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.     

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.     

Generation mean analysis of resistance to downy mildew in adult muskmelon plants

Downy mildew (Pseudoperonospora cubensis) is the most devastating disease in muskmelon (Cucumis melo). A generation mean analysis study was designed to determine the types of gene action and estimate the heritability for resistance to downy mildew in four selected crosses of muskmelon. Generation mean analysis revealed that genetic dominance may be of greater importance for expression of resistance to downy mildew in both greenhouse and field experiments and in all the crosses. The F₁ mean was significantly lesser than the mid-parent value and skewed towards resistant parent in all the crosses. Negative sign associated with gene effects indicated, in those crosses, that disease level could be decreased in relation to midparent. All the crosses expressed significant and positive additive (d) gene effects. Dominance (h) and dominance × dominance (l) gene effects had opposite sign in all crosses and both experiments, which implied duplicate type of gene action. High mid-parent heterosis in all the crosses indicated strong dominance effects (as combination of parental alleles) for resistance to downy mildew. In all the crosses, both resistant and susceptible parent contributed one or more dominant/partially dominant factors for resistance. Estimates of broad-sense heritability were high and relatively consistent in both experiments. The two different screening experiments showed that fixable gene effects (d + i) were lower than the non-fixable gene effects (h + l) in all the crosses indicating greater role of non-additive effects in the inheritance of resistance to downy mildew. Resistance to downy appeared to be controlled mainly by dominance effects, therefore the inbred lines IIHR 121 and IIHR 122 could be used strategically to exploit heterotic effects.     

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.     

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.     

Gene action and heritability for photosynthetic activity in two wheat crosses

Summary Gene action and heritability for photosynthetic activity were estimated from generation means in two wheat crosses during two stages (5 ^th leaf and flag leaf between 2 and 5 days after anthesis). Six generations were available for each cross: parents (P₁ and P₂), F₁, F₂ and backcrosses (BC₁ and BC₂).Correlations between some morphophysiological characters and photosynthetic activity of the flag leaf was also determined. The joint scaling test described by Mather & Jinks was used to determine the gene action. It showed that them; [d]; [h]; [i], [l] (mean, additivity, dominance, additive x additive interallelic interaction effects, dominance x dominance interallelic interaction effects) model fits the two crosses at both measurement times. All the model genetic components were significant for the flag leaf, however for the 5 ^th leaf only [h]; [i] and [l] were significant. The presence of additive and additive x additive effects suggested the possibility of selecting for this character using the flag leaf so as to obtain pure inbred lines. Dominance effects [h] were negative and dominance x dominance effects [l] were positive. Broad sense heritability values were medium to low. There were no correlations between the studied morphophysiological characters and the photosynthetic activity.     

Genetics of resistance to early blight (Alternaria solani Sorauer) in tomato (Lycopersicon esculentum L.)

The genetic nature of early blight resistance in tomato was studied in three crosses at seedling and adult plant stages. A six generation mean analysis of the cross Arka Saurabh (susceptible) × IHR1939 (resistance) and its reciprocal cross revealed that the resistance to early blight was conferred by recessive polygenes at both seedling and adult plant stages. This polygenic early blight resistance revealed the importance of additive and additive × additive gene effects at seedling stage and higher magnitude of dominance and dominance× dominance gene effects at adult plant stage. Evaluation of parents, F₁, F₂ and backcross generations of IHR1816 (resistance) × IHR1939 (resistance) revealed that the early blight resistance genes in IHR1816 (Lycopersicon esculentum NCEBR-1) and IHR1939 (Lycopersicon pimpinellifolium L4394) are independent. This revised version was published online in July 2006 with corrections to the Cover Date.     

Triple test cross analysis for some morphological traits in mungbean (Vigna radiata (L.) Wilczek)

Thirty progenies of mungbean were produced by crossing 10 true-breeding genotypes with three testers (NM 92, 6601, and their F₁) in a Triple Test cross (TTC) fashion and evaluated with parents in the kharif (July-October) and spring/summer (March-June) seasons. The data on parents and F₁s were analysed for pod clusters on main stem, pod clusters on branches, node of the first peduncle, nodes on main stem and average internode length to detect epistasis and estimate additive and dominance components of genetic variation. Epistasis was observed for node of the first peduncle and nodes on main stem in the kharif season. Partitioning of total epistasis revealed that both additive × additive (i type), and additive × dominance, and dominance × dominance (j and l types) interactions were significant with prevalent influence of i type interactions on these traits. Both additive and dominance components of genetic variation were significant for all those traits not significantly influenced by epistasis in either or both seasons. The additive component was predominant for pod clusters on main stem, pod clusters on branches and average internode length in the kharif season, and for the node of the first peduncle and nodes on main stem in spring/summer season whereas dominance component was important for pod clusters on main stem, pod clusters on branches, and average internode length in spring/summer season. These results suggested that particular generation of segregating population and specific breeding method for selection might be adopted in each season for the improvement of these traits in mungbean. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Inheritance of β-carotene concentration in durum wheat (Triticum turgidum L. ssp. durum)

Despite being one of the important characteristics in determining pasta quality in durum wheat (Triticum turgidum ssp. durum), there is no direct report on inheritance of β-carotene concentration. The objectives of this study were to determine the inheritance of β-carotene concentration and the number of genes involved in six crosses of durum. For the cross PDW-233 (P₁) × Bhalegaon-4 (P₂), F₁, F₂, BCP₁ and BCP₂ populations were developed. For all other crosses, only the F₁ and F₂ populations were developed. β-carotene concentration was determined for all populations and parents of each cross grown at Hol, Maharastra, India. The cross PDW-233 × Bhalegaon-4 was also evaluated at Dharwad, Karnataka, India. Low β-carotene concentration was partially dominant in most of the crosses. Broad sense heritability was 67 and 91% at Dharwad and Hol, respectively, for the cross PDW-233 × Bhalegaon-4 and varied from 74 to 93% for the other five crosses indicating the presence of additive gene effects. The frequency distributions of the trait in the F₂ populations were not normal and were skewed towards the lower parent. Segregation of β-carotene concentration in the six F₂ populations indicated that at least two major genes and two or three minor genes with modifying effects govern the trait. Analysis of variance indicated that environment had comparatively little influence on the trait and this should allow for easy selection. The joint scaling test revealed additive × additive, additive × dominance and dominance × dominance epistatic interactions in the cross PDW-233 × Bhalegaon-4. These authors contributed equally.     

Comparison of intra- and inter-pool crosses in faba beans (Vicia faba L.). II. Genetic effects estimated from generation means in Mediterranean and German environments

Genetic diversity between parents is important for hybrid breeding and for maximum usefulness of a cross in pure line breeding. However, wide crosses may suffer from poor adaptation in the target environments and recombination losses owing to disruption of favourable epistatic gene combinations. In this study we investigated the importance of various gene effects for different traits in the parents and generations F¹ to F⁵ derived from 99 intra- and inter-pool crosses among 20 lines of the Minor, Major, and Mediterranean germplasm pools of faba bean in Mediterranean and German environments. Yield performance of parental inbreds and progenies of crosses in different selfing generations was high in the region of adaptation of the respective germplasm pool. Heterosis for yield was greatest in interpool crosses of Minor × Mediterranean germplasm, amounting to 106% in Mediterranean and 72% in German environments. Interpool crosses exhibited significantly greater heterosis compared with intrapool crosses in both mega-environments. Generation means analysis revealed the importance of additive gene effects and dominance × dominance epistasis in Mediterranean and German environments. Dominance effects and additive × additive epistasis for yield were negligible in most crossing groups. Since intrapool crosses of Mediterranean germplasm in Mediterranean environments and crosses of Minor and Major germplasm in the German environments were most promising, we suggest crosses among adapted germplasm for hybrid breeding and for establishing new base material in line breeding in faba bean.     

Estimation of genetic parameters for in vitro traits in wheat immature embryo cultures involving high X low regeneration capacity genotypes

Summary Estimates of gene actions were obtained for five in vitro traits of immature wheat (Triticum aestivum L.) embryo cultures from a cross of two wheat cultivars and the resulting reciprocal, F₁, F₂ and backcross populations. The contribution of additive gene effects to in vitro traits was not as important as the dominance gene effects. Epistatic gene effects were relatively more important than either additive or dominance gene effects. Of the individual types of digenic epistatic effects, the dominance x dominance estimates were relatively larger in magnitude for all in vitro culture traits measured. The maternal effect played a minor role in the inheritance of the in vitro studied traits since the difference among the reciprocal values was not significant. It is shown from the generation mean method that epistasis played a major role in the inheritance of most of the traits under study. The negative values of additive and dominance genetic variance were estimates of zero. Heritability estimates, in broad sense, were relatively high for the in vitro studied traits. In some cases, heritability estimates in broad and narrow senses are almost equal since the estimation of dominance genetic variance led to negative values. According to the results of the gene effects, dominance and epistasis were important for the shoot formation trait. Selection would be effective among the isolated genotypes on individual basis.     

Inheritance of internode length,plant form,and annual habit in a cross of cabbage and broccoli (Brassica oleracea var. capitata L. and var. italica Plenck.)

Summary When an inbred line of cabbage, Brassica oleracea L. var. capitata L., was crossed with an inbred line of broccoli B. oleraceae var. italica, the F₁ progeny were vigorous late annuals. All F₁ × broccoli backcross plants and 92% of the 3260 F₂ plants were annuals, while 40% of the F₁ × cabbage backcross plants were biennials. Annual habit is thus dominant and controlled by more than a single gene. Number of days to bud appearance in annuals varied continuously, and was primarily additive in inheritance. F₁ data suggested partial dominance for lateness but this was not supported by the F₂. Internode length was also continuous in distribution and primarily additive in inheritance, but with some dominance for short internodes in the F₁. Cabbage head forming ability was recessive and multigenic, with 2% of the F₂ plants forming heads, of which none were of commercial type and about half bolted as annuals. There was a significant chi square association between biennial habit and tendency for cabbage head formation. Clasping habit of terminal leaves was recessive to open leaves, multigenic, and associated with both cabbage heading and biennial habit.Technical Paper 4836, Oregon Agricultural Experiment Station; from an M.S. thesis by the senior author.     

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.     

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.     

Variation of harvest index in several wheat crosses

Summary Harvest index was studied in F₁ and F₂ generations of eight wheat (Triticum aestivum L.) crosses and their reciprocals. The parental varieties involved in the crosses represented a fairly wide range of character expression for plant height, tillering potential, grain yield per plant and harvest index. Differences between reciprocal crosses were not evident for the expression of harvest index of the crosses under study. Means and degrees of dominance of F₁ and F₂ populations suggested partial dominance of high harvest index over low harvest index. The pattern of variation among F₂ segregates was quantitative and the distribution was normal. The gene action governing the expression of harvest index was largely additive. Evidence was obtained for non-additive gene action in some crosses. There was complete absence of high parent heterosis for harvest index in the F₁'s but midparent heterosis was found to be present in all crosses. Estimates of heritability and genetic advance were moderate to high. Usefulness of selecting for harvest index as a measure of yield efficiency particularly in early generations is discussed.     

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.     

Inheritance of Carbon Isotope Discrimination in Bread Wheat (Triticum Aestivum L.)

Summary Reliable selection of families with increased grain yield is difficult in breeding programs targeting water-limited environments. Carbon isotope discrimination (Δ) is negatively correlated with transpiration efficiency, and low Δ is being used for indirect selection of high wheat yield in rainfed environments. Yet little is known of genetic control and opportunities for improving selection efficiency of Δ in wheat. Half-diallel and generation means mating designs were undertaken to provide estimates of the size and nature of gene action for Δ in a range of wheat genotypes varying for this trait. Significant (P < 0.01) differences were observed for leaf tissue Δ among parents (19.3 to 20.7‰) and F₁ progeny (19.4 to 20.9‰) in the half-diallel. General (GCA) and specific combining ability (SCA) effects were significant (P < 0.05), while Baker's GCA/SCA variance ratio of 0.89 was close to unity, indicating largely additive gene effects. GCA effects varied from −0.38 to + 0.34‰ for low and high Δ genotypes `Quarrion' and `Gutha', respectively. GCA effects and parental means were strongly correlated (r = 0.95, P < 0.01) while directional dominance and epistasis contributed to small, non-additive gene action for Δ. Smaller Δ in F₁ progeny was associated with accumulation of recessive alleles from the low Δ parent. Narrow-sense heritability was high (0.86) on a single-plant basis. Generation means analysis was undertaken on crosses between low Δ genotype Quarrion and two higher Δ genotypes `Genaro M81' and `Hartog'. The F₁, F₂ and midparent means were not statistically (P > 0.05) different, whereas backcrossing significantly changed Δ toward the mean of the recurrent parent. Gene action was largely additive with evidence for additive × additive epistasis in one cross. Narrow-sense heritabilities were moderate in size (0.29 to 0.43) on a single-plant basis. Genetic gain for Δ in wheat should be readily achieved in selection among inbred or partially inbred families during the later stages of population development.     

Inheritance of downy mildew resistance in Indian cauliflower (group III)

Summary Inheritance of downy mildew (Peronospora parasitica) resistance in three resistant x susceptible crosses, one susceptible x susceptible and one resistant x resistant cross were studied in Indian cauliflower (Group III) over the two years (1990 and 1991). No significant difference was observed between the years for various estimates and hence pooled data are presented. Downy mildew resistance in crosses cc×HR 5-4 and 3-5-1-1×244 (R×S) is governed by single dominant gene PPA3 but in cross cc×244 (R×S), recessive epistasis was observed. The resistance level was not improved in both the cc×3-5-1-1 (R×R) and 244×267-6-9 (S×S) crosses. Exploitation of downy mildew resistance from cc and 3-5-1-1 in F₁ hybrid is explained in detail.