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.     

Allelic relationship between spontaneous and induced mutant genes for stem fasciation in chickpea

Stem fasciation is a morphological abnormality observed in plants where the stem is widened and leaves and flowers or pods are clustered at the apex. Several spontaneous mutants and one induced mutant for stem fasciation are found in chickpea (Cicer arietinum L.). This study was aimed at determining allelic relationship between spontaneous and induced mutant genes controlling stem fasciation and effects of stem fasciation on grain yield. Two spontaneous (ICC 2042 and ICC 5645) and one induced (JGM 2) stem fasciation mutants were crossed in all combinations, excluding reciprocals. The F₁ and F₂ plants from a cross between the two spontaneous mutants had fasciated stem. This indicated the presence of a common gene (designated fas1) for stem fasciation in the two spontaneous mutants. The F₁s of the crosses of the induced mutant JGM 2 with both spontaneous mutants had normal plants and segregated in a ratio of 9 normal : 7 fasciated plants in F₂. Thus, the gene for stem fasciation in the induced mutant JGM 2 (designated fas2) is not allelic to the common gene for stem fasciation in spontaneous mutants. The two genes in dominant condition produced normal non‐fasciated stem. The fasciated and the non‐fasciated F₂ plants did not differ significantly for number of pods per plant, number of seeds per plant, grain yield per plant and seed size, suggesting that it is possible to exploit the fasciated trait in chickpea breeding without compromising on yield.     

An induced brachytic mutant of chickpea and its possible use in ideotype breeding

Mutations were induced in chickpea (Cicer arietinum L.) cultivar ‘JG 315’ through treatment of seeds with ethyl methane sulphonate (EMS). One of the mutants, named JGM 1, had brachytic growth (compact growth), characterized by erect growth habit, thick and sturdy stem, short internodal and interleaflet distances and few tertiary and later order branches. It was isolated from M₂ derived from seeds treated with 0.6% EMS for 6 h. Segregation analyses in F₂ progenies of its crosses with normal chickpea genotypes (JG 315, ICC 4929, and ICC 10301) suggested that a single recessive gene controlled brachytic growth in JGM 1. This gene was not allelic to the br gene for brachytic growth in spontaneous brachytic mutant E100YM. Thus, the gene for brachytic growth in JGM 1 was designated br2 and the br gene of E100YM was redesignated br1. Efforts are being made to use JGM 1 in development of a plant type with short internodes and erect growth habit. Such plant type may resist excessive vegetative growth in high input (irrigation and fertility) conditions and accommodate more plants per unit area.     

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.     

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.     

Genetics of thermosensitive genic male sterility in rice

Summary Inheritance of thermosensitive genic male sterility (TGMS) in Norin PL12 and IR32364TGMS and their allelic relationship were studied from F₁, F₂ testcross (TC) and F₃ generations of the crosses made with the two mutants and several fertile tester parents. F₂, TC and F₃ segregation behavior for pollen and spikelet fertility indicated that the TGMS trait in the two mutants was controlled by a single recessive gene. Allelic relationship studies indicated that TGMS genes of the two mutants were different. Since TGMS gene in Norin PL12 has been designated as tms ₂ , the TGMS gene present in IR32364TGMS is tentatively designated as tms ₃ (t) until allelic test is done with another TGMS gene (tms ₁ ) reported from China in a line 5460S seeds of which were not available.     

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.     

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 whitebacked planthopper,Sogatella furcifera (Horvath) in some rice cultivars

Summary The genetics of resistance to whitebacked planthopper, Sogatella furcifera (Horvath) in ten resistant cultivars was studied. The reactions of the F₁, F₂ and F₃ populations of resistant varieties with Taichung Native 1, a suspectible check, showed that WBPH resistance is monogenic in nature and governed by dominant gene(s) in Ptb 19 and IET 6288 and recessive gene in eight cultivars viz. ARC 5838, ARC 6579, ARC 6624, ARC 10464, ACR 11321, ARC 11320, Balamawee and IR 2415-90-4-3. Allelic relationship of resistance gene(s) in the test cultivars revealed recessive gene in IR 2415-90-4-3, ARC 5838 and ARC 11324 to be allelic but it was non allelic to the resistance gene in ARC 6624. Cultivars ARC 6579, ARC 11321 and Balamawee have identical gene among themselves but their relationship with IR 2415-90-4-3, ARC 5838, ARC 11324 and ARC 6624 is unknown. The recessive gene in ARC 10464 is non-identical to all other cultivars having the recessive gene except ARC 6624 with which its relationship needs further investigation.     

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 two components of early maturity in groundnut (Arachis hypogaea L.)

Summary Knowledge of inheritance of early maturity or its components is important to groundnut breeders in developing short-duration cultivars. This study was conducted to determine the inheritance of two components of early maturity: days to first flower from sowing, and days to accumulation of 25 flowers from the appearance of first flower, using three groundnut genotypes. Two early-maturing (Chico and Gangapuri) and one late-maturing (M 13) genotypes were crossed in all possible combinations, including reciprocals. The parents, F₁, F₂, F₃, and backcross populations were evaluated for days to first flower from sowing, and for days to accumulation of 25 flowers. The data suggest that days to first flower in the crosses studied is governed by a single gene with additive gene action. Chico and Gangapuri possess the same allele for this component of earliness. Three independent genes with complete dominance at each locus appear to control the days to accumulation of 25 flowers. In crosses between late (M 13) and early (Chico or Gangapuri) parents, a segregation pattern suggesting dominant-recessive epistasis (13 late:3 early) was observed for this component. Segregation in the F₂ generation (1 late:15 early) of both early parents (Chico x Gangapuri) indicated that the genes for early accumulation of flowers in these two parents are at different loci.Submitted as ICRISAT J.A. No. 1557.     

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.     

Two independent gene loci controlling non-brittle pedicels in buckwheat

The shattering habit in buckwheat occurs because of brittle or weak pedicels. Brittle pedicels are observed in wild buckwheat but not in cultivated buckwheat. Using 2 self-compatible lines, 01AMU2 with brittle pedicels and Kyukei SC2 (KSC2) with non-brittle pedicels, produced by an interspecific cross between Fagopyrum esculentum cv Botansoba (non-brittle) andF. homotropicum (brittle), we investigated the inheritance of brittle pedicels. F₁ plants derived from crosses between Botansoba × 01AMU2 and Botansoba × KSC2 had brittle pedicels. The F₂ population derived from the cross between Botansoba × 01AMU2 showed segregation of brittle and non-brittle pedicels that fit the expected 3:1 ratio, suggesting that non-brittle pedicel in Botansoba is controlled by a single recessive gene (sht1). Another F₂ population, derived from the cross between Botansoba × KSC2, showed segregation of brittle and non-brittle pedicels that fit an expected ratio of 9:7,suggesting that non-brittle pedicel in KSC2is controlled by a different single recessive gene (sht2). Thus, brittle pedicel is achieved by 2 complementary genes Sht1 and Sht2. The sht1 locus is linked to the S locus with a recombination frequency of5.46±1.18 (%). We investigated whether common buckwheat has the allele sht2by crossing 6 common buckwheat lines withKSC2. An analysis of the preliminary data showed that some of the F₁ had brittle pedicels and others had non-brittle pedicels, suggesting that some common buckwheat lines possess both the allelesSht2 and sht2. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Genetics of colour traits in common vetch (Vicia sativa L.)

Five parents of common vetch (Vicia sativa L.) having orange/beige cotyledon colour, brown/white testa colour, purple/green seedling colour and purple/white flower colour were crossed as a full diallele set. The inheritance patterns of cotyledon, testa or seed coat colour, flower and seedling colour, were studied by analyzing their F₁, F₂, BC₁ and BC₂ generations. The segregation pattern in F₂, BC₁ and BC₂, showed that cotyledon colour was governed by a single gene with incomplete dominance and it is proposed that cotyledon colour is controlled by two allelic genes, which have been designated Ct₁ and Ct₂. Testa colour was governed by a single gene with the brown allele dominant and the recessive allele white. This gene has been given the symbol H. Two complementary genes governed both flower and seedling colours. These flower and seedling colour genes are pleiotropic and the two genes have been given the symbols S and F.     

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.     

Fertility improvement in tetraploid pearl millet

Summary Autotetraploid (2n=4x=28) pearl millet inbred lines, Tift 23BE and Tift 239DB, have been developed for use in crosses with other polyploid Pennisetum species. Each line set less than 1% and 3% selfed and open-pollinated seed, respectively. Seed germination was usually less than 17%. First generation (F₁) hybrids between the two inbreds set up to 61% seed while up to 40% of the seed from hybrids germinated.Seed weight per inflorescence for two planting dates averaged 0.32 g and 3.06 g for the two inbreds and the second generation F₂ progeny, respectively. One hundred seed weight was also significantly higher (0.78 g vs 0.48 g) in the F₂ progeny, probably due better endosperm development. Chromosome behavior and pollen stainability were similar in the inbred parents and hybrids. However, the hybrids shed more pollen than the inbred parents.Heterosis was evident in the F₁ hybrids and F₂ progeny which showed significant increases in plant height, leaf length, leaf width, and inflorescence length in addition to fertility improvement.     

RAPD and SCAR markers for resistance to acochyta blight in lentil

Resistance to ascochyta blight of lentil (Lens culinaris Medikus),caused by the fungus Ascochyta lentis, is determined by a single recessive gene, ral ₂, in the lentil cultivar Indian head. Sixty F₂ individuals from a cross between Eston (susceptible) and Indian head (resistant) lentil were analyzed for the presence of random amplified polymorphic DNA (RAPD) markers linked to the ral ₂gene, using bulked segregant analysis (BSA). Out of 800 decanucleotide primers screened, two produced polymorphic markers that co-segregated with the resistance locus. These two RAPD markers, UBC227₁₂₉₀and OPD-10₈₇₀, flanked and were linked in repulsion phase to the gene ral ₂ at 12 cm and 16 cm, respectively. The RAPD fragments were converted to SCAR markers. The SCAR marker developed from UBC227₁₂₉₀ could not detect any polymorphism between the two parents or in the F₂. The SCAR marker developed from OPD-10₈₇₀ retained its polymorphism. The polymorphic RAPD marker UBC227₁₂₉₀ and the SCAR marker developed from OPD-10₈₇₀ can be used together in a marker assisted selection program for ascochyta blight resistance in lentil. This revised version was published online in July 2006 with corrections to the Cover Date.