Hybrid breeding for biomass yield in winter triticale: II. Combining ability and hybrid prediction

Accurate hybrid prediction and knowledge about the relative contribution of general (GCA) and specific combining ability (SCA) are of utmost importance for efficient hybrid breeding. We therefore evaluated 91 triticale single-cross hybrids in field trials at seven environments for plant height, heading time, fresh biomass, dry matter content and dry biomass. Fresh and dry biomass showed the highest proportion (23%) of variance due to SCA. Prediction accuracies based on GCA were slightly higher than based on mid-parent values. Utilizing parental kinship information yielded the highest prediction accuracies when both parental lines have been tested in other hybrid combinations, but still moderate-to-low prediction accuracies for two untested parents. Thus, hybrid prediction for biomass traits in triticale is currently promising based on mid-parent values as emphasized by our simulation study, but can be expected to shift to GCA-based prediction with an increasing importance of GCA due to selection in hybrid breeding. Moreover, the performance of potential hybrids between newly developed lines can be predicted with moderate accuracy using genomic relationship information.     

Heterosis and combining ability for grain yield and other agronomic traits in winter triticale

Hybrid breeding is a widely discussed alternative for triticale. Heterosis as well as general (GCA) and specific combining ability (SCA) effects were estimated for eight agronomic traits. The experiment comprised 24 F₁ hybrids, produced by a chemical hybridizing agent, together with their six female and four male parents, grown in drilled plots in two locations. In comparison with the mid‐parent values, hybrids averaged a 6.4 dt/ha (10.1%) higher grain yield, 8.4% more kernels per spike, a 6.8% higher 1000‐kernel weight, 9.7% lower falling number (FN) and 4.4% greater plant height. SCA effects for grain yield were significant and ranged from 4.5 to 6.9 dt/ha for grain yield. Together with GCA x location interactions, they explained most of the variation. For 1000‐kernel weight, GCA effects were predominant. SCA and interactions with location accounted for most of the variation in FN, whereas interactions were negligible for plant height. Correlations between mid‐parent and hybrid performance and between GCA and per se performance of parents were tight for all traits except grain yield, which allows for pre‐selection of parental lines. Although the amount of heterosis in triticale at present is closer to wheat than to rye, by selecting parents for combining ability and identifying heterotic patterns, grain yield heterosis of up to 20% appears sufficiently encouraging to embark on hybrid breeding.     

Heterosis for yield and other agronomic traits of winter triticale F1 and F2 hybrids

Triticale is generally treated as a self‐pollinating crop and line breeding is practised. Hybrid breeding has been discussed for some time, but there is little information for winter triticale. This study investigated heterosis for eight agronomic traits in F₁ and F₂ hybrids grown together with their parents as drilled plots in three environments. On average, grain yield heterosis was 12.5 dt/ha (a relative 10.5%) compared with the mid‐parent value for F₁ hybrids, and 6.2 dt/ha (5.0%) for F₂ hybrids and withawide range of 4.4–17.1 dt/ha for F₁ hybrids. A positive contribution to the heterosis of yield was made by kernels/spike and 1000‐kernel weight, whereas spikes/m² showed negative heterosis. Hybrid plants in F₁ and F₂ were taller than mid‐parents (8.3 cm and 5.3 cm, respectively), with a tendency to earlier heading. The negative heterosis for falling number in F₁ and F₂ hybrids could be a problem for commercial production of triticale hybrids. By selecting parents for combining ability and the identification of heterotic patterns, grain yield heterosis of 20% appears feasible.     

Relationship between grain yield and grain nitrogen concentration in winter triticale

Summary Ten hexaploid winter triticale lines were grown for two cropping periods at three locations in western Switzerland. Averaged across the six environments, the differences between lines were statistically significant (P=0.05) for grain yield, above-ground biomass, N uptake, grain N yield, nitrogen harvest index, grain N concentration and straw N concentration. There were significant line x environment interactions for all traits. Grain yield and grain N concentration were inversely related (r=–0.74^**). Diagrams in which grain yields were plotted against grain N concentration were used to identify lines with a consistently unusual combination of grain yield and grain N concentration. Despite comparable grain yields, Line 3 had a high grain N concentration, while that of Line 7 was low. Line 3 was superior to Line 7 in both N uptake and N harvest index. Averaged across environments and lines, the N harvest index was 0.73 which corresponds to N harvest indices reported for bread wheat in the same region. We considered the feasibility of developing triticale lines which would outperform the best recent ones in N uptake and partitioning. However, we doubted that this would bring about a marked increase in grain N concentration, because, in the long run, the expected genetic progress in grain yield will lead to a dilution of grain protein by grain carbohydrate increments.Abbreviations GNC grain N concentration - GNY grain N yield - GY grain yield - HI above-ground dry matter harvest index - NHI nitrogen harvest index - SNC straw N concentration - TB total above-ground biomass - TPN total plant N     

Prospects for hybrid breeding in winter triticale: II. Relationship between parental genetic distance and specific combining ability

Significant relative midparent heterosis (MPH%) for grain yield in triticale (×Triticosecale Wittm.) has generated interest in the development of hybrid cultivars. The objectives of this study were to (i) examine the association between parental genetic distance (GD) and specific combining ability (SCA), (ii) investigate the existence of genetically distant heterotic groups in elite germplasm, and (iii) draw conclusions for future hybrid breeding in winter triticale. Genetic distance between 61 lines was estimated, based on 93 polymorphic simple sequence repeat (SSR) marker loci and 10 AFLP (amplified fragment length polymorphism) primer‐enzyme combinations (PEC). Agronomic data of 206 F₁ crosses and their 61 parental lines grown in six German environments were published recently in a companion study. Correlations were made between SCA for grain yield, number of spikes/m², 1000‐kernel weight and number of kernels per spike with GD estimates of the 56 female and five male parents (testers). Principal co‐ordinate analyses (PCoA) based on SSR data revealed no distinct subgroups in the germplasm. Correlations between GD and SCA were low for all traits (|r| ≤ 0.31), which hampers the prediction of SCA from molecular data. A multi‐stage procedure is recommended for future hybrid breeding in triticale by applying a pragmatic approach for the grouping of germplasm following the early history of hybrid breeding of maize.     

Combining ability and heterosis over environments for yield and yield components in two-line hybrids involving thermosensitive genic male sterile lines in rice (Oryza sativa L.)

Presence of substantial heterosis and economic hybrid seed production are two most desirable components for success of any commercial hybrid breeding programme. Thermosensitive genic male sterile (TGMS) lines of rice, in this regard, have tremendous potential in realizing further quantum jump in yield and economical hybrid seed cost. Analyses for combining ability and heterosis over optimum (120N : 60P₂O₅ : 40K₂O kg/ha) and high (200N : 90P₂O₅ : 60K₂O kg/ha) fertility environments for six traits were made in 2 years (2001 and 2002) using 120 hybrids of inter‐ and intra‐subspecific nature derived from hybridization of 30 elite indica TGMS lines and four cultivars, viz., ‘Pant Dhan 4’ and ‘Ajaya’ (I = indica), ‘Taichung 65’ (J = japonica) and ‘IR 65598‐112‐2’ (TJ = tropical japonica) in line × tester mating design. Predominance of non‐additive genetic variance suggested good prospects of hybrid breeding. Pooled analysis revealed highly significant variances for lines, general combining ability (GCA), specific combining ability (SCA) and line x tester. TGMS line 365‐8S was the best general combiner for all the six traits including grain yield. Trend of relative mid‐parent heterosis for grain yield, panicle length, grain number per panicle and earliness in flowering was I/TJ > I/J > I/I. For panicle number per plant and 1000‐grain weight, trends were I/TJ > I/I > I/J and I/I > I/TJ > I/J, respectively. Grain yield recorded heterosis of 49.3%, 71.9% and 92.7% for I/I, I/J and I/TJ hybrid groups respectively. Effect of environments on the hybrid performance indicated better response of hybrids at high fertilizer dose. Study suggests greater prospects of combining improved japonica and tropical japonica germplasms having wide compatible gene with indica TGMS lines for exploitation of intersubspecific heterosis.     

Inheritance of yield components and yield in relation to evidence for heterosis in F1 barley hybrids

Summary Yield components and yield were studied in F₁ barley hybrids produced by hand pollination or male sterility. Grain number exhibited only partial dominance but grain weight showed dominance or overdominance and contributed to the heterotic situation particularly in 2×6-row crosses. For the commercial exploitation of heterosis it is essential that hybrids should be found which show greater dominance for high grain number.     

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.     

A fast generation cycling system for oat and triticale breeding

A fast generation cycling system (FGCS) was developed in oat and triticale, allowing the production of pure line populations of the crops within 1‐year timeframe. Twelve oat and 12 triticale cultivars were tested using the FGCS, and most of them completed a generation cycle within 48–61 days under the system achieving up to seven generations a year. This system involves growing plants under stressed conditions to promote reproduction and in vitro culture of immature embryos bypassing full seed maturation. The developed system could be widely adopted in breeding and genetic studies of crops for producing desired segregating pure line populations, which could significantly shorten the breeding cycles.     

Heterosis breeding in Indian mustard (Brassica juncea L. Czern & Coss): Analysis of component characters contributing to heterosis for yield

Summary Divergence of 25 accessions of Brassica juncea of Indian, CIS (Commonwealth of Independent States, former USSR) and synthetic origin was studied by D² analysis. On the basis of divergence, ten accessions were selected and crossed in a diallel fashion without reciprocals to study the combining ability and heterosis. None of the accessions was found to be a good general combiner for all the nine quantitative characters that were studied. Significant heterosis over better parent for single plant yield was recorded in CIS x Indian and synthetic x CIS crosses (5 each) followed by Indian x synthetic types (3). The analysis of component characters showed that the mean performance of the majority of hybrids was intermediate for five out of six yield attributing traits, thus exhibiting dominance or partial dominance effect. To estimate the contribution of such yield attributing traits towards heterosis for yield, a comparison was made among three parameters viz. heterosis over mid parent (MP), better parent (BP) and better yielding parent (BYP) of the concerned hybrid. It was observed that estimation of heterosis from BYP was a more accurate method to determine the contribution of component characters towards yield heterosis than the analysis based on MP and BP. From the component character analysis, it was concluded that characters like number of primary and secondary branches, number of siliqua per plant and siliqua density contributed significantly towards heterosis in yield. Plot level yield trials of two selected hybrids (Skorospieka II x RH30 and Donskaja IV x Varuna) over two growing seasons revealed 29.4 to 91.8% heterosis over BYP.     

The potential use of heterosis in Beta vulgaris L. II. Yield performance of pure lines and their hybrids

Summary The yield performance is described of pure lines obtained from doubled monoploids and their corresponding polycross and pair-cross hybrids. The total dry matter yields of fifteen pure lines averaged 60–90% of the yields of the progenitor and control varieties. Seven of the resultant polycross hybrids gave dry matter yields similar to or higher than the progenitor varieties, whilst the remaining hybrids were less productive than their parental lines. In pair-cross hybrids significant reciprocal differences were found in some cases. This was attributed to differences in self-fertility and the degree of self-fertilization in reciprocal crosses. Based on the frequency of offspring with white or coloured roots, self-fertilization frequencies ranging from 14% to 92% were found. It is estimated that in the event of complete cross-fertilization, the yields of certain pair-cross hybrids would have exceeded those of the control varieties by 9–29%.     

Multiple‐line cross QTL mapping for grain yield and thousand kernel weight in triticale

Grain yield and its component trait thousand kernel weight are important traits in triticale breeding programmes. Here, we used a large mapping population of 647 doubled haploid lines derived from four families to dissect the genetic architecture underlying grain yield and thousand kernel weight by multiple‐line cross QTL mapping. We identified 3 QTL for grain yield and 13 for thousand kernel weight which cross‐validated explained 5.2% and 48.2% of the genotypic variance, respectively. Both traits showed a positive phenotypic correlation, and we found two QTL overlapping between them. Full two‐dimensional epistasis scans revealed epistatic QTL for both traits, suggesting that epistatic interactions contribute to their genetic architecture. Based on QTL identified in our results, we conclude that the potential for marker‐assisted selection is limited for grain yield but more promising for thousand kernel weight.     

Using yield quantitative trait locus targeted SSR markers to study the relationship between genetic distance and yield heterosis in upland cotton (Gossypium hirsutum)

The objective of this study was to examine the relationship between heterosis and genetic distance in upland cotton (Gossypium hirsutum L.). Heterosis in the different environments was evaluated and the relationship between heterosis and genetic distance (GD) was determined based on SSR markers for yield quantitative trait loci (QTL). Yields of seed cotton and lint showed a linear relationship with mid‐parent heterosis (MPH) and better‐parent heterosis (BPH). The variation in heterosis for other traits and their correlation with GD may be due to environmental factors or the effort of QTLs tested in the present study may vary in different environments. The present findings provide a foundation for heterotic grouping of parental lines and breeding of new cotton hybrids with improved seed cotton yield. This study calls for more research with stable QTLs as well as advance molecular marker techniques may be used in predicting yield heterosis in a more precise and reliable manner.     

Hybrid performance and heterosis in spring bread wheat, and their relations to SSR-based genetic distances and coefficients of parentage

Development of hybrids is considered to be a promising avenue to enhance the yield potential of crops. We investigated (i) the amount of heterosis observed in hybrid progeny, (ii) relative importance of general (GCA) versus specific (SCA) combining ability, and (iii) the relationship between heterosis and genetic distance measures in four agronomic traits of spring bread wheat. Eight male and 14 female lines, as well as 112 hybrids produced in a factorial design were grown in replicated trials at two environments in Mexico. Principal coordinate analysis based on Rogers' distance (RD) estimates calculated from 113 SSRs revealed three different groups of parents. Mid-parent heterosis (MPH) for grain yield averaged 0.02 t ha⁻¹ (0.5%) and varied from −15.33% to 14.13%. MPH and hybrid performance (F1P) were higher for intra-group hybrids than for inter-group hybrids, with low values observed in inter-group crosses involving two non-adapted Chinese parents. Combined analyses of variance revealed significant differences among parents and among hybrids. Estimates of GCA variances were more important than SCA variances for all traits. Tight correlations of GCA with line per se performance, and mid-parent value with F1P were observed for all traits. In contrast, correlations of MPH with RD and coefficient of parentage were not significant. It was concluded that the level of heterosis in spring wheat was too low to warrant a commercial exploitation in hybrids. SSRs proved to be a powerful tool for the identification of divergent groups in advanced wheat breeding materials.     

Prediction of heterosis for grain yield in rice using 'key' informative EST-SSR markers

This study was undertaken to assess the comparative potential of 25 Expressed Sequence Tag derived simple sequence repeats (EST-SSRs) and 25 genomic SSRs in the prediction of grain yield heterosis using a set of nine cytoplasmic male sterile (CMS) lines and 32 restorer lines of rice. EST-SSRs and genomic SSRs exhibited an average Polymorphism Information Content value of 0.37 and 0.45, respectively. The coefficient of marker polymorphism among parental lines with respect to a set of hypervariable EST and genomic SSRs was correlated with standard heterosis for grain yield of six public bred rice hybrids. EST-SSRs gave a better correlation (r = 0.75) as compared with genomic SSRs (r = 0.09). When 10 'key' informative EST-SSR markers which showed a higher positive correlation with grain yield heterosis were validated in a new set of 14 experimental hybrids, the markers exhibited a higher correlation (r = 0.79), indicating the predictive value of these EST-SSRs. We recommend these 10 'key' informative EST-SSR markers for analysis of genetic diversity of parental lines and prediction of heterosis in hybrid rice breeding programmes.     

Estimates of additive and dominance effects for Fusarium head blight resistance of winter triticale

The most effective strategy to control Fusarium head blight (FHB), a devastating disease of small‐grain cereals, is breeding resistant cultivars. This resistance study of F₁ crosses, F₂ and backcross generations of triticale estimates heterosis, general and specific combining ability (GCA, SCA), additive and dominance effects and compares parents with segregating generations. The genetic material consisted of 10 parents with their 45 F₁ crosses and of six parents with their 15 F₂ progeny and backcrosses to each parent. Genotypes were grown in various environments and artificially inoculated with an aggressive isolate of F. culmorum. FHB was assessed, by visual rating, as the mean of four to five individual ratings of disease development. Heterosis for FHB was of little importance. The correlation between the FHB rating of F₁ crosses and their mid‐parent performance was close. GCA was the predominant source of variation, although the significance of the SCA variance also implied non‐additive allelic interaction. The preponderance of additive gene effects is encouraging for increasing resistance by a recurrent selection programme. The relationship between the GCA effect of a parent and its per se performance was close, which gives the possibility of predicting FHB resistance in F₁ crosses. Additive effects were predominant in the F₂ progeny and also in the backcrosses. Transgressive segregants could not be detected. Searching for them should be postponed to the F₃ or later generations.     

Agronomic performance of anther‐derived doubled haploid and single seed descent lines in crosses between primary and secondary winter triticale

Octoploid (8x) and hexaploid (6x) primary triticales (xTriticosecale Wittm.) can be used as crossing parents with secondary 6x triticales to enlarge the genetic basis of a breeding programme or introgress traits. Doubled haploid (DH) production permits to develop homozygous lines more rapidly from a segregating generation than other breeding methods such as single seed descent (SSD). Both anther‐derived DH and SSD lines were produced from reciprocal cross‐combinations between 8x and 6x primary and 6x secondary triticales. Field experiments of DH and SSD lines were conducted in three environments as two‐replicate lattices to measure seven agronomic traits. A tendency for higher grain yield, taller plants and a higher 1000‐kernel weight of SSD lines compared with DH lines was found. Significant genetic variation for all traits in both breeding methods was revealed, indicating their suitability to select superior genotypes. Hexaploid and even more so 8x primary triticales can profitably be included as crossing parents in a commercial breeding programme. In such crosses, the primary triticales should be used as the male parents if followed by DH method.     

Heterosis for leaf photosynthesis,grain yield and yield components in maize

Summary One of the primary avenues of improving the biological efficiency of crop plants is through the improvement of the leaf and canopy photosynthetic rates. However, the question whether the superiority of hybrids in respect of productivity potential could be traced, in retrospective fashion, to the photosynthetic parameters should be answered first. Once established and standardized, such indices could be streamlined in innovative breeding to predict the heterotic combinations for final yield formation. To answer this question, the photosynthetic rate and other components of photosynthetic efficiency were monitored among eight inbred stocks of maize (selected for variable photosynthetic rate from a previous study) and their all possible one-way crosses. The results demonstrated extensive heterosis in respect of photosynthetic rate and other photosynthetic indices which, in turn, was also realized in terms of higher biomass productivity and yield. Nonetheless, high leaf photosynthesis alone did not result in higher grain yield. On the contrary, component interaction among the photosynthetic indices like photosynthetic rate, leaf area/plant, number of leaves and chlorophyll content on one hand and the complementary gene action on the other, could be held responsible for higher yields in hybrids. Since the components of photosynthetic efficiency can predict heterosis for biomass and grain yield upto a reasonable extent, judicious incorporation of such indices in selection parameters for applied genetic protocols can add yet another dimension to the strategies for future yield improvements.     

Genetic analysis of rapeseed self-incompatibility lines reveals significant heterosis of different patterns for yield and oil content traits

Self‐incompatibility is one of the most effective approaches to utilizing heterosis in oilseed rape around the world. To evaluate the heterosis of double low self‐incompatibility, the possibility of combining seed yield and oil content, and the genetic effects of parents on their hybrid progenies, a 2‐year field trial using a 3 × 22 NC II mating design was conducted during the 1999‐2001 growing seasons in Wuhan, China. Significant differences in seed yield per plant and seed oil content were observed among the F1 hybrids and between F1 progenies and their parents. However, the heterosis for seed yield per plant was much greater than that for seed oil content. Mid‐parent heterosis and high‐parent heterosis of seed yield per plant ranged from 5.50 to 64.11% and from –2.81 to 46.02%, while those of seed oil content ranged from –1.55 to 7.44% and –3.61 to 6.55%, respectively. Non‐additive genetic effects were a major mechanism that accounted for the yield heterosis in addition to additive effects. In contrast, seed oil content heterosis was mainly dependent on an additive genetic effect. General combining ability (GCA) determined the stability of hybrid cultivars. In hybrid breeding, parental materials might be selected by the sum of GCAs and variances of special combining abilities (SCAs) of female and male parents for traits affected by both additive and non‐additive effects, and by the sum of GCAs of two parents for traits controlled mainly by additive effects. Primary branches and their siliques were the most important yield traits.     

Doubled haploid production in winter wheat and triticale genotypes,using the Hordeum bulbosum system

Summary An attempt was made to produce doubled haploids on 16 winter wheat and six spring and winter triticale genotypes thought to carry genes for interspecific incompatibility. The potential for haploid production was maximized by the use of Hordeum bulbosum genotypes selected for high crossability on crossable wheat genotypes, the use of two post-pollination applications of gibberellic acid and by the pollination of immature florets.A low frequency of seed was set on both the wheat and the triticale genotypes, having mean seed sets of 0.20 per cent and 0.27 per cent respectively. Although the frequency of embryos (seed quality) was high, doubled haploid production was further limited by poor embryo differentiation and regeneration. Haploid plantlets were obtained from the wheat cultivars Moulin and Renard, although successful chromosome doubling and doubled haploid production was achieved in Moulin only.