Cold Stress Injury during the Pod-Filling Phase in Chickpea (Cicer arietinum L.): Effects on Quantitative and Qualitative Components of Seeds

Chilling stress (<10 °C) is detrimental for chickpea, especially at the reproductive phase and leads to abortion of flowers, pods and impaired seed filling, causing severe reduction in yield. The information on the effects of low temperature during different pod-filling stages on quality and quantity of developing seeds is lacking in chickpea and hence this study. In this study, chickpea plants growing under warm conditions of the glasshouse were subjected to cold conditions of the field at the two stages, (a) early pod-filling and (b) late pod-filling, and subsequently analysed for stress injury in terms of electrolyte leakage (EL), 2,3,5-triphenyl tetrazolium chloride reduction, relative leaf water content and total chlorophyll content in the leaves of control and cold-stressed plants. Cold stress caused elevation of EL but reduced all the other parameters. Sucrose content decreased significantly in the leaves of cold-stressed plants. The differences between the effects of stress at two stages on the total plant dry weight were small and insignificant. The seed growth rate, seed fill duration, seed number, and average seed weight and size decreased greatly in the plants cold-stressed at the late pod-filling stage than those stressed at the early pod-filling stage. Greater reduction was observed in starch, proteins, soluble sugars, fat, crude fibre and storage protein fractions in the seeds of the plants cold-stressed at the late pod-filling stage. This coincided with a larger decrease in sucrose content, the activities of sucrose synthase, invertase and starch synthase observed at this stage. The germination and growth potential were, however, inhibited to a greater extent in seeds of plants stressed at the early pod-filling stage.     

Inheritance of Seed Coat Thickness in Chickpea (Cicer arietinum L.) and its Evolutionary Implications

The desi and kabuli chickpeas are characterized, among other things, by their seed coats being thicker in the desi than in the kabuli type. The inheritance of seed coat thickness, and its relation to flower colour and seed size, was studied. Seed coat thickness exhibits monogenic inheritance, the thin kabuli seed coat being the recessive character. Linkage was found between seed coat thickness and flower colour, the recombinant fraction being 0.19. No relationship was found between seed coat thickness and seed size. The role of these characters in the evolution of the chickpea is discussed.     

Salinity Effects on the Growth and Ion Contents of Some Chickpea (Cicer arietinum L.) and Lentil (Lens culinaris Medic.) Varieties

The effect of salinity on seed germination, plant yield parameters, and plant Na, Cl and K concentrations of chickpea and lentil varieties was studied. Results showed that in both crops percentage emergence was significantly reduced by increasing NaCl levels (0–8dSm^?1). From the plant growth studies it was found that differences existed among chickpea and lentil varieties in their response to NaCl application. In chickpea, the variety Mariye showed the comparatively lowest germination percentage and the lowest seedling shoot dry weight in response to salinity and was also among the two varieties which had the lowest relative plant height, shoot and root dry weight and grain yield at maturity. Similarly, variety DZ-10-16-2, which was the second best in germination percentage and the highest in terms of seedling shoot dry weight, also had the highest relative plant height, shoot and root dry weights, and grain yield at maturity. In lentil, however, such relationships were less pronounced. Chloride concentration (mg g^?1) in the plant parts at salt levels other than the control was about 2–5 times that of Na. K concentration in the plants was significantly reduced by increasing NaCl levels. Chickpea was generally more sensitive to NaCl salinity than lentil. While no seeds were produced at salinity levels beyond 2dSm^?1 in chickpea (no seeds were produced at this salt level in the most sensitive variety, Mariye), most lentil varieties could produce some seeds up to the highest level of NaCl application. Overall, varieties R-186 (lentil) and Mariye (chickpea) were the most sensitive of all varieties. On the other hand, lentil variety NEL-2704 and chickpea variety DZ-10-16-2 gave comparatively higher mean relative shoot and root dry weights, and grain yield, thus showing some degree of superiority over the others. The observed variations among the varieties may be useful indications for screening varieties of both crops for salt tolerance.     

Relationship Between Cold Severity and Yield Loss in Chickpea (Cicer arietinum L.)*

Seed yield in chickpea (Cicer arietinum L.) is substantially increased by advancing sowing date from the traditional spring to early winter at low to medium elevation areas around the Mediterranean Sea. This shift, however, increases the probability of the exposure to subzero temperatures as low as -10 °C for up to 60 days in a year. These low temperatures often reduce seed yield of cold-susceptible cultivars. Yield losses from cold were estimated in two experiments conducted at Tel Hadya, Syria. In experiment 1, of 96 genotypes sown on nine dates ranging from autumn to spring during the 1981–82 season, those lacking tolerance to cold were killed and produced no yield in autumn sowing, whereas lines with cold tolerance produced nearly 4 t/ha which corresponds to a four-fold increase over spring sowing. Moderately cold-tolerant genotypes sown during early winter produced substantially more seed yield than the normal spring-sown crop. Seedlings were more cold tolerant than the plants in early or late vegetative stages. In experiment 2, in which yield loss due to cold in the field was estimated in 12 yield trials comprising 288 newly bred lines in the 1989–90 season, the regression of cold susceptibility on seed yield in each of the trials was highly significant and negative. On average, winter-sown trials produced 67 % more seed yield than spring-sown trials, but 125 out of 288 genotypes produced yield more than double in winter sowing. Early maturing lines suffered severe cold damage and many lines produced no seed.     

Genotypic Variation in Root Systems of Chickpea (Cicer arietinum L.) across Environments

Root systems of various chickpea genotypes were studied over time and in diverse environments, – varying in soil bulk density, phosphorus (P) levels and moisture regimes. In a pot study comparing a range of chickpea genotypes, ICC 4958 and ICCV 94916‐4 produced higher root length density (RLD) and root dry weight (RDW), which were better expressed under P stress conditions. In two field experiments in soils of intermediate and high soil bulk densities, ICC 4958 also had greater RLD and RDW, particularly under soil moisture stress conditions. The expression of greater rooting ability of ICC 4958 under a wide range of environmental conditions confirms its suitability as a parent for genetically enhancing drought resistance and P acquisition ability. The superiority of ICC 4958 over other genotypes was for root proliferation expressed through RLD. Thus, the variation in RLD can be the most relevant root trait that reflects chickpea's potential for soil moisture or P acquisition.     

Interactive Effects of Salinity and Biological Nitrogen Fixation on Chickpea (Cicer arietinum L.) Growth

The effect of salinity on the nodulation, N-fixation and plant growth of selected chickpea- Rhizobium symbionts was studied- Eighteen chickpea rhizobial strains were evaluated for their growth in a broth culture at salinity levels of 0 to 20 dS m⁻¹ of NaCl + Na₂SO₄. Variability in response was high. Salinity generally reduced the lag phase and/or slowed the log phase of multiplication of Rhizobium. Nine chickpea genotypes were also evaluated for salt tolerance during germination and early seedling growth in Petri dishes at five salinity levels (0–32 dS m⁻¹). Chickpea genotypes ILC-205 and ILC-1919 were the most salt-tolerant genotypes. The selected rhizobial strains and chickpea cultivars were combined in a pot experiment aimed at investigating the interactive effect of salinity (3, 6 and 9 dS m⁻¹) and N source (symbiosis vs. inorganic N) on plant growth. Symbiotic plants were more sensitive to salinity than plants fed mineral N. Significant reductions in nodule dry weight (59.8 %) and N fixation (63.5 %) were evident even at the lowest salinity level of 3 dS m-¹. Although nodules were observed in inoculated plants grown at 6 dS m-¹, N-fixation was completely inhibited. The findings indicate that symbiosis is more salt-sensitive than both Rhizobium and the host plant, probably due to a breakdown in one of the processes involved in symbiotic-N fixation. Improvement of salinity tolerance in field grown chickpea may be achieved by application of sufficient amounts of mineral nitrogen.     

Effect of Weed Removal on Productivity of Chickpea (Cicer arietinum L.) and Lentil (Lens culinaris Med.) in a Mediterranean Environment

Three field experiments were conducted on chickpea ( Cicer arietinum L.) and four on lentil ( Lens culinaris Med.) at different winter-sown rainfed locations in Jordan from 1988/89 to 1990/91 to study the effect of the duration of weed-free and weed-infested conditions on yields and yield components of the crops. Chickpea seed yields were reduced on average by 81 % and straw yields by 63 % when fields remained weed infested until harvest compared with weed-free conditions throughout the growing season. The corresponding lentil seed and straw yield decreases were 63 % and 55 %. As the duration of weed-free period increased and the duration of weed-infested period decreased, yields increased. However, the critical period of weed interference was between 35 and 49 days after emergence in chickpea and between 49 and 56 days after emergence in lentil, when these crops were at an advanced stage of vegetative growth. There were significant negative correlations between the weed dry weight and the seed or straw yields. The reduction in seed yields in both crops because of weed interference occurred mainly through the reduced number of pods /plant, which in turn was partly the result of reduced number of secondary branches. In chickpea, some reduction also occurred through reduced 100-seed weight.     

Two new traits - open flower and small leaf in chickpea (Cicer arietinum L.)

Two new traits – open flower and small leaf in chickpea are discussed. Open flower, a natural mutant in a good agronomic background is reported for the first time, small leaf trait has been reported earlier, and has now been studied by breeders. Both useful traits were found to be monogenic recessive. The joint F₂ segregation data revealed no linkage between flower colour and flower type, but flower type and leaf size showed some linkage. Open flower could contribute to a higher rate of cross pollination and utilization of heterosis. The small leaf allows light to penetrate the crop canopy, and could be useful in designing a physiologically efficient plant type in chickpea. This revised version was published online in July 2006 with corrections to the Cover Date.     

Exogenous Application of Abscisic Acid Improves Cold Tolerance in Chickpea (Cicer arietinum L.)

Chickpea suffers cold stress (<10 °C) damage especially during reproductive phase resulting in the abortion of flowers and pods, poor pod set, and reduction in seed yield and seed quality. One of the ways in modifying cold tolerance involves exogenous treatment of the plants with chemicals having established role in cold tolerance. In the present study, the chickpea plants growing under optimum temperature conditions (28/12 °C, as average maximum and minimum temperature) were subjected to cold conditions of the field (10–12/2–4 °C; day/night as average maximum and minimum temperature) at the bud stage. Prior to exposure, these plants were treated exogenously with 10 μm abscisic acid (ABA) and thereafter again after 1 week of exposure. The stress injury measured in terms of increase in electrolyte leakage, decrease in 2,3,5-triphenyl tetrazolium chloride reduction %, relative leaf water content and chlorophyll content was observed to be significantly mitigated in ABA-applied plants. A greater pollen viability, pollen germination, flower retention and pod set were noticed in ABA-treated plants compared with stressed plants. The seed yield showed considerable improvement in the plants treated with ABA relative to the stressed plants that was attributed to the increase in seed weight, greater number of single seeded pods and reduction in number of infertile pods. The oxidative damage measured as thiobarbituric acid-reactive substances was lesser in ABA-treated plants that was associated with greater activities of superoxide dismutase, catalase, ascorbate peroxidase, ascorbic acid, glutathione and proline in these plants. It was concluded that cold stress effects were partly overcome by ABA treatment because of the improvement in water status of the leaves as well as the reduction in oxidative damage.     

Evaluation of Yield Criteria for Drought and Heat Resistance in Chickpea (Cicer arietinum L.)

The chickpea (Cicer arietinum L.) is usually grown under rainfed, rather than irrigated conditions, where drought accompanied by heat stress is a major growth constraint. The aim of this study was to select chickpea genotypes having resistance to drought/heat stress and to identify the most appropriate selection criteria for this. A total of 377 chickpea accessions were sown 2 months later than normal for the Antalya region (Turkey) to increase their exposure to the drought and high‐temperature conditions of a typical summer in this part of the world. Interspersed between every 10 test genotypes as benchmark genotypes, were plants of the two known genotypes ILC 3279 (drought‐susceptible) and ILC 8617 (drought‐susceptible), while ICC 4958 (known drought‐resistant) and ICCV 96029 (known very early, double‐podded) were also sown for confirmation. All plants were subsequently screened for drought and heat stress resistance. Soon after the two known susceptible genotypes had died, evaluations of the entire trial were made visually on a scale from ‘1’ (free from drought/heat damage) to ‘9’ (all plants died from drought/heat). Yield loss in many of the test genotypes and in the two known susceptible genotypes (ILC 3279 and ILC 8617) rose to 100 %. The desi chickpeas (smaller, dark seeds) were generally more drought‐ and heat‐resistant than the kabuli chickpeas (larger, pale seeds). Two desi chickpeas, ACC 316 and ACC 317, were selected for drought and heat (>40 °C) resistance under field conditions. Seed weight was the trait least affected by adverse environmental conditions and having the highest heritability, and it should be used in early breeding selections. When breeding drought‐ and heat‐resistant chickpeas, path and multivariate analyses showed that days to the first flowering and maturity to escape terminal drought and heat stresses should be evaluated ahead of many other phenological traits, and harvest index, biological yield and pods per plant for increased yield should also be considered.     

Consistent Variation Across Soil Types in Salinity Resistance of a Diverse Range of Chickpea (Cicer arietinum L.) Genotypes

Chickpea is considered sensitive to salinity, but the salinity resistance of chickpea germplasm has rarely been explored. This study aimed to (i) determine whether there is consistent genetic variation for salinity resistance in the chickpea minicore and reference collections; (ii) determine whether the range of salinity resistance is similar across two of the key soil types on which chickpea is grown; (iii) assess the strength of the relationship between the yield under saline conditions and that under non‐saline conditions; and (iv) test whether salinity resistance is related to differences in seed set under saline conditions across soils and seasons. The seed yield of 265 chickpea genotypes in 2005–2006 and 294 cultivated genotypes of the reference set in 2007–2008 were measured. This included 211 accessions of the minicore collection of chickpea germplasm from the International Crops Research Institute for the Semi‐Arid Tropics (ICRISAT). The experiments were conducted in a partly controlled environment using a Vertisol soil in 2005–2006 and an Alfisol soil in 2007–2008, with or without 80 mm sodium chloride (NaCl) added prior to planting. In a separate experiment in 2006–2007, 108 genotypes (common across 2005–2006 and 2007–2008 evaluations) were grown under saline (80 mm NaCl) and non‐saline conditions in a Vertisol and an Alfisol soil. In 2005–2006 in the Vertisol and 2007–2008 in the Alfisol, salinity delayed flowering and maturity, and reduced both shoot biomass and seed yield at maturity. There was a large variation in seed yield among the genotypes in the saline pots, and a small genotype by environment interaction for grain yield in both soil types. The non‐saline control yields explained only 12–15 % of the variation of the saline yields indicating that evaluation for salinity resistance needs to be conducted under saline conditions. The reduction in yield in the saline soil compared with the non‐saline soil was more severe in the Alfisol than in the Vertisol, but rank order was similar in both soil types with a few exceptions. Yield reductions due to salinity were closely associated with fewer pods and seeds per pot (61–91 %) and to lesser extent from less plant biomass (12–27 %), but not seed size. Groups of consistently salinity resistant genotypes and the ones specifically resistant in Vertisols were identified for use as donor sources for crossing with existing chickpea cultivars.     

Effect of Sulphur and Foliar-applied Chemicals on Cold Tolerance in Chickpea (Cicer arietinum L.)

In a field study it was observed that sulphur fertilization of chickpea at 100 kg S ha⁻¹ imparted cold tolerance under low temperature stress conditions. Further, foliar sprays of DMSO, H₂SO⁴, KCl and H³BO³ proved effective in alleviating cold injury. Glucose spray also showed efficacy in this regard. The effects of sulphur fertilization and foliar applied DMSO and H²SO⁴ were largely associated with improved sulphur nutrition of plants, while improvement in K content under KCl treatment and B content under H₃BO₃ treatment was responsible for cold tolerance effects. Improvement in overall soluble carbohydrate and protein status of plants was held responsible for glucose effects possibly associated with osmoregulation.     

Pod volume and pod-filling percentage as additional traits for the characterization of chickpea (Cicer arietinum L.)

Thirty short- to medium-duration chickpea germplasm accessions from diverse geographic origins and with a wide range of physiological and morphological traits were grown in three environments at ICRISAT Asia Center, Patancheru, during 1992/93. Data were recorded on time to flowering, leaf area, 100-seed mass, pod volume, and pod-filling percentage. Quantitative data on the last two traits were recorded for the first time to examine their relevance to the characterization of germplasm accessions and their use as selection criteria in breeding. The accessions exhibited considerable variation for the traits. The broad-sense heritabilities were 0.98 for pod volume and 0.85 for pod-filling percentage. The two traits showed consistent relationships with other morphological characters indicating that the pod volume and pod-filling percentage traits can be utilized in genotype characterization of chickpea.     

Induction and inheritance of determinate growth habit in chickpea (Cicer arietinum L.)

Summary The character of determinate plant growth has not been reported for chickpea and has not been observed in the world germplasm collection at ICRISAT, Patancheru, India. A determinate growth habit would be desirable where growing conditions often lead to excessive vegetative growth. We attempted to generate this trait by mutation breeding. Seeds of the cultivar ICCV 6 were exposed to varying irradiation treatments, M₁ and M₂ populations were raised, and in the latter one plant was detected that showed the determinate growth habit and female sterility. The character of determinate growth segregated in a postulated digenic epistatic 3:13 fashion in the F₂ and confirmed its digenic mode of inheritance in the F₃ and F₄. The symbol cd is proposed for the allele conditioning for determinancy and Dt for the allele expressing the determinate trait. Continued mutation breeding with this and other material may result in identifying fully fertile, determinate plant types.Abbreviations DT - determinate - IDT - indeterminate ICRISAT Journal Article No. 1396.     

Leaf types and their genetics in chickpea (Cicer arietinum L.)

Summary Commonly the chickpea leaf is uni-imparipinnate, having 9–15 leaflets. However, certain variants have been reported; these are available in the chickpea collection at ICRISAT and were re-examined. Based on the lamina differentiation, three major classes of leaf type can be recognized: uni-imparipinnate (normal), multipinnate and simple (leaf). (Certain other leaf forms reported earlier are not classes of leaf type though they are distinct variants). It was determined that the leaf type differences are governed by two genes (mlsl), which show supplementary gene action. The multipinnate leaf is formed when the first gene is dominant (ml⁺sl/.sl). Whereas the simple leaf occurs when the first gene is recessive and the second gene is in either form (ml./ml.), the normal leaf is expressed when both dominant genes are present (ml⁺sl⁺/..).Submitted as J.A. No. 814 by the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT).     

Alterations in Photochemical and Physiological Activities of Chickpea (Cicer arietinum L.) Cultivars under Drought Stress

In this study, some morphological, physiological and biochemical parameters of two chickpea cultivars, cv. Gökçe and Canıtez, were analysed to understand their tolerance to drought stress. Twenty-day-old plants were subjected to three different regimes of drought stress by withholding water for 3, 5 or 7 days, and then rewatering for 2 days after the initial 7 days of drought stress. Drought treatments only reduced shoot elongation in the Canıtez cultivar. Leaf production and fresh biomass decreased in both cultivars under all drought treatments, however to a greater extent in Canıtez. In both cultivars, malondialdehyde, proline and anthocyanin accumulation increased significantly, whereas relative water content declined under drought stress. The total chlorophyll and carotenoid contents of Gökçe were not affected by drought stress, whereas the chlorophyll content of Canıtez increased greatly at the end of the treatments. Using chlorophyll a fluorescence measurements, we found that extended drought treatment caused photoinhibition of PSII activity in both cultivars. However, this was greater in Canıtez, especially under severe drought stress. Although Canıtez recovered quickly from drought stress and exhibited a good ability to overcome drought stress, via activation of many protection mechanisms such as increasing antioxidant enzymes and proline and anthocyanin accumulation during vegetative stage, our results show that Canıtez is less drought tolerant than Gökçe.     

The Effects of Temperature and Soil Moisture on Chickpea (Cicer arietinum L.) Genotype Sensitivity to Isoxaflutole

Isoxaflutole at 75 g ai ha^?1 is registered in Australia for the control of several broadleaf weeds in chickpea (Cicer arietinum L.). Although isoxaflutole provides satisfactory control of problematic weeds, under certain conditions crop injury can occur. Higher air temperature and moisture content of soil are reported to affect the metabolism of soil applied herbicide. Controlled environment experiments were used to determine the tolerance of chickpea to isoxaflutole under a range of temperature and soil moisture levels. For the soil moisture study, the variables examined were two desi chickpea genotypes (Kyabra as a tolerant cultivar and Yorker as a sensitive cultivar), three soil moisture levels [50 % field capacity (FC), 75 % FC and FC] with three herbicide rates [0, 75 (recommended rate) and 300 g ai ha^?1]. For the temperature by soil moisture study, the variables examined were two other desi chickpea genotypes (97039‐1275 as a tolerant line and 91025‐3021 as a sensitive line), three temperature regimes (20/5, 30/15 and 35/25 °C), two soil moisture conditions (50 % FC and FC) with the same three herbicide rates. The results demonstrated that the chickpea genotypes exhibited differential tolerance to isoxaflutole, but that differences in response were affected by rate, temperature and soil moisture. Increasing temperature and soil moisture content made the susceptible chickpea genotype more vulnerable to isoxaflutole damage. Injury to the susceptible genotype in terms of increased leaf chlorosis and reduction in shoot height and dry matter production increased as soil moisture increased from 50 % FC to FC and temperature increased from 20/5 to 35/25 °C. Overall damage of the sensitive genotype from increasing rates of isoxaflutole also increased when soil moisture content increased from 50 % FC to FC within the fixed temperature regime of 30/15 °C. The sensitivity of chickpea to isoxaflutole depends on existing temperature and moisture content and the chances of crop damage were enhanced with increasing temperature and moisture levels.     

Cultivar identification and genetic relationship among selected breeding lines and cultivars in chickpea (Cicer arietinum L.)

Twenty two RAPD and 22 ISSR markers were evaluated for their potential use in determination of genetic relationships in chickpea (Cicer arietinum L.) cultivars and breeding lines. We were able to identify six chickpea cultivars/breeding lines by cultivar-specific markers. All of the cultivars tested displayed a different phenotype generated either by the RAPD or ISSR primers. Though ISSR primers generated less markers than RAPD primers, the ISSR primers produced higher levels of polymorphism (% of polymorphic markers per primer) than RAPD primers. A high level of within cultivar homogeneity was observed in chickpea. Cultivars/breeding lines originating from a common genetic background showed closer genetic relationship. Chickpea lines with similar seed type(kabuli or desi) had a tendency to cluster together. This revised version was published online in August 2006 with corrections to the Cover Date.