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.     

Combining ability and correlations for yield components in early generations of potato breeding

A partial diallel set of crosses was made between 14 potato cultivars chosen for their fertility, from those included in a potato breeding programme at the NEIKER – Basque Institute for Agricultural Research. The progeny were grown in completely randomized trials from 1997 to 1999. Performance for yield, tuber number and average tuber weight was analysed in seedling and two clonal generations. Variance estimates due to both general combining ability (GCA) and specific combining ability (SCA) were significant in all generations for all traits under study. However, SCA was more important than GCA in almost all cases. Correlation coefficients among characters, generations, GCA and SCA effects were examined. For tuber yield no relation was obtained between generations; however, average tuber weight and yield were positively associated in all generations. The results indicate that appropriate selection criteria depend strongly on the particular cross. The implication for a breeding strategy are discussed.     

New techniques for breeding maize (Zea mays) varieties with fall armyworm resistance and market-preferred traits for sub-Saharan Africa

Deploying maize varieties with fall armyworm (Spodoptera frugiperda [J.E. Smith]; FAW) resistance, desirable product profiles (PPs) and climate resilience is fundamental for food and economic security in sub-Saharan Africa (SSA). This study reviewed and identified challenges and opportunities for effective and accelerated breeding of demand-led maize hybrids with FAW resistance and adaptation to the diverse agro-ecologies of SSA. Lessons were drawn on improving breeding efficiency through adequate genetic variation delivered via prebreeding programmes, speed breeding and a reduced breeding stage plan. Appropriate PPs aligned with demand-led breeding approaches were highlighted as foundations for variety design and commercialization. Challenges to accelerated FAW resistance breeding in maize included inadequate funds and modern tools; poor adaptation of some exotic donor parental lines; lack of information on FAW resistance among local varieties; lack of integration of molecular markers associated with FAW resistance and agronomic traits into selection plans; and limited infrastructure for FAW rearing and germplasm screening. Integration of modern breeding tools and scientific innovations were recommended for accelerated development and release of FAW resistant and market-preferred maize varieties.     

Combining ability of maize inbred lines containing genes from Zea diploperennis for resistance to Striga hermonthica (Del.) Benth

Maize ( Zea mays ) is a staple food crop in sub-Saharan Africa, but its production is threatened by Striga hermonthica (Del.) Benth. Transfer of resistance genes from wild relatives may increase resistance to S. hermonthica in tropical maize. The objective of this study was to determine the combining ability of resistance to S. hermonthica among lines containing Zea diploperennis and tropical germplasm. Forty-five diallel crosses of 10 inbred lines were evaluated in an alpha-lattice design with and without artificial Striga infestation at two locations each in the Republic of Benin and Nigeria for 3 years. Results of analyses showed that only general combining ability (GCA) mean square was significant (P = 0.01) for number of emerged Striga plants (NESPP), while both GCA and specific combining ability (SCA) mean squares were significant for host damage score (HDS) and grain yield under Striga infestation. The ratio of GCA to SCA mean squares for the three traits varied from 3.5 to 57.5. Although GCA × environment interaction was significant for the three traits, two inbred lines containing Z. diploperennis (ZD 551) and tropical (TZL TC 87) germplasm had negative and significant GCA effects for NESPP and HDS and positive GCA effects for grain yield under Striga infestation in the two countries. Correlation between NESPP and HDS was strong and significant ( r  = 0.87, P = 0.01). Our results highlight the importance of harnessing useful genes from wild relatives to improve resistance to S. hermonthica in adapted maize germplasm.     

Combining ability for beet armyworm, Spodoptera exigua, resistance and horticultural traits of selected Lycopersicon pennellii-derived inbred backcross lines of tomato

Ten selected inbred backcross lines (IBL), from a Lycopersicon esculentum cv.‘Peto 84’×Lycopersicon pennellii IBL population, with resistance to beet armyworm (BAW), Spodoptera exigua, higher fruit mass and fruit yield, were crossed with eight elite cultivated L. esculentum inbred lines in a Design II mating design. Three elite inbreds were also crossed to ‘Peto 84′, the IBL recurrent parent, as a control for combining ability. Field plots of all resulting F₁ progenies and control cultivars were inoculated with BAW eggs and evaluated for resistance to BAW, fruit mass, fruit yield, vine size and maturity at three field locations. Reductions in fruit damage by BAW were found in four of the 10 IBL F₁ progenies. Significant male and female general combining ability (GCA) estimates for BAW resistance were observed, but significant specific combining ability for BAW resistance was not detected. The fruit mass of F₁ hybrids was significantly lower than large-fruited controls, but was not significantly different from elite inbred by ‘Peto 95’F₁ hybrids. Selection based on inbred performance identified IBL with positive GCA for BAW resistance and yields in Design II hybrids. BAW resistance in the 09 selected IBL and IBL- derived F₁ progeny was associated with two undesirable traits, later maturity and larger vine size. Index selection of IBL was more effective at identifying IBL with positive GCA for fruit mass and fruit yield than GCA for BAW resistance.     

Combining ability of common bean (Phaseolus vulgaris) genotypes for root traits across diverse environments

Root system architecture is important for common bean (Phaseolus vulgaris) adaptability to diverse environments. Beans employ complex adaptive root mechanisms for coping with multiple stresses in production environments. Understanding genetic control of root traits is central to improvement of common bean for adaptation to marginal environments. The objectives of this study were to (i) determine combining ability of root and agronomic traits and (ii) estimate the heritability and genetic correlation of root and agronomic traits in common bean. Four bean lines with superior root traits were crossed with four locally adapted varieties in a North Carolina II mating scheme to generate 16 crosses. The 16 F₁s were selfed and advanced to F₂ generation. Eight parents and their F₂ progenies were evaluated in an alpha-Lattice design with two replications. General and specific combing ability mean squares were significant (p ≤ .05) for all traits measured. General predictability ratios ranged from .47 to .68 across locations suggesting that both additive and non-additive gene action modulate root traits and seed yield. Positive and significant (p ≤ .05) phenotypic and genetic correlations revealed significant association between root traits and yield. Moderate to high heritability estimates of between .43 and .67 were realized. Such estimates point to possible deployment of a successful selection programme. Genotype AFR398 displayed significant positive GCA effects among its crosses for both root and agronomic traits hence a potential candidate genotype for inclusion in a bean genetic improvement programme for marginal environments.     

Combining ability for seed yield traits of Chenopodium quinoa breeding lines

For a better understanding of the inheritance of seed yield traits in Quinoa (Chenopodium quinoa) Willd., a half-diallelic crossing experiment with six diverse but uniform breeding lines was conducted. True hybrid plants were detected by means of differences in panicle colour in 14 out of the 15 crosses performed. The agronomic performance of 14 F₂ populations and six parental lines was evaluated in field trials on fertile clay soil. General and specific combining ability of the lines (GCA and SCA) were estimated. Highly significant differences in GCA effects were found between the lines for plant height at maturity, early flowering, early maturity, seed yield and thousand seed weight. Significant SCA effects were only found for plant height at maturity. However, SCA effects were rather small and accounted for 7% of the total variance. The finding of large GCA effects and low SCA effects suggests that best selection results could be expected from crosses between the agronomic best performing genotypes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genomic prediction of sunflower hybrid performance

Predicting single‐cross performance is of high importance to improve the efficiency of sunflower (Helianthus annuus L.) hybrid breeding programmes. We used experimental data from inter‐ and intragroup sunflower hybrids and their parental lines adapted to Central Europe to (i) study the genetic diversity and combining ability and (ii) examine the accuracy to predict hybrid performance based on phenotypic and genomic data. We evaluated 133 intragroup and 104 intergroup hybrids with their parental lines in replicated trials at four environments for grain yield, oil yield and oil content. Furthermore, the parental lines were fingerprinted with 572 AFLP markers. Variance due to specific combining ability was comparable for intergroup and intragroup crosses. This suggested a lack of clearly defined heterotic groups for the sample of studied sunflower lines. Prediction accuracy of hybrid performance based on general combining ability effects was high and could not be increased using genomic selection approaches. For situations where no information on GCA effects of parental lines was available, hybrid prediction based on genomic selection methods was accurate for groups of related lines. For groups of unrelated lines, however, we observed a strong decrease in the prediction accuracy. This suggests that prediction of hybrid performance for crosses based on genetically distant parents remains challenging.     

Combining ability and patterns of inheritance for plant and fruit traits in Japanese quince (Chaenomeles japonica)

Phenotypic variation and pattern ofinheritance was investigated for plant andfruit traits in Japanese quince (Chaenomeles japonica, Maloideae). Japanesequince is a minor fruit crop in Latvia andLithuania, and is presently being developedfor production of juice, aroma and fruitfibre in Sweden and Finland. General andspecific combining ability was estimatedfor five plant vegetative traits, eightfruit yield and morphology traits, and fivefruit biochemistry traits in a studypopulation consisting of 14 hybridfamilies. Thorniness seems to be controlledprimarily by a single locus. All othertraits studied appear to be governedprimarily by additive genes, as judged fromgeneral combining ability estimates.Significant specific combining ability(SCA) was detected for yield, plant heightand some fruit traits. However, therelative contribution of SCA to the overallgenetic variance was in general low. Anefficient breeding strategy for Japanesequince could therefore be based onrecurrent selection.     

Identifying quantitative trait loci for the general combining ability of yield-relevant traits in maize

Maize is the most important staple crop worldwide. Many of its agronomic traits present with a high level of heterosis. Combining ability was proposed to exploit the rule of heterosis, and general combining ability (GCA) is a crucial measure of parental performance. In this study, a recombinant inbred line population was used to construct testcross populations by crossing with four testers based on North Carolina design II. Six yield-relevant traits were investigated as phenotypic data. GCA effects were estimated for three scenarios based on the heterotic group and the number of tester lines. These estimates were then used to identify quantitative trait loci (QTL) and dissect genetic basis of GCA. A higher heritability of GCA was obtained for each trait. Thus, testing in early generation of breeding may effectively select candidate lines with relatively superior GCA performance. The GCA QTL detected in each scenario was slightly different according to the linkage mapping. Most of the GCA-relevant loci were simultaneously detected in all three datasets. Therefore, the genetic basis of GCA was nearly constant although discrepant inbred lines were appointed as testers. In addition, favorable alleles corresponding to GCA could be pyramided via marker-assisted selection and made available for maize hybrid breeding.     

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.     

Trends in genetic variance components during 30 years of hybrid maize breeding at the University of Hohenheim

The ratio of variance due to specific vs. general combining ability (GCA) (σ²_SCA:σ²_GCA) is of central importance for predicting hybrid performance from GCA effects. The objectives of our study were to (1) analyse the changes in estimates of σ²_GCA, σ²_SCA and their ratio during 30 years of hybrid maize breeding and (2) compare the observed trends in genetic variances with those expected under a simple genetic model. We analysed multilocation yield trials based on the North Carolina Design II conducted in the maize breeding programme of the University of Hohenheim from 1975 to 2004 for grain yield (GY) and dry matter content (DMC). GY showed a significant (P < 0.05) annual increase of 0.17 Mg/ha, but no linear trend was found for DMC. Since the beginning of hybrid breeding at the University of Hohenheim, the sum of estimates of σ²_GCA of the flint and dent heterotic groups were higher than the estimates of their σ²_SCA. This predominance did not change with ongoing inter‐population improvement. Consequently, superior hybrids can be identified and selected mainly based on their prediction from GCA effects.     

印水型杂交粳稻产量及产量构成因素的配合力分析

为确定新育成印水型杂交粳稻亲本的潜力,并为印水型杂交粳稻育种提供理论基础,本研究以4个不育系和4 个恢复系为亲本进行不完全双列杂交,研究产量及其构成因素的配合力。结果表明：产量性状一般配合力(GCA)方差均达到极显著水平,除结实率外,父本的一般配合力方差大于母本,单株产量、有效穗和结实率的特殊配合力(SCA)方差达到显著或极显著水平。139A和晚轮422 的GCA效应较高,产量性状以Ⅱ类为主,139A×晚轮422 和辽粳10A×晚轮422 的单株产量水平及组合SCA效应较高。父本GCA效应和组合SCA效应对产量性状的影响较大,母本GCA效应对产量性状的影响相对较小。本研究中,父本的遗传差异大于母本,对产量性状的影响也大于母本,139A和晚轮422 有较大的应用潜力。     

Combining ability for rancidity and associated traits in pearl millet (Pennisetum glaucum)

Pearl millet occupies an important place in attaining nutritional security in marginal areas; however, as it develops off-flavours, it is less preferred by food industry. The objective of the current study was to determine the variations in rancidity-related traits and estimate the combining ability of inbreds and hybrids for these traits under varied environmental conditions. In this study, 32 hybrids were developed from eight lines and four testers using line × tester mating design and evaluated along with parents and checks 86M86 and HHB 67 for yield and rancidity-associated traits in three environments. The hybrids L7T2 and L6T1 are identified as promising hybrids for grain yield and rancidity. Higher variance due to specific combining ability and predictability ratio for grain yield and rancidity-associated characters indicate the predominance of non-additive gene action. The tester T1 recorded a significant negative GCA effect with low alcoholic acidity. A significant and positive SCA for grain yield was observed in L3T1 and L7T2, whereas for rancidity in L5T4. Alcoholic acidity showed a significant positive association with 1000 seed weight, lipase and lipoxygenase in parents and negative association with 1000 seed weight in hybrids.     

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.     

Heterosis and combining ability for plant height and its components in hybrid wheat with Triticum timopheevi cytoplasm

The heterosis and combining ability for plant height and its components of hybrid wheat were investigated in an incomplete diallel experiment including 5 CMS lines and 4 restorer lines. The results showed that heterosis (HS) and heterobeltiosis (HBS) occurred in plant height (PH) and length of the first internode (LFI), second internode (LSI), third internode (LTI), basal internode (LBI) and the spike (LS) of hybrids, but their values varied among crosses and characters; the HS and HBS of LBI were larger than those of other characters, the HS and HBS of LSI and LTI contributed a lot to those of PH. There were significant relationships between internode lengths and PH for specific combining ability (SCA) and general combining ability (GCA), and among lengths of the adjacent internodes for SCA and/or GCA effects. However, the relationships of LS with the lengths of internodes and PH were insignificant for GCA, SCA, HS and HBS. The SCA effects were more important than GCA effects for LFI, the reverse was true for LSI, LTI, LS and PH, and the SCA effects was nearly equal to the GCA effects for LBI. So, LFI was mainly influenced by non-additive effect of genes, while LSI, LTI, LS and PH were mainly controlled by additive gene effects, LBI was controlled equally by additive and non-additive effects of genes. The genes that control the length of specific internode not only affect PH, but also the length of the adjacent internode. The genetic system in charge of lengths of internodes and plant height is independent of that for length of spike. Thus, it is possible to develop new wheat cultivars or hybrid combinations having long spike but dwarf plant height. This revised version was published online in July 2006 with corrections to the Cover Date.     

The breeding values of potato parents for field resistance to late blight measured by whole seedlings

Summary Parents used in a potato breeding programme were intercrossed in a multiple mating scheme. The resulting progenies were assessed for resistance to late blight, using as criteria the size and frequencies of lesions on leaves, on petioles and on stems, and an overall score. Analysis of variance showed that all the statistically significant genetic variation was attributable to general combining ability (GCA) differences. The seven variates were highly correlated.     

A diallel crossing approach aimed on selection for ripening time and yield in breeding of new strawberry (Fragaria × ananassa Duch.) cultivars

The extension of the ripening season in open field production is of high economic interest for strawberry growers. Therefore, targeted breeding for extreme early or late ripening cultivars with high yield potential is of particular interest. Thirteen strawberry cultivars were crossed in a reciprocal way without selfing, and the 144 resulting F₁ populations were evaluated in a field trial over a period of two consecutive years. The data were analysed using a mixed‐model approach adapted for diallel crossing designs using SAS 9.3. The variability in the crossing approach is mainly based on the general combining ability (GCA) of the cultivars (additive effects). Specific and reciprocal combining abilities (non‐additive effects) appear less important. The highest GCAs for the trait Marketable Yield were found for the cultivars ‘Polka’ and ‘Yamaska’. The trait Earliness is bilateral with significantly low GCAs for early ripening in ‘Clery’ and ‘Daroyal’ and significantly high GCAs for late ripening in ‘Yamaska’ and ‘Florence’. Crosses with these cultivars are likely to deliver populations with both high yield and an extended ripening period.     

羽衣甘蓝主要观赏性状配合力与遗传力分析

旨在为观赏羽衣甘蓝的杂种优势利用与优良新品种选育提供科学依据。以羽衣甘蓝2个雄性不育系为母本,7个高代自交系为父本,按NCⅡ不完全双列杂交设计,配制了14个杂交组合,对株高、外叶数、心叶数、外叶开展度、心叶开展度、外叶叶形指数、心叶叶形指数7个观赏性状进行了配合力和遗传力的分析。结果表明：对于一般配合力,母本C6BZ在心叶数和外叶开展度的表现优于CD10;父本中,HZ1631和DFZ在心叶数、外叶开展度和心叶开展度的一般配合力较高。根据特殊配合力效应值选出较优良的4个杂交组合为CD10×DFZ、CD10×HZ1631、C6BZ×HZ1631和C6BZ×F0819。7个性状的广义遗传力均较高,均在50%以上。外叶叶形指数和心叶叶形指数的狭义遗传力均高于50%,这2个性状可在早期进行选择;而其他5个性状的狭义遗传力均较低,需在高代选择。通过本试验得出的参试母本系和自交系的一般配合力和特殊配合力,以及7个重要观赏性状的广义遗传力和狭义遗传力,可为利用杂种优势选育出优良的观赏羽衣甘蓝杂交种提供依据。     

Resistance breeding strategy for Stenocarpella maydis and Fusarium graminearum cob rots in tropical maize

Maize cob rot caused by Fusarium graminearum and Stenocarpella maydis affects grain yield and quality. The objective of this study was to investigate the appropriateness of multiple infection as a selection and breeding strategy for multiple resistance to F. graminearum and S. maydis. Twelve tropical inbred lines with varying resistance to either or both pathogens were mated in a full diallel and the progeny and their parents evaluated for reaction to single or multiple infection. Under multiple inoculation, S. maydis suppressed colonization of cobs by F. graminearum. General combining ability (GCA) effects indicated that inbred WL 118–10 effectively transmitted resistance to both diseases. Hybrids' resistant to S. maydis was also resistant to F. graminearum, but the reverse was not true. Therefore, efficient screening should initially involve screening for S. maydis followed by F. graminearum. Overall, the suppression of F. graminearum by S. maydis shows that multiple infection cannot be used as an appropriate breeding strategy to obtain multiple resistance. The use of F. graminearum and S. maydis separately is, therefore, the best breeding strategy.