Biotic Stresses of Okra: Present Status and Future Breeding

Strategies

B Singh and SK Sanwal

Indian Institute of Vegetable Research (IIVR), Varanasi

India is the largest producer of okra, grown in an area of 0.49 m hectares

with annual production of 5.78 million tones and a productivity of 11.6
MT/ha (Indian Hort. Data Base, 2011). It is considered as an important fruit
vegetable crop of the tropical and sub-tropical regions of the world. It is
cultivated on a wider scale in warmer parts of temperate Asia, southern
Europe, northern Africa, the United States of America, and almost all parts of
the tropics. Okra production in tropical regions is constrained by several
adverse factors. The yield losses due to diseases are quite substantial. This
problem has been compounded further with increasing spread of very few
superior cultivars and hybrids leading to development of disease infestations
of epidemic proportions which may pose disastrous consequences.
Furthermore, in countries where the yielding capacity and quality
requirements of crops have been subjected to deliberate and intensive
breeding over many generations and yield appears to be stagnating, further
advances in performance can be more economically achieved by removing
factors which limit maximal yield, such as disease susceptibility.

Viruses pose serious constraints to okra production. Okra is susceptible to at

least 19 plant viruses (Brunt et al. 1990; Swanson & Harrison 1993). These
viruses severely affect okra production in terms of yield and fruit quality.
Among these viruses yellow vein mosaic virus (YVMV) causes significant
losses in the okra production.

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Yellow Vein Mosaic Virus

YVMV disease of okra was first reported in 1924 (Kulkarni 1924) during
the erstwhile Bombay Presidency in India and later studied by Capoor and
Verma (1950) and Verma (1952). This is the earliest report of this virus,
implying that BYVMV might have originated in India. Further Uppal et al.
(1942) established the viral origin of the disease and coined the name yellow
vein mosaic (YVM). It has been shown to be a geminivirus based on its
morphology and its serological relationship with African cassava mosaic
virus (Harrison et al., 1991). Inoculation of bhendi plants with cloned
BYVMV DNA, a monopartite begomovirus, produced mild symptoms;
typical vein yellowing symptoms were produced only in association with the
cognate betasatellite (Jose and Usha, 2003), possibly due to the silencing
suppression activity of the βC1, reported later (Gopal et al. 2007). The CP
showed nuclear localization, whereas the βC1 localized to the cell periphery
(Kumar et al. 2006). Since 2003, it was assumed that the disease is caused by
a complex consisting of the monopartite begomovirus Bhendi yellow vein
mosaic virus (BYVMV) and a betasatellites (Jose and Usha, 2003). But
recently a detail survey on the disease and causal virus revealed that
minimum 16 different begomoviruses and 4 different betasatellites are
associated with the disease in different combination under different agro-
ecological zones (Venkataravanappa et al., 2011).
In the recent past, frequent break down of the YVMV resistance have
been observed in popular varieties like Parbhani Kranti, P-7, Arka Anamika,
Arka Abhey in all over the country as they were in all probability
symptomless carriers or else new strains of virus have evolved. The
hypothesis of evolution of new strains of virus seems to be one of the factors
leading to break-down of tolerance as the tolerance in most of the cases has
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been reported to be location specific. The emergence of the polyphagous ‘B’
biotype of B. tabaci with its increased host range of more than 600 plant
species, has resulted in geminiviruses infecting previously unaffected crops.
Whiteflies and YVMV are largely influenced by weather conditions. In
north India, YVMV severity is pronounced in rainy season crops due to high
temperature and humidity. It has been reported that okra sown in June and
pods reaching to marketable stage in July-August were least susceptible to
YVMV (4.1 %) as compared to 92.3 % infection in okra sown in July and
maturing in August-September (Roychaudhary et al., 1997). In Kalyani, the
population dynamics of whitefly was monitored throughout the seasons and
found to be remarkably low during February to 1st fortnight of April and
reached its peak in the month of August (Chattopadhyay et al; 2011).
Under Bangalore conditions, the occurrence of whitefly and that of
YVMV is the highest during March to June. In contrast to this, the vector
population and YVMV incidence are less during the cooler months. This
happens as the hot and dry weather conditions favour fast multiplication of
whitefly and also these conditions facilitate easy movement of whiteflies
transmitting the YVMV. Cooler weather with high relative humidity and
rainfall were detrimental to whitefly population and spread. (Singh, 1980;
1990).

Hot-Spots for YVMV

The occurrence of YVMV is severe in certain locations in certain seasons

and accordingly these locations are called as hot-spot locations for field
screening of genotypes against YVMV. In north India, in general, rainy
season is most conducive for occurrence and spread of YVMV. In rainy
season also, certain locations are most ideal for screening against okra
YVMV. These locations include Karnal, Tarai region of Uttarakhand, Nadia
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district of West Bengal and Varanasi area of Uttar Pradesh. In central/south
India, the disease is pronounced in summer season at several locations, the
prominent ones being Guntur in Andhra Pradesh, Jalgaon in Maharashtra,
Surat in Gujarat and Coimbatore in Tamil Nadu. For western Maharashtra,
summer season is conducive for YVMV than the rainy season (Prabu et al.,
2007). In Pune, the disease occurs severely if the crop is planted in mid-April
and harvested in beginning of June. For screening at hot-spot locations, Pusa
Sawani is sown all around the experimental plots and within the experimental
plots at regular intervals to provide inoculums in abundance. No spraying
should be done to control insects and YVMV.

Resistant sources

In India, inter-varietal hybridization followed by pedigree selection produced

the widely cultivated, high yielding, yellow vein mosaic virus tolerant
cultivar, Pusa Sawani. This was called as symptomless carrier and is no more
tolerant. Kashi Pragati (VRO 6), Kashi Mohini (VRO 3), Varsha Uphar,
Hisar Unnat came later on as a result of inter varietal crosses and were
reported to be tolerant to YVMV. Inter-specific hybridization (with
Abelmoschus manihot L. Medikus subsp. Manihot) has been followed in
development of YVMV tolerant cultivars like Punjab Padmini, P 7, Parbhani
Kranti, Arka Anamika, Arka Abhaya, etc. In case of inter-specific
hybridization, 1-2 backcrosses have normally been used and then the material
has been handled as per pedigree method of breeding. However, at present,
none of the above cultivars are showing stable and durable tolerance to
YVMV due to break-down of tolerance as they were in all probability
symptomless carriers or else new strains of virus have evolved. The
hypothesis of evolution of new strains of virus seems to be one of the factors

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leading to break-down of tolerance as the tolerance in most of the cases has
been reported to be location specific.

Eight Abelmoschus species occur in India. Out of these, A. esculentus is the

only known cultivated species. Accessions resistance to YVMV are A.
manihot, A. angulosus, A. crinitus, A. vitifolius, A. tuberculatus, A.
panduraeformis, A. pungens and A. tetraphyllus (Dhankar & Mishra 2004;
Singh et al. 2007). A. manihot ssp. manihot is a good source of resistance to
YVMV (Sharma & Sharma 1984b) and has been widely used in resistance
breeding. A. manihot ssp. manihot, A. manihot ssp. manihot var. ‘Ghana’ and
West African okra were symptomless carriers of YVMV and may be useful
in developing YVMV-resistant hybrids (Dhankar & Mishra 2004).

Enation Leaf Curl of Okra

This disease was first observed at Indian Institute of Horticultural Research,

Hessarghatta, Bangalore as reported by Singh (1984) and Singh and Dutta
(1986). Later on, Chakraborty et al. (1997) reported the occurrence of this
disease from Varanasi in Uttar Pradesh. Initial symptoms of this disease
include small, pin-headed enations on leaves, leaf curling, followed by warty
and rough texture of the leaves. Later on, leaves begin to curl in adexial
direction. The undersurface of the leaves is characterized by mild, bold and
prominent enations. There is twisting of main stem, lateral branches and leaf
petiole. The leaves become thick and leathery. Leaf curling and enations are
more prominent in middle aged leaves. In severe cases, there are enations,
leaf thickening and curling even in the young leaves. At times, the twisting
and the bending of the stem are so severe that the entire plant looks spreading
on the soil surface. The infected plants either do not produce fruits or
produce few deformed and small fruits unfit for marketing and consumption.

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The virus is transmitted by whitefly (Bemisia tabaci). Electron microscopic
observations of ultrathin sections of leaf mid-rib of okra plants infected with
enation leaf curl disease revealed the presence of bacilliform particles
measuring approximately 85 x 300 nm in size (Singh, 1990). Depending
upon the stage of infection, yield losses from 30 to 100 % have been reported
(Singh, 1986). A minimum of 15 minutes of acquisition feeding on the virus
source is required by the whiteflies to acquire the virus from the infected
plant. However, increased acquisition feeding period enhances the percentage
of transmission. The inoculation threshold period is reported to be 15
minutes. Whiteflies fed on virus source continuously for 12 hrs remain
viruliferous throughout the life time (Singh, 2005). Perhaps no information is
available on sources of resistance and breeding programmes for resistance to
enation leaf curl virus of okra.

Cercospora Leaf Spot (Cercospora abelmoschi, C. malayensis)

In India, two species have been reported to cause this disease. C. abelmoschi
causes sooty black, angular leaf spots while C. malayensis produces brown,
irregular spots. The severely affected leaves roll, wilt and fall. Under severe
infection during hot and humid seasons, there may be defoliation of leaves.
No information is reported on sources of resistance and breeding for
resistance to this disease.

Fusarium Wilt (Fusarium oxysporum f. sp. vasinfectum)

Fusarium wilt is a serious disease of okra. The fungus persists in soil for a
long time. Initially the plants show temporary wilting symptoms which
become permanent and progressive affecting more plant parts. The leaves of
the affected plants show yellowing, loose turgidity and show drooping
symptoms. Ultimately the plant dies. In older plants, leaves wilt suddenly and

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vascular bundles in collar region become yellow or brown. The fungus
invades the roots, colonizes the vascular system and thereby restricts water
translocation. Cutting the base of stem reveals a dark woody portion along
with dark brown streak underside of bark. Disease is soil borne and spreads
through inter-culture operation. In this case, no breeding reports are
available.

Powdery Mildew (Erysiphe cichoracearum, Sphaerotheca fuliginea)

The disease caused by E. cichoracearum is most common in major okra

growing areas of India while the powdery mildew caused by Sphaerotheca
fuliginea has been reported from Bangalore area. The disease affects mainly
the older leaves, petiole and even the stem. The yield loss is due to premature
foliage loss due to foliage defoliation. The severity of the disease gets
increased under high humidity. The symptoms appear as blotches of white
powdery coating mainly on the lower surface of the leaves but may appear on
the upper surface of the leaves as well.

Future strategies

Identification of stable sources of resistance to YVMV & OELCV

Development of pre-breeding lines using identified resistant sources
To develop a liens combined resistance to fusarium wilt and powdery mildew
for fresh market and processing type.
Genetic and cytological investigations in inter-specific hybrids and
inheritance pattern specific to wild species and their derivatives
Identification and characterization of predominant viruses/virus strains for
horizontal breeding programme
Host-Virus-Vector relationships in okra

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