Accuracy of within‐ and among‐family genomic prediction in triticale

Genomic prediction has emerged as a powerful genomic tool to assist breeding of complex traits. In this study, we employed a population of 647 triticale doubled haploid lines derived from four families to assess the potential of this approach for triticale breeding. All lines were phenotyped for grain yield, thousand‐kernel weight, biomass yield, plant height, frost tolerance and Fusarium head blight resistance. The obtained prediction accuracies were moderate to high and consisted to varying degrees of within‐ and among‐family variance, in line with the different degrees of phenotypic differences between family means. The prediction accuracy within individual families also varied with the genetic complexity of the traits and was generally highest based on effect estimation with lines from the respective family, whereas the prediction accuracy decreased with decreasing relatedness among the families. Taken together, our results illustrate the potential of genomic prediction to increase selection gain in triticale breeding, but the composition of the training set is of utmost importance, and consequently, the implementation of this approach in applied breeding programmes is not straightforward.     

Modified full-sib selection and best linear unbiased prediction of progeny performance in a European F2 maize population

Four cycles of modified recurrent full‐sib (FS) selection were conducted in an intermated F₂ population of European flint maize. The objectives of our study were to monitor trends across selection cycles in the estimates of population mean, inbreeding coefficients and variance components, and to investigate the usefulness of best linear unbiased prediction (BLUP) of progeny performance under the recurrent FS selection scheme applied. We used a selection rate of 25% for a selection index, based on grain yield and dry matter content. A pseudo‐factorial mating scheme was used for recombination. In this scheme, the selected FS families were divided into an upper‐ranking group of parents mated to the lower‐ranking group. Variance components were estimated with restricted maximum likelihood (REML). Average grain yield increased 1.2 t/ha per cycle, average grain moisture decreased 20.1 g/kg per cycle, and the selection index relative to the F₂ check entries decreased 0.3% per cycle. For a more precise calculation of selection response, the four cycles should be tested together in multi‐environmental trials. We observed a significant decrease in additive variance in the selection index, suggesting smaller future selection response. Predictions of FS family performance in Cn + 1 based on mean performance of parental FS families in Cn were of equal precision as those based on the mean additive genetic BLUP of their parents, and corresponding correlations were of moderate size for grain moisture and selection index.     

Identification and characterization of new S‐alleles associated with self‐incompatibility in almond

Almond is a highly heterozygous species with a high number of S‐alleles controlling its gametophytic self‐incompatibility system (GSI). In this work, we have analysed 14 Spanish local almond cultivars for S‐RNase allele diversity. Five new S‐RNase alleles were identified by cloning and sequencing, S₃₁ (804 bp) in ‘Pou de Felanitx’ and ‘Totsol’, S₃₂ (855 bp) in ‘Taiatona’, S₃₃ (1165 bp) in ‘Pou d’Establiments’ and ‘Muel’, S₃₄ (1663 bp) in ‘Pané‐Barquets’ and S₃₅ (1658 bp) in ‘Planeta de les Garrigues’. Additionally, seven already known almond alleles could be recognized in the local cultivars studied. The high number of new alleles identified reveals the wide diversity of almond germplasm still existing and requiring characterization, and points to the possibility of new findings by a wider study focusing on other provenances. The almond S‐RNases have been compared to those of other Prunus species, showing a high identity and confirming that the S‐RNase gene in this genus presents a probable common ancestor.     

Choice of shrinkage parameter and prediction of genomic breeding values in elite maize breeding populations

Genomic selection (GS) is a promising alternative to marker‐assisted selection particularly for quantitative traits. In this study, we examined the prediction accuracy of genomic breeding values by using ridge regression best linear unbiased prediction in combination with fivefold cross‐validation based on empirical data of a commercial maize breeding programme. The empirical data is composed of 930 testcross progenies derived from 11 segregating families evaluated at six environments for grain yield and grain moisture. Accuracy to predict genomic breeding values was affected by the choice of the shrinkage parameter λ², by unbalanced family size, by size of the training population and to a lower extent by the number of markers. Accuracy of genomic breeding values was high suggesting that the selection gain can be improved implementing GS in elite maize breeding programmes.     

Identification of novel alleles from wild barley for the improvement of alpha‐amylase and related malt quality traits

Germplasms consisting of 64 wild barley accessions (Hordeum vulgare ssp. spontaneum) were examined for polymorphisms in α‐amylase using both isoelectric focusing (IEF) and thermostability assays. Wide variation was found for the high pI α‐amylase with 20 IEF band patterns identified. Enzyme activity and thermostability assays showed large differences among α‐amylase isoenzymes. Two wild accessions Tel‐Shoket CPI 77146‐32 and Afiq CPI 77128‐41 showed superior enzyme activity and thermostability compared with commercial varieties such as ‘Baudin’, ‘Flagship’ or ‘Navigator’. The functionality of the Tel‐Shoket allele was validated in backcross lines with ‘Flagship’ as the recurrent parent. The Tel‐Shoket allele at the amy1 locus increased α‐amylase thermostability at 75°C by 8.4% and α‐amylase activity in kilned malt by 18.7%. The introgression of the wild allele also led to significant improvements in fermentability, hot water extract and viscosity. Gene sequencing showed that there are three single nucleotide polymorphisms in the Tel‐Shoket amy1 sequence, which can be used as diagnostic markers for the selection of this allele in breeding programmes.     

Mitigating the impact of selective phenotyping in training populations on the prediction ability by multi-trait pedigree and genomic selection models

Training populations for pedigree and genomic prediction in plant breeding programmes are largely updated with superior genotypes from multi-environment trials, where they are tested with the goal of variety development. Such a selective phenotyping has, however, a negative impact on prediction abilities, especially when only a subset of breeding lines can be tested, as for laborious and costly to phenotype traits. This study focused, thus, on investigating the impact of selective phenotyping in the training population of an applied wheat breeding programme, and assessing the potential to mitigate this impact by pedigree and genomic multi-trait prediction models as well as bi-directionally selected training populations for several baking quality parameters. Combining both pedigree and genomic information in multi-trait prediction models with pre-existing phenotypic information for protein content and sedimentation value compensated for the observed loss in prediction ability, while entering few inferior breeding lines into the training population further mitigated the impact of selective phenotyping and even led to a slight increase of prediction ability in comparison to a randomly chosen training population.     

Genetic architecture of plant height in maize phenotype‐selected introgression families

This study aimed at developing, characterizing and evaluating two maize phenotypic‐selected introgression libraries for a collection of dominant plant height (PHT)‐increasing alleles by introgressing donor chromosome segments (DCS) from Germplasm Enhancement of Maize (GEM) accessions into elite inbred lines: PHB47 and PHZ51. Different backcross generations (BC₁‐BC₄) were developed and the tallest 23 phenotype‐selected introgression families (PIFs) from each introgression library (PHB47 or PHZ51) were selected for single nucleotide polymorphism genotyping to localize DCS underlying PHT. The result shows that most PIFs carrying DCS were significantly (α = 0.01) taller than the respective recurrent parent. In addition, they contained larger donor genome proportions than expected in the absence of selection or random mating across all BC generations. The DCS were distributed over the whole genome, indicating a complex genetic nature underlying PHT. We conclude that our PIFs are enriched for favourable PHT‐increasing alleles. These two libraries offer opportunities for future PHT gene isolation and allele characterization and for breeding purposes, such as novel cultivars for biofuel production.     

Genetic variation of the rate of leaf appearance in maize: Possible yield prediction at the early stage

Summary Evidence of genetic variation for early vigour is presented using maize line × tester crosses. The leaf appearance rate and associated variance components are affected by a physiological stress attributable to the transition to autotrophic nutrition by the plant. At this stage, specific combining ability plays an important role in the total genetic variation. It is concluded that differing genetic controls exist in early and late material, and that this difference is also manifested in leaf initiation and elongation rates. The ground coverage rate, as a component of plant development, is genetically correlated to total dry matter yield. The genetic correlations vary according to the growth stage.     

Detection of epistatic and environmental interaction QTLs for leaf orientation‐related traits in maize

Leaf architecture traits in maize are quantitative and have been studied by quantitative trait loci (QTLs) mapping. However, additional QTLs for these traits require mapping and the interactions between mapped QTLs require studying because of the complicated genetic nature of these traits. To detect common QTLs and to find new ones, we investigated the maize traits of leaf angle, leaf flagging‐point length, leaf length and leaf orientation value using a set of recombinant inbred line populations and single nucleotide polymorphism markers. In total, 19 QTLs contributed 4.13–13.52% of the phenotypic effects to the corresponding traits that were mapped, and their candidate genes are provided. Common and major QTLs have also been detected. All of the QTLs showed significant additive effects and non‐significant additive × environment effects in combined environments. The majority showed additive × additive epistasis effects and non‐significant QTL × environment effects under single environments. Common and major QTLs provided information for fine mapping and gene cloning, and SNP markers can be used for marker‐assisted selection breeding.     

Identification,distribution and effects on agronomic traits of the semi-dwarfing <Emphasis Type="Italic">Rht</Emphasis> alleles in Bulgarian common wheat cultivars

Bulgarian common wheat cultivars released in the period 1925–2003 were studied using the gibberellic acid (GA) test and microsatellite analysis of the Xgwm261 locus on chromosome 2DS to identify the semi-dwarfing (Rht) genes. The old cultivars, isolated through selection from landraces, carried rare alleles (211- and 215-bp) at Xgwm261 locus, and those developed by hybridisation to foreign cultivars, carried the 165- and 174-bp alleles. Forty-two (55.3%) of 76 modern cultivars were GA-responsive. The 192-bp allele, diagnostic for Rht8, was observed in 64 (84.2%) modern cultivars, of which 37 carry Rht8 alone, and 27 possess a combination of Rht8 and a GA-insensitive allele viz. Rht-B1d (17); Rht-D1b (6) and Rht-B1b (4). The 174-bp allele is present in seven cultivars, only one of which is photoperiod-sensitive, and the rest are day-length insensitive. The 203-bp allele was found in six modern cultivars. Cultivars carrying the Rht8 allele are the most widespread and some of them have been cultivated for a long period. Cultivars with the `Saitama 27' allele (Rht-B1d) are the most productive and are second in distribution in the country. The recently observed trend for increasing the proportion of cultivars with GA-insensitive Rht genes is probably due to their combination with the 192-bp allele of Xgwm261 locus tightly linked to the Ppd-D1, to the break of the link between the 174-bp allele and ppd-D1, and to the introduction of other genes influencing flowering time.     

Influence of rare alleles of β‐carotene hydroxylase and lycopene epsilon cyclase genes on accumulation of provitamin A carotenoids in maize kernels

Vitamin A deficiency is one of the major health problems worldwide. Traditional yellow maize possesses very low provitamin A (proA) concentration in endosperm. The influence of rare alleles of β‐carotene hydroxylase (crtRB1) and lycopene epsilon cyclase (lcyE) genes capable of enhancing proA concentration was studied in four BC₂F₂ populations generated using subtropical inbreds and CIMMYT‐HarvestPlus lines. The occurrence of severe segregation distortion for crtRB1 gene was observed, while lcyE gene was segregated as per Mendelian ratio. Genotype with favourable allele of crtRB1 (CC) had a significant effect on β‐carotene (BC) (7.9‐fold), β‐cryptoxanthin (BCX) (twofold) and proA (5.5‐fold) accumulation, compared to unfavourable genotype (C⁺C⁺). Genotype with favourable allele of lcyE (LL) showed 2.1‐fold, 1.6‐fold and twofold significant enhancement in BC, BCX and proA, respectively, over unfavourable genotype (L⁺L⁺) in pooled analysis. Of the nine genotypes, double homozygote (CC/LL) had the highest mean BC (12.60 μg/g), BCX (4.44 μg/g) and proA (14.82 μg/g), and combined effect was significantly better than individual gene effects or any other combinations. The information generated here would be useful in designing strategy for proA enrichment in subtropical maize.     

Dissection of the genetic architecture for grain quality‐related traits in three RIL populations of maize (Zea mays L.)

To manipulate the composition of the maize kernel to meet future needs, an understanding of the molecular regulation of kernel quality‐related traits is required. In this study, the quantitative trait loci (QTL) for the concentrations of grain protein, starch and oil were identified using three sets of RIL populations in three environments. The genetic maps and the initial QTL were integrated using meta‐analyses. A total of 38 QTL were identified, including 15 in population 1, 12 in population 2 and 11 in population 3. The individual effects ranged from 2.87% to 13.11% of the phenotypic variation, with seven QTL each contributing over 10%. One common QTL was found for the concentrations of grain protein and starch in bin 3.09 in the three environments and the three RIL populations. Of the 38 initial QTL, 22 were integrated into eight mQTL by meta‐analysis. mQTL3 and mQTL8 of the key mQTL in which the initial QTL displayed R² > 10% included six and three initial QTL for grain protein and starch concentrations from two or three populations, respectively. These results will provide useful information for marker‐assisted selection to improve the quality of the maize kernel.     

Mapping and validation of the quantitative trait loci for leaf stay‐green‐associated parameters in maize

Functional stay‐green is generally regarded as a desirable trait of varieties in major crops including maize. In this study, we used an F_3:4 recombinant inbred line population with 165 lines from a cross between a stay‐green inbred line (Zheng58) and a model inbred line (B73) using 211 polymorphic simple sequence repeat markers to map quantitative trait loci for three stay‐green‐associated parameters, chlorophyll content, photosystem II photochemical efficiency and stay‐green area, at maturity stage, detected a total of 23 quantitative trait loci (QTL) on nine chromosomes. Single QTL explained 3.7–13.5% of the phenotypic variance. Additionally, we validated some important stay‐green QTL using a heterogeneous inbred family approach and found that the stay‐green‐associated parameters were significantly correlated with the plant yield. This study may contribute to a better insight into the regulatory mechanism behind leaf stay‐green in maize and a novel development of elite maize varieties with delayed leaf senescence through molecular marker‐assisted selection.     

Genetic association of within‐boll yield components and boll morphological traits with fibre properties in upland cotton (Gossypium hirsutum L.)

Knowledge of genetic relationships between within‐boll yield components and fibre quality is essential for simultaneous improvement of lint yield and fibre quality in upland cotton (Gossypium hirsutum L.). Nine parents and their F₁ progeny with reciprocals from a 3 × 6 factorial mating design were grown in 2008 and 2009. Seven within‐boll yield components and two boll morphological traits and the three fibre quality parameters were analysed based on a conditional additive/dominance (AD) genetic model. Results showed that boll length contributed to the largest proportion of phenotypic, additive and dominance variances for UHM length; seed index contributed to the largest proportion of phenotypic and additive variances for fibre strength; boll width made the largest contribution to phenotypic and additive variances and the second largest contribution to dominance‐by‐environment interaction variance for micronaire, indicating that they played an important role than the other traits for fibre length, strength and fineness/maturity, respectively. It is worthy of note that those correlations between boll shape and fibre quality apply only to the nine parents and the resultant hybrids in this study and do not imply a cause and effect relationship.     

Exogenous 6‐benzyladenine improves antioxidative system and carbon metabolism of summer maize waterlogged in the field

A field experiment was performed to study the effects of spraying exogenous 6‐benzyladenine (6‐BA) after waterlogging for 6 days at the third leaf stage and the sixth leaf stage on the activities of key enzymes in carbon metabolism, leaf gas exchange parameters, fluorescent characteristic, antioxidant enzymes activities and MDA content of summer maize hybrids Denghai605 (DH605) and Zhengdan958 (ZD958), in order to illuminate the regulations of exogenous spraying 6‐BA to antioxidative system and carbon metabolism of waterlogged summer maize. Results showed that exogenous application of 6‐BA effectively alleviated the adverse effects of waterlogging on carbon metabolism by improving photosynthetic characteristics and the activities of RuBPCase (in the range of 48%–81%) and PEPCase (in the range of 14%–46%). Treatments with spraying exogenous 6‐BA resulted in significant and substantial increases in net photosynthetic rate (around 49%) and Φ_PSII (around 16%) compared to those of waterlogging treatments. In addition, spraying 6‐BA effectively alleviated waterlogging damages on antioxidative enzyme activity and decreased MDA content, delaying leaf senescence, resulted in significant and substantial (around 19%) increases in grain yield. Visibly, spraying exogenous 6‐BA effectively alleviated waterlogging damages on antioxidative system and carbon metabolism of summer maize, resulting in yield increase in waterlogged maize.     

Converting restriction fragment length polymorphism to single‐strand conformation polymorphism markers and its application in the fine mapping of a trichome gene in cotton

A well‐characterized and systematically organized collection of genetic markers is crucial in the study of any crop species. It is the basis of map‐based gene cloning and crop improvements through marker‐assisted selections. Single‐strand conformation polymorphism (SSCP) has been a robust way of discovering new polymorphisms in marker development without the requirement of sequencing. Here, we report the first approach of applying SSCP marker discovery methods in the genetic map construction and gene mapping of cotton species. A total of 80 restriction fragment length polymorphism (RFLP) markers were selected from a region on published cotton genetic maps around the T1 gene related to cotton trichome. Among the 80 RFLPs, 28 showed polymorphisms through SSCP, showing a polymorphic rate of approximately 35%, which is much higher than that of simple sequence repeat (SSR) markers in the same region (7.8%). By integrating these newly generated SSCP markers, a detailed genetic map was reconstructed around this region using an F₂ population derived from a cross between Gossypium arboreum and G. herboceum. The reconstructed region comprises 22 SSCP markers, eight SSR markers and the T1 gene, spanning 21.6 cM. The marker order of the new map agrees well with published reference RFLP maps. The above results suggest that SSCP method can be applied very efficiently and reliably to the marker development of cotton genomes. It will prove to be even more valuable and robust after the public release of cotton whole‐genome sequences.     

Alleles on the two dwarfing loci on 4B and 4D are main drivers of FHB‐related traits in the Canadian winter wheat population “Vienna” × “25R47”

Fusarium head blight (FHB), leaf rust and stem rust are among the most destructive wheat diseases. High‐yielding, native disease resistance sources are available in North America. The objective of this study was to map loci associated with FHB traits, leaf rust, stem rust and plant height in a “Vienna”/”25R47” population. DArT markers were used to generate a genetic map, and quantitative trait loci (QTL) analysis was performed by evaluating 113 doubled haploid lines across three environments in Ontario, Canada. FHB resistance QTL were identified on chromosomes 4D, 4B, 2D and 7A, while a QTL for leaf and stem rust resistance was identified on chromosome 1B. The dwarfing alleles of both Rht‐B1 and Rht‐D1 were associated with increased FHB index and DON content.