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.     

Linkage relationships of genes for leaf morphology,flower color,and root nodulation in chickpea

Summary The allelic and linkage relationships among five chickpea (Cicer arietinum L.) morphological markers were investigated. When crossed with purple-flowered line ICC 640 and with each other, white flowered variety UC5 and mutant line PM974 were shown to carry non-allelic, single recessive genes for white flower color, provisionally designated w1 and w2, respectively. The single recessive gene conferring simple leaves in mutant PM299 was allelic to the previously described slv gene carried by variety Surutato 77, line ICC 10301, and other simple leaf chickpea mutants. In mutant 756M, a filiform leaf trait was controlled by a single recessive gene, fil, which was non-allelic to slv.The fil and w2 genes were linked, with recombination frequencies of 0.05 and 0.14 estimated from results of coupling and repulsion phase crosses, respectively. fil and w1 segregated independently. Root nodulation gene rn3 was closely linked to slv: recombination frequencies of 0.05 and 0.11 were estimated from results of coupling and repulsion phase crosses, respectively. A loose linkage detected between the w2-fil and the rn3-slv linkage groups will be the subject of further scrutiny.     

Direct and indirect selection for yield in chickpea

Summary The efficiency of indirect selection for seed yield was compared with direct selection for yield per se in chickpea. A total of 2500 single F₂ plants, derived from 50 crosses with 50 plants from each cross, were divided into five sub-populations (SP1 to SP5) of 500 plants each by including 10 plants from each of the 50 crosses. The five sub-populations were advanced upto F₆ by exercising 10% selection intensity for four successive generations for number of pods per plant in SP1, number of seeds per pod in SP2, seed weight in SP3, seed yield in SP4 and random selection in SP5. The efficiency of direct and indirect selection for yield was evaluated by comparing groups of 50 F₆ lines from each sub-population. SP1 and SP3 F₆ lines showed higher mean grain yield than the other three methods. SP1 and SP3 were found to be almost equally efficient in developing F₆ lines which were significantly superior to the check. This suggests that indirect selection for yield via pod number and seed weight is more efficient than direct selection for yield.     

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).     

Fasciation in chickpea: genetics and evaluation

Summary A spontaneous fasciated mutant was detected in the chickpea cv. Amethyst. It was characterised by broad, strap-like stems, irregular leaf arrangement and clustering of pods towards the stem apices. F₂ and F₃ segregations showed that fasciation was controlled by a single, recessive gene for which the symbol fas is proposed. Field trials comparing the fasciated mutant with its non-fasciated isoline showed that fasciation was associated with lower yield, larger seeds, delayed flowering and increased lodging.     

Allelic relationships of genes controlling number of flowers per axis in chickpea

Genetic variation for number of flowers per axis in chickpea (Cicer arietinum L.) includes single-flower, double-flower, triple-flower and multi-flower traits. A double-flowered (DF) line ICC 4929, a triple-flowered (TF) line IPC 99-18 and a multi-flowered (MF) line JGM 7 were intercrossed in all possible combinations and flowering behavior of parents, F₁s and F₂s was studied to establish allelic relationships, penetrance and expressivity of genes controlling number of flowers per axis in chickpea. The F₁ from ICC 4929 (DF) × IPC 99-18 (TF) cross were double-flowered, whereas F₁ from ICC 4929 (DF) × JGM 7 (MF) and IPC 99-18 (TF) × JGM 7 (MF) crosses were single-flowered. The F₂ from ICC 4929 (DF) × IPC 99-18 (TF) cross gave a good fit to a 3:1 ratio for double-flowered and triple-flowered plants. The F₂ from ICC 4929 (DF) × JGM 7 (MF) cross segregated in a ratio of 9:3:3:1 for single-flowered, double-flowered, multi-flowered and double-multi-flowered plants. The F₂ from IPC 99-18 (TF) × JGM 7 (MF) cross segregated in a ratio of 9:3:4 for single-flowered, triple-flowered and multi-flowered plants. The results clearly established that two loci control number of flowers per axis in chickpea. The double-flower and triple-flower traits are controlled by a single-locus (Sfl) and the allele for double-flowered trait (sfl ^d ) is dominant over the allele for triple-flower trait (sfl ^t ). The three alleles at the Sfl locus has the dominance relationship Sfl > sfl ^d > sfl ^t . The multi-flower trait is controlled by a different gene (cym). Single-flowered plants have dominant alleles at both the loci (Sfl_ Cym_). The double-flower, the triple-flower and the multi-flower traits showed complete penetrance, but variable expressivity. The expressivity was 96.3% for double-flower and 76.4% for double-pod in ICC 4929, 81.2% for triple-flower and 0.0% for triple-pod in IPC 99-18, and 51.3% for multi-flower and 24.7% for multi-pod in JGM 7. Average number of flowers per axis and average number of pods per axis were higher in JGM 7 than double-flowered line ICC 4929 and triple-flowered line IPC 99-18. The results of this study will help in development of breeding strategies for exploitation of these flowering and podding traits in chickpea improvement.     

Detection of epistasis in chickpea

Summary Triple test cross-analysis was used to detect epistasis in chickpea. None of the characters investigated exhibited epistasis. In the absence of epistasis, additive and dominance effects were estimated. The results indicated the importance of additive genetic variance for seed yield, biological yield, number of primary branches, number of secondary branches, 100-seed weight, days to flower, and number of seeds per pod; dominance genetic variance for days to mature; and both additive and dominance genetic variances for plant height. Selection methods, such as pedigree and bulk, are suggested for the improvement of most characters.Joint contribution from ICARDA, P.O. Box 5466, Aleppo, Syria and ICRISAT (International Crops Research Institute for the Semi-Arid Tropics), Patancheru P.O. 502 324, A.P., India.     

A gene producing one to nine flowers per flowering node in chickpea

Chickpea (Cicer arietinum L.) has a racemose type of inflorescence and at each axis of the raceme usually one or two and rarely three flowers are borne. Plants producing 3 to 9 flowers, arranged in acymose inflorescence, at many axis of the raceme, were identified in F₂ of an interspecific cross ICC 5783 (C. arietinum) × ICCW 9 (C. reticulatum)in which both the parents involved were single-flowered. A spontaneous mutation in one of the two parents or in the F₁was suspected. However, the possibility for establishment of a rare recombination of two interacting recessive genes could not be ruled out. The number of pods set varied from 0 to 5 in each cyme. Inheritance studies indicated that a single recessive gene, designated cym, is responsible for cymose inflorescence. The allelic relationship of cym with sfl, a gene for double-flowered trait, was studied from a cross involving multi flowered plants and the double-flowered line ICC 4929. Thecym gene was not allelic to sfl, suggesting that two loci control the number of flowers per peduncle in chickpea. The cym locus segregated independently of the locus sfl, ifc (inhibitor of flower color) and blv (bronze leave). This revised version was published online in August 2006 with corrections to the Cover Date.     

Induction,genetics and possible use of glabrousness in chickpea

Summary A giabrous mutant was identified from progenies of chickpea seeds that were treated with ethyl methane sulphonate (EMS). The mutant has no shoot hairs in contrast to the dense hairs on normal chickpeas. The character is governed by a single recessive gene. This mutant can be useful in certain pathological and eniomological studies.     

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 a plant type mutant and its utilization in chickpea

Summary A macro-mutant, E 100Y(M) in chickpea (Cicer arietinum L.) was found to affect several plant and seed characters. For plant type monogenic inheritance was observed. A single pair of recessive genes pt/pt was ascribed to this mutant. The mutant locus seemed to be exerting pleiotropic action. The utilization of this mutant for chickpea improvement has been discussed.     

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.     

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.     

SSR-based genetic diversity assessment among Tunisian winter barley and relationship with morphological traits

For studying genetic diversity caused byselection for adaptation and end-use, 17microsatellites (SSR), representative ofthe barley genome, were used in 26 barley(Hordeum vulgare L.) accessions andcultivars in Tunisia. Theaccessions/cultivars originate fromdifferent geographic regions and are ofdifferent end-use. For the 15 polymorphicSSR, the mean number of alleles per locuswas 3.6 and the average polymorphisminformation content was 0.45. Clusteranalysis based on SSR data and onmorphological data clearly differentiatethe genotypes according to their type(local landraces vs. varieties), row-numberand end-use. The correlation between bothdiversity measures was highly significant(r = 0.25, p<10^-5) and thecorrespondence between the clustering basedon SSR and morphological data wasrelatively good. Our results show the largegenetic diversity of the Tunisian barleycultivars and the association of thisdiversity with adaptation traits.     

Pollen selection for chilling tolerance at hybridisation leads to improved chickpea cultivars

The potential of pollen selection as part of the breeding efforts to increase chilling tolerance in chickpea was investigated. This alternative approach to apply selection pressure at the gametophytic stage in the life cycle has been proposed widely, but there are no reports of the technique being implemented in a crop improvement program. In this paper, we describe how we developed a practical pollen selection technique useful for chickpea improvement.Pollen selection improved chilling tolerance in crossbreds compared with the parental chickpea genotypes and compared with progeny derived without pollen selection. This is backed up by controlled environment assessments in growth rooms and by field studies. We also clearly demonstrate that chilling tolerant pollen wins the race to fertilise the ovule at low temperature, using flower color as a morphological marker. Overall, pollen selection results in a lower threshold temperature for podding, which leads to pod setting two to four weeks earlier in the short season Mediterranean-type environments of Western Australia. Field testing at multiple sites across Australia, as part of the national crop variety testing program, shows that these breeding lines have a significant advantage in cool dryland environments.The major factors which affected the success of pollen selection are discussed in the paper, from generation of variability in the pollen to a means to recover hybrids and integration of our basic research with an applied breeding program. We conclude that chilling tolerance observed in the field over successive generations are the result of pollen selection during early generations.     

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.     

GBS高密度遗传连锁图谱定位甘蓝型油菜粉色花性状

甘蓝型油菜花瓣颜色是重要的观赏性状,花瓣颜色的选育和改良已成为材料创制的主要研究方向。目前对甘蓝型油菜粉色花性状定位研究未见报道。本研究以甘蓝型油菜纯系黄花62和甘蓝型油菜纯系粉花77为亲本构建双单倍体(doubled haploid,DH)群体。利用简化基因组测序技术(genotyping-by-sequencing,GBS)筛选出3253个单核苷酸多态性(single nucleotide polymorphism,SNP)标记,构建了全长1766.06 cM甘蓝型油菜连锁图谱,标记间平均遗传距离为0.54 cM;使用WinQTL Cartographer复合区间作图方法对粉色花性状进行数量性状座位(quantitative trait locus,QTL)定位,检测到位于A07和C03染色体上各1个QTL;将定位区间内基因与甘蓝和白菜同源基因进行线性比对,在这些区间中找到了一些存在于3个物种的同源基因;对粉色花定位区间内基因进行可变剪切分析发现,2个花色相关基因BnaA07g15980D和BnaA07g17500D在亲本粉色花瓣中发生内含子保留可变剪切。上述研究结果为甘蓝型油菜粉色花相关基因精细定位和分子连锁标记开发提供了更多线索。     

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.