Genomic selection in hybrid breeding

While hybrid breeding is widely applied in outbreeding species, for many self‐pollinating crop plants, it has only recently been established. This may have had its reason in the limitations of methods available for hybrid performance prediction, in particular when established heterotic pools were absent. Genomic selection has been suggested as a promising approach to resolve these limitations. In our review, we briefly introduce the principles of genomic selection as an extension of marker‐assisted selection using genome‐wide high‐density molecular marker data and discuss the advantages and limitations of currently used algorithms. Including the outcome from a recent extended approach to hybrid wheat as a timely example, we summarize current progress in empirical studies on the application of genomic selection for prediction of hybrid performance. Here, we put emphasis on the factors affecting the accuracy of prediction, pointing in particular to the relevance of relatedness, genotype x environment interaction and experimental design. Finally, we discuss future research needs and potential applications.     

Genetic relatedness and the ratio of subpopulation‐common alleles are related in genomic prediction across structured subpopulations in maize

Genomic prediction (GP), which could predict the breeding value of crop plants genotyped with molecular markers, has been carried out in multiple species. Prediction accuracy (PA) of GP depends on various factors, including genetic relatedness and genetic basis. In this study, we examined the rationale for the low PA of GP when the training and validation populations were distinct using 170 temperate inbred lines and 210 tropical and subtropical inbred lines, respectively. All inbred lines were evaluated for 17 traits and genotyped with 550K high‐density markers. The results show that: (a) the influences of heritability and marker number on PA reflected variations in phenotypic variance captured by the genetic information; (b) the low PA of GP when the training and validation populations represent structured subpopulation is related to the ratio of subpopulation‐common alleles (RSCA) and the genetic relatedness between the two subpopulations; (c) RSCA and PA increased with the increase of genetic relatedness, suggesting that these three factors were related. Our findings would provide references when performing GP, and guidance when designing breeding populations.     

Accelerated development of quality protein maize hybrid through marker‐assisted introgression of opaque‐2 allele

Vivek Maize Hybrid 9‐ a popular single‐cross hybrid developed by crossing CM 212 and CM 145 was released for commercial cultivation in India. The parental lines, being deficient in lysine and tryptophan, were selected for introgression of opaque‐2 allele using CML 180 and CML 170 as donor lines through marker‐assisted backcross breeding. The opaque‐2 homozygous recessive genotypes with >90% recovery of the recurrent parent genome were selected in BC₂F_2, and the seeds with <25% opaqueness in BC₂F₃ were forwarded for seed multiplication. Vivek Quality Protein Maize (QPM) 9, the improved QPM hybrid, showed 41% increase in tryptophan and 30% increase in lysine over the original hybrid. The grain yield of the improved hybrid was on par with the original hybrid. The newly improved QPM maize hybrid released in 2008 will help in reducing the protein malnutrition because its biological value is superior over the normal maize hybrids. This short duration QPM maize hybrid has been adopted in several hill states of North Western and North Eastern Himalayan regions.     

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.     

The potential of genomic‐assisted breeding to improve Fusarium head blight resistance in winter durum wheat

Durum wheat is the most important tetraploid wheat mainly used for semolina and pasta production, but is notorious for its high susceptibility to Fusarium head blight (FHB). Our objectives were to identify and characterize quantitative trait loci (QTL) in winter durum and to evaluate the potential of genomic approaches for the improvement of FHB resistance. Here, we employed an international panel of 170 winter and 14 spring durum lines, phenotyped for Fusarium culmorum resistance at five environments. Heading date, plant height and mean FHB severity showed significant genotypic variation with high heritabilities and FHB resistance was negatively correlated with both heading date and plant height. The dwarfing gene Rht‐B1 significantly affected FHB resistance and the genome‐wide association scan identified eight additional QTL affecting FHB resistance, explaining between 1% and 14% of the genotypic variation. A genome‐wide prediction approach yielded only a slightly improved predictive ability compared to marker‐assisted selection based on the four strongest QTL. In conclusion, FHB resistance in durum wheat is a highly quantitative trait and in breeding programmes may best be tackled by classical high‐throughput recurrent phenotypic selection that can be assisted by genomic prediction if marker profiles are available.     

A model calculation approach towards the optimization of a standard scheme of seed‐parent line development in hybrid rye breeding

The importance of hybrid rye (Secale cereale L.) breeding has steadily increased over the last decades. This paper presents results of model calculations aiming to optimize the number of candidates, testers to assess combining ability and test locations at each selection stage of a standard scheme of seed‐parent line development. Two variants of the scheme differing in the number of stages and the cycle length are investigated. Optimization criterion is the expected selection gain per year under the restriction of a fixed budget. Prediction of selection gain rests on quantitative–genetic and economic parameters estimated from breeders’ data. Optimization covers different genetical and economical situations. Results show that the optimum number of testers to assess combining ability depends on the relative amount of dominance variance. The efficiency of a breeding scheme strongly increases with decreasing cycle length. A larger budget should mainly be used to increase the number of candidates at all selection stages. Recommendations for practical breeding schemes are given. We conclude that model calculations are a valuable tool for the optimization of breeding schemes.     

Detection of donor effects in a rye introgression population with genome‐wide prediction

Introgression populations are developed to make genetic resources for breeding purposes available. In the case that the number of donor segments exceeds the number of lines, genome‐wide prediction (GWP) methods are suggested as promising for the analysis of such populations. Our objectives were to characterize a rye introgression population with the Rye5K SNP assay and to apply a GWP model with a modification of the restricted maximum likelihood procedure that yields heteroscedastic variances to detect significant donor effects. The introgression lines (ILs) carried on average 4.6 donor segments with a mean length of 27 cM and represented 94% of the donor genome. Two donor effects were detected that significantly increased thousand‐kernel weight. We found four donor effects for protein, total pentosan and starch content that can improve baking quality. Three donor effects for protein content were observed for improving feeding purposes and one donor effect for starch content to improve ethanol production. The effects were localized to small genomic regions. Consequently, these ILs can improve rye breeding by directly employing them in breeding programmes for variety development.     

Development of AS‐PCR marker based on a key mutation confirmed by resequencing of Wx‐mp in Milky Princess and its application in japonica soft rice (Oryza sativa L.) breeding

Soft rice with low amylose content (AC) ranging by 5–15% is a unique type with special eating and appearance quality and has become popular in the rice market. We resequenced the Wx‐mp, a key allele from Milky Princess, a Japanese low AC variety, and found that the +473 mutation in exon 4 is the key mutation in both Wx‐mp and its ancestor allele, Wx‐mq from Milky Queen. Based on this functional mutation, an allele‐specific PCR (AS‐PCR) marker was developed and proven in a breeding population derived from a cross between a Chinese late variety Nan Keng 46 (Wx‐mp/Wx‐mp) and an early line Ning 63121(Wx‐b/Wx‐b). Based on the marker‐aided selection by our newly developed AS‐PCR marker for Wx‐mp and the known ST10 marker for Stvb‐i, a total of 12 Wx‐mp homozygotes were selected from 198 F₂ progenies, and four of them were immune to rice stripe virus (RSV) with averagely 11.3 days earlier heading than Nan Keng 46 without significant change in grain yield.     

Construction of chromosomal segment substitution lines and genetic dissection of introgressed segments associated with yield determination in the parents of a super‐hybrid rice

Heterosis is a phenomenon whereby hybrids of inbred lines produce favourable phenotypes that exceed those of their parents. Traits of interest are higher yield and stronger stress tolerance. The two‐line super‐hybrid rice ‘Liangyoupei9’ (LYP9) shows superiority to both its elite inbred line ‘93‐11’ and ‘Pei'ai64s’ (‘PA64s’) parents and conventional hybrids. However, the genetic basis of its hybrid vigour, especially yield determination, remains elusive. In the present study, a set of 156 chromosome segment substitution lines (CSSLs) carrying overlapping segments from ‘PA64s’ in a genetic background of ‘93‐11’ were constructed and planted in six environments. Three major agronomic traits, viz. panicle length (PL), heading date (HD) and plant height (PH), and five yield‐related traits, viz. grain weight per panicle (GWP), number of grains per panicle (GPP), 1000‐grain weight (TGW), seed set (SS) and number of panicles of per plant (PPP), were evaluated over 3 years. Quantitative trait loci (QTL) analysis was conducted using a likelihood ratio test based on stepwise regression. Forty‐six putative QTL distributed on 11 chromosomes were detected in more than one year. Remarkably, GWP of four CSSLs carrying positive yield QTL outperformed the recurrent parent ‘93‐11’ by more than 15%, in at least two environments. These results indicate that CSSLs are effective in identifying yield‐associated traits, and lines harbouring such QTL will be rich in resources for future molecular breeding programmes.