Exploitation of wild annual Cicer species for widening the gene pool of chickpea cultivars

Introgression of unadapted genes from the wild Cicer species could contribute to the widening of genetic base of important traits such as yield, yield attributes and resistance to major biotic and abiotic stresses. An attempt was made successfully to intercross two wild annual Cicer species with three cultivated chickpea cultivars. Four interspecific cross‐combinations were made, and their true hybridity was ascertained through morphological and molecular markers. These cross‐combinations were also studied for some important quantitative traits under real field conditions. The range, mean and coefficient of variation of agro‐morphological traits were assessed in the parental lines, their F₁ and F₂ generations to determine the extent of variability generated in cultivated chickpea varieties. A high level of heterosis was recorded for number of pods/plant and seed yield/plant in F₁ generation. Three cross‐combinations of ‘Pusa 1103’ × ILWC 46, ‘Pusa 256’ × ILWC 46 and ‘Pusa 256’ × ILWC 239 exhibited substantially higher variability for important yield‐related traits. The present research findings indicate that these wild annual Cicer species can be easily exploited to broaden the genetic base of cultivated gene pool for improving seed yield as well as adaptation.     

Genetic diversity estimates in Cicer using AFLP analysis

Amplified fragment length polymorphism (AFLP) analysis was used to evaluate the genetic variation among cultivated chickpea and wild Cicer relatives. In total, 214 marker loci were assessed, of which 211 were polymorphic (98.6%) across the 95 accessions that represented 17 species of Cicer. The genetic variation within a species was highest in C. pinnatifidum followed by C. reticulatum and lowest in C. macracanthum. Three main species groups were identified by UPGMA clustering using Nei's pair‐wise distance calculations. Group I included the cultivated species C. arietinum, C. reticulatum and C. echinospermum. Within this group, C. reticulatum accessions were clustered closest to the C. arietinum cultivars ‘Lasseter’, ‘Kaniva’ and ‘Bumper’, supporting the hypothesis that C. reticulatum is the most probable progenitor of the cultivated species. Cicer bijugum, C. judaicum and C. pinnatifidum were clustered together creating group II. Group III contained all nine perennial species assessed and two annual species C. yamashitae and C. cuneatum. The genetic distance detected between group I and group III (0.13) was equivalent to the genetic distance detected between group I and group II (the primary and annual tertiary species, respectively; 0.14). This indicated that the perennial tertiary species may be as valuable for increasing variation to incorporate novel germplasm in the cultigen as the annual tertiary species.     

Genetic Variations and Adaptations in Chickpea

A total of 329 chickpea lines, comprising 130 kabiili and 199 desi types, originating from six regions, viz., Indian sub-continent, Middle East, North Africa, America, Europe and USSR, were sown in augmented design. The data recorded on agronomic characters on all entries from each region were evaluated in terms of group (region) means. Among the six regions, greater plant height and lateness in flowering were specific to USSR entries. Group means of USSR accessions, both desi and kabuli, differed significantly from those of Indian varieties for a majority of the characters. These observations indicate that the degree of expression of certain characters could be ascribed to specific areas, which led to area-specific adaptations. Furthermore, type-specific differentiating characters, differing in degree of expression, were observed in desi and kabuli types. For example, high mean values of characters, like branches, pods, grain yield and harvest index, were associated with desi types, and 100-seed weight with kabulis. History of cultivation and selection for specific purpose has been cited as cause for area-specific adaptations and type-specific differentiating characters.     

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.     

Pollen grain germination and pollen tube growth following in vivo and in vitro self and interspecific pollinations in annual Cicer species

Summary Germination of pollen grains and growth of pollen tubes were studied to determine the cause of barreness in crosses among annual Cicer species. In vivo and in vitro time-course studies and fluorescent microscopy revealed no pollination incompatibility among the selfs, crosses and reciprocals of C. arietinum L., C. reticulatum Lad. and C. cuneatum Rich. In general, Cicer pollen grains germinated and grew on styles of Cicer species. Pollen tube growth was characterized by irregularly spaced and intermittent callose deposits. Failure of seed formation in interspecific pollinations may be attributed to the slowness of pollen tube growth or collapse of fertilized ovules. In addition to these causes, shortness of stamens and sparsity of pollen grains were responsible for flower drop in natural selfs. Although the number of pollen tubes entering the micropyle in interspecific pollinations was low, it may be possible to grow the fertilized ovules on an artificial medium to obtain F₁ plants.     

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.     

Low Temperature Effects during Seed Filling on Chickpea Genotypes (Cicer arietinum L.): Probing Mechanisms affecting Seed Reserves and Yield

Chickpea is sensitive to cold conditions (<15 °C), particularly at its reproductive phase and consequently it experiences significant decrease in the seed yield. The information about the effects of cold stress on chickpea during the seed filling phase is lacking. Moreover, the underlying metabolic reasons associated with the low temperature injury are largely unknown in the crop. Hence, the present study was undertaken with the objectives: (i) to find out the possible mechanisms leading to low temperature damage during the seed filling and (ii) to investigate the relative response of the microcarpa (Desi) and the macrocarpa (Kabuli) chickpea types along with elucidation of the possible mechanisms governing the differential cold sensitivity at this stage. At the time of initiation of the seed filling (pod size ∼1 cm), a set of plants growing under warm conditions of the glasshouse (temperature: 17/28 ± 2 °C as average night and day temperature) was subjected to cold conditions of the field (2.3/11.7 ± 2 °C as average night and day temperature), while another set was maintained under warm conditions (control). The chilling conditions resulted in the increase in electrolyte leakage, the loss of chlorophyll, the decrease in sucrose content and the reduction in water status in leaves, which occurred to a greater extent in the macrocarpa type than in the microcarpa type. The total plant weight decreased to the same level in both the chickpea types, whereas the rate and duration of the seed filling, seed size, seed weight, pods per plant and harvest index decreased greatly in the macrocarpa type. The stressed seeds of both the chickpea types experienced marked reduction in the accumulation of starch, proteins, fats, crude fibre, protein fractions (albumins, globulins, prolamins and glutelins) with a larger decrease in the macrocarpa type. The accumulation of sucrose and the activity levels of the enzymes like starch synthase, sucrose synthase and invertase decreased significantly in the seeds because of the chilling, indicating impairment in sucrose import. Minerals such as calcium, phosphorous and iron as well as several amino acids (phenylalanine, tyrosine, threonine, tryptophan, valine and histidine) were lowered significantly in the stressed seeds. These components were limited to a higher extent in the macrocarpa type indicating higher cold sensitivity of this type.     

Distribution of qualitative traits in the world germplasm of chickpea (Cicer arietinum L.)

Summary The chick pea germplasm collection maintained at ICRISAT Center, Patancheru, India, is the largest collection of this crop available in one place. This collection was grown in instalments and described for qualitative and agronomical traits. The importance and distribution of six qualitative traits, namely flower colour, plant colour, growth habit, seed shape, seed surface and seed colour have been discussed.Approved as J. A. No. 365 by the International Crops Research Institute for the Semi-Arid Tropics (ICRI-SAT).     

Cross compatibility between chickpea and its wild relative,Cicer echinospermum Davis

Summary Cicer echinospermum, a wild relative of chickpea (Cicer arietinum L.), has traits that can be used to improve the cultivated species. It is possible to obtain successful crosses between the two species, even though their cross progenies have reduced fertility. The reasons for this low fertility could be due to the two species differing in small chromosome segments or at genic level. Another limitation to the use ofC. echinospermum at ICRISAT Asia Center is that the species is not adapted to the short photoperiod which prevails during the chickpea cropping season at Patancheru, Andhra Pradesh, India. Future work will include screening the segregating progenies for monitoring traits from both the species through isozyme analysis and to incorporate these into good agronomic backgrounds following backcrosses.Submitted as JA 1669 by ICRISAT     

Inheritance of Dry Root Rot (Rhizoctonia bataticola) Resistance in Chickpea (Cicer arietinum)

The inheritance of resistance to dry root rot of chickpea caused by Rhizoctonia bataticola was studied. Parental F₁ and F₂ populations of two resistant and two susceptible parents, along with 49 F₁ progenies of one of the resistant × susceptible crosses were rested for their reaction to dry root rot using the blotting-paper technique. All F, plants of the resistant × susceptible crosses were resistant; the F₂ generation fitted a 3 resistant: 1 susceptible ratio indicating monogenic inheritance, with resistance dominant over susceptibility. F³ family segregation data confirmed the results. No segregation occurred among the progeny of resistant × resistant and susceptible × susceptible crosses.     

Genetic resources of salt tolerance in wild Hordeum species

Salt tolerance was evaluated in 340 accessions of Hordeum, consisting of 41 brittle-rachis forms of Hordeum vulgare L. subsp. vulgare (H. agriocrithon) accessions, 154 H. vulgare L. subsp. spontaneum (H. spontaneum) accessions, and 145 accessions of ten other species or subspecies of wild Hordeum. Germination was carried out at concentrations of 171, 257, and 342 mM NaCl. The levels of salt tolerance for seed germination in wild Hordeum species were generally lower than those found by Mano et al. (1996) in cultivated barley; the NaCl tolerance level of the different species were as follows: H. agriocrithon > H. spontaneum > other wild Hordeum species. In addition, leaf injury index was used to assess tolerance at the seedling stage after treatment with 500 mM NaCl solution for four weeks. The levels of salt tolerance at the seedling stage in wild Hordeum species were generally higher than those found by Mano & Takeda (1995) in cultivated barley. Most wild Hordeum species showed high NaCl tolerance at the seedling stage and are considered good sources of germplasm for salt tolerance breeding. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic diversity and phylogenetic relationships among the annual Cicer species as revealed by isozyme polymorphism

Summary There are few estimates of genetic variability within and among populations of the nine annual Cicer species and for the wild species this information is based on few accessions. The present study was undertaken to examine genetic variation within and between annual Cicer species. One hundred and thirty-nine accessions of nine annual Cicer species were used for electrophoretic analysis at ICARDA. High levels of polymorphism in all eight wild annual Cicer species was found. This is in contrast to earlier research which had shown high polymorphism only in C. reticulatum. Cicer reticulatum had the highest proportion of polymorphic loci. However, for the cultigen, among 14 loci assayed, only two were polymorphic, ADH and PGD2. The nine species formed four phylogenetic groups based on the neighbor-joining method. The first group comprised C. arietinum, C. Reticulatum and C. echinospermum, the second C. bijugum, C. judaicum and C. pinnatifidum, the third C. chorassanicum and C. yamashitae; and the fourth group consisted of one species, C. cuneatum. The phylogenetic tree developed from the neighbor-joining technique illustrated that C. reticulatum is the probable progenitor of C. arietinum and that C. echinospermum split off from a common ancestor at an earlier stage in the evolutionary history of Cicer. Genetic diversity data showed that the greatest diversity was within C. reticulatum and the lowest with the cultigen, C. arientinum. With the exception of C. reticulatum, genetic diversity increased with genetic distance from the cultigen. Little geographic variation in genetic diversity was found.     

Response of accessions within tomato wild species,Solanum pimpinellifolium to late blight

Late blight (LB), caused by the oomycete Phytophthora infestans, is one of the most devastating diseases of the cultivated tomato (Solanum lycopersicum) worldwide. Most commercial cultivars of tomato are susceptible to LB. Previously, three major LB resistance genes (Ph‐1, Ph‐2, Ph‐3) were identified and incorporated into a few commercial cultivars of tomato. Reduced effectiveness and potential breakdown of the resistance genes has necessitated identification, characterization and utilization of new sources of resistance. We evaluated the response of 67 accessions of the wild tomato species, S. pimpinellifolium to LB, under multiple field and greenhouse (GH) conditions and compared them with six control genotypes. Sixteen accessions were identified with strong LB resistance in both field and GH experiments. However, 12 accessions exhibited resistance similar to a control line which was homozygous for Ph‐2 + Ph‐3. Genotyping accessions with molecular markers for Ph‐2 and Ph‐3 were not conclusive, indicating that resistance in these accessions could be due to these or other resistance genes. Strong correlations were observed between field and GH disease response and between foliar and stem infection.     

Two genes and linked RAPD markers involved in resistance to Fusarium oxysporum f. sp. Ciceris race 0 in chickpea

The inheritance of resistance to fusarium wilt race 0 of chickpea and linked random amplified polymorphic DNA (RAPD) markers were studied in two F₆:₇ recombinant inbred line (RIL) populations. These RILs were developed from the crosses CA2156 × JG62 (susceptible × resistant) and CA2139 × JG62 (resistant × resistant), and were sown in a field infected with fusarium wilt race 0 in Beja (Tunisia) over 2 years. A1:1 resistant to susceptible ratio was found in the RIL population from the CA2156 × JG62 cross, indicating that a single gene with two alleles controlled resistance. In the second RIL population (CA2139 × JG62) a 3:1 resistant to susceptible ratio indicated that two genes were present and that either gene was sufficient to confer resistance. Linkage analysis showed a RAPD marker, OPJ20600, linked to resistance in both RIL populations, which is present in the resistant parent JG62.     

Screening of Kabuli chickpea germplasm for resistance to Fusarium wilt

A total of 1915 Kabuli chickpea lines were screened in a wilt sick plot containing Fusarium oxysporum f.sp. ciceri race 0 at Béja, Tunisia. Complete resistance was found in 110 lines and this result was confirmed by a laboratory screening method. Principal components analysis showed that > 80% of the variation of the resistant lines was explained by hundred seed weight and days to maturity. Cluster analysis divided the resistant lines into four groups: 21 had high seed weight (48.25 ± 3.81 g) and early maturity (95.09 ± 2.50 d), 24 had high seed weight (46.84 ± 2.10 g) and late maturity (117.00 d), 34 had low seed weight (22.35 ± 4.72 g) and early maturity (92.97 ± 3.97 d) and 31 had low seed weight (19.62 ± 5.37 g) and late maturity (112.09 ± 4.51 d). This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of seed weight and growth habit in 10 intercross chickpea (Cicer arietinum) nested association mapping populations

Objective of investigation

Chickpea is a major global food legume for which seed weight and plant growth habit are important yield and harvestability components for plant breeding. This study tested seed weight and plant growth habit inheritance and identified quantitative trait loci (QTL).

Experimental material

A 10 nested association mapping (NAM) populations of chickpea were created from crosses between ‘Gokce’, a cultivar and wild crop relative accessions of Cicer reticulatum and Cicer echinospermum. Families were then developed to the F2:4 generation.

Method of investigation

A 10 families were grown at the Field Experiment Station, Harran University near Şanlıurfa, Turkey during 2019.

Data collection

A 100-seed weight and prostrate or erect growth habit was scored in the field. Two families were genotyped for 60 single-nucleotide polymorphisms (SNP).

Result and conclusions

A 100-seed weight showed polygenic control, and three QTLs were found. Growth habit was controlled by one or two QTLs. The two traits were significantly correlated for five populations. The crop wild relatives of chickpea contain variations at novel loci affecting seed weight compared to the literature.     

Investigations into the Barrier(s) to Interspecific Hybridization between Cicer arietinum L. and eight other Annual Cicer species

Barrier(s) to interspecific hybridization between the cultivated chickpea, Cicer arietinum L., and eight other annual wild species, i.e. C. reticulatum Lad., C. echinospermum Dav., C. pinnatifidum J. and S., C. judaicum Boiss., C. bijugum Rech., C. chorassanicum (Bge) M. Pop., C. yamashitae Kit. and C. cuneatum Rich., were investigated. In general, good pollen germination and pollen tube growth were observed in all eight crosses and their reciprocals. En spite of a few pollen cube growth abnormalities in most crosses, pollen tube penetration into the ovule and, thus, fertilization was observed in all cross combinations. However, differences were observed in the time from pollination to fertilization, not only between different interspecific crosses but also between reciprocals of a particular interspecific cross. The crossability barrier is, therefore, believed to be due to factor(s) operating after fertilization.     

Introgression from wild Cicer reticulatum to cultivated chickpea for productivity and disease resistance

Interspecific hybridization is known to improve productivity and resistance to diseases in many crops. Therefore, an attempt was made to introgress productivity and disease resistance into chickpea from wild Cicer species. The true F₁ hybrids of cultivated chickpea genotypes ‘L550’ and ‘FGK45’ with C. reticulatum were backcrossed twice to their cultivated female parents to minimize the linkage drag of undesirable wild traits. The pedigree method was followed to advance the segregating populations from straight crosses (without backcross) and BC₁/BC₂ generations to F₅–F₇. The interspecific derivatives recorded up to a 16.9% increase over the check cultivars and a 25.2% increase over the female parent in a preliminary yield evaluation trial. Of the 22 interspecific derivatives thus derived, four desi and two kabuli lines were further evaluated for seed yield in replicated trials at three diverse locations. These lines possess a high degree of resistance to wilt, foot rot and root rot diseases, and recorded a 6.1–17.0% seed yield increase over the best check cultivars.