The genetics of hard seed coat in the genus Lens

Summary Seeds of the cultivated lentil are capable of germinating shortly after maturation. The seed dormancy of wild lentil species is due to a hard seed coat. In crosses between the cultivated species L. culinaris and its wild progenitor L. orientalis the hard seed coat of the wild species was controlled by a single recessive gene in homozygous condition. In a cross between the wild species L. ervoides and L. culinaris the hard seed coat of L. ervoides was controlled by a single dominant gene. The significance of the genetics of seed coat hardness in the domestication of lentil is briefly discussed.     

Inheritance of ‘domestication’ traits in bambara groundnut (Vigna subterranea (L.) Verdc.)

Controlled crosses in bambara groundnut were attempted between a range of thirty-six bambara groundnut landraces (thirty domesticated (V. subterranea var. subterranea) and six wild (V. subterranea var. spontanea)). Ten F₁ seed were produced. Of these, eight germinated producing F₂ populations. On seed set, four populations could be unambiguously confirmed as true crosses by F₃ seed coat colour. A single F₂ population, derived from a domesticated landrace from Botswana (DipC; female parent) crossed with a wild accession collected in Cameroon (VSSP11; male parent) was used to study a range of agronomic and domestication traits. These included; days to emergence, days to flowering, internode (fourth) length at harvest, number of stems per plant, leaf area, Specific Leaf Area (SLA), Carbon Isotope Discrimination (CID), 100 seed weight, testa colour and eye pattern around the hilum. On the basis of variation for internode length and stems per plant, 14 small F₃ families were selected and grown under field conditions to further investigate the genetic basis of the ‘spreading’ versus ‘bunched’ plant character, a major difference between wild and cultivated bambara groundnut. Results presented suggest that traits including leaf area, SLA, CID and 100 seed weight are controlled by several genes. In contrast, the variation for traits such as internode length, stems per plant, days to emergence and seed eye pattern around the hilum are likely to be under largely monogenic control. The results of this work are discussed in relation to the domestication of bambara groundnut.     

Evidence of a partial reproductive barrier between wild and cultivated pearl millets (Pennisetum glaucum)

Summary The occurrence of seed malformation in association with reduced thousand grain weight and germination ability has been observed in crosses between cultivated female plants and wild male plants. A survey of 16 cultivated accessions (P. glaucum subsp. glaucum) and 11 wild accessions (P. glaucum subsp. monodii) ranging over the whole species diversity showed this postzygotic incompatibility was general, but its intensity varied greatly with the cultivated female accession used and very little with the wild male parent origin. About 15% of the 123 cultivated x wild crosses observed gave normal seeds. Seed malformation has never been observed in crosses between cultivated accessions and appeared independent of genetic distances between the parents. The reciprocal crosses between wild female plants and cultivated male plants gave normal-looking seeds with good germination but consistently reduced thousand grain weight. Both seed malformation and seed small size are an expression of a genetic imbalance. These slight reproductive barriers seem to have been built during the domestication process.Abbreviation ICRISAT International Crop Research Institute for the Semi-Arid Tropics     

大豆结荚习性、荚色和种皮色相关野生片段分析

结荚习性、荚色和种皮色是大豆的重要形态性状,与进化密切相关。利用由151个家系组成的野生大豆(Glycinne soja Sieb et Zucc.)染色体片段代换系(CSSL)群体(SojaCSSLP1),通过不同表型CSSL组间比对,分别检测到与结荚习性、荚色和种皮色相关的1个、3个和2个野生片段(基因)。其中,5个野生片段(基因)分别与前人在栽培豆中检测到的无限结荚Dt₁、荚色L₁、荚色L₂、绿种皮G和黑种皮i基因相对应,说明野生大豆与栽培大豆间、栽培大豆与栽培大豆间在这些片段上均存在等位变异分化,是与大豆进化相关的基因/片段。另一个与荚色相关的Satt273野生片段能使大豆表现黑荚,可能是本研究的新发现,但还需进一步验证。     

Independent selection for seed free tryptophan content and vernalization response in chickpea domestication

Chickpea shows a distinct domestication trajectory vis‐a‐vis pod dehiscence and growth cycle mediated by vernalization insensitivity compared with its companion Near Eastern legumes. Our objectives were: (i) to map the quantitative trait loci (QTLs) associated with vernalization response and seed free tryptophan in domesticated × wild chickpea progeny and (ii) estimate the genetic correlation between vernalization response and free tryptophan content. A domesticated × wild chickpea cross was used to document phenotypic segregation in both traits and to construct a skeletal genetic map for QTL detection. A number of vernalization response and seed free tryptophan content QTLs were documented in both F₂ and F₃ generations. No significant genetic correlation between these two traits was observed. Epistatic relationship between two free tryptophan loci was documented. It is evident that selection for high seed tryptophan is easier to accomplish relative to selection for vernalization insensitivity. This suggests that the two traits were selected independently in antiquity, thereby corroborating earlier claims for conscious selection processes associated with chickpea domestication.     

Phaseolin Diversity in the Tepary Bean, Phaseolus acutifolius A. Gray

The tepary bean (Phaseolus acutifolius A. Gray) is grown mainly in and regions of Mexico and the southwestern U.S. as a subsistence crop by small farmers. It is also a store of genetic variability for traits such as disease and pest resistance and stress tolerance to improve the common bean (P. vulgaris L.). To determine geographic patterns of variation and the influence of domestication on genetic variability, the genetic diversity of phaseolin, the major seed storage protein, was characterized among 55 wild and 8 cultivated teparies using polyacrylamide gel electrophoresis. Fifteen electrophoretic phaseolin patterns were identified among wild forms, whereas only one pattern was exhibited by cultivars. This result suggests a single domestication in this species leading to a strong reduction in diversity. An additional finding is the divergence m phaseolin types between populations east and west of the Sierra Madre Occidental mountains.     

Cross compatibility between Abelmoschus esculentus and A. moschatus

Interspecific cross compatibility between cultivated and wild okra (Abelmoschus esculentus and A. moschatus) and pollen tube growth behaviour in the crosses among a local cultivar of A. esculentus, A. moschatus and their F₁s were studied. Fruit set was observed in all the crosses except one and seed setting was absent in two of the crosses which set fruit. All seed produced were shrivelled but F₁ plants were obtained from two crosses where cultivated okra was used as the seed parent. The F₁ plants were perennial in nature with very low pollen viability and seed set. A high percentage of pollen germination and profuse pollen tube penetration in the style were observed in the cross A. esculentus × A. moschatus but low pollen tube penetration with abnormal pollen tubes was observed in the reciprocal cross. The number of pollen tubes was very low but they appeared to be normal in the backcross A. esculentus × F₁, but were generally abnormal in the reciprocal cross. Both pre- and postzygotic barriers seemed to occur in crosses between the two species. The present studies indicate that these barriers can be overcome and desirable characters from A. moschatus transferred to cultivated okra using conventional hybridisation techniques. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effect of quantitative trait loci for seed shattering on abscission layer formation in Asian wild rice Oryza rufipogon

Asian cultivated rice Oryza sativa L. was domesticated from its wild ancestor, O. rufipogon. During domestication, the cultivated rice lost its seed-shattering behaviour. Previous studies have shown that two major quantitative trait loci (QTLs; qSH1 and sh4) are responsible for the seed-shattering degree. Here, we produced introgression lines carrying non-functional alleles from O. sativa ‘Nipponbare’ at the two major QTLs in the genetic background of wild rice O. rufipogon W630, and examined the effects of the two QTLs on seed shattering and abscission layer formation. The introgression lines, with Nipponbare alleles at either or both loci, showed complete or partial abscission layer formation, respectively, indicating that other unknown loci might be involved in enhancing seed shattering in wild rice. We detected a single QTL named qSH3 regulating seed-shattering degree using an F₂ population between Nipponbare and the introgression line carrying Nipponbare alleles at the two QTLs. Although we generated an introgression line for qSH3 alone, no effects on seed shattering were observed. However, a significant effect on seed-shattering degree was observed for the introgression line carrying Nipponbare alleles at qSH3 and the two QTLs, suggesting an important role of qSH3 on seed shattering in coordination with the two QTLs.     

Overcoming interspecific barriers in Sorghum and molecular characterization of the derivatives

Crop improvement in sorghum has progressed extensively availing the variation available in the primary gene pool of S. bicolor. Crossing further, the yield plateau can be accomplished by bringing in the genes from tertiary gene pool into the cultivated gene pool. Earlier efforts for wide hybridization in sorghum with tertiary gene pool were not successful due to inhibition of wild pollen germination on the stigma of cultivated species. In the current study, elite lines of cultivated Sorghum species were pollinated with wild species using different floral treatments. The results showed that spray with the synthetic auxin, 2,4-dichlorophenoxyacetic acid, has promoted the seed set. Overall, 76% crosses showed seed set, but many seeds were underdeveloped and deformed. F₁ progeny showed traits of both cultivated and wild parents. Hybrids were confirmed through PCR analysis. Many (80%) F₁ plants were sterile, and 20% of F₁s were advanced to F₂ generation. Wide variation was observed in F₂ progeny, and the phenotype selections were made and are being evaluated in the field.     

Reproductive behaviour and broomrape resistance in interspecific hybrids of sunflower

Interspecific hybrids and backcross generations between the wild perennial species Helianthus resinosus, Helianthus paucifiorus, Helianthus laevigatus, Helianthus nuttallii ssp. nuttallii T. & G. and Helianthus giganteus, resistant to broomrape (Orobanche cernua) and susceptible inbred lines were obtained to study crossability to cultivated sunflower and the transmission and expression of resistance to this parasitic weed. Conventional crosses with all the species tested were successful except for the crosses with diploid H. giganteus, for which embryo rescue techniques were needed to overcome hybrid incompatibility. Pollen viability and seed set were highest for F₁ hybrids with hexaploid species and lowest for those with the diploid H. giganteus. We evaluated F₁, BC₁F₁, some BC₂F₁ plants and the wild and cultivated parents. The wild species and interspecific hybrids were resistant to broomrape infection except for H. nuttallii, which showed segregation, indicating that the resistance is dominant. The crossability and resistance of F₁, and back-cross generations of species with different ploidy levels indicate that the transfer of broomrape resistance to cultivated sunflower is feasible.     

Variation in progenies of an Arachis hypogaea x diploid wild species hybrid

Summary Derivatives of a cross between cultivated peanuts, Arachis hypogaea L. (2n=40), and the wild species collection GKP 10017 (2n=20) were compared morphologically, for leafspot resistance and for yield. The objective of the study was to determine the effects of wild species germplasm on the A. hypogaea genome. The sterile F₁ hybrid which resulted from crossing the two species was treated with colchicine to restore fertility at the 6x ploidy level. The resulting hexaploid was cytologically unstable and progeny lost chromosomes until stability was regained at the 2n=40 chromosome level. Forty-seven characters were used to analyze the variation among plants in the tetraploid interspecific hybrid population. The plants were compared to four cultivated lines plus GKP 10017. Many hybrids were intermediate to the two parents in morphology. Individual traits such as growth habit, pod and seed size, elongation of the constricted area between pods, nodulation and leaflet size were altered by the presence of GKP 10017 germplasm in many of the hybrid plants. Cercospora arachidicola Hori and Cercosporidium personatum (Berk. & Curt.) Deighton resistances were evaluated for all plants. Several hybrids had few lesions due to either leafspot pathogen. In addition, 24 largeseeded interspecific hybrid selections were compared to the cultivated variety NC 5 for yield. Five selections were superior to both parents at p=0.01. Morphology, disease resistance and yields appeared to be greatly influenced by the wild species GKP 10017 germplasm in plants of the interspecific hybrid population. The potentials of using wild species for improvement of the cultivated peanut are discussed.Paper number 5948 of the journal series of the North Carolina Agricultural Research Service, Raleigh, NC 27650. The investigation was supported in part by ICRISAT and SEA-CR grant no. 701-15-51.     

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

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

Identification of QTLs for domestication-related and agronomic traits in an Oryza sativa×O. rufipogon BC1F7 population

Asian cultivated rice was domesticated from the wild rice, Oryza rufipogon and throughout the domestication process, a wide range of morphological and physiological changes altered the ancestral form. This study was conducted to identify the genetic basis of changes associated with the domestication process. An recombinant Inbred line (RIL) population consisting of 120 lines was developed from a cross between the Juponica cultivar.‘Hwayeongbyeo’and a presumed wild progenitor. O. rufipogon Griff. Acc.01944. The population was genotyped with 124 simple sequence length repeat (SSR) markers, providing an average interval size of 15 cM, and also evaluated for 20 traits related to domestication and agricultural performance. A total of 63 quantitative trait locus (QTLs) and one locus associated with qualitative variation for pericarp coloration were identified using single point and composite interval analysis. The number of QTLs per trait ranged from one to seven. Phenotypic variation associated with each QTL ranged from 3.7 to 40.4%. with an average of 15.3%. The results indicated that most domestication‐related traits clustered in chromosomal blocks, and the positions of many of these clusters were consistent with those reported in previous studies and with skewed segregation ratios in these BC₁,F₇ RILs. For 13 (20.6%) of the QTLs identified in this study. the O. rufipogan ‐derived allele contributed a desirable agronomic effect despite the overall undesirable characteristics of the wild phenotype. Favourable alleles from O. rufipogan were detected for panicle length, spikelets per panicle, days to heading and leaf discoloration associated with cold stress. When compared with previous studies involving interspecific crosses, it can be concluded that O. rufipogon is useful as a source of valuable alleles for rice improvement and that many of the introgressed regions contain genes that have a favourable impact on phenotype in different genetic backgrounds and different environments.     

Production and characterization of hybrids between Cajanus cajan × C. reticulatus var. grandifolius

Cajanus reticulatus var. grandifolius, endemic to Australia and a wild relative of the cultivated species, C. cajan, was successfully crossed with the latter as the female parent. The major wild species characters such as persistent stipules, long pod hairs, pod shattering, brown seeds with grey speckles, and presence of seed strophiole were dominant in the hybrid. For growth and branching habit, and leaflet, flower, pod, and seed size, the hybrid was intermediate between the parents. The meiotic cells of the hybrid were found to have quadrivalents, trivalents, univalents,and showed reduced chromosome pairing as revealed by the increased number of rod bivalents per cell at metaphase-I, and stickiness and precocious movement of chromosomes to poles in the second division. In comparison to the parents, the hybrid had fewer pods and seeds. However,these anomalies in the interspecific hybrid are not significant enough to preclude the gene transfer from C. reticulatus to the cultivated species through a sexual route. This revised version was published online in August 2006 with corrections to the Cover Date.     

Broadening the genetic base of lentil cultivars through inter‐sub‐specific and interspecific crosses of Lens taxa

Wild Lens taxa are invaluable sources of useful traits for broadening genetic base of cultivated lentil. Nine inter‐sub‐specific and interspecific crosses were made successfully between cultivated (Lens culinaris ssp. culinaris) and wild lentils (L. culinaris ssp. orientalis, odemensis, lamottei and ervoides). The effect of species groups, day length and temperature on crossability in lentils was evident under normal winter sowing in New Delhi and in summer Himalayan nursery at Sangla in Himachal Pradesh, India, although pollen fertility assessed in all the cross‐combinations showed no significant variation. True hybridity of nine inter‐sub‐specific and interspecific crosses was confirmed through morphological and molecular (ISSR) markers, in which three of 120 primers could confirm the hybridity of all the crosses. All cross‐combinations were also studied for important quantitative traits related to yield. The range, mean and coefficient of variation were estimated in parental lines, F₁ and F₂ generations to determine the extent of variability generated in cultivated lentils through the introgression of genes from wild L. taxa. A high level of heterosis was observed in F₁ crosses for important traits studied. Substantially higher variations for seed yield and its attributing traits were exhibited in F₂ generations indicating transgressive segregation. The results of the present investigation revealed that wild L. taxa can be successfully exploited for lentil improvement programmes, and the variations generated could be easily utilized for broadening the genetic base of cultivated lentil gene pool for improving the yield as well as wider adaptation.     

Introgression of Cajanus platycarpus genome into cultivated pigeonpea, C. cajan

Summary Cajanus platycarpus, an incompatible wild species from the tertiary gene pool of pigeonpea (C. cajan (L.) Millspaugh), has many desirable characteristics for the improvement of cultivated varieties. To necessitate such transfers, embryo rescue techniques were used to obtain F₁ hybrids. The F₁ hybrids were treated with colchicine to obtain tetraploid hybrids, that were selfed to obtain F₂, F₃ and F₄ progenies. All of the hybrids and subsequent progenies had an intermediate morphology between the two parents. Backcrossing of the tetraploid hybrids with cultivated pigeonpea was not possible given embryo abortion, with smaller aborted embryos than those obtained in the F₀ parental cross.As a route of introgression, diploid F₁ hybrids were backcrossed with cultivated pigeonpea and BC₁ progeny obtained by in vitro culture of aborting embryos. BC₂ plants were obtained by normal, mature seed germination. Although embryo rescue techniques had to be used to obtain F₁ and BC₁ plants, it was possible to produce BC₂ and subsequent generations through direct mature seed. Every backcross to cultivated pigeonpea increased pollen fertility and the formation of mature seeds.Special project assistant till December, 2003.     

Genetic studies in four interspecific hybrids of the genus oryza

The paper reports the results of genetic studies made in four interspecific hybrids of the genus Oryza, and the data are employed to discuss the inter-relationships between the different species. The characters of wild parents were found to be dominant in the F₁ hybrids.In O. sativa (variety Fukoku) × O. perennis subsp. balunga, the inheritance of the eight characters studied was found to be of the same nature as that of complementary, modifying, duplicating and polygenic types involving two or three genes. In O. perennis (Cuban type) × O. perennis subsp. balunga, the interesting feature was the occurrence of recessive characters (typical of cultivated rices) in the F₂ population, even though the two parents carried the dominant wild characters. This should be possible only by the operation of duplicate genes and such a mode of inheritance was observed for three characters. No definite Mendelian ratios could be got for the remaining five characters studied. The appearance of characters of cultivated rices has been suggested to point to the ability of O. perennis to give rise to O. sativa through natural crossing and selection. In O. glaberrima × O. perennis subsp. balunga, the ten characters studied segregated in monogenic or digenic fashion. When digenic, these characters exhibited complementary or inhibitory factor interactions. In O. glaberrima × O. breviligulata, the simple mode of inheritance in the F₂ generation has been suggested to indicate the close relationship of the two species.Some of the general features of the hybrids and segregating populations studied were the vigorous and fertile F₁s followed by the appearance of weak and unthrifty plants in the F₂ generation and the tendency for the parental characters to inherit together. The appearance of plants looking similar to O. sativa var. fatua in the F₂ populations of O. sativa × O. perennis, and of O. stapfii in the F₂s of O. glaberrima × O. perennis has been supposed to provide further evidence on the hybrid origin of these two forms.     

Molecular mapping of QTLs for domestication-related traits in cowpea (V. unguiculata (L.) Walp.)

Understanding the genetic basis underlying domestication-related traits (DRTs) of cowpea (Vigna unguiculata (L.) Walp.) is important since the genome has experienced divergent domestication and in addition it is also useful to utilize the wild germplasm efficiently for improving different traits of the cultivated cowpea. Quantitative trait loci (QTLs) for DRTs were identified in a population of 159 F₇ recombinant inbred lines derived from a cross between a domesticated cowpea (V. unguiculata (L.) Walp.) variety, 524B, and a wild accession, 219. Using the constructed linkage map, QTLs for 10 DRTs were analysed and mapped. QTLs for seed, pod and flower related traits were detected. Subsequently, QTL for ovule number was also identified. To our knowledge, this is the first time a QTL for this trait has been observed. QTLs for DRTs show co-localization on three linkage groups and pleiotropy or close linkage of genes for the traits is suggested in these chromosome regions. The information gained in this study can be used for marker-assisted selection of domestication-related QTLs in cowpea and enhance understanding of domestication in the genus Vigna.     

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

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

Chromosomal Stability in the Backcross Progenies of Pentaploid Hybrids between Avena sativa L. and A. maroccana Gdgr.

In a backcrossing programme to transfer desirable characters from wild Avena maroccana Gdgr. to cultivated oats, A. sativa L., meiotically stable plants in BC₁F₃ and BC₂F₂ progenies were isolated. The recovery of stable genotypes with 2n = 6×= 42 chromosomes indicated that two backcrosses are enough for such a programme. The cytological observations in various backcross generations are presented and discussed.