and alkalinity (pH >11.0) higher than
biomass yield. other cereal crops such as wheat, maize and
Three prominent examples where sorghum. This remarkable plasticity makes
replacement of old genotypes by new ideo- rice one of the most interesting crop species
types has been very rapid in the 20th cen- for studying crop adaptation. This also
tury are maize, wheat and rice, the principal makes the job of defining a rice ideotype
food crops of the world. There are several more difficult, as the ideal type of culti-
other examples, where plant types have vars will vary according to the environment.
considerably been changed to suit the agri- To augment anatomical structures to these
cultural system or to adapt to higher envi- ideotype descriptions is a mammoth task
ronments. Needless to say, the structural requiring in-depth study of structural
anatomy of the ideotypes has also changed changes of major plant parts under such
considerably, and played a significant role environments.
in adaptation of new genotypes in the pro- Rice was predominantly cultivated in
duction system. In some cases, ideotype Asian countries as photosensitive, tall
construction has been directed by selective plants producing few tillers. The Asian cul-
anatomical features, while in other cases tivated rice is differentiated in three eco-
anatomy has played an underlying but geographic races, indica, japonica and
important role in formation and selection of javanica. The race differentiation culmi-
plant ideotype. The criterion for this ideal nated from climatic adaptation and human
plant type was fixed by Donald as a plant selection. Indica rice is adapted to the
type that would ‘make a minimum demand humid, tropical to subtropical environment
on resources per unit of dry matter pro- of South Asia, particularly the Indian sub-
duced’. This led to an important conclusion continent. It is characterized by tall plants
that the sink must be large enough to store with many tillers spreading from the base.
all the photosynthetic reserves produced The grains are borne on tall panicles, are
by the plant. Although Donald proposed medium to long and are easily threshed.
ideotypes for wheat and barley that will The japonica rice is short and adapted to a
contain few tillers, erect leaves, small can- temperate climate, bears a high number of
opy size and higher seed number, the ideo- tillers and short panicles with short, bold
type breeding has been most successful in grains. The javanica rice is adapted to the
another important cereal crop, rice (Virk tropical climate of Indonesia, being taller
et al., 2004). but bearing fewer panicles than indica rice.
The anatomical differences among these
three types are prominently observed in
culm and leaves.
14.2.1 Ideotype for increasing Early rice ideotype in the first half of
productivity in rice 20th century could be described by geno-
types that matured late, as the predominant
Rice is probably the only crop to be grown cropping system was single crop (rice fol-
in diverse ecological conditions, starting lowed by fallow land). As the reproductive
from above 2000 m in small villages of the growth phase in rice is fixed at around
Himalayas in India and Nepal to below sea 25–35 days, late maturity types had higher
level in southern India and South-east Asia. vegetative growth and photosynthesis, this
As a crop it withstands extreme dry condi- being channelled into grain development
tions and high temperature in western India for higher productivity. Longer vegetative
and Pakistan. On the other hand, it is also growth also led to higher cellular growth
grown under moderately low temperatures and maturation, providing better strength to
302 Chapter 14
the tillers. The anatomical structure of stem solar energy. Although anatomical features
and leaves of these types reveals higher were not the basis of selection of semi-dwarf
maturity of vascular bundle tissue and rice, they served to develop the ideotype,
higher deposition of lignin, suberin and sil- which was selected by the plant breeders at
ica in the stem. The leaves of these geno- morphological level.
types are also thicker with a higher number A new plant type (NPT) was developed
of cell layers and increased thickness of cell in the late 1980s by modifying the features
wall. Resistance to blast disease (causal of the semi-dwarf varieties (Khush, 1995).
organism – Pyricularia oryzae) is also con- The semi-dwarf varieties produce many
sidered as a character for rice ideotype in unproductive tillers with small panicles,
this period, although the formal concept of limiting the sink size as well as contributing
ideotype was developed much later. to unproductive vegetative growth (low HI).
A second ideotype was developed by The NPT for irrigated tropical indica rice,
combining the desirable features of indica which is capable of yielding 10–14 t/ha,
and japonica rice. Both these ecotypes have consists of a semi-dwarf variety with 8–10
their inherent problems of low productivity. tillers bearing heavy panicles with a high
Development of an ideal type combining number of grains per panicle (200–250), a
the features of both of these ecotypes (long plant height of 90–100 cm, high grain weight
panicle with higher grains per panicle, (25 g/1000 grains) and increased HI (Virk
semi-dwarf non-lodging stature with high et al., 2004). The NPT is designed to have
tillering) was targeted by the breeders of higher leaf growth at the vegetative stage
many countries to improve productivity, and higher translocation of food reserves
including Korea, Indonesia and India. Since leading to more grain filling with a longer
the two ecotypes exhibit high sterility in reproductive phase. To sustain such heavy
intercrosses, limited success was obtained, panicles with high grains the stem strength
except in Korea. At structural anatomy should be very high with vigorous root
level, strong, well-developed vascular bun- growth. Thus the morpho-anatomical struc-
dle was combined with higher panicle ture of NPT rice should bear dense vascular
extensibility, rapid cellular growth, better bundles, with higher cell wall thickness.
translocation of soil nitrogen to leaves and Better translocation of food reserve during
structures favouring higher translocation of reproductive growth phase depends on the
food reserves from source to sink. activity of phloem tissues, which should be
Development of semi-dwarf, high well developed in NPT rice.
fertilizer-responsive rice varieties in the
1960s in the International Rice Research
Institute (IRRI) in the Philippines transfor-
med the rice production system, uplifting 14.3 Changes in Crop Anatomy in
the yield potential of rice from 6 t/ha to Vegetative Structures of Economic
10 t/ha (Virk et al., 2004). The plant features Importance
of these new semi-dwarf varieties were
short stature, sturdy stem with profuse till- Apart from the grain crops, a substantial
ering, dark green and erect leaves (Jennings, amount of food requirement is supplied by
1964). The semi-dwarf rice types could sup- starchy and sugary crops, where seed is not
port rapid cellular growth and development the economic product. These include potato
by the uptake of more nitrogen from the (stem tuber), sweet potato (root tuber) and
soil and channelling it for further vegetative cassava (root tuber), which form the staple
growth. This led to an increase in the number food crop of millions of people in different
and thickness of cell layers in the stem and countries. There are two other major crops,
leaves, higher cellular growth, and more sugarcane (stem) and sugarbeet (root tuber),
cell layer formation in the root. The leaves where vegetative structures are the princi-
of semi-dwarf rice exhibit better developed pal source of sugar. An interesting distinc-
chloroplasts with a higher capacity to trap tion of these species from the seed crop
Crop Ideotype for Higher Productivity 303
species is that in most of these crops, seeds where raised nodes with buds are present. It
are formed through sexual reproduction, is attached to the main stem with a narrow
but the food reserve is channelled in special- attachment known as a stolon. The growth
ized vegetative structures. As these vege- and size of the tuber depends on the number
tative structures are economic products to and size of the stolons. Genotypes with
humans, selection has been practised for many stolons produce numerous small
the enhancement of the size and shape of tubers. This leads to low productivity, as
these vegetative structures. In natural popu- the plant’s energy and food reserve is
lations, these structures have primarily been divided among many translocation paths.
evolved for asexual reproduction, where Cellular density in the tuber is yet
new clones are developed from specialized another important anatomical feature. Large
tissues (e.g. scale or eye in potato, which is cells pack larger starch granules, while in
a dormant bud). The new plants derive their small cells the size of the starch grains are
initial food requirement from the food smaller. The rate of enlargement of starch
reserve in these storage tissues. Efficient grains in the tuber cells is also important to
channelling of the starch and sugar in these increase productivity without extending
reserve tissues depends primarily on the the period of tuberization and maturation.
metabolic processes leading to food reserve A high rate of starch granule formation is
development and structural anatomy of the thus a selection criterion for early maturing
plant for translocation of the reserve to varieties without sacrificing productivity.
storage tissues. The late maturing varieties have higher cel-
lular density. The rate of cell division is
higher at the pith than the cortical region in
the initial stages of tuber development. In
14.3.1 Potato the later stages, extensive growth takes place
between the region of cortex and the pith.
Potato (Solanum tuberosum subsp. tubero-
sum) is by far the most important tuber crop
and is grown in about 150 countries. A
related species S. tuberosum subsp. andi- 14.3.2 Sugarbeet
gena is cultivated to some extent in Chile.
Although the crop originated in South About 25% of the total sugar produced in
America, its domestication is one of the the world comes from sugarbeet (Beta vul-
most recent events in the history of crop garis L. ssp. vulgaris). It is a crop that came
domestication. Although potato cultivation under cultivation only in the 19th century
dates back to 13,000 BP in Chile (Ugent et al., and consequently, the domestication pro-
1982), the tetraploid S. tuberosum subsp. andi- cess of sugarbeet as a crop is very recent and
gena was brought to Europe by the Spanish the genetic base of present-day beet varie-
invaders during the discovery of the New ties is narrow. However, it is possibly the
World. Only during the mid-1800s, when all only crop where scientific breeding started
the cultivated clones of S. tuberosum subsp. as early as the process of domestication.
andigena were devastated by late blight The process of mass selection and progeny
disease (causal organism – Phytophthora testing was first carried out in this crop by
infestans), a resistant S. tuberosum subsp. Louis de Vilmorin during the 1850s, which
tuberosum clone (Rough Purple Chili) was is considered as a pioneer work in scientific
introduced to save the European countries plant breeding.
from food crisis. The present-day potato A significant change in crop architec-
cultivation spread very quickly in the rest of ture of sugarbeet has been obtained by
the world within a very short period of inducing autopolyploidy by colchicine. The
time. tetraploid varieties (2n = 4x = 36) produced
The potato tuber is a shortened fleshy better root shape, larger leaves with stron-
stem. Botanically it is an enlarged branch ger petioles with better photosynthesis.
304 Chapter 14
References
Ceballos, H., Okogbenin, E., Pérez, J.C., López-Valle, L.A.B. and Debouck, D. (2010) Cassava. In: Bradshaw,
J.E. (ed.) Handbook of Plant Breeding: Root and Tuber Crops. Springer, New York, pp. 53–96.
Cock, J.H. (1984) Cassava. In: Goldsworthy, P.R. and Fisher, N.M. (eds) The Physiology of Tropical Field Crops.
Wiley, New York, pp. 529–549.
Donald, C.M. (1968) The breeding of crop ideotypes. Euphytica 17, 385–403.
FAOSTAT (2012) Available at: http://faostat.fao.org/site/567/DesktopDefault.aspx?PageID=567#ancor
(accessed 9 March 2012).
Heinzen, F., Ramos, J. and Tivano, J.C. (2002) Comparative quantitative anatomy of grass species in Santa Fe
province. Revista Fave Seccion Ciencias Agrarias 1, 57–63.
Jennings, P.R. (1964) Plant type as a rice breeding objective. Crop Science 4, 13–15.
Khush, G.S. (1995) Breaking the yield frontier of rice. GeoJournal 35, 329–332.
Reilly, K., Gomez-Vasquez, R., Buschman, H., Tohme, J. and Beeching, J.R. (2003) Oxidative stress responses
during cassava post-harvest physiological deterioration. Plant Molecular Biology 53, 669–685.
Saunders, J.W., McGrath, J.M., Halloin, J.M. and Theurer, J.C. (1999) Registration of SR94 sugar beet germ-
plasm with smooth root. Crop Science 1, 297.
Ugent, D., Pozorski, S. and Pozorski, T. (1982) Archeological potato tuber remains from the Casma Valley of
Peru. Economic Botany 36, 182–192.
Virk, P.S., Khush, G.H. and Peng, S. (2004) Breeding to enhance yield potential of rice at IRRI: the ideotype
approach. International Rice Research Notes 29, 5–9.
Index
abscisic acid (ABA) 14, 275, 279 archaeology see ancient civilizations
abscission, cotton leaves and bolls 167 aster yellows virus, flax 195–196
actinobacterial nodules 289, 290 aubergine (eggplant, brinjal,
aerenchyma 9 Solanum melongena)
rice leaves, physiological anatomy
effects 49, 49, 50, 283 leaves 223–224, 224
rice roots and stems 47, 47, 48, 282–283 petiole 225
African rice (Oryza glaberrima) 44, 45 roots 220–222, 221
aleurone layer, seeds 50, 62, 79, 90–91, 91 seed coat 225, 226
allergic reactions 219 stems 222–223, 223
aluminium toxicity 291 cultivation and origin 218–219
anatomy effects of chromium 224–225
current applications 4–5, 6–14 morphology 220, 220
definition and scope 1–2, 21 shoot/fruit borer
historical studies 2–4, 3 susceptibility 223, 225
ancient civilizations uses and medicinal value 219–220
beginnings of agriculture 300 awns, functions 298
beliefs and traditions, crop plants 51, 52
crop productivity (harvest index) 288
depictions of crops 70 bark, screening method for fibre 179–180
documented plant studies 2, 232 bean, common (Phaseolus vulgaris)
evidence for spread of cultivation 119, 142, anatomy 105–106
156, 226 domestication, molecular biology
plant products, archaeological evidence 104
remains 11–12, 191–192, 196 drought tolerance, genotypic
animal feed see fodder crops variation 106, 281–282
anthesis morphology 104–105
cereals 46, 65 production and food importance 104
legumes 97–98 bees, and crop pollination 127, 202, 295,
antioxidants 295–296
enzymatic activity 133, 171, 281 biofuels see fuels, crops as sources
in vegetable foods 207, 213, 232, 263 biotic stress
aphid resistance 249 disease, pest and weed threats 269
Arabidopsis thaliana (model plant) 12, 39, 271 host defence mechanisms
307
308 Index
deep water rice cultivars 283 ecogeographic races (rice) 44, 45, 47, 301–302
desi (Indian chickpea type) 95, 96, 98 eggplant see aubergine
dicot plants electron microscopy 17–18
dorsiventral leaves 33–35, 34 embedding and dehydration, specimens 16
roots 26–27, 27, 28 embryo (germ) anatomy 62–63, 69, 80, 91
stems 30, 30–32, 31, 128–129 embryonic development 30, 41, 63, 70, 175–176
differentiation in pollinated and unpollinated
lint and fuzz, cotton 174 fruit 235–236
phases of plant development 21–22 endodermis
plasticity of plant cells 10 root 26, 28, 54, 105
root nodule bacteroids 100, 100, 117 stem 31, 113
secondary fibre elements 186 endophytes, N-fixing 289
vascular elements, from procambium 193 endosperm (seeds) 39, 40
digestibility, animal fodder 13, 66–67, 74, 305 corneous and floury 62, 77, 78,
diseases, crop plant 79–80, 90–91
antagonism with root nodule embryonic development 70
symbionts 108 nutritional quality 63
definition 269 environmental stress
fungal infections 118–119, 270, anatomical adaptations 9–10
271–272, 303 flooding submergence 282–283
host responses, resistant water stress and drought 278–282
and susceptible 137–138 development of adapted genotypes
viral (rice) 301–302
abnormal phloem symptoms 195–196 tolerance of poor conditions 81, 107,
and cell sap flow rate 272 115, 189, 278
insect transmission, control 210 enzymes
domestication process 51, 64, 80 activity and drought stress 133
allelic changes 70 expression analysis 13
investigation by molecular biology role in fibre cell lignification 194–195
(bean) 104 epicuticular wax
selection and genetic resources 285–286, amount, correlation with seed yield 146
300, 303 bloomless mutants (sorghum) 75
sites, for individual crops form of deposits 67–68, 76, 130, 262
chickpea 95 pest and disease resistance
cowpea 115 properties 262, 270
flax 191 and water loss by transpiration 279
groundnut 134 epidermal peel (impression) technique 18
maize 51 epidermis
rice 44 leaves 33, 35, 141, 144–145
sesame 142 root (rhizodermis) 26, 28, 105
sugarcane 305 stem 30, 32, 105
sunflower 125–126 see also cuticle; trichomes
tomato 206 esterification, cell wall 272
wheat 64 ethanol fuel production 52, 71, 305
dormancy, seeds 14, 40, 133 evolution
drought tolerance anatomical modifications 10–11
anatomical adaptations 9, 35, 36, floral structure and pollination 293,
152, 278–280 295, 296
cereals 55, 59–60, 69, 80, 280–281 pest and crop co-evolution 269
cotton 158, 159, 163, 167–168 photosynthetic systems 286, 287
legumes, arid region 106, 114, timescale, maize origins 51
119, 137, 281–282
vegetable genotypes 211, 215, 224, 231
effects on fibre yield 196, 199 fertilization process
effects on seed size and number 146 effects of weather conditions 112
escape, by early maturity 95, 107 efficiency of pollination
physiological adaptations 133 mechanism 293, 294
Index 311
tossa jute (Corchorus olitorius) vegetable crops see under individual crop species
compared with white jute 182, 188 vegetative storage organs, crops 285, 302–305
genotypic variation 177, 184, 185, 185, 186 vitamin C, vegetable sources 207, 259
origins and centres of diversity 181
transgenic plants 12
trichomes water flux, roots 159
cotton leaves 163, 164–165, 165, 166, 167 water stress see drought tolerance
effects of length and density 118, waterlogging
132–133, 137 hypoxia and toxicity stresses 282
glandular tolerance, anatomical adaptations 9, 235,
active antibiotic compounds 210 282–283
aphid deterrence, cucumber 249 watermelon (Citrullus vulgaris)
chemical exudates 102–103, 273–274 adaptations to flooding 235
secretory activity development anatomy
(hemp) 201 leaves 233–234
inheritance patterns 274–275 petioles 234, 234–235
pest susceptibility increase, examples 275 roots 233
physical defence function 98, 106, 114 stems 233, 233
pest oviposition consumption and uses 231, 232
deterrence 211, 230, 273 morphology 232, 232–233
sorghum leaves 75, 76, 76–77, 273, 275 origins of cultivation 232
sunflower, genotypic variation 129, 130, pollination and fruit development 235, 236
131, 132, 132–133 waxes see epicuticular wax
tubers, food storage 285, 302–304 wheat (Triticum spp.)
anatomy
leaves 67–69, 68
ultimate fibre cell, form roots 65, 66
and dimensions 177, 178 seed development 69–70
flax 194, 194 stems 66–67, 67
hemp 200–201 cultivated species 63–64
jute 186, 186, 188 morphology 64, 64–65
kenaf 190, 191 origin, domestication and uses 64, 298
ramie 197–198, 198 soil aluminium tolerance 291
sunnhemp 202–203 white jute (Corchorus capsularis)
ultraviolet (UV-B) radiation 165, 207 bundle structure and fibre quality 177,
179–180
compared with tossa jute 182, 188
vascular tissues cultivation origins 181
bundle arrangement genotypic variation 185, 186, 187
bundle cap length and peg strength, whitefly, tomato (Bemisia tabaci) 210, 211, 213
groundnut 135 wild crop plant relatives
fibrous sheaths, stems current distribution
and leaves 48, 58, 73, 85 maize 51
strands, in floral parts 39 rice 44
structure in C3 plants 68–69 sesame 141–142
structure in C4 plants 36, 58–59, drought resistance 282
59, 75, 85–86 identification of crop ancestors 134
in cotyledons, taxonomic variation 42, 42 pest and disease resistance 118, 210,
history of early observations 2–3 269, 274
leaf veins 34–35, 35–36, 102 variation, and character selection 8–9, 50,
in roots 26–27, 28–29 285–286
in stems 30, 31, 32, 33, 102
types, classification 26
vessel size variation, and water stress 56, xerophytic plants 9, 34, 59
149, 150, 151, 159
see also phloem
Vedic literature, plant studies 2, 232 yarn quality, composites 188