Genetics of resistance to ascochyta blight in chickpea (Cicer arietinum L.)

Summary Six chickpea lines resistant to Ascochyta rabiei (Pass.) Lab. were crossed to four susceptible cultivars. The hybrids were resistant in all the crosses except the crosses where resistant line BRG 8 was involved. Segregation pattern for diseases reaction in F₂, BCP₁, BCP₂ and F₃ generations in field and glasshouse conditions revealed that resistance to Ascochyta blight is under the control of a single dominant gene in EC 26446, PG 82-1, P 919, P 1252-1 and NEC 2451 while a recessive gene is responsible in BRG 8. Allelic tests indicated the presence of three independently segregating genes for resistance; one dominant gene in P 1215-1 and one in EC 26446 and PG 82-1, and a recessive one in BRG 8.Research paper No. 3600     

Resistance to Ascochyta blight in chickpea - Genetic basis

Summary Genetic regulation of host resistance in chickpea-Ascochyta rabiei interaction system is governed by two dominant complementary genes each in the genotypes GLG 84038 and GL 84099, whereas the resistance in a black seeded genotype ICC 1468 was controlled by one dominant and one recessive independent gene. In all the genotypes, resistance is operated by inter-allelic interactions. The genes conferring resistance in GLG 84038 were found to be different to those operating in GL 84099 and ICC 1468. Among the five dominant genes dispersed in 3 genotypes under study, at least one has been reported for the first time, as to date, only three dominant genes have been reported in the literature.The four identified dominant genes in GLG 84038 and GL 84099 have been named as Arc₁, Arc₂ (in GLG 84038) and Arc₃, Arc₄ (in GL 84099). The undistinguished dominant gene in ICC 1468 has been named as Arc_5(3,4) as it could not be equated or differentiated from Arc₃ or Arc₄. The recessive gene in ICC 1468 has been named as Arc₁.Generation mean analysis of the 6 resistant × susceptible crosses involving the same genotypes, revealed that the genes conferring resistance in any of the 3 genotypes did not follow simple Mendelian inheritance but were influenced by inter allelic interactions. Additive gene effect along with dominance were operative in all the 3 genotypes under study in conferring resistance. However, the mechanism of resistance in GLG 84038 and GL 84099 were primarily additive in nature while that in ICC 1468, dominance as well as dominance × dominance interactions were more important than additive gene action.     

Leaf type is not associated with ascochyta blight disease in chickpea (Cicer arietinum L.)

The three major leaf types in chickpea are normal compound leaf, simple leaf and multipinnate. Simple leaf types are less commonly cultivated worldwide and are often reputed to be susceptible to ascochyta blight disease, whereas other leaf types range from resistant to susceptible. This study determined the association between host plant resistance to ascochyta blight and different leaf types in segregating populations derived from crosses between disease resistant and susceptible chickpea genotypes. In addition, the inheritance of disease resistance and leaf type was investigated in intraspecific progeny derived from crosses between two resistant genotypes with normal leaf type (ICC 3996 and Almaz), one susceptible simple leaf type (Kimberley Large) and one susceptible multipinnate leaf type (24 B-Isoline). Our results showed that, in these segregating populations, susceptibility to ascochyta blight was not linked to multipinnate or simple leaf types; resistance to ascochyta blight depended more on genetic background than leaf shape; leaf type was controlled by two genes with a dihybrid supplementary gene action; normal leaf type was dominant over other leaf types; and inheritance of ascochyta blight resistance was controlled by two major genes, one dominant and one recessive. Since there was no linkage between ascochyta blight susceptibility and leaf type, breeding various leaf types with ascochyta blight resistance is a clear possibility. These results have significant implications for chickpea improvement, as most current extra large seeded kabuli varieties have a simple leaf type.     

Methods for screening resistance in chickpea to Ascochyta blight

Summary In Morocco, Ascochyta blight is a major limiting factor in chickpea production. The best long term solution to the problem seems to be the production of chickpea lines with durable resistance to the disease. Because of the nature of durable resistance, screening methods assessing resistance quantitatively had to be developed. Four methods are described: a seedling test, a germination test, a score of the percentage infected pods and a hair density score. With these screening methods a quantitative assessment of resistance in chickpea to blight appeared possible.Mr Pieters is with the FAO Plant Protection and Production division. Mr Tahiri is with the Service de Contrôle des Semences et Plants in Morocco.     

Genetics for speed of plumule emergence in chickpea (Cicer arietinum L.)

Summary Genetics for speed of plumule emergence was studied using six generations (P₁, P₂, F₁, BC₁(P₁), BC₂(P₂) and F₂) in three crosses. Two of the crosses which had parents of different emergence speed were controlled by two genes with duplicate epistasis. The third cross which involved parents of little difference for speed, indicated incomplete dominance for one gene of bit fast parent over the slow one. In all the crosses F₂ segregation pattern was confirmed by the segregation pattern of back crosses. The gene symbols were designated as Sp1Sp1 Sp2Sp2 for fast speed parents: sp1sp1 sp2sp2 for slow parent and sp1sp1 Sp2Sp2 for the parent with bit fastness for speed of plumule emergence.     

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).     

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.     

Two major genes for seed size in chickpea (Cicer arietinum L.)

Summary Seed size as determined by seed weight, is an important trait for trade and component of yield and adaptation in chickpea (Cicer arietinum L.). Inheritance of seed size in chickpea was studied in a cross between ICC11255, a normal seed size parent (average 120 mg seed⁻¹) and ICC 5002, a small seed size parent (average 50 mg seed⁻¹). Seed weight observations on individual plants of parents, F₁, F₂, and backcross generations, along with reciprocal cross generations revealed that the normal seed size was dominant over small seed size. No maternal effect was detected for seed size. The numbers of individuals with normal, small and medium (average 150 mg seed⁻¹) seed sizes in F ₂ population were 1237, 323 and 111 fitting well to the expected ratio of 12:3:1 (χ² = 0.923, P = 0.630). The segregation data of backcross generations also indicated that seed size in chickpea was controlled by two genes with dominance epistasis. We designate the genotype of ICC 11255 as Sd ₁ Sd ₁ sd ₂ sd ₂, and ICC 5002 as sd ₁ sd ₁Sd₂ Sd ₂ wherein Sd ₁ is epistatic to Sd ₂ and sd ₂ alleles.     

Inheritance of resistance to pea blight (Ascochyta pinodella) and induction of resistance in pea (Pisum sativum L.)

Summary Pea blight caused by Assochyta pinodella does considerable damage to the pea crop every year. To ascertain the inheritance of resistance to pea blight and incorporate resistance in the commercial cultivars, crosses were made between Kinnauri resistant to pea blight and four highly susceptible commercial pea cultivars — Bonneville, Lincoln, GC 141 and Sel. 18. Studies of the F₁'s, F₂'s, back crosses and F₃'s indicated that Kinnauri carries a dominant gene imparting resistance to pea blight.     

Inheritance of resistance to Ascochyta rabiei in 15 chickpea germplasm accessions

Inheritance of resistance to race 4 of Ascochyta rabiei was studied in fifteen chickpea accessions known internationally for Ascochyta blight (AB) resistance. Resistance in ILC 200, ILC 5921, ILC 6043 and ILC 6090 was governed by a single recessive gene. Resistance in ILC 202 and ILC 2956 was conferred by two recessive complementary genes. In the case of ILC 5586, resistance was controlled by two dominant complementary genes and in the case of ILC 2506, two recessive genes with epistasis interaction were responsible for resistance. Resistance in ILC 3279, ILC 3856 and ILC 4421 was controlled either by three recessive genes or two recessives duplicated genes and in ILC 72, ILC 182 and ILC 187 resistance was polygenic in nature. The study provided insights into the genetics of Ascochyta blight resistance, and these could be used in crossing programmes to develop durable resistance. While the virulence spectrum of the pathogen in a region plays a crucial role in the deployment of resistance, ILC72, ILC182, ILC200, ILC442 and ILC6090 could provide acceptable level of resistance if incorporated into commercial cultivars.     

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).     

Diallel analyses reveal the genetic control of resistance to ascochyta blight in diverse chickpea and wild Cicer species

Ascochyta blight is a major fungal disease affecting chickpea production worldwide. The genetics of ascochyta blight resistance was studied in five 5 × 5 half-diallel cross sets involving seven genotypes of chickpea (ICC 3996, Almaz, Lasseter, Kaniva, 24B-Isoline, IG 9337 and Kimberley Large), three accessions of Cicer reticulatum (ILWC 118, ILWC 139 and ILWC 184) and one accession of C. echinospermum (ILWC 181) under field conditions. Both F₁ and F₂ generations were used in the diallel analysis. The disease was rated in the field using a 1–9 scale. Almaz, ICC 3996 and ILWC 118 were the most resistant (rated 3–4) and all other genotypes were susceptible (rated 6–9) to ascochyta blight. Estimates of genetic parameters, following Hayman’s method, showed significant additive and dominant gene actions. The analysis also revealed the involvement of both major and minor genes. Susceptibility was dominant over resistance to ascochyta blight. The recessive alleles were concentrated in the two resistant chickpea parents ICC 3996 and Almaz, and one C. reticulatum genotype ILWC 118. The wild Cicer accessions may have different major or minor resistant genes compared to the cultivated chickpea. High narrow-sense heritability (ranging from 82% to 86% for F₁ generations, and 43% to 63% for F₂ generations) indicates that additive gene effects were more important than non-additive gene effects in the inheritance of the trait and greater genetic gain can be achieved in the breeding of resistant chickpea cultivars by using carefully selected parental genotypes.     

Breeding for resistance to lentil Ascochyta blight

Ascochyta blight, caused by Ascochyta lentis, is one of the most globally important diseases of lentil. Breeding for host resistance has been suggested as an efficient means to control this disease. This paper summarizes existing studies of the characteristics and control of Ascochyta blight in lentil, genetics of resistance to Ascochyta blight and genetic variations among pathogen populations (isolates). Breeding methods for control of the disease are discussed. Six pathotypes of A. lentis have been reported. Many resistant cultivars/lines have been identified in both cultivated and wild lentil. Resistance to Ascochyta blight in lentil is mainly under the control of major genes, but minor genes also play a role. Current breeding programmes are based on crossing resistant and high‐yielding cultivars and multilocation testing. Gene pyramiding, exploring slow blighting and partial resistance, and using genes present in wild relatives will be the methods used in the future. Identification of more sources of resistance genes, good characterization of the host‐pathogen system, and identification of molecular markers tightly linked to resistance genes are suggested as the key areas for future study.     

Mapping and validation of QTLs for resistance to an Indian isolate of Ascochyta blight pathogen in chickpea

Ascochyta blight (AB) caused by Ascochyta rabiei, is globally the most important foliar disease that limits the productivity of chickpea (Cicer arietinum L.). An intraspecific linkage map of cultivated chickpea was constructed using an F₂ population derived from a cross between an AB susceptible parent ICC 4991 (Pb 7) and an AB resistant parent ICCV 04516. The resultant map consisted of 82 simple sequence repeat (SSR) markers and 2 expressed sequence tag (EST) markers covering 10 linkage groups, spanning a distance of 724.4 cM with an average marker density of 1 marker per 8.6 cM. Three quantitative trait loci (QTLs) were identified that contributed to resistance to an Indian isolate of AB, based on the seedling and adult plant reaction. QTL1 was mapped to LG3 linked to marker TR58 and explained 18.6% of the phenotypic variance (R ²) for AB resistance at the adult plant stage. QTL2 and QTL3 were both mapped to LG4 close to four SSR markers and accounted for 7.7% and 9.3%, respectively, of the total phenotypic variance for AB resistance at seedling stage. The SSR markers which flanked the AB QTLs were validated in a half-sib population derived from the same resistant parent ICCV 04516. Markers TA146 and TR20, linked to QTL2 were shown to be significantly associated with AB resistance at the seedling stage in this half-sib population. The markers linked to these QTLs can be utilized in marker-assisted breeding for AB resistance in chickpea.     

The origin of chickpea Cicer arietinum L.

Summary Species relationship between the cultivated chickpea Cicer arietinum and the two newly discovered wild species C. echinospermum and C. reticulatum were assessed through breeding experiments and cytological examination of the hybrids.The two wild species differed from each other by a major reciprocal translocation and their hybrid was completely sterile. The wild species C. echinospermum also differed from the cultivated species by the same translocation and their hybrid was highly sterile. The other wild species, C' reticulatum, was crossed readily with the cultivated chickpea. Meiosis of the hybrids, involving 4 different C. arietinum lines, was normal, and they were fertile. This wild species therefore can be considered as the wild progenitor of the cultivated chickpea.     

Inheritance of resistance to Botrytis cinerea Pers. in Cicer arietinum L.

Summary Chickpea (Cicer arietinum L.) line ICC 1069 was selected as resistant parent after screening for resistance to grey mould (Botrytis cinerea Pers.) under artificial inoculation conditions. It was crossed with four high yielding susceptible varieties of chickpea. Crosses ICC 1069 × BGM 413 and ICC 1069 × BG 256 showed monogenic dominant resistance in ratio of 3R (resistant): 1S (susceptible). However, in crosses, ICC 1069 × BGM 419 and ICC 1069 × BGM 408, a ratio of 13S (susceptible) : 3R (resistant) was obtained indicating the presence of epistatic interaction. The results pointed towards the presence of a type of major gene resistance to grey mould in chickpea.     

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.     

Molecular mapping of QTLs for resistance to Fusarium wilt (race 1) and Ascochyta blight in chickpea (Cicer arietinum L.)

Fusarium wilt (FW) and Ascochyta blight (AB) are two important diseases of chickpea which cause 100 % yield losses under favorable conditions. With an objective to validate and/or to identify novel quantitative trait loci (QTLs) for resistance to race 1 of FW caused by Fusarium oxysporum f. sp. ciceris and AB caused by Ascochyta rabiei in chickpea, two new mapping populations (F_2:3) namely ‘C 214’ (FW susceptible) × ‘WR 315’ (FW resistant) and ‘C 214’ (AB susceptible) × ‘ILC 3279’ (AB resistant) were developed. After screening 371 SSR markers on parental lines and genotyping the mapping populations with polymorphic markers, two new genetic maps comprising 57 (C 214 × WR 315) and 58 (C 214 × ILC 3279) loci were developed. Analysis of genotyping data together with phenotyping data collected on mapping population for resistance to FW in field conditions identified two novel QTLs which explained 10.4–18.8 % of phenotypic variation. Similarly, analysis of phenotyping data for resistance to seedling resistance and adult plant resistance for AB under controlled and field conditions together with genotyping data identified a total of 6 QTLs explaining up to 31.9 % of phenotypic variation. One major QTL, explaining 31.9 % phenotypic variation for AB resistance was identified in both field and controlled conditions and was also reported from different resistant lines in many earlier studies. This major QTL for AB resistance and two novel QTLs identified for FW resistance are the most promising QTLs for molecular breeding separately or pyramiding for resistance to FW and AB for chickpea improvement.     

Genetics of resistance to 3 isolates of Ascochyta fabae on Faba bean (Vicia faba L.) in controlled conditions

Summary Genetics of resistance to Ascochyta fabae Speg. in Vicia faba L. was studied with a final objective to develop resistant faba bean varieties to Ascochyta blight. The study was conducted separately on 3 single spore isolates (AF10-2 and AF13-2 from Tunisia and AF4-3 from France) and belonging to different groups of virulence (GV1 and GV2). Important general combining ability (GCA) effects were found especially with isolates AF10-2 and AF4-3. Specific combining ability (SCA), although significant for the 3 isolates, was important only with AF13 -2, but less important than GCA. Additive gene effects were predominant to non-additive effects. Lines 29H and A8817 transmitted to their progenies resistance to the 3 isolates, whereas 14–12 and 19TB conferred resistance to their progenies only with isolates AF13-2 and AF4-3, respectively. In the material studied, resistance was generally controlled by dominant genes but also could be attributed to recessive genes although less frequent. Analysis of segregation in the F2 of 2 crosses between the resistant lines (A8817 and 29H) and the susceptible line (14–12) with isolate AF4-3 revealed dominant monogenic control at the level of leaves in the 2 resistant lines and, in addition, a recessive gene controlling resistance of stems. Non-allelic interactions were occasionally manifested and their origin appeared to be due to line 19TB. A recurrent selection scheme was proposed with the objective to develop improved open-pollination populations and synthetic varieties responding to the objective of the national Tunisian research programme on faba bean.     

Identification of resistant sources and DNA markers linked to genomic region conferring dry root rot resistance in chickpea (Cicer arietinum L.)

A set of 520 chickpea germplasm lines was screened under laboratory conditions using blotter paper technique for reaction to dry root rot caused by Rhizoctonia bataticola (Taub.) Butler. The lines PG06102, BG2094 and IC552137 were identified as resistant for dry root rot. Phenotyping the mapping population consisting of 129 F_2:3 progeny derived from the cross L550 × PG06102 during 2013 winter indicated monogenic inheritance of dry root rot resistance. Fifty‐two of 381 simple sequence repeat (SSR) primers polymorphic between the two parents were used to genotype F₂ resistant and susceptible bulks prepared on the basis of reaction of F_2:3 progeny. Four markers differentiated the resistant and susceptible bulks. All the four polymorphic markers were then assayed on the entire F₂ population. Linkage analysis using 129 F₂ plants revealed that two markers ICCM0299 and ICCM0120b were co‐segregating with resistance to dry root rot. These two markers appeared to have additive effects on resistance and could be potentially utilized in dry root resistance breeding programme.