Beruflich Dokumente
Kultur Dokumente
Dr Kiran K. Sharma
Genetic Transformation
Laboratory
ICRISAT (International Crops
Research Institute for the Semi-
Arid Tropics)
Patancheru
Andhra Pradesh - 502 324,
INDIA
E-mail: k.sharma@cgiar.org
Dr Reto J. Strasser
Laboratory of Bioenergetics
University of Geneva
1254 Jussy, SWITZERLAND
E-mail:
reto.strasser@bioen.unige.ch
RATIONALE
Only about 13% of the total pulse crop yield is produced under irrigation. Pulses are
produced to a large extent in marginal agricultural areas with poor soil quality, since
their intrinsic characteristics allow production under harsh environmental conditions.
As a result, the crop plants have acquired primitive traits such as long duration,
indeterminate growth, bushy habit and flower drop, etc. rather than high yield. There
is also general agreement that the production of pulses is very much limited by
abiotic stress factors evoked by soil salinity, drought, and low temperature. A shift
from rainfed to irrigated cultivation is very unlikely, since pulses are not an important
cash crop and thus have a secondary status in the farming system.
Four different genes will be transferred into chickpea to increase the resistance to
salt, drought, and low temperature by metabolic pathway engineering.
• P5CSF129A (mutant gamma-pyrrolidine-5-carboxylate synthetase) ->
glutamate to gamma-pyrrolidine-5-carboxylate -> proline
• mtID (mannitol-1-phosphate dehydrogenase) -> mannitol-1-phosphate
dehydrogenase to mannitol
• codA (cholin oxidase) -> choline to glycinebetaine
• annexin -> annexin protein
Being a good source of protein especially for vegetarians, chickpea (Cicer arietinum
L.) is grown as the most important pulse crop in India . Crops grown in over 90% of
arable land are exposed to abiotic stress. Chickpea is sensitive to drought, salinity
and frost/low temperature stresses, and the overall yield of chickpea is often
curtailed significantly due to its exposure to these stresses. Therefore, investigations
carried out in this project were aimed at enhancing the tolerance of chickpea to
drought, salinity and frost/low temperature stress by introducing the genes for
compatible solutes (the codA gene for glycinebetaine biosynthesis and the P5F129A
gene for proline biosynthesis), which are well established to impart abiotic stress
tolerance, through modern genetic manipulation technology. Both these genes could
be successfully introduced independently into chickpea through Agrobacterium
mediated transformation using half-embryonic axis with single cotyledon as explants,
following the protocols that were fine-tuned by us earlier. The putative transformants
selected after rigorous selection for over three months with regular (15 day)
subculture in the presence of 50 mg/l kanamycin were rooted in the presence of 50
mg/l kanamycin and/or 20 mM choline following a short treatment (3 sec) of the cut
end of the shoots with 100 mM IBA. A number of putative codA transgenic genotypes
could be successfully established in pots filled with garden soil and manure in the
ratio of 3:1. PCR analysis carried out independently with nptII and hpt primers
revealed that only five out of twenty six independent T0 genotypes well-established in
pots showed the presence of both nptII as well as hpt marker genes. Healthy
seedlings obtained from the seed of five independent T0 lines germinated in the
presence of 50 mg/l kanamycin and 20 mg/l hygromycin and were established in
pots. All the T1 plants established in the pots showed the presence of both nptII as
well as hpt marker genes suggesting the presence of the codA gene in them.
Western analysis confirmed the presence of the protein choline oxidase in all the
independent T0 and T1 lines that possessed npt II as well as hpt marker genes,
demonstrating that the prokaryotic codA gene had not only been integrated into the
genome of these transgenic lines, but could also be translated successfully. In the
subsequent season, seeds of nine independent codA transgenic genotypes (five of T2
generation and four of T1 generation were collected).
51 putative transgenic lines with P5CSF129A gene were advanced to T3 generation
and their confirmatory molecular analysis is currently underway. The seed material
was harvested from all 51 lines. PCR analysis showed over 75% of the selected
plants to be positive for nptII gene and showed transcription in RT-PCR analysis of
the mRNA. The P5CSF:129A positive plants showed the overproduction of proline in
the range of 2.0 to 5.0 fold in individual transgenic lines (10 lines tested) of
chickpea. The transgenic lines were advanced to T3 generations and will be used for
physiological, molecular and biochemical studies under glasshouse (for drought in
dry down experiments) during the second phase of this project.
Chlorophyll and fluorescence analysis is recognized as a useful tool for quick and
easy detection of the influence of many stress types on plants such as drought, heat,
cold etc.. Protocols were fine-tuned to determine abiotic stress tolerance, in
particular drought and salinity, using the Plant Efficiency Analyzer (PEA), a technical
set-up developed at the University of Geneva . These methods were adapted to
chickpea plants and experiments confirmed their utility in analyzing 9 chickpea
varieties under normal and stress conditions. Drought stress was the predominant
interest of the studies. It was shown that all 9 varieties suffered under drought
conditions, but recovered after rewatering. Depending on the variety, a recovery
between 65% and over 100 % in comparison to the unstressed control variety was
observed.
With the aim to develop a quick test, preliminary experiments with detached leaves
were carried out (mimicking salt stress and drought conditions on filter paper and
with salt solutions). A first comparative analysis of putative transgenic chickpea lines
carrying the P5CSF129A gene revealed that the performance index (indication of
photosynthetic efficiency) of transgenic lines was significantly superior to that of wild
type.
***********************************************************************
Cicer arietinum L.
Contributors: F.J. Muehlbauer and Abebe Tullu<
Copyright © 1997. All rights Reserved. Quotation from this document should cite and
acknowledge the contributors.
1. Common Names
2. Scientific Names
3. Uses
4. Chemistry
1. Traditional Medicinal Uses
5. Origin
6. Botany
1. Taxonomy, Morphology and Floral Biology
2. Ecology
7. Crop Culture
1. Field Cultivation
2. Harvesting
8. Yields and Economics
9. Germplasm
10. Biotic Factors
11. References
12. Selected Experts
Common Names
Bengal gram (Indian), Chickpea (English), Garbanzo (Latin America),
Hommes, Hamaz (Arab world), Nohud, Lablabi (Turkey), Shimbra
(Ethiopia)
Scientific Names
Species: Cicer arietinum L.
Family: Leguminosae
Uses
Chickpea is grown in tropical, sub-tropical and temperate regions.
Kabuli type is grown in temperate regions while the desi type chickpea
is grown in the semi-arid tropics (Muehlbauer and Singh, 1987;
Malhotra et al., 1987). Chickpea is valued for its nutritive seeds with
high protein content, 25.3-28.9 %, after dehulling (Hulse, 1991).
Chickpea seeds are eaten fresh as green vegetables, parched, fried,
roasted, and boiled; as snack food, sweet and condiments; seeds are
ground and the flour can be used as soup, dhal, and to make bread;
prepared with pepper, salt and lemon it is served as a side dish
(Saxena, 1990). Dhal is the split chickpea without its seedcoat, dried
and cooked into a thick soup or ground into flour for snacks and
sweetmeats (Saxena, 1990; Hulse, 1991). "Sprouted seeds are eaten
as a vegetable or added to salads. Young plants and green pods are
eaten like spinach. A small proportion of canned chickpea is also used
in Turkey and Latin America, and to produce fermented food. Animal
feed is another use of chickpea in many developing countries. An
adhesive may also be prepared; although not water-resistant, it is
suitable for plywood. Gram husks, and green or dried stems and leaves
are used for stock feed; whole seeds may be milled directly for feed.
Leaves are said to yield an indigolike dye. Acid exudates from the
leaves can be applied medicinally or used as vinegar. In Chile, a
cooked chickpea-milk (4:1) mixture was good for feeding infants,
effectively controlling diarrhea. Chickpeas yield 21% starch suitable for
textile sizing, giving a light finish to silk, wool, and cotton cloth" (Duke,
1981).
Chemistry
Chickpea seed has 38-59% carbohydrate, 3% fiber, 4.8-5.5% oil, 3%
ash, 0.2% calcium, and 0.3% phosphorus. Digestibility of protein varies
from 76-78% and its carbohydrate from 57-60%. (Hulse, 1991,
Huisman and van der poel, 1994). Raw whole seeds contain per 100 g:
357 calories, 4.5-15.69% moisture, 14.9-24.6 g protein, 0.8-6.4 % fat,
2.1-11.7 g fiber, 2-4.8 g ash, 140-440 mg Ca, 190-382 mg P, 5.0-23, 9
mg Fe, 0-225 µ g β-carotene equivalent, 0.21-1.1 mg thiamin, 0.12-
0.33 mg riboflavin, and 1.3-2.9 mg niacin (Duke, 1981; Huisman and
van der Poel, 1994). "Boiled and roasted seeds contain similar
amounts. Sprouting is said to increase the proportionate amounts of
ascorbic acid, niacin, available iron, choline, tocopherol, pantothenic
acid, biotin, pyridoxine, inositol, and vitamin K. Malic and oxalic acid
exudation from the leaves of the plant may soil and damage trousers
and shoes. Wild species often have similar glandular secretions" (Duke,
1981). The limiting amino acid concentrations are 0.52 for methionine,
1.45 for lysine and cystine, 0.71 for threonine and 0.16 for tryptophan
(Williams et al., 1994). The amino acid composition of seeds with
19.5% protein, 5.5% oil is (per 16 g N): 7.2 g lysine, 1.4 g methionine,
8.8 g arginine, 4.0 g glycine, 2.3 g histidine, 4.4 g isoleucine, 7.6 g
leucine, 6.6 g phenylalanine, 3.3 g tyrosine, 3.5 g threonine, 4.6 g
valine, 4.1 g alanine, 11.7 g aspartic acid, 16.0 g glutamic acid, 0.0 g
hydroxyproline, 4.3 g proline, and 5.2 g serine" (Duke, 1981; Huisman
and van der poel, 1994; and Williams et al., 1994). "Percent fatty acid
compositions are: 'Desi': oleic 52.1, linoleic 38.0, myristic 2.74, pactic
5.11, and steatic 2.05; 'Kabuli': oleic 50.3, linoleic 40.0, myristic 2.28,
palmitic 5.74, stearic 1.61, and arachidic 0.07%. The leaves contain 4-
8% protein" (Duke, 1981).
Origin
van der Maesen (1972) believed that the species originated in the
southern Caucasus and northern Persia. However, Ladizinsky, (1975)
reported the center of origin to be southeastern Turkey. van der
Maesen (1987) recognized the southeastern part of Turkey adjoining
Syria as the possible center of origin of chickpea based on the
presence of the closely related annual species, C. reticulatum
Ladizinsky and C. echinospermum P.H. Davis. Wild C. reticulatum is
interfertile with the cultivated pulse and morphologically closely
resembles cultivated C. arietinum. It is regarded as the wild progenitor
of chickpea (Ladizinsky, 1975). "Botanical and archeological evidence
show that chickpeas were first domesticated in the Middle East and
were widely cultivated in India, Mediterranean area, the Middle East,
and Ethiopia since antiquity. Brought to the New World, it is now
important in Mexico, Argentina, Chile, Peru and the U.S. Also important
in Australia. Wild species are most abundant in Turkey, Iran,
Afghanistan, and Central Asia" (Duke, 1981).
Botany
Taxonomy, Morphology and Floral Biology
Cicer, which was classified under Vicieae Alef., was later reported to
belong to the monogeneric tribe, Cicereae (Kupitcha, 1977). The Genus
includes 9 annuals and 34 perennial herbs (van der Maesen, 1972; and
Muehlbauer, 1993). Crossability and fertility of hybrids in interspecific
crosses have been used as a basis to classify the annuals into 4
crossability groups. The first group includes the cultivated chickpea
(Cicer arietinum L.) (Ladizinsky et al., 1988) and C. reticulatum.
Chickpea plants can be described as "stems are branched, erect or
spreading, sometimes shrubby much branched, 0.2-1 m tall, glandular
pubescent, olive, dark green or bluish green in color. Root system is
robust, up to 2 m deep, but major portion up to 60 cm. Leaves
imparipinnate, glandular-pubescent with 3-8 pairs of leaflets and a top
leaflet (rachis ending in a leaflet); leaflets ovate to elliptic, 0.6-2.0 cm
long, 0.3-1.4 cm wide; margin serrate, apex acuminate to aristate,
base cuneate; stipules 2-5 toothed, stipules absent. Flowers solitary,
sometimes 2 per inflorescence, axillary; peduncles 0.6-3 cm long,
pedicels 0.5-1.3 cm long, bracts triangular or tripartite; calyx 7-10 mm
long; corolla white, pink, purplish (fading to blue), or blue, 0.8-1.2 cm
long. The staminal column is diadelphous (9-1) and the ovary is sessile,
inflated and pubescent" (Duke, 1981; Cubero, 1987; van der Maesen,
1987). Pod rhomboid ellipsoid, 1-2 with three seeds as a maximum,
and inflated, glandular-pubescent. Seed color cream, yellow, brown,
black, or green, rounded to angular, seedcoat smooth or wrinkled, or
tuberculate, laterally compressed with a median groove around two-
thirds of the seed, anterior beaked; germination cryptocotylar (Duke,
1981; Cubero, 1987 van der Maesen, 1987).
Ecology
Chickpea is a self-pollinated crop. Cross-pollination is rare; only 0-1 %
is reported (Singh, 1987; Smithson et al., 1985). Grown usually as a
rainfed cool-weather crop or as a dry climate crop in semi-arid regions.
Optimum conditions include 18-26°C day and 21-29°C night
temperatures and annual rainfall of 600-1000 mm (Duke, 1981;
Muehlbauer et al., 1982; Smithson et al., 1985). The Palouse region of
the states Washington and Idaho, appears to be well suited to chickpea
and can be characterized as having 18-25°C during the day and 5-10°C
during the night and a sufficiently long growing season (Muehlbauer et
al., 1982). California is very suited to the chickpea crop and it has
thrived in the coastal areas and in the Central Valley. Thrives on a
sunny site in a cool, dry climate on well-drained soils and grows on a
residual moisture in the post-rainy seasons of sub tropical winter or
spring of the northern hemisphere (Smithson et al., 1985). "Generally
grown on heavy black or red soils pH 5.5-8.6. Frost, hailstones, and
excessive rains damage the crop. Though sensitive to cold, some
cultivars can tolerate temperatures as low as -9.5°C in early stages or
under snow cover. Daily temperature fluctuations are desired with cold
nights with dewfall. Relative humidity of 21-41% is optimum for seed
setting. In virgin sandy soils or for the first planting in heavier soils,
inoculation is said to increase yield by 10-62%" (Duke, 1981). Although
spoken of as "day-neutral," chickpea is a quantitative long-day plant,
but flowers in every photoperiod (Smithson et al., 1985).
Crop Culture
Field Cultivation
Harvesting
Biotic Factors
The main fungi that affect chickpea are Fusarium oxysporum
Schlechtend.:Fr. f. sp. ciceris (Padwick) Matuo & K. Sato, causing the
plant to wilt and Ascochyta blight caused by Ascochyta rabiei.
Ascochyta blight is the most serious disease in North India, Pakistan,
the U.S. and the Middle East (sometimes causing l00% losses)
(Smithson et al., 1985). Blight causes brown spots on leaves, stems,
pods and seeds (Kaiser, 1992). Other fungi known to attack chickpea
include leaf spot (Alternaria sp.), Ascochyta pisi, rust (Uromyces
ciceris-arientini), gray mould (Botrytis cinera), powdery mildew
(Leviellula taurica), Pythium debar-yanum, P. ultimum, dry root rot
(Rhizoctonia bataticola), R. solani, foot rot (Sclerotium rolfsii),
Sclerotinia sclerotiorum, wilt (Verticillium albo-atrum). Some of these
fungi may become of economic importance. Viruses isolated from
chickpea include alfalfa mosaic, pea enation mosaic, pea leaf roll, pea
streak, bean yellow mosaic, and cucumber mosaic (Duke, 1981; Kaiser,
1988; Smithson et al., 1985; van Emden et al., 1994). Pod borer
(Helicoverpa armigera), the most important pest, and feeds on leaves
and developing seeds (Smithson et al., 1985). Cutworms (Agrotis sp.),
lesser armyworms (Spodoptera exigua) and leaf minor. Groundnut
aphid (Aphis craccivora), pea aphid (Acyrthsosiphon pisum), cowpea
bean seed beetle (Callosobruchus maculatus), and Adzuki bean seed
beetle (C. chinensis) are also important. "Many storage insects
specifically Bruchid sp. are a serious pest of stored chickpea.
Chickpeas stored as dhal harbor fewer bruchids. Callosobruchus
chinensis lowers seed viability. For control of bruchids, dusting with
BHC, DDT, derris, lindane, or pyrethrum or fumigation with methyl
bromide, have been recommended" (Duke, 1981). In general,
estimates of yield losses by individual pests, diseases or weeds range
from 5-10 % in temperate regions and 50-100 % in tropical regions
(van Emden, 1988).
Among the abiotic factors, drought stands to be the number one
problem in major chickpea growing regions because the crop is grown
on residual moisture and the crop is eventually exposed to terminal
drought (Johansen et al., 1994). In west Asia and North African
countries, low temperature causing freezing injury or death or delayed
onset of podding reduces yield tremendously (Singh, 1987). Heat and
salinity problems are relatively important following drought and cold
stresses (Singh et al., 1994).
Germplasm
Chickpea germplasm is maintained at two International centers
(ICRISAT in India and ICARDA in Syria) and at National centers including
the Vavilov institute in Russia, the USDA-ARS Regional Plant
Introduction Station at Pullman in the U.S. and other gene banks.
Tremendous variation for economically important traits has been
documented and improved cultivars have been developed and
released. Variation for Flower and seed color and size, growth duration,
yield, and biomass, disease resistance, quality traits (cooking time,
amino acid content, flatulence and digestibility) are recorded. 'Kabuli'
type chickpeas (Mediterranean and Middle Eastern origin) generally
have the largest seeds, and grow well under irrigation. Desi chickpeas
(Indian distribution) have smaller seeds, and yield better in Indian
subcontinent, Ethiopia and often elsewhere. Hybrids between Kabuli
and Desi have produced strains with medium-size seeds and fair yields.
The bulk of chickpeas grown in developing countries are from
unselected land races. Germplasm with resistance to major diseases
has been identified and genes for important diseases have been
named (Muehlbauer and Singh, 1987).
References
• Cubero, J.I. 1987. Morphology of chickpea. p. 35-66. In: M.C. Saxena and K.B.
Singh (eds.), The Chickpea. CAB. International, Wallingford, Oxon, OX10 8DE,
UK.
• Duke, J.A. 1981. Handbook of legumes of world economic importance. Plenum
Press, New York. p. 52-57.
• Food and Agriculture Organization of the United Nations. 1976 Production
Yearbook. Rome, Italy.
• Food and Agriculture Organization of the United Nations. 1994. Production
Yearbook. Rome, Italy.
• Geervani, P. 1991. Utilization of chickpea in India and scope for novel and
alternative uses. p. 47-54. In: Uses of Tropical Grain Legumes: Proceedings of
Consultants' Meeting, 27-30 March, 1989. ICRISAT Center, Patancheru, Andhra
Pradesh, India.
• Hawtin, G.C., K.B. Singh and M.C. Saxena. 1980. Some recent development in
the understanding and improvement of Cicer and Lens. p. 613-623. In: R.J.
Summerfield and A.H. Bunting (eds.), Advances in Legume Science. Proceedings
of the International Legume Conference, Kew, 31 July-4 August 1978, held under
the auspices of the Royal Botanic Garden, Kew, the Missouri Botanical Garden,
and the University of Reading, UK.
• Huisman, J. and A.F.B. van der Poel. 1994.Aspects of the nutritional quality and
use of cool season food legumes in animal feed. p. 53-76. In: F.J. Muehlbauer and
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Legumes. Kluwer Academic Publishers. Dordrecht, The Netherlands.
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In: Uses of tropical Legumes: Proceedings of a Consultants' Meeting, 27-30
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Malik, A. Ahad Miah and S.N. Silim. Biotic and abiotic stresses constraining
productivity of cool season food legumes in Asia, Africa and Oceania. p. 175-194.
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