Comparison of intra- and inter-pool crosses in faba beans (Vicia faba L.). I. Hybrid performance and heterosis in Mediterranean and German environments

Two recently detected, promising CMS systems in faba bean have opened up new possibilities in hybrid breeding. This study was conducted to determine hybrid performance and heterosis of 33 intra-pool and 66 inter-pool crosses from a factorial of six European small-seeded (Minor), six European large-seeded (Major), and eight Mediterranean faba bean lines. The parent lines and F₁ crosses were evaluated at two German and seven Mediterranean environments for anthesis, maturity, yield, yield components and plant height. Yield in the Mediterranean environments was 60% of that in the German environments. European Minor lines and their intra-pool crosses were unadapted to Mediterranean environments and yielded only 31 % compared with the Mediterranean lines and their crosses, whereas European Major genotypes yielded 58%. In German environments the European crosses yielded 107% of the Mediterranean crosses. Heterotic yield increase over the parental mean was largest for European Minor x Mediterranean inter-pool crosses, amounting to 95% in the Mediterranean and 73% in German environments. Our results confirmed the expectation of an increased heterosis in inter-pool over intra-pool hybrids and poor adaptation of European genotypes in Mediterranean environments. In the latter, Mediterranean intra-pool hybrids were the most promising, whereas in Germany inter-pool hybrids of the European Minor × Mediterranean type were most promising.     

Genetic analysis of photosynthesis‐related traits in faba bean (Vicia faba) for crop improvement

Increasing productivity through improvement of photosynthesis in faba bean breeding programmes requires understanding of the genetic control of photosynthesis‐related traits. Hence, we investigated the gene action of leaf area, gas exchange traits, canopy temperature, chlorophyll content, chlorophyll fluorescence parameters and biomass. We chose inbred lines derived from cultivars 'Aurora' (Sweden) and 'Mélodie' (France) along with an Andean accession, ILB 938, crossed them (Aurora/2 × Mélodie/2, ILB 938/2 × Aurora/2 and Mélodie/2 × ILB 938/2), and prepared the six standard generations for quantitative analysis (P₁, P₂, F₁, F₂, B₁, and B₂). Gene action was complex for each trait, involving additive and dominance gene actions and interactions. Additive gene action was important for SPAD, photosynthetic rate, stomatal conductance and F_v/F_m. Dominance effect was important for biomass production. It is suggested that breeders selecting for productivity can maximize genetic gain by selecting early generations for canopy temperature, SPAD and F_v/F_m, then later generations for biomass. The information on genetics of various contributing traits of photosynthesis will assist plant breeders in choosing an appropriate breeding strategy for enhancing productivity in faba bean.     

Breeding for wide adaptation in faba bean

Summary Broadening agronomic adaptation will improve yield stability in the grain legume Vicia faba L. We gathered information on the adaptation of European and Mediterranean material to European and Mediterranean environments. The material comprised 20 inbred lines (12 European and 8 Mediterranean lines) and 99 intra- and interpool-crosses in generation F₁. These were evaluated in 9 environments: two spring-sown Southern German environments (SGermE), and seven autumn-sown Mediterranean environments (MedE) in Sicily, Puglia, Andalucia and South Africa. Standard ANOVA, stability analyses and AMMI analysis were performed. Mean yield in F₁ was 257 g/row, the overall parental mean was 144 g/row. The range of environmental means was from 94 g/row to 411 g/row. The average regression coefficient in F₁ was b _i =1.07, being significantly greater than for the parents (b _i =0.68). The opposite was true for the relative magnitude of the deviations from the regressions, which were highly correlated to the AMMI-PC1-results. The AMMI analysis clearly separated the SGermE from the MedE, as well as the germplasm pools. Though the superiority of the F₁-hybrids over their parents was striking, their pattern of interactions with the environments strictly reflected that of their parents. A number of promising crosses was identified as a nucleus of a widely adapted faba bean genepool.     

Detection and characterization of two new CMS systems in faba bean (Vicia faba) *

Wide crosses were made to identify new cytoplasmic male sterility (CMS) systems in faba beans, based on the interaction of cytoplasm with restorer and maintainer alleles. A total of 330 F₁ hybrids were produced in both reciprocal forms. Male sterile segregates were observed in one reciprocal version in the F₂ generation of six crosses. Two of these crosses with female parents originating from Afghanistan and Egypt expressed stable male sterility in subsequent backcross generations. Based on the female parents of the two crosses, these two CMS systems were designated CMS 199 and CMS 297. CMS 199 was more stable than CMS 297 during backcross generations and across different environments. Maintainer and restorer lines for both CMS systems were identified. Lower expression of male sterility occurred in CMS 297 in the greenhouse during the winter generations than in isolation cages during the summer generations, which may be utilized to maintain male sterile lines by selfing. Regarding practical applications, the CMS 199 shows great promise for hybrid breeding in faba beans.     

The inheritance of quantitative traits in Brassica napus ssp. rapifera (swedes): Augmented triple test cross analyses of production characters

Two F₂ triple test crosses, augmented with F₃s, produced from crosses between different inbred lines of swedes (Brassica napus ssp.rapifera L.) were assessed in field trials at Dundee in 1988 and 1989,respectively. This paper reports the analyses of resistance to powdery mildew, neck length, growth cracks, sugar content and hardness; analyses of yield have been published previously. Additive genetical variation was found for all traits while non-additive variation was less important, the highest degree of dominance being 0.44 for hardness. There was evidence of additive × dominance and dominance × dominance epistasis for mildew and additive × additive epistasis for neck length and hardness. Significant,consistent reciprocal differences were found and these were particularly large for neck length and growth cracks. Sugar determination was carried out on the basic generations of the second cross, the parental lines of which showed large differences in concentration of glucose, fructose and sucrose. Directional dominance was found for high glucose but not for fructose or sucrose. The implications of these results for swede breeding are discussed and it is proposed that inbred cultivars would be a more practical option than F₁ hybrids. This revised version was published online in August 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.     

Estimation of additive, dominance and digenic epistatic interaction effects for certain yield characters in Vigna sesquipedalis Fruw

Inheritance of yield and yield contributing characters were investigated using generation mean analysis, utilising the means of six basic populations viz., P₁, P₂, F₁, F₂, BC₁P₁ and BC₁P₂ in four crosses of Vigna sesquipedalis. The analysis reiterated that the importance of dominance (h) gene effects for pod yield/plant and pods/plant as compared to additive (d) gene effects. However, significant and positive additive effects were noticed for pod yield/plant, pods/plant, pod weight and seed weight in different crosses. The three types of gene interactions (additive, dominance and epistasis) were significantly involved for pods/plant in cross KU 7 ×KU 8. Among the digenic epistatic interactions, both additive ×additive (i) and dominance × dominance (l) contributed more for pod yield/plant and pods/plant, however, it varied among the crosses. Populations having earliness can be developed as indicated by reducing dominance effects. Pedigree selection and heterosis breeding is suggested to exploit the fixable and non fixable components of variation respectively in Vigna sesquipedalis. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of culm number in two uniculm x multiculm wheat crosses

Summary Effective utilization of uniculm wheat (Triticum aestivum L.) germplasm in breeding programs requires an understanding of the inheritance of the uniculm character. The parents, F₁, F₂, and first generation backcrosses (B₁ and B₂) of two crosses, each utilizing a uniculm spring wheat line and a locally adapted winter wheat cultivar, were space planted in an experiment to acquire information regarding the genetic control of culm number. Significant F₁ deviations from midparent values revealed the presence of substantial non-additive gene effects. The scaling tests of Mather and the joint scaling test detected the presence of epistasis. Hayman's six-parameter model revealed that a negative dominance effect provided the major contribution to variation in culm number, while additive x additive and dominance x dominance gene effects were of considerable importance.     

Inheritance of drought resistance related traits in two crosses of groundnut (<Emphasis Type="Italic">Arachis hypogaea</Emphasis> L.)

Groundnut (Arachis hypogaea L.) is an important oilseed crop grown in more than 100 countries across wide range of environments. Frequent occurrence of drought is one of the limiting factors adversely affecting groundnut productivity, especially in rainfed areas, and hence genotypes having high water use efficiency (WUE) under limited available water need to be developed. In groundnut, WUE is correlated with SPAD chlorophyll meter reading (SCMR) and specific leaf area (SLA). These two traits, SCMR and SLA, can be used as surrogate traits for selecting for WUE. In order to improve SCMR and SLA, and in turn WUE in groundnut, a good knowledge of the genetic system controlling the expressions of these traits is essential for the selection of the most appropriate and efficient breeding procedure. The present investigation was conducted to determine the gene action controlling the inheritance of SCMR and SLA in two crosses, ICG 7243 × ICG 9418 and ICG 6766 × Chico, and their reciprocals. Six generations of each cross (P₁, P₂, F₁, F₂, BC₁P₁, and BC₁P₂) were evaluated for SCMR and SLA at two stages of the crop growth viz., 60 and 80 days after sowing (DAS). For SCMR at 80 DAS, additive effects were important in both the crosses whereas predominance of dominance effects with duplicate epistasis was observed for SCMR at 60 DAS and SLA at both stages in both the crosses. Predominance of additive effect for SCMR at 80 DAS suggested effective selection could be practiced even in early generations whereas for SCMR at 60 DAS and SLA at both stages in both crosses, it would be better to defer selection to later generations. Further, recording of SCMR and SLA should be done between 60 and 80 DAS for screening the germplasm lines for drought tolerance.     

Inheritance of grain filling duration in spring wheat

The genetic basis of grain-filling duration (GFD, days from anthesis to maturity) in six spring wheat hybrids involving nine varieties (‘Son-alika’/‘Bobwhite’, ‘Sonalika’/‘Glennson 70’, ‘Lelija’/‘Bobwhite’, ‘Lelija’/‘Mitacore’, ‘Buckbuck’/‘Dugoklasa’, and ‘Vesna’/‘Radu?a’) and their F₁, F₂, BC₁, and BC₂ generations was studied in the field near Sarajevo, Yugoslavia. Parental means differed in four of the six crosses. Generation mean analyses of genetic effects indicated that an additive-dominance model was sufficient for only two crosses: Lel/Bow and Lel/Mco. One or more types of epistasis were significant in the remaining crosses. The F₁ and F₂ means were either intermediate, closer to the mean of the parent with the longer GFD, or closer to the mean of the Parent with the shorter GFD. Even though different modes of gene action controlled GFD among the six crosses, the heritabilities were reasonably high (narrow sense, 39-59) range for six crosses), indicating that progress could be made from selection in these crosses for either long or short GFD. The parents were selected to have a range in days from planting to anthesis and to maturity. The relationship between dates of anthesis or maturity and GFD was not consistent, but the two latest-maturing varieties had the longest GFD, indicating that anthesis or maturity dates are not a good criteria for choosing parents for modifying GFD. Additive genetic effects predominated in the crosses studied here, but epistasis involving dominance gene action was sufficiently important. To eliminate confounding epistatic dominance effects and to take advantage of favourable additive × additive effects during selection for GFD, a breeding strategy involving rapid approach to homozygosity followed by selection after the achievement of homozygosity was suggested.     

Genetic studies for seed size and grain yield traits in kabuli chickpea

Seed size, determined by 100-seed weight, is an important yield component and trade value trait in kabuli chickpea. In the present investigation, the small seeded kabuli genotype ICC 16644 was crossed with four genotypes (JGK 2, KAK 2, KRIPA and ICC 17109) and F₁, F₂ and F₃ populations were developed to study the gene action involved in seed size and other yield attributing traits. Scaling test and joint scaling test revealed the presence of epistasis for days to first flower, days to maturity, plant height, number of pods per plant, number of seeds per plant, number of seeds per pod, biological yield per plant, grain yield per plant and 100-seed weight. Additive, additive?×?additive and dominance?×?dominance effects were found to govern days to first flower. Days to maturity and plant height were under the control of both the main as well as interaction effects. Number of seeds per pod was predominantly under the control of additive and additive?×?additive effects. For grain yield per plant, additive and dominance?×?dominance effects were significant in the cross ICC 16644?×?KAK 2, whereas, additive?×?additive effects were important in the cross ICC 16644?×?JGK 2. Additive, dominance and epistatic effects influenced seed size. The study emphasized the existence of duplicate epistasis for most of the traits. To explore both additive and non-additive gene actions for phenological traits and yield traits, selection in later generations would be more effective.     

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.     

中国蚕豆种质资源ISSR标记遗传多样性分析

利用ISSR标记对中国18个省(区)的527份春播区和秋播区蚕豆资源的遗传多样性与亲缘关系进行分析。结果表明,11个ISSR引物共扩增出278条清晰的条带,其中多态性条带268条(占96%)。不同地理来源蚕豆资源群间的基因多样性指数在0.1267~0.2509之间,平均为0.1716;Shannon指数范围为0.1932~0.3767,平均为0.2608。两指数均以内蒙古资源群最高,云南和甘肃资源群次之,江西资源群体最低。非加权配对算术平均法(UPGMA)的聚类分析和二维主成分分析(PCA)结果表明,中国春播区和秋播区蚕豆资源明显不同,浙江和四川的资源形成独立的组群,明显不同于其他省份,其遗传背景独特,有待进一步研究。中国蚕豆种质资源遗传多样性差异和遗传关系与其生长习性、生态分布及地理来源密切相关。     

Genetics of Six Components of Autofertility in Vicia faba

The inheritance of six components of autofertility was studied in faba beans in two consecutive years (1989 and 1990) by analysing parents, Fl and segregating generations of 14 crosses between preselected pure lines. Genetic analysis was performed by a joint scaling test proposed by Cavalli (1952). Additive gene effects were more important than dominance effects in flowers/node, pods/flower and ovules/ovary. In contrast, characters directly related with the seed such as pods/node, seeds/pod and, to a lesser extent, seeds/ovule showed high positive values of dominance. The three latter traits in general exhibited strong heterosis, probably due to genetic interactions. In order to produce selection gain, any breeding programme has to consider that dominance and heterosis vary according to the character involved. The present study reveals different degrees of dominance expression depending on cross specificity as well as on environmental conditions. The advantages of studying segregating generations instead of diallel crosses and the need for a simultaneous study of many crosses are emphasized in this paper.     

Inheritance of Bolting in Three Sugar Beet Crosses after Different Periods of Vernalization

The induction of bolting after different periods of vernalization was studied in the parental, F₁, F₂ and backcross generations (F₂× P₁ and F₂× P₂) of three crosses between five sugar beet lines. The parental lines represented different levels of bolting resistance. The populations were evaluated in replicated field or greenhouse trials. Generation mean analysis was used to quantify the genetic effects of bolting. The additive (d) parameter was predominant for those levels of cold treatment that stimulated the bolting to occur. The dominance (h) parameter was also shown to be important in most cases. In one cross, the additive × additive (i) type of epistasis was significant. The chances of detecting genetic effects are increased by exposing the plants to vernalization conditions which maximize the difference between two parental lines of each cross. Hence, it would be sensible to exploit both additive and non-additive genetic effects in any selection programme.     

Triple Test Cross Analysis in F2 Populations of Four Promising Crosses of Field Pea

The components of genetic variation were studied in four F₂ crosses (PG₃× 5064, T₁₆₃× B.V., T₁₆₃× Arkel and 5064 × ED) for quantitative characters such as days to flowering, plant height, pod number, seed number, test weight and yield/plant. The overall epistasis (T type) was, in general, a major contributor for genetically controlling the expression of the characters in all the- four crosses except for the character seed number in crosses 1 and JV, where the j + l component at epistasis played a significant role in determining the expression of this attribute:. Though the ‘j + l’ component of the epistasis was significant, it was relatively less important than the ‘i’ type epistasis. Both the additive and dominance components of the genetic variation were highly significant for all the trans studied in all die four sets of crosses. The expression of the dominance was directional only for a few characters in certain crosses. The degree of dominance lies in the partial range, and heritability estimates obtained were high for most of the trails. The possible application of cross prediction in the isolation of superior recombinant inbred lines in pea is discussed.     

Effectiveness of early generation selection under two plant densities in faba bean (Vicia faba L.)

The effectiveness of early generation selection under low and high plant density was evaluated in faba bean (Vicia faba L.). For this, the faba bean cv. 'Polycarpe' was pollinated by eight higher yielding cultivars of different origin. The F₁'s and F₂'s together with 'Polycarpe' and the pollinator with the highest yielding ability (cv. 'Effe') were evaluated under high plant density with adjacent controls and under low plant density in a honeycomb design. It was found that the eight hybrids had a different ranking at low than at high plant density for both generations (F₁and F₂). Hybrid Pol.x A-58 had the best combined yield performance in the F₁and F₂under high and hybrid Pol.x A-37 under low plant density. Application of individual plant selection in the F₂and F₃generation of the two crosses (Pol.x A-58 and Pol.x A-37), under both low plant density according to UNR-O honeycomb design and high plant density using a moving average, resulted in the selection of four groups with 20 selected F₃plants each. Within each group, equal numbers of seeds from each plant were mixed to establish four mixtures. These mixtures together with two controls ('Polycarpe' and 'Effe') were evaluated in a randomized complete block design (22 plants/m²) in one location for two years. It was found that the material selected continuously under low plant density had a significantly higher yield than the material selected under high plant density. In addition, the material selected from Pol.x A-37 (H₁R₂) under high plant density had a lower mean yield than that selected continuously under low plant density (H₁H₂). Their average yield over years was significantly higher than the yield of the material selected from Pol.x A-58 under both plant densities. Furthermore, the material selected from Pol.x A-58 under low plant density, had a significantly higher mean yield over years than its sibs selected under high plant density. It was concluded that honeycomb selection at low plant density can be used effectively for early generation selection in faba bean.     

Genetics of spike length in durum wheat

Gene effects were analyzed using mean spike length of 12 populations, viz., both parents, F₁, F₂, first back cross generation, BC₁ and BC₂, second backcross generations, BC₁₁,BC₁₂, BC₂₁ and BC₂₂ along with BC₁ self and BC₂ self derived by selfing BC₁ and BC₂populations of three crosses involving six diverse cultivars of Triticum durumto determine the nature of gene actions governing spike length through generation mean analysis under normal and late sown environments. The six-parameter model was adequate in most of the cases to explain genetic variation among the generation means under both the sowing environments. Additive (d) gene effect was significant in all the cases, whereas dominance (h) gene effect was not so frequently observed significant. Epistatic effects, particularly digenic types were predominant over additive and dominance effects in most of the cases under both normal and late sown environments except in the cross Cocorit 71 × A-9-30-1 (normal sown).Additive × dominance × dominance (y), trigenic interaction played significant role in controlling the inheritance of this trait in the cross HI 8062 × JNK-4W-128under late sown condition. Duplicate epistasis was observed in the cross HI 8062× JNK-4W-128 (normal sown). Non-fixable gene effects were of higher magnitude than fixable gene effects in almost all cases, confirmed the major role of non-additive gene effects to control the inheritance of spike length in durum wheat. Significant heterosis over better parent was not observed. Similarly, inbreeding depression was not commonly observed. Favourable and suitable environment must be considered before finalizing breeding programme for its simple inheritance to get desirable improvement for high grain yield. Hybridization systems, such as biparental mating and / or diallel selective mating, which exploit both additive and non-additive gene effects, simultaneously, could be useful in the improvement of spike length in durum wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Identification of stable restorers and genetics of fertility restoration in late‐maturing pigeonpea [Cajanus cajan (L.) Millspaugh]

A total of sixty‐six germplasm lines were crossed with five CMS lines, where two belong to A₄ cytoplasm, while other three belong to A₂ cytoplasm. On the basis of pollen fertility test as well as good pod setting, of 330 hybrids, 34 restorer lines were observed in ICPA 2043 and 19 in ICPA 2092. Thirteen germplasm lines restored fertility in both the A₄ CMS lines, viz. ICPA 2043 and ICPA 2092; however, none of the lines restored fertility in A₂ CMS lines. For confirmation of result, restoration competence of identified lines tested subsequently 2 years at two different temperatures. The segregation patterns for fertility restoration studied in F₂ and BC₁F₁ generations of selected ten crosses. Six crosses indicated the involvement of two major genes with recessive epistasis, three crosses confirmed dominant epistasis, and one cross indicated the involvement of duplicate recessive epistasis. The obtained results from this study will hasten the future three‐line breeding programme and lead the hybrid technology to the farmers' field with the better exploitation of CMS lines.     

Hybrid performance and AFLP- based genetic similarity in faba bean

Successful prediction of heterosis and performance of F₁-hybrids from the genetic similarity of their parents based on molecular markers has been reported in several crops and can be very helpful in hybrid breeding. The relationship between genetic similarities based on amplified fragment length polymorphism (AFLP) of 18 European faba bean lines and their hybrid performance and heterosis was investigated. Parental lines, 62 F₁-hybrids and their F₂-progenies were evaluated in field trials in four environments in Germany for seed yield, 1,000-seed weight and plant height. Results clearly demonstrated a stable superiority of the hybrids over their inbred parents and elite check cultivars, and showed a marked and varying amount of heterosis. Parental seed yield and F₂-hybrid yield were promising as predictors for F₁-hybrids. AFLP analysis of the 18 inbred lines using 26 EcoRI/MseI primer combinations resulted in 1202 polymorphic fragments. Cluster analysis based on genetic similarity estimates unambiguously identified pedigree-related inbred lines. No clear separation of the 18 inbred lines into subgroups was detected. Correlation coefficients between genetic similarity estimates and either heterosis or F₁-hybrid performance were small and not useful. Also correlations between specific genetic similarity and specific combining ability were too small for all traits to be of predictive value. Results showed that AFLP-based genetic similarities are not useful to predict the performance of hybrids or heterosis within the elite European faba bean gene pool.