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.     

Studies on the inheritance of five qualitative characteristics in clusterbean (Cyamopsis tetragonoloba (L.) Taub)

Summary The inheritance of five characteristics namely branching behaviour, clustering pattern, growth habit, leaf size and hairiness was studied in clusterbean. All the traits were controlled by single pairs of genes except branching behaviour which exhibited digenic inheritance. Complementary gene effects were involved in the expression of branching in the F₁ and F₂ generations; the F₂ showing a ratio of 9:7 for branched to unbranched genotypes in the cross FS 277 × NC 70 where both the parents were single shooted. The alleles governing branching, discontinuous clustering, indeterminate growth habit, small leaf size and hairiness were dominant over the alleles controlling non-branching, continuous clustering, determinate growth habit, broad leaf and glabrous leaf and stem.     

Yield stability in common bean (Phaseolus vulgaris L.) genotypes

An experiment was undertaken to determine the stability of seed yield in 21 common bean genotypes representing three growth habits. Seven genotypes in each growth habit (determinate bush, indeterminate bush and indeterminate prostrate) were evaluated in replicated trials at three locations for three years under rain fed conditions in Ethiopia. A combined analysis of variance, stability statistics and rank correlations among stability statistics and yield-stability statistic were determined. The genotypes differed significantly for seed yield and genotype × environment (year by location) interaction (GE). The different stability statistics namely Type1, Type 2 and Type 3 measured the different aspects of stability. This was substantiated by rank correlation coefficient. There were strong rank correlations among S_i ²d, W_i ²,σ_i ² and S_i ², where as there was weak correlation between b_iand R_i ², S_i ²d, W_i ², σ_i ² and S_i ². R² was significantly and negatively correlated with W_i ², σ_i ² and S_i ². σ_i ² is significantly correlated with W_i ².Yield is significantly correlated with b_i and R_i ².None of the statistics per se was useful for selecting high yielding and stable genotypes except the YS(yield-stability statistic). Most of the high yielding genotypes were relatively stable. Of the 21 genotypes, only 11genotypes were selected for their high yielding and stable performance. Genotypes with growth habit III and I (in determinate prostrate and determinate bush) were generally more stable than in determinate bush. This revised version was published online in July 2006 with corrections to the Cover Date.     

Segregation Pattern of Decumbent, Climbing and Bushy Growth Habits in Vigna unguiculata (L.) Walp.

F₁ and F₂ data from crosses of cowpea accessions with three different growth forms showed that the inheritance of growth habit is governed by two allelic pairs, AA and BB. The genetic ratio obtained implicated a modifier gene interaction in which the genotype A-B- has the decumbent growth habit; the genotype aaB- has the climbing and aabb has the bushy habit. While the decumbent growth habit was dominant and epistatic over the climbing habit, the decumbent and the climbing growth forms were each monogenically dominant over the bushy habit.     

Morphology and genetics of a new found determinate genotype in chickpea

Chickpea (Cicer arietinum L.) is an indeterminate plant and produce excessive vegetative growth that acts as a competitive sink for developing pods resulting in reduced fruit set whenever soil moisture and temperature conditions are favorable. Determinate genotype was so far not available in the world chickpea germplasm. Determinate chickpea is needed to change its plant architecture in order to achieve a breakthrough in its productivity and stabilize yields in cool and long-season sub-tropical environments of semi-arid tropics. A true breeding determinate chickpea genotype was developed for the first time in the breeding programme. The objectives were to study: (i) the new found determinate genotype BGD 9971 for important plant characteristics and (ii) the genetics of determinate growth habit in it. Determinate genotype was bushy, compact and dwarf in its morphology; the stem growth terminated by a flower and produced 1–4 seeds per pod. The segregation patterns in the F₂ and F₃ of the two crosses (BGD 72 × BGD 9971 and BGD 128 × BGD 9971) involving indeterminate and determinate parents have shown that the determinate growth habit in BGD 9971 was governed by two recessive genes. The genes for determinacy in BGD 9971 were designated as dt1 and dt2. The homozygous recessive for both alleles (dt1dt1dt2dt2) produced a determinate phenotype. The utilization of genes identified for determinacy in the newly developed BGD 9971 has the major impact on chickpea breeding for better adaptation to cool climate, high fertility and irrigated environments.     

The genetics of naked oats (Avena nuda L.)

Summary The inheritance of the character complex of naked grain and multiflorous spikelets was studied in a diallel set of crosses comprising five varieties of naked oats (Avena nuda)-As 78, Manu, Caesar, Nuprime and AJ86/2/1-and one variety of husked oats (A. sativa), Maris Oberon. In the F₁ generation the distribution of multiflorous spikelets was in all cases similar to that of the mid-parent. Crosses between the varieties of A. nuda produced only naked grain on plants in the F₁ and F₂ generations, indicating that nakedness in the varieties studied was determined by the same loci.The three-gene model proposed by Moule (1972) for the determination of A. nuda characters was inadequate to account for the observed F₂ segregation in naked x husked crosses. An extension of this model is proposed to include a third modifying gene, N₃, which in the homozygous dominant condition produces the husked phenotype when the principal gene, N, is heterozygous. The model assumes complex epistatic relationships between the three modifying genes N₁, N₂ and N₃. Published information and further experimental data suggest that the genotype NN--N₃N₃ is uncommon. The expression of the genes determining nakedness was greatly influenced by the environment.     

Validation of the two-gene epistatic model for vernalization response in a winter × spring barley cross

A two gene epistatic model in which a dominant “winter growth habit” allele at Vrn-H2 encodes a repressor with a corresponding binding site in a recessive vrn-H1 allele explains the vernalization response phenotypes in an array of barley germplasm. In order to validate the model genetically, we developed an F ₂ population (and F ₂-derived F ₃ families) from the cross of Hardy (winter) × Jubilant (spring). Using gene-specific primers, we determined the Vrn-H1 and Vrn-H2 allele architecture of each F ₂ plant and we measured the growth habit phenotype of each F ₂ plant via phenotyping of its F ₃ progeny under controlled environment conditions. We used a set of treatments involving plus/minus vernalization under long photoperiod and vernalization under short photoperiod. Alleles at the two loci showed expected patterns of segregation and independent assortment. Under long day conditions, the two Vrn genes were the primary determinants of heading date, regardless of the vernalization treatment. Under short photoperiod, the effects of these loci were not significant. There was incomplete dominance at Vrn-H1: heterozygotes were significantly later to head than Vrn-H1Vrn-H1 genotypes. Vrn-H2 genotypes were also significantly later to head, even when plants were vernalized. These results validate the two-gene epistatic model for vernalization response under long-day conditions. The results under short photoperiod, and the variance in flowering with vernalization, confirm that while the two Vrn genes are the primary determinants of vernalization response, they are part of a larger interactome that determines the timing of the vegetative to reproductive transition.     

Interrelationships among height,lodging and yield in determinate and indeterminate soybeans

Summary Seventy-three determinate and 93 indeterminate random F₉ lines from a cross between determinate and indeterminate soybean [Glycine max (L.) Merr.] parents were evaluated for seed yield, mature plant height, and lodging. Regression analyses indicated that among the determinate lines, seed yield increased by 350 kg/ha and lodging score increased by 0.008 for every 10 cm increase in plant height. Among indeterminate lines seed yield increased by 112 kg/ha and lodging score by 0.3 for every 10 cm increase in plant height. The highest yielding indeterminate lines were tall and very susceptible to lodging. The tall determinate lines had the best combination of high seed yield and excellent lodging resistance.Cooperative Investigations of the Science and Education Administration, U.S. Department of Agriculture and the Purdue University Agricultural Experiment Station.     

Identification of alleles at two loci controlling resistance to Phomopsis stem blight in narrow-leafed lupin (Lupinus angustifolius L.)

The genetics of resistance to Phomopsis stem blight caused by Diaporthe toxica Will., Highet, Gams & Sivasith. in narrow-leafed lupin (Lupinus angustifolius L.) was studied in crosses between resistant cv. Merrit, very resistant breeding line 75A:258 and susceptible cv. Unicrop. A non-destructive glasshouse infection test was developed to assess resistance in the F₁, F₂, selected F₂-derived F₃ (F_2:3) families, and in selfed parent plants. The F₁ of Unicrop × 75A:258 (and reciprocal cross) was very resistant, and the F₂ segregated in a ratio of 3:1 (resistant: susceptible), which suggested the presence of a single dominant allele for resistance in 75A:258. In Merrit × Unicrop (and reciprocal), the F₁ was moderately resistant, and the F₂ segregated in a ratio of 3:1 (resistant: susceptible). Thus Merrit appeared to carry an incompletely dominant resistance allele for resistance. The F₁ of Merrit × 75A:258 (and reciprocal) was very resistant and the F₂ segregated in a ratio of 15:1 (resistant: susceptible), which supported the existence of independently segregating resistance alleles for resistance in 75A:258 and Merrit. Alleles at loci for early flowering (Ku) and speckled seeds (for which we propose the symbol Spk) segregated normally and independently of the resistance alleles. Resistant F₂ plants gave rise to uniformly resistant or segregating F_2:3 families, whereas susceptible F₂ plants gave rise only to susceptible F_2:3 families. However, the variation in resistance in the F₂ and some F_2:3 families of crosses involving 75A:258, from moderately to extremely resistant, was greater than that expected by chance or environmental variation. We propose the symbols Phr1 to describe the dominant resistance allele in 75A:258, and Phr2 for the incompletely dominant resistance allele in Merrit. Phr1 appears to be epistatic to Phr2, and expression of Phr1 may be altered by independently segregating modifier allele(s). This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of seed size in cowpea (Vigna unguiculata (L.) Walp.)

Summary The inheritance of seed weight in cowpea was examined in a field planting of the parents, reciprocal F₁s, F₂s and backcrosses to both parents of a cross between TVu 1977-OD (small seeded) and ACC 70002 (large).Seed weight was inherited quantitatively and small seed was partially domiminant to large seed size. Gene action was predominantly additive but dominance and additive × additive epistatic effects were also significant.Broad and narrow sense heritabilities were 85.1±5.3% and 75.4±18.6% respectively. The minimum number of loci involved in the inheritance of seed size was eight, and each gene pair contributed up to 1.02 g increase to seed weight. The estimate of genetic advance from F₂ to F₃ generations with 5% selection intensity was 3.58 g.International Institute of Tropical Agriculture, Semi-Arid Food Grain Research and Development Project/National Cowpea Improvement Program, B.P. 1783, Ouagadougou, Upper Volta.Department of Primary Industries, Hermitage Research Station, Warwick 4370, Queensland, Australia.International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), ICRISAT Center, Patancheru, P.O. 502 324, A.P., India.     

Possibility of genetic improvement of pigeonpea (Cajanus cajan (L.)Millsp.) utilizing wild gene sources

Summary Various wild relatives of pigeonpea,Cajanus cajan, namely some species ofAtylosia andRhynchosia, possess desirable characteristics that could be utilized for effecting genetic improvement of this crop. In total 73 cross combinations among two cultivars ofC. cajan and one accession each of eightAtylosia species and one ofRhynchosia were attempted. Twelve hybrids were obtained. Seven of these were analysed for F₁ fertility and their utility for agronomic improvement of theC. cajan. Fertility behaviour of the different F₁ hybrids varied and indicated that potential of gene transfer between the two genera,Atylosia andCajanus, was as good as within the genusAtylosia. From F₂ and F₃ families ofC. cajan × A. scarabaeoides andC. cajan × A. albicans, plants were selected with greater physiological efficiency and agronomic superiority. The prospects of transferring pod borer resistance and higher seed protein content from someAtylosia species to pigeonpea are discussed.     

Inheritance of resistance to angular leaf spot (Isariopsis griseola Sacc.) in french bean (Phaseolus vulgaris L.)

Summary Angular leaf spot (Isariopsis griseola Sacc.) is a serious disease of French bean in the hills of India and 40 to 70 per cent of the green pods are damaged and rendered unmarketable. Crosses were made between PLB 257, (Phaseolus coccineus L.), a red flowering pole tope, resistant to angular leaf spot, and Contender (Phaseolus vulgaris L.), a highly susceptible commercial cultivar. Studies of the F₁, F₂, and F₃ progenies indicated that PLB 257, carries a recessive gene imparting resistance to angular leaf spot.     

Chloroplast-DNA variation in the wild and cultivated olives (Olea europaea L.) of Morocco

The male sterile plants that segregated in a BC₅F₂ of `C. sericeus × C. cajan var. TT-5' population were maintained by sib mating. The male sterile plants were crossed with ICPL-85012.Approximately 50% of the F₁ plants were sterile. F₂ plants derived from the fertile F₁ plants did not segregate for male sterility. The reciprocal hybrid i.e. ICPL-85012 × Fertile derivatives from C. sericeus × TT-5, did not express male sterility. However, among the 12 F₂ plant to row progenies, two segregated 25% male sterile plants and remaining 10 did not segregate. The segregation pattern in subsequent progenies revealed that the sterility was under control of a single recessive allele. Studies on the backcross and their BC₁F₂ and BC₁F₃progenies revealed another sterility gene which was found to be dominant in inheritance. This paper shows that what was thought to be cytoplasmic male sterility from C. sericeus cytoplasm is actually a single dominant gene possibly acting in concert with a single recessive gene to mimic cytoplasmic male sterility. This revised version was published online in July 2006 with corrections to the Cover Date.     

Involvement of higher order interactions addressing complex polygenetically controlled inheritance of downy mildew [Sclerospora graminicola (Sacc.) Schrot] resistance in pearl millet [Pennisetum glaucum (L.) R.Br.]

Inheritance of downy mildew [Sclerospora graminicola (Sacc.) Schrot]resistance was studied using generation mean analysis in pearl millet [Pennisetum glaucum (L.) R.Br.]. Eleven basic generations, namely, P₁, P₂, F₁, F₂, B₁, B₂, B₁F₂, B₂F₂, L₁, L₂ and L₃ of three crosses involving six diverse lines for downy mildew incidence were evaluated under artificial epiphytotic conditions over two environments. The downy mildew incidence was best fitting for digenic, trigenic and tetragenic ratios when fitted into classical Mendelian ratios demonstrating involvement of two or more genes. Digenic and trigenic interaction models were adequate in the case of crosses I and III respectively, to account for the total variability in generation means. Unlike severity, comparative estimates of gene effects over two environments were mostly consistent in all crosses for prevalence. Most of the epistatic and major gene effects were found significant in all crosses for both the disease traits. Non-allelic interactions particularly at three-gene loci viz., w (additive × additive × additive) and y (additive × dominance × dominance) in cross II and all trigenic interactions in cross III were predominant. Duplicate dominance (cross I) and complementary epistasis (crosses II and III) were observed for both the traits revealing inconsistency of gene effects over crosses. The gd₁ (interaction of additive gene effect with e₁) and gh₁(interaction of dominant gene effect with e₁) were significant in crosses I and II, indicating interaction of additive and dominance gene effects with environments. Thus a breeding method that can mop up the resistant genes to form superior gene constellations interacting in a favorable manner against pathotype I would be more suitable to accelerate the pace of resistance improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Yield, yield components and plant architecture in the F3 generation of common bean (Phaseolus vulgaris L.) derived from a cross between the determinate cultivar ‘Prelude’ and an indeterminate landrace

Growth trials were conducted outdoors in the UK to determine the yield, yield components and plant architectural differences between determinate and indeterminate Phaseolus vulgaris bean plants. F3 lines derived from crosses between ‘Prelude’, a determinate cultivar and ‘V8’, an indeterminate landrace, were grown together with the parents and ‘Carioca’, an indeterminate landrace from Brazil. Data were recorded on flowering date, number of nodes on main stem at flowering, plant height at maturity, number of pods/plant, number of seeds/pod, number of seeds/plant, 100-seed weight, seed mass/plant, percentages of diseased and healthy seeds/plant. Determinate F3 lines had significantly lower (P < 0.05) seed mass/plant, fewer pods/plant, fewer seeds/pod, fewer seeds/plant, lower harvest index, shorter stems, earlier date of flowering, fewer nodes at flowering and fewer healthy seeds/plant than indeterminate F3 lines. However, determinate genotypes had significantly larger (P < 0.05) pods to accommodate their larger seeds compared with indeterminate lines. A similar situation was found when the yield and yield components of ‘V8’, ‘Prelude’ and ‘Carioca’ were evaluated; the highest seed mass/plant was given by ‘V8’ and the lowest by ‘Prelude’. Furthermore, some indeterminate genotypes with Type IV growth habit yielded significantly higher (P < 0.05) than the high-yielding parent, ‘V8’, indicating a positive influence on seed yield by crossing different genotypes.     

Inheritance of resistance to Fusarium wilt in Indian lentil germplasm (Lens culinaris Medik.)

Summary Three lentil genotypes resistant to Fusarium oxysporum f.sp. lentis viz. Pant L 234, JL 446 and LP 286 were crossed with two susceptible ones. The hybrid plants were all resistant in the eight crosses evaluated. Segregation pattern for wilt reaction in F₂, BC(P₁), BC(P₂) and F₃ generations in field and glasshouse conditions indicated that resistance to Fusarium wilt is under the control of two dominant duplicate genes in Pant L 234 and two independent dominant genes with complementary effects in JL 446 and LP 286. A third dominant gene complementary to the dominant genes in JL 446 and LP 286 is present in two susceptible lines. Allelic tests suggest the presence of five independently segregating genes for resistance. Duplicate dominant genes in Pant L 234 are non-allelic to two dominant genes with complementary effects in LP 286 and JL 446 and the third gene complementary to the two genes in JL 446 and LP 286 in susceptible lines JL 641 and L 9–12. Gene symbols among parental genotypes have been designated.     

Increasing biodiversity of irrigated rice in Africa by interspecific crossing of Oryza glaberrima (Steud.) × O. sativa indica (L.)

The African rice Oryza glaberrima, traditionally cultivated since more than 3.500 years, is of poor agronomic performance but resistant/tolerant to various stresses and diseases. The introduction of these characters into O. sativa cultivars is difficult since crossing barriers cause spikelet sterility in F1. Backcrossing can restore fertility and recently facilitated the development of fertile O. glaberrima × O. sativa ssp. japonica hybrid progenies for rain fed systems. With the objective to gain access to African rice germplasm for improvement of irrigated rice, crosses were performed with eighteen O. glaberrima and twenty O. sativa ssp. indica accessions. In total about one hundred F1-hybrid grains were obtained. The F1 plants were all completely sterile and backcrossing (BC) to O. sativa was performed in order to restore spikelet fertility. Monitoring of Tog5681 × IR64 hybrid progenies under field conditions revealed a broad genetic diversity within the BC₁ and BC₂ populations. Some BC₁ and BC₂ progeny plants outperformed the O. sativa parent, indicating that the heterozygocity level and complementary gene action after two backcrosses are still sufficient to positively influence plant vigor. Spikelet fertility of progenies was highly variable, but almost complete fertility was already observed within the BC₁F₂ population. High spikelet fertility was preserved in one out of two analyzed BC₁F₃ families and inmost of the BC₂F₃ families. The ability to restore spikelet fertility within few generations and the potential of the genetic diversity present in interspecific progenies facilitates the development of plant types specifically designed for the African irrigated and lowland environment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetics of stem elongation ability in rice (Oryza sativa L.)

Summary The genetics of stem elongation ability in rice was studied in parents, F₁, F₂ and backcross generations of six crosses. Segregation analysis indicated dominance for stem elongation ability. Estimation of genetic parameters under epistatic model indicated more than one locus control stem elongation ability and both additive and nonadditive gene effects were important. Epistatic effects were predominant over additive and dominance effects with an important role of duplicate type of epistasis. The occurrence of significant additive and additive x additive types of genetic variation and the moderately high broad sense heritability indicated the possibility of selection for an increased manifestation of stem elongation ability.     

Variation in pigment content of flour color in bread wheat (Triticum aestivum L.)

Summary Variation in pigment content of the flour of bread wheats (Triticum aestivum L.) was studied in the progenies of F₁ and F₂ of three crosses and their reciprocals. Reciprocal differences in pigment content were observed in the F₁ and F₂ means. Low pigment content was found to be partially dominant or over dominant in the crosses studied. There was evidence of substantial mid-parent F₁ heterosis in all crosses and betterparent F₁ heterosis in three crosses. In the F₂, heritability estimates were moderate to high. The F₂ frequency distributions were not normal. Estimation of effective factor pairs indicated the presence of one or two major gene pairs involved in the expression of pigment content in the flour. Action of modifiers was also assumed in one cross and its reciprocal. A factorial approach to metrical character suggested that the F₂ segregation ratios of low pigment content to high pigment content were 3:1, 15:1, 13:3 and 9:7 for the different crosses. Utilization of the findings in a wheat breeding program is briefly discussed.     

A major QTL associated with preharvest sprouting in rapeseed (<Emphasis Type="Italic">Brassica napus</Emphasis> L.)

Preharvest sprouting (PHS) is one of the most important factors affecting the cereal production worldwide, in regions characterized by rainfall and high humidity during harvest season. It is sometimes a problem in rapeseed (Brassica napus L.), especially in production of commercial F₁ hybrids. To detect quantitative trait loci (QTL) controlling PHS, a F₂ population consisting of 269 F_2:3 lines was created from the cross between a PHS-tolerant line (117AB) and a PHS-susceptible line (7,605). A linkage map was constructed using 35 Simple Sequence Repeat markers and 242 Amplified Fragment Length Polymorphism markers. PHS was measured as a percentage of sprouted seeds on the mother plant, 7 days after physiological maturity. Five putative QTLs for PHS were detected and located on LG2 (N11) and LG7 (N3), respectively. Phenotypic variance explained by each QTL ranged from 4.11 to 50.78% and the five putative QTLs explained about 75.63% of the total phenotypic variance. A major QTL was identified on LG2 (N11) flanked by P3C4180 and C6C13160, which explained 50.78% of the total phenotypic variance. Meanwhile, we detected four significant epistatic interactions with a total contribution of 17.16% of the total phenotypic variance.