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.     

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     

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.     

Genetic basis of pod borer (Helicoverpa armigera) resistance and grain yield in desi and Kabuli Chickpea (Cicer arietinum L.) under unprotected conditions

A series of half-diallel crosses involving early, medium and late maturity desi and kabuli type chickpea (Cicer arietinum L.) genotypes with stable resistance to Helicoverpa pod borer, along with the parents, were evaluated at two locations in India to understand the inheritance of pod borer resistance and grain yield. Inheritance of resistance to pod borer and grain yield was different in desi and kabuli types. In desi type chickpea, the additive component of genetic variance was important in early maturity and dominance component was predominant in medium maturity group, while in the late maturity group, additive as well as dominance components were equally important in the inheritance of pod borer resistance. Both dominant and recessive genes conferring pod borer resistance seemed equally frequent in the desi type parental lines of medium maturity group. However, dominant genes were in overall excess in the parents of early and late maturity groups. In the kabuli medium maturity group, parents appeared to be genetically similar, possibly due to dispersion of genes conferring pod borer resistance and susceptibility, while their F₁s were significantly different for pod borer damage. The association of genes conferring pod borer resistance and susceptibility in the parents could be attributed to the similarity of parents as well as their F₁s for pod borer damage in kabuli early and late maturity groups. Grain yield was predominantly under the control of dominant gene action irrespective of the maturity groups in desi chickpea. In all the maturity groups, dominant and recessive genes were in equal frequency among the desi parental lines. Dominant genes, which tend to increase or decrease grain yield are more or less present in equal frequency in parents of the early maturity group, while in medium and late maturity groups, they were comparatively in unequal frequency in desi type. Unlike in desi chickpea, differential patterns of genetic components were observed in kabuli chickpea. While the dominant genetic component was important in early and late maturity group, additive gene action was involved in the inheritance of grain yield in medium duration group in kabuli chickpea. The dominant and recessive genes controlling grain yield are asymmetrically distributed in early and medium maturity groups in kabuli chickpea. The implications of the inheritance pattern of pod borer resistance and grain yield are discussed in the context of strategies to enhance pod borer resistance and grain yield in desi and kabuli chickpea cultivars.     

Inheritance of resistance to the chlorosis component of tan spot of wheat caused by Pyrenophora tritici-repentis, races 1 and 3

The fungus Pyrenophora tritici-repentis, causal agent of tan spot of wheat, produces two phenotypically distinct symptoms, tan necrosis and extensive chlorosis. The inheritance of resistance to chlorosis induced by P. tritici-repentis races 1 and 3 was studied in crosses between common wheat resistant genotypes Erik, Hadden, Red Chief, Glenlea, and 86ISMN 2137 and susceptible genotype 6B-365. Plants were inoculated under controlled environmental conditions at the two-leaf stage and disease rating was based on presence or absence of chlorosis. In all the resistant × susceptible crosses, F₁ plants were resistant and the segregation of the F₂ generation and F₃ families indicated that a single dominant gene controlled resistance. Lack of segregation in a partial diallel series of crosses among the resistant genotypes tested with race 3␣indicated that the resistant genotypes possessed␣the same resistance gene. This resistance gene was effective against chlorosis induced by P.␣tritici-repentis races 1 and 3.     

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).     

Genetical analysis of resistance to Mycosphaerella pinodes in pea seedlings

Summary In studies of the inheritance of resistance, pea seedlings of seven lines in which stems and leaves were both resistant to Mycosphaerella pinodes were crossed with a line in which they were both susceptible. With seven of the crosses resistance was dominant to susceptibility. When F₂ progenies of five crosses were inoculated on either stems or leaves independently, phenotypes segregated in a ratio of 3 resistant: 1 susceptible indicating that a single dominant gene controlled resistance. F₂ progenies of one other cross gave ratios with a better fit to 9 resistant: 7 susceptible indicating that two co-dominant genes controlled resistance. The F₂ progeny of another cross segregated in complex ratios indicating multigene resistance.When resistant lines JI 97 and JI 1089 were crossed with a susceptible line and leaves and stems of each F₂ plant were inoculated, resistance phenotypes segregated independently demonstrating that leaf and stem resistance were controlled by different genes. In two experiments where the F₂ progeny of the cross JI 97×JI 1089 were tested for stem and leaf resistance separately, both characters segregated in a ratio of 15 resistant:1 susceptible indicating that these two resistant lines contain two non-allelic genes for stem resistance (designated Rmp1 and Rmp2) and two for leaf resistance (designated Rmp3 and Rmp4). Evidence that the gene for leaf resistance in JI 1089 is located in linkage group 4 of Pisum sativum is presented.     

Leaf Waxes of Cassava (Manihot Esculenta Crantz) in Relation to Ecozone and Resistance to Xanthomonas Blight

Summary To elucidate the role of leaf surface structures as first barriers to confer resistance to bacterial blight, leaf stomata and their occlusion with leaf waxes were studied in cassava genotypes. For the first time, cassava leaf waxes were quantitatively and qualitatively analysed. Comparing the susceptible and resistant standard genotypes BEN86052 and TMS30572, respectively, the total quantity of triterpenes was significantly higher in the resistant genotype, grown in three ecozones of Benin. In cuticular leaf waxes of seven cassava genotypes the triterpenes beta amyrins, epi-taraxerol, taraxerone and taraxerol were dominant constituents across genotypes, and alkanes (C₂₅-C₃₃) and acids (C₃₀ and C₃₂) occurred in minor concentrations. Comparing seven genotypes, no clear relation between resistance or ecozones and total quantities of the major wax constituents was observed. Only the highly resistant genotype TMS30572 showed high triterpene levels irrespective of ecozone. Scanning electron-microscopy revealed a regular distribution of waxes at the abaxial leaf surface, covering and occluding stomatal pores of susceptible and resistant genotypes, while on the adaxial leaf surface waxes were in form of crystalloids and did not occlude the stomata. The number of stomata on the abaxial surfaces was about 7–11 fold higher than on the adaxial surfaces, where stomata were located along the midrib and major veins. No significant differences in stomata number were observed between genotypes varying in resistance to bacterial blight. It is suggested, that stomata on the adaxial surface might be portals of entry for the bacteria.     

Association between advanced generations and genealogy in inbred lines of snap bean by the Ward-Modified Location Model

Genetics of resistance to ascochyta blight was studied using different generations of fifteen crosses of chickpea (Cicer arietinum L.). Six parents comprising two susceptible varieties GL 769, C 214 and four resistant lines GG 1267, GL 90168, GL 96010 and GL 98010 were used to develop one S × S, eight S × R and six R × R crosses and some of the back crosses and F₃ generations were developed. Field screening technique was used to evaluate the different generations for disease reaction using mixture of ten prevalent isolates (ab₁–ab₁₀) of ascochyta blight (Ascochyta rabiei). Inheritance study showed digenic recessive control of resistance in the cross GL 769 × C 214, whereas monogenic recessive control of resistance was found in the crosses GL 769 × GL 98010 and C 214 × GL 98010. Digenic dominant and recessive control of resistance was found in the crosses GL 769 × GG 1267 and C 214 × GG 1267 while the crosses GL 769 × GL 90168 and C 214 × GL 96010 showed the monogenic dominant control of resistance. Trigenic dominant and recessive control of resistance was observed in the crosses GL 769 × GL 96010 and C 214 × GL 90168. Allelic relationship studies showed that three resistant parents viz., GG 1267, GL 96010 and GL 90168 possessed allelic single dominant gene for resistance. Besides, GG 1267 possessed two minor recessive genes for resistance, one of them was allelic to the minor recessive gene possessed by GL 90168 and other with GL 96010. The resistant parents GL 90168 and GL 96010 possessed non-allelic minor gene for resistance. The resistant parent GL 98010 possessed two minor recessive genes for resistance which were allelic to respective single recessive gene for resistance possessed by the susceptible parents GL 769 and C 214. The susceptible parents GL 769 and C 214 also possessed single independent inhibitory dominant susceptibility gene. The inhibitory gene was epistatic to the corresponding recessive gene for resistance.     

Selection of wild Cicer accessions for the generation of mapping populations segregating for resistance to ascochyta blight

In order to determine genetically diverse parents for the generation of mapping populations segregating for resistance to ascochyta blight in wild Cicer species, the genetic diversity between a selection of resistant and susceptible accessions was assessed using molecular markers. Twenty Cicer accessions — comprising eight C. reticulatum accessions, six C. echinospermum accessions, five C. bijugum accessions, and one C. arietinum accession — were compared using a combination of seven RAPD primers and seven ISSR primers. A total of 231 polymorphic bands were scored and used to determine the genetic distances between accessions using Jaccard similarity coefficients. The most genetically diverse parents for the generation of intraspecific and interspecific populations segregating for resistance to ascochyta blight are reported. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of resistance to angular leaf spot in common bean and validation of the utility of resistance linked markers for marker assisted selection out side the mapping population

Inheritance of resistance to angular leaf spot (ALS) disease caused by Phaeoisariopsis griseola (Sacc.) Ferr was investigated in two common bean cultivars, Mexico 54 and BAT 332. Both Andean and Mesoamerican backgrounds were used to determine the stability of the resistance gene in each of the two cultivars. Resistance to P. griseola was phenotypically evaluated by artificial inoculation with one of the most widely distributed pathotypes, 63–39. Evaluation of the parental genotypes, F₁, F₂ and backcross populations revealed that the resistance to angular leaf spot in the cultivars Mexico 54 and BAT 332 to pathotype 63–39 is controlled by a single dominant gene, when both the Andean and Mesoamerican backgrounds were used. Allelism test showed that ALS resistance in Mexico 54 and BAT 332 to pathotype 63–39 was conditioned by the same resistance locus. Resistant and susceptible segregating populations generated using Mexico 54 resistant parent were selected for DNA extraction and amplification to check for the presence /absence of the SCAR OPN02 and RAPD OPE04 markers linked to the Phg-2 resistance gene. The results indicated that the SCAR OPN02 was not polymorphic in the study populations and therefore of limited application in selecting resistant genotypes in such populations. On the other hand, the RAPD OPE04 marker was observed in all resistant individuals and was absent in those scored susceptible based on virulence data. Use of the RAPD OPE04 marker in marker-assisted selection is underway.     

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.     

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.     

Genetical relationships between reactions to bacterial leaf spot,yellow mosaic and Cercospora leaf spot diseases in mungbean (Vigna radiata)

Summary Studies on the inheritance pattern of bacterial leaf spot (BLS), yellow mosaic (YM) and Cercospora leaf spot (CLS) reactions in crosses of BLS and YM resistant/tolerant but CLS susceptible × CLS resistant but BLS and YM susceptible parents indicated that resistances to BLS and CLS were governed by single dominant genes, whereas YM tolerance was a monogenic recessive character. The studies also indicated that these three genes were inherited independently. The simple inheritance pattern and independent assortment of the genes governing resistance/tolerance to these diseases suggest that the usual breeding methods will be adequate to develop multi-disease resistant mungbean cultivars.Paper XII in the series Studies on resistance in crops to bacterial diseases in India.     

Evaluation of resistance to verticillium wilt in diploid,wild potato interspecific hybrids

Summary Verticillium wilt (V. albo-atrum Reinke & Berthold or V. dahliae Kleb) threatens potato (Solanum tuberosum L.) production in most growing areas of the world. Genetic resistance offers the most cost-effective and environmentally-sound control measure. However, there is a dearth of genetic and breeding information on resistance to verticillium wilt in potato, because of obscure parentage of some standard cultivars and the complex segregation at the tetraploid level. The wide range of genetic variability in wild relatives of potatoes can be potentially useful as a source of disease resistance, such as verticillium wilt resistance. Six diploid, wild, interspecific Solanum hybrids involving resistant x resistant and susceptible x resistant crosses, were assayed for verticillium wilt resistance under greenhouse conditions to evaluate their potential as sources of verticillium wilt resistance. The cross between S. gourlayi Oka. and S. chacoense Bitt. and its reciprocal had the most resistant progenies based on mean colony counts. No simple mode of inheritance can be proposed based on the observed segregation ratios. However, the continuous distributions observed on verticillium wilt disease response among hybrid families indicate that inheritance of resistance may be polygenic and complex. In addition, skewness of colony count distributions toward the resistance parents were characteristic of all resistant x susceptible crosses suggesting that resistance may be dominant. By contrast, the susceptible x susceptible cross showed a more normal distribution. Overall, the cross between S. gourlayi and S. chacoense showed the most potential as a source of verticillium wilt resistance.     

DAF marker tightly linked to a major locus for Ascochyta blight resistance in chickpea (Cicer arietinum L.)

Resistance of chickpea against the disease caused by the ascomycete Ascochyta rabiei is encoded by two or three quantitative trait loci, QTL1, QTL2 and QTL3. A total of 94 recombinant inbred lines developed from a wide cross between a resistant chickpea line and a susceptible accession of Cicer reticulatum, a close relative of cultivated chickpea, was used to identify markers closely linked to QTL1 by DNA amplification fingerprinting in combination with bulked segregant analysis. Of 312 random 10mer oligonucleotides, 3 produced five polymorphic bands between the parents and bulks. Two of them were transferred to the population on which the recent genetic map of chickpea is based, and mapped to linkage group 4. These markers, OPS06-1 and OPS03-1, were linked at LOD-scores above 5 to markers UBC733B and UBC181A flanking the major ascochyta resistance locus. OPS06-1 mapped at the peak of the QTL between markers UBC733B (distance 4.1 cM) and UBC181A (distance 9.6 cM), while OPS03-1 mapped 25.1 cM away from marker UBC733B on the other flank of the resistance locus. STMS markers localised on this linkage group were transferred to the population segregating for ascochyta resistance. Three of these markers were closely linked to QTL1. Twelve of 14 STMS markers could be used in both populations. The order of STMS markers was essentially similar in both populations, with differences in map distances between them. The availability of flanking STMS markers for the major resistance locus QTL1 will help to elucidate the complex resistance against different Ascochyta pathotypes in future. This revised version was published online in August 2006 with corrections to the Cover Date.     

Allele-specific amplification for the detection of ascochyta blight resistance in chickpea

A new co-dominant molecular marker, CaETR, was developed based on allelic sequence length polymorphism in an ethylene receptor-like gene located in the genomic region of a QTL (QTL_AR1) conferring ascochyta blight resistance in chickpea. This marker not only discriminated resistance and susceptible phenotypes of chickpea to ascochyta blight, but also easily detected heterozygous genotypes. Using the CaETR marker in combination with a previously developed co-dominant SCAR marker (SCY17₅₉₀) linked to another QTL (QTL_AR2) it was possible to detect resistance alleles in 90?% of resistant accessions in a collection of landraces, advances breeding lines and cultivars, and also detected susceptible alleles in all cases. The results of this study offer a scope for improving the efficiency of conventional chickpea breeding by carrying out negative selection for QTL_AR1 and QTL_AR2 in early generations without relaying directly on the phenotype. This PCR-based approach using both co-dominant markers is proposed as an efficient tool for selecting blight-resistant genotypes in breeding programs.     

Inheritance of resistance to wildfire and angular leaf spot derived from Nicotiana rustica var. Brasilea

Summary The introgression of wildfire (races 0 and 1) and angular leaf spot (ALS) resistance from N. rustica var. Brasilea into N. tabacum has proved economically useful in Zimbabwe although the mode of inheritance of, and genetic relationships between the resistance are unknown. This study was undertaken to (1) examine the mode of inheritance of the resistance to races 0 and 1 of wildfire, and ALS, (2) determine the genetic relationship between the resistances and (3) establish whether the N. rustica-derived wildfire race 0 resistance is allelic to that obtained from N. longiflora. Inheritance was examined under greenhouse and field conditions by studying disease reactions in the parental, F1, F2 and backcross generations derived from crosses of three susceptible lines to a resistant line Nr-7. Three-point backcrosses to the susceptible parent were examined for linkage and segregating generations from a cross of Nr-7 to Burley 21 which carries the N. longiflora race 0 resistance were used to test for allelism. In general, we observed that all resistances are determined by a single dominant gene although some incosistent ratios were obtained likely due to misclassification of disease reactions and erratic transmission. All resistances showed linkage although pleiotropism cannot be ruled out. Allelism tests demonstrated that the N. rustica race 0 resistance is not allelic to that obtained from N. longiflora. Our findings are examined in relation to the efficacy of indirect selection for resistance.     

Mid-parent heterosis and combining ability of European corn borer resistance in maize

Success in breeding maize resistant to the European corn borer has been limited, with the exception of leaf feeding resistance. The inheritance of resistance to leaf, sheath-collar and ear damage in four maize germplasms and their six F₁ crosses was evaluated by diallel analysis. Plants in a completely randomized design were artificially infested at the whorl, anthesis or full silk stage of plant development and were evaluated in the field for insect damage. A damage index based on size, number and location of lesions was calculated for each stage. Stowell's Evergreen (susceptible) had a mean damage index three to six times that of Maiz Amargo (resistant) at the whorl stage and the progeny plants were more resistant than the susceptible parent. Maiz Amargo and its crosses had significantly lower mean indices than Stowell's Evergreen for sheath-collar damage in Year 1 but not Year 2. Zapalote Chico, Maiz Amargo and their cross were significantly less damaged than other genotypes at the full silk stage. Heterosis values indicated an increase in resistance of crosses over the midparent average at all three stages of development. General combining ability (GCA) was highly significant for all types of damage, but specific combining ability was significant only for leaf damage. Based on estimates of GCA, Maiz Amargo was the best source of resistance to leaf and sheath-collar damage and both Zapalote Chico and Maiz Amargo would be good parents for ear damage resistance. Results suggest that resistance at different plant development stages can be combined.     

Inheritance of early blight resistance in diploid potatoes

Early blight (Alternatia solani) is a fungal disease in hot and humid environments, which causes leaf, stem and tuber lesions. Early blight resistance should be incorporated into potato cultivars because the fungicide spraying is an expensive solution for developing countries. The diploid cultivated species Solanum tuberosum group Phureja and group Stenotomum are sources of resistance alleles. The elucidation of the inheritance for early blight resistance must help to decide what could be the best breeding procedure to improve this diploid germplasm and transfer the resistance to the tetraploid level. Three experiments were carried out under controlled and field conditions to determine the heritability of this trait using nested and diallel mating designs with haploid, species and haploid-species hybrids. The narrow-sense heritability estimates were relatively high (0.64–0.78). This means that additivity was the most important type of gene action for determining resistance to early blight at the diploid level. The results suggested that diploid parents showing highest levels of resistance, throughout the cycle of disease development, can be used in 4x×2x crosses to obtain resistant tetraploid progenies to this fungal disease.