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The man-made cereal triticale was fully resistant to the biotrophic diseases powdery mildew, leaf rust, yellow rust, and stem rust from its introduction in Europe in the mid-1970s until about 1990. In the following years, new races that were able to infect at least some triticale genotypes developed in all four pathogen populations, and resistance breeding came into focus. Here, we analyzed 656 winter triticale cultivars from 12 countries for resistance to these biotrophic diseases and Fusarium head blight (FHB) at up to 8 location-year combinations (environments). FHB ratings were corrected for plant height and heading stage by comparing three statistical methods. Significant (p < 0.001) genetic variances were found for all resistances with moderate to high entry-mean heritabilities. All traits showed a normal distribution, with the exception of stem rust, where the ratings were skewed towards resistance. There were no substantial correlations among the five disease resistances (r = −0.04 to 0.26). However, several genotypes were detected with multi-disease resistance with a disease rating below average for all five diseases simultaneously. In future, such genotypes must be selected primarily to cope with future challenges of less pesticide use and global climate change.

Key message The phenomic predictive ability depends on the genetic architecture of the target trait, being high for complex traits and low for traits with major QTL. Abstract Genomic selection is a powerful tool to assist breeding of complex traits, but a limitation is the costs required for genotyping. Recently, phenomic selection has been suggested, which uses spectral data instead of molecular markers as predictors. It was shown to be competitive with genomic prediction, as it achieved predictive abilities as high or even higher than its genomic counterpart. The objective of this study was to evaluate the performance of phenomic prediction for triticale and the dependency of the predictive ability on the genetic architecture of the target trait. We found that for traits with a complex genetic architecture, like grain yield, phenomic prediction with NIRS data as predictors achieved high predictive abilities and performed better than genomic prediction. By contrast, for mono- or oligogenic traits, for example, yellow rust, marker-based approaches achieved high predictive abilities, while those of phenomic prediction were very low. Compared with molecular markers, the predictive ability obtained using NIRS data was more robust to varying degrees of genetic relatedness between the training and prediction set. Moreover, for grain yield, smaller training sets were required to achieve a similar predictive ability for phenomic prediction than for genomic prediction. In addition, our results illustrate the potential of using field-based spectral data for phenomic prediction. Overall, our result confirmed phenomic prediction as an efficient approach to improve the selection gain for complex traits in plant breeding.

The quantitatively inherited trait plant height is routinely evaluated in triticale breeding programs as it substantially influences lodging and disease susceptibility, is a main contributor to biomass yield, and is required to improve hybrid seed production by fine-tuning plant height in the female and male parental pools in hybrid breeding programs. In this study, we evaluated a panel of 846 diverse Central European triticale genotypes to dissect the genetic architecture underlying plant height by genome-wide association mapping. This revealed three medium- to large-effect QTL on chromosomes 5A, 4B, and 5R. Genetic and physical fine-mapping of the putative QTL revealed that the QTL on chromosome 5R most likely corresponds to Ddw1 and that the QTL on chromosome 5A is likely to be Rht12. Furthermore, we observed a temporal trend in registered cultivars with a decreasing plant height during the past decades, accompanied by an increasing use of the height-reducing alleles at the identified QTL. In summary, our results shed new light on the genetic control of plant height in triticale and open new avenues for future improvement by breeding.

Key message The comparably low genotype-by-nitrogen level interaction suggests that selection in early generations can be done under high-input conditions followed by selection under different nitrogen levels to identify genotypes ideally suited for the target environment. Abstract Breeding high-yielding, nitrogen-efficient crops is of utmost importance to achieve greater agricultural sustainability. The aim of this study was to evaluate nitrogen use efficiency (NUE) of triticale, investigate long-term genetic trends and the genetic architecture, and develop strategies for NUE improvement by breeding. For this, we evaluated 450 different triticale genotypes under four nitrogen fertilization levels in multi-environment field trials for grain yield, protein content, starch content and derived indices. Analysis of temporal trends revealed that modern cultivars are better in exploiting the available nitrogen. Genome-wide association mapping revealed a complex genetic architecture with many small-effect QTL and a high level of pleiotropy for NUE-related traits, in line with phenotypic correlations. Furthermore, the effect of some QTL was dependent on the nitrogen fertilization level. High correlations of each trait between N levels and the rather low genotype-by-N-level interaction variance showed that generally the same genotypes perform well over different N levels. Nevertheless, the best performing genotype was always a different one. Thus, selection in early generations can be done under high nitrogen fertilizer conditions as these provide a stronger differentiation, but the final selection in later generations should be conducted with a nitrogen fertilization as in the target environment.

Breeding exploits the phenomenon of heterosis for an improved productivity, but negative effects reduce heterosis in wide crosses., The genetics underlying heterosis, the difference in performance of crosses compared with midparents, is hypothesized to vary with relatedness between parents. We established a unique germplasm comprising three hybrid wheat sets differing in the degree of divergence between parents and devised a genetic distance measure giving weight to heterotic loci. Heterosis increased steadily with heterotic genetic distance for all 1903 hybrids. Midparent heterosis, however, was significantly lower in the hybrids including crosses between elite and exotic lines than in crosses among elite lines. The analysis of the genetic architecture of heterosis revealed this to be caused by a higher portion of negative dominance and dominance-by-dominance epistatic effects. Collectively, these results expand our understanding of heterosis in crops, an important pillar toward global food security.

Restoration of fertility in the cytoplasmic male sterility-inducing Triticum timopheevii cytoplasm can be achieved with the major restorer locus Rf3 located on chromosome 1B, but is also dependent on modifier loci. Hybrid breeding relies on a hybrid mechanism enabling a cost-efficient hybrid seed production. In wheat and triticale, cytoplasmic male sterility based on the T. timopheevii cytoplasm is commonly used, and the aim of this study was to dissect the genetic architecture underlying fertility restoration. Our study was based on two segregating F2 triticale populations with 313 and 188 individuals that share a common female parent and have two different lines with high fertility restoration ability as male parents. The plants were cloned to enable replicated assessments of their phenotype and fertility restoration was evaluated based on seed set or staining for pollen fertility. The traits showed high heritabilities but their distributions differed between the two populations. In one population, a quarter of the lines were sterile, conforming to a 3:1 segregation ratio. QTL mapping identified two and three QTL in these populations, with the major QTL being detected on chromosome 1B. This QTL was collinear in both populations and likely corresponds to Rf3. We found that Rf3 explained approximately 30 and 50% of the genotypic variance, has a dominant mode of inheritance, and that the female parent lacks this locus, probably due to a 1B.1R translocation. Taken together, Rf3 is a major restorer locus that enables fertility restoration of the T. timopheevii cytoplasm, but additional modifier loci are needed for full restoration of male fertility. Consequently, Rf3 holds great potential for hybrid wheat and triticale breeding, but other loci must also be considered, either through marker-assisted or phenotypic selection.

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.

Genome-wide association mapping of resistance against the novel, aggressive ‘Warrior’ race of yellow rust in triticale revealed a genetic architecture with some medium-effect QTL and a quantitative component, which in combination confer high levels of resistance on both leaves and ears. Yellow rust is an important destructive fungal disease in small grain cereals and the exotic ‘Warrior’ race has recently conquered Europe. The aim of this study was to investigate the genetic architecture of yellow rust resistance in hexaploid winter triticale as the basis for a successful resistance breeding. To this end, a diverse panel of 919 genotypes was evaluated for yellow rust infection on leaves and ears in multi-location field trials and genotyped by genotyping-by-sequencing as well as for known Yr resistance loci. Genome-wide association mapping identified ten quantitative trait loci (QTL) for yellow rust resistance on the leaves and seven of these also for ear resistance. The total genotypic variance explained by the QTL amounted to 44.0% for leaf and 26.0% for ear resistance. The same three medium-effect QTL were identified for both traits on chromosomes 1B, 2B, and 7B. Interestingly, plants pyramiding the resistance allele of all three medium-effect QTL were generally most resistant, but constitute less than 5% of the investigated triticale breeding material. Nevertheless, a genome-wide prediction yielded a higher predictive ability than prediction based on these three QTL. Taken together, our results show that yellow rust resistance in winter triticale is genetically complex, including both medium-effect QTL as well as a quantitative resistance component. Resistance to the novel ‘Warrior’ race of this fungal pathogen is consequently best achieved by recurrent selection in the field based on identified resistant lines and can potentially be assisted by genomic approaches.

Triticale is a promising crop for agricultural biomass production but breeding has until now mainly focused on grain yield. Here, we evaluated the potential of marker-assisted simultaneous improvement of grain yield and biomass yield. To this end, we employed a large triticale doubled haploid population with 647 individuals derived from four families that were phenotyped for grain yield and biomass yield, as well as thousand-kernel weight, tiller density, and plant height in multi-environment field trials. Employing an association mapping approach, we identified quantitative trait loci (QTL) for all the five traits. The phenotypic correlation between grain yield and biomass yield was low, and we detected only one overlapping QTL suggesting different genetic architectures underlying both traits. Our results indicate that a marker-based selection for either grain yield or biomass yield does not adversely affect the other traits. Furthermore, an improvement of the multiplicative yield traits can to some extent also be achieved by selection for QTL identified for the component traits. Taken together, our results suggest that marker-assisted breeding can assist the establishment of dual-purpose triticale cultivars with high grain and biomass yield.

Knowledge of the extent of phenotypic variability, genetic diversity and the realized breeding progress is central for the optimum design of breeding programmes, but little information is available for triticale (× Triticosecale Wittmack). In this study, a collection of 885 diverse European triticale lines was evaluated in multilocation field trials in 2 years. We observed significant genotypic variances and high heritabilities for several agronomic and morphological traits and significant correlations among different traits including grain yield. Based on a subset of the population of 121 varieties registered in Europe between 1983 and 2014, we observed a substantial breeding progress for grain yield with a significant rate of increase of 53 kg/ha or 0.67% per year. All lines were genotyped by a genotyping-by-sequencing approach yielding 58 888 polymorphic markers. Our analyses revealed the absence of major population structure but a certain grouping of lines according to their origin. Taken together, our results on triticale germplasm and its breeding history provide important information for breeding programmes and future selection gain in this crop.

Triticale (×Triticosecale Wittmack) holds great potential as a source for biomass production for industrial applications in Central Europe and hybrid breeding in particular appears promising owing to the higher vigour of hybrids compared to lines. In this study, a set of 178 winter triticale genotypes, including 91 hybrids, their 10 male and 23 female parental lines, as well as 54 varieties were evaluated for biomass yield and other agronomically relevant traits in 2 years at five locations in Germany. We observed a large variation of dry biomass yield as well as significant genotypic variances and high heritabilities for all traits. For the hybrids, a moderate correlation was observed between biomass and plant height and between biomass and grain yield. Mid-parent heterosis of biomass yield ranged from −13.6 to 16.5% with an average of 4.8%, and the maximum commercial heterosis was 9.1%. Taken together, our results illustrate the potential of hybrid breeding of triticale for biomass yield to diversify our portfolio of crops for biomass production.

BACKGROUND The influence of nitrogen fertilisation and genotype on the amino acid (AA) digestibility of triticale grain was investigated in caecectomised laying hens. Three genotypes, Grenado, EAW6002 and Lasko, were cultivated with and without nitrogen fertilisation at the end of the heading stage. The six triticale variants as well as a basal diet were each used to feed seven laying hens in a 7 × 7 Latin square design. RESULTS Nitrogen fertilisation influenced the digestibility of Cys, Glu, Phe and Ser in some triticale genotypes and reduced Ala, Ile, Lys, Met and Val digestibility in all genotypes (P < 0.05). Nitrogen fertilisation increased the concentration of all AAs in the grain. Consequently, the concentration of digestible AAs in the grains was increased for most AAs upon nitrogen fertilisation. Overall, Lys had the lowest digestibility, whereas that of Glu and Pro was the highest. For the triticale genotypes, the level of AA digestibility was highest for EAW6002 followed by Lasko and Grenado, with significant differences (P < 0.05) between genotypes for some but not all AAs. CONCLUSION The results indicated that the accuracy of the digestible AA supply for hen feeding might benefit from considering fertilisation and genotype-specific digestibility data in feed formulation. © 2016 Society of Chemical Industry.

Fusarium head blight (FHB) is one of the most serious diseases in small-grain cereals including triticale (× Triticosecale Wittmack). The disease reduces yield and accumulates mycotoxins which are harmful to human and animal health. Triticale grain is almost exclusively used on-farm in feed formulations for swine and other animals, and swine is the most susceptible farm animal to Fusarium mycotoxins. In order to evaluate the potential of genomics-assisted breeding to FHB, we performed the first genome-wide association study for FHB resistance in triticale. QTL for FHB resistance were identified on chromosomes 2A, 2B, 5B and 3R with an explained genotypic variance ranging from 0.28 to 30.23% and a total explained genetic variance of 56.64%. A QTL on chromosome 3R that explained 15.38% of the genotypic variance was identified for the first time. Association mapping was complemented by genome-wide prediction, which yielded a high prediction accuracy of 0.78 for FHB resistance when weighted genomic selection was performed. Collectively our findings highlight the potential of genomics-assisted approaches to improve Fusarium resistance in triticale in early generations.

The mechanisms of aluminum (Al) resistance in wheat and rye involve the release of citrate and malate anions from the root apices. Many of the genes controlling these processes have been identified and their responses to Al treatment described in detail. This study investigated how the major Al resistance traits of wheat and rye are transferred to triticale (x Tritosecale Wittmack) which is a hybrid between wheat and rye. We generated octoploid and hexaploid triticale lines and compared them with the parental lines for their relative resistance to Al, organic anion efflux and expression of some of the genes encoding the transporters involved. We report that the strong Al resistance of rye was incompletely transferred to octoploid and hexaploid triticale. The wheat and rye parents contributed to the Al-resistance of octoploid triticale but the phenotypes were not additive. The Al resistance genes of hexaploid wheat, TaALMT1, and TaMATE1B, were more successfully expressed in octoploid triticale than the Al resistance genes in rye tested, ScALMT1 and ScFRDL2. This study demonstrates that an important stress-tolerance trait derived from hexaploid wheat was expressed in octoploid triticale. Since most commercial triticale lines are largely hexaploid types it would be beneficial to develop techniques to generate genetically-stable octoploid triticale material. This would enable other useful traits that are present in hexaploid but not tetraploid wheat, to be transferred to triticale.

Fusarium head blight (FHB) in triticale (× Triticosecale Wittmack) results in yield losses and mycotoxin contamination, for example, by deoxynivalenol (DON). This study aimed to analyse the correlation between FHB severity and DON content in a DH population of 146 entries across environments. Additionally, Fusarium damaged kernel (FDK) rating, heading stage and plant height were recorded. Highly significant (P

Doubled haploid technology is an important tool in plant breeding and research, but routine application requires the establishment of efficient protocols. Micro- spore culture is an attractive approach although its effi- ciency is strongly dependent on the genotype. In this study we evaluated the effects of the three ethylene inhibitors aminooxyacetic acid, 2,5-norbornadiene, and silver thio- sulphate (STS) on embryogenesis, regeneration rate and green plant rate in triticale microspore culture. Our results show that STS at 20 lM in particular had a positive effect on the embryogenesis rate, suggesting that for many genotypes ethylene accumulates in the culture vessel above a critical threshold where it becomes inhibitory for embryogenesis. In addition, STS also appears to positively influence the green plant rate. Taken together, our results show that the addition of ethylene inhibitors at defined concentrations can counteract the negative effect of ethy- lene, increase the embryogenesis rate and potentially also the green plant rate in microspore culture.

Lodging tolerance is an important agronomic trait as it can have a severe negative impact on grain yield and quality. Here, we used a large mapping population of 647 doubled haploid triticale lines derived from four families to dissect the genetic architecture underlying lodging tolerance and to assess different approaches for a genomics-based improvement of the trait. The plants were evaluated for lodging in two environments and genotyped with 1710 genomewide DArT markers. We observed a large genotypic variation for lodging and transgressive segregation in all families. Employing two complementary QTL mapping approaches, we identified both main effect and epistatic QTL. Using cross-validation, we showed that the proportion of genotypic variance explained by the detected QTL is low, thus limiting the efficiency of marker-assisted selection to improve this trait. By contrast, the cross-validated predictive ability of genomic prediction was approximately twice as high as that of the QTL-based selection approaches. In conclusion, our results show that lodging tolerance is a complex trait that can be improved by classical breeding but also assisted by marker-based approaches.

Like other small grain cereals, triticale (×Triticosecale Wittmack) is also affected by Fusarium head blight (FHB) which is a serious plant disease leading to yield losses and to a contamination of feed and food by harmful mycotoxins. Use of resistant varieties is the most effective way to manage FHB infections. More recently, commercial hybrid breeding in autogamous cereals has received increased interest. Therefore, we built a bridge between hybrid and resistance breeding and conducted a multi-environmental field trial using deliberate inoculation with Fusarium culmorum to: (i) evaluate different hybrid prediction approaches based on phenotypic and/or genomic data; (ii) compare the effect of cytoplasmic male sterility (CMS) inducing cytoplasm and normal (N) cytoplasm on FHB resistance. We evaluated 226 single cross hybrids, their paternal and maternal lines in both CMS- and N-cytoplasm. Our analysis revealed a large genetic variation for FHB resistance. Single cross hybrids tended to be more resistant than their parental lines. We observed a close association between N-lines and CMS-lines for FHB resistance. We found a predominance of σ 2 over σ 2 for FHB resistance. Correlation between the sum of general combining ability (GCA) effects and hybrid performance was high for FHB resistance. Mid-parent performance served as a moderate indicator for FHB resistance of single crosses. Best linear unbiased prediction of hybrid performance based on relatedness between tested and untested genotypes estimated by marker data showed great potential for hybrid prediction when no reliable per se information or GCA estimates are available.

Fusarium head blight (FHB) is a devastating disease that causes significant reductions in yield and quality in wheat, rye, and triticale. In triticale, knowledge of the genetic architecture of FHB resistance is missing but essential due to modern breeding requirements. In our study, four doubled-haploid triticale populations (N = 120 to 200) were evaluated for resistance to FHB caused by artificial inoculation with Fusarium culmorum in four environments. DArT markers were used to genotype triticale populations. Seventeen quantitative trait loci (QTL) for FHB resistance were detected across all populations; six of them were derived from rye genome and located on chromosomes 4R, 5R, and 7R, which are here reported for the first time. The total cross-validated ratio of the explained phenotypic variance for all detected QTL in each population was 41 to 68%. In all, 17 QTL for plant height and 18 QTL for heading stage were also detected across all populations; 3 and 5 of them, respectively, were overlapping with QTL for FHB. In conclusion, FHB resistance in triticale is caused by a multitude of QTL, and pyramiding them contributes to higher resistance.

Many biologically and agronomically important traits are dynamic and show temporal variation. In this study, we used triticale (× Triticosecale Wittmack) as a model crop to assess the genetic dynamics underlying phenotypic plasticity of adult plant development. To this end, a large mapping population with 647 doubled haploid lines derived from four partially connected families from crosses among six parents was scored for developmental stage at three different time points. Using genome-wide association mapping, we identified main effect and epistatic quantitative trait loci (QTL) at all three time points. Interestingly, some of these QTL were identified at all time points, whereas others appear to only contribute to the genetic architecture at certain developmental stages. Our results illustrate the temporal contribution of QTL to the genetic control of adult plant development and more generally, the temporal genetic patterns of regulation that underlie dynamic traits.

The rye-derived dwarfing gene Ddw1 on chromosome 5R acts in triticale in considerably reducing plant height, increasing FHB severity and delaying heading stage. Triticale, an amphiploid hybrid between durum wheat and rye, is an European cereal mainly grown in Germany, France, Poland, and Belarus for feeding purposes. Dwarfing genes might further improve the genetic potential of triticale concerning lodging resistance and yield. However, they might have pleiotropic effects on other, agronomically important traits including Fusarium head blight. Therefore, we analyzed a population of 199 doubled haploid (DH) lines of the cross HeTi117-06 × Pigmej for plant height, heading stage, and FHB severity across 2 locations and 2 years. The most prominent QTL was detected on chromosome 5R explaining 48, 77, and 71 % of genotypic variation for FHB severity, plant height, and heading stage, respectively. The frequency of recovery in cross validation was ≥90 % for all three traits. Because the markers that detect dwarfing gene Ddw1 in rye are also in our population the most closely linked markers, we assume that this major QTL resembles Ddw1. For FHB severity two, for plant height three, and for heading stage five additional QTL were detected. Caused by the considerable genetic variation for heading stage and FHB severity within the progeny with the dwarfing allele, short-strawed, early heading and FHB-resistant lines can be developed when population size is large enough.

Doubled haploid (DH) production is a key technology in plant breeding and research. One emerging method of choice for DH production is microspore culture, which requires reprogramming of the microspores from their normal gametophytic development to a sporophytic development resulting in embryo formation. This commonly requires the application of stress such as cold, heat, or starvation. Here, we report the effect of different stress treatments on embryo formation and the proportion of green plants in triticale microspore culture. We observed different responses to the applied stress treatments among three studied genotypes. In general, a 3-wk cold stress treatment performed best with regard to the two criteria. For one genotype, the application of a 24- or 48-h heat stress gave similar or slightly better results and consequently may be an alternative for genotypes that are recalcitrant to the cold stress treatment.

Abiotic stress can enhance the heterogeneity of fields, which leads to imprecise estimates of genotypic effects in variety trials. Our study is based on multilocation field trials of triticale (×Triticosecale Wittm.). Two of the six locations were affected by drought stress and showed increased field heterogeneity. At Willstatt, drought stress intensity was visually scored twice, and at Issoudun, overall impression of a plot was scored once. We investigated if the visual scorings can be used as covariates in an analysis of covariance (ancova) to eliminate the influence of different drought stress intensity on plot yields. For evaluation of the ancova models, we examined if the covariates were independent from the genotypic effects and linearly associated with grain yield. In addition, ancova models were compared with baseline and spatial models based on AIC and phenotypic correlation between genotype means estimated with the model under investigation in a drought stress location with genotype means calculated across the remaining locations. We found that both scorings in Willstatt fulfilled the requirements of an ancova and led to an increase in broad-sense heritability (h2) and efficiency. ancova with the second scoring increased h2 from 0.03 for the baseline model to 0.60, whereas the best spatial model increased h2 only up to 0.50. The scoring at Issoudun was not independent from the genotypic effects and reduced phenotypic correlations. We concluded that environmental factors causing spatially differing yield potential can be scored or measured and used as covariates to obtain more precise genotypic estimates.

Triticale, an anthropogenic hybrid grain, is increasing in importance as a feed grain for laying hens. However, our limited knowledge of its nutritional qualities and their impact on hen performance prevents optimization of its use. The present study investigated the digestibility of amino acids ( AA: ) in triticale grain in laying hens, and additionally examined relationships between AA digestibility and chemical and physical characteristics of the grain. Twenty genotypes of triticale were grown under standardized agronomic and environmental conditions and were characterized according to their physical properties (thousand-seed weight, test weight, falling number, extract viscoelasticity), chemical composition (proximate nutrients, non-starch polysaccharides, AA, minerals, inositol phosphates) and gross energy concentration. Additionally, the in vitro solubility of nitrogen was determined. The animal trial comprised 4 Latin Squares (6 × 6) distributed among 2 subsequent runs. Twelve cecectomized LSL-Classic hens were individually housed in metabolism cages and either fed a basal diet containing 500 g/kg cornstarch or one of 20 triticale diets, each replacing the cornstarch with one triticale genotype, for 8 d. During the last 4 d, feed intake was recorded and excreta were collected quantitatively. Amino acid digestibility of the triticale genotypes was calculated by linear regression. The digestibility of all AA differed significantly between the 20 genotypes, including Lys (digestibility range 68 to 80%), Met (77 to 86%), Thr (68 to 78%) and Trp (74 to 83%). However, AA digestibility only correlated with characteristics of the grain in few cases, without a consistent pattern among AA. Equations to predict AA digestibility based on the grain's physical and chemical characteristics were calculated by multiple linear regression. The explanatory power (adjusted R2;) of these prediction equations was below 0.7 for most AA and thus not sufficiently precise to be suitable for practical application. In conclusion, AA digestibility of triticale grain is high overall in laying hens but varies significantly between crop genotypes. This variation could not be well explained by physical and chemical characteristics of the grain.

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.

Family mapping is based on multiple segregating families and is becoming increasingly popular because of its advantages over population mapping. Athough much progress has been made recently, the optimum design and allocation of resources for family mapping remains unclear. Here, we addressed these issues using a simulation study, resample model averaging and cross-validation approaches. Our results show that in family mapping, the predictive power and the accuracy of quatitative trait loci (QTL) detection depend greatly on the population size and phenotyping intensity. With small population sizes or few test environments, QTL results become unreliable and are hampered by a large bias in the estimation of the proportion of genotypic variance explained by the detected QTL. In addition, we observed that even though good results can be achieved with low marker densities, no plateau is reached with our full marker complement. This suggests that higher quality results could be achieved with greater marker densities or sequence data, which will be available in the near future for many species.

Predicting single-cross performance is of special interest in hybrid breeding of triticale. We used molecular and phenotypic data of factorial triticale crosses and compared several approaches to predict their single-cross performance. Twenty-three inbred lines and their 76 incomplete factorial crosses were field evaluated for grain yield, plant height, and heading time at five locations in Central Europe. In addition, the parental lines were genotyped with 52 SSR markers. Plant height and heading time were predicted with high accuracy based on mid-parent performance. In contrast, prediction of hybrid perfor- mance based on mid-parent value was not accurate for grain yield. Using general combining ability effects led to an enhanced prediction accuracy of hybrid grain yield performance. This accuracy could be slightly improved using best linear unbiased prediction approaches. The prediction accuracy was considerably high even if the number of tested hybrids was small. Consequently, best linear unbiased prediction of hybrid performance is a promising tool for hybrid triticale breeding programs.

Hybrid breeding in autogamous cereals has a long history of attempts with moderate success. There is a vast amount of literature investigating the potential problems and solutions, but until now, market share of hybrids is still a niche compared to line varieties. Our aim was to summarize the status quo of hybrid breeding efforts for the autogamous cereals wheat, rice, barley, and triticale. Furthermore, the research needs for a successful hybrid breeding in autogamous cereals are intensively discussed. To our opinion, the basic requirements for a successful hybrid breeding in autogamous cereals are fulfilled. Nevertheless, optimization of the existing hybridization systems is urgently required and should be coupled with the development of clear male and female pool concepts. We present a quantitative genetic framework as a first step to compare selection gain of hybrid versus line breeding. The lack of precise empirical estimates of relevant quantitative genetic parameters, however, is currently the major bottleneck for a robust evaluation of the potential of hybrid breeding in autogamous cereals.

Root system architecture (RSA) is seldom considered as a selection criterion to improve yield in maize breeding, mainly because of the practical difficulties with their evaluation under field conditions. In the present study, phenotypic profiling of 187 advanced-backcross BC(4)F(3) maize lines (Ye478 × Wu312) was conducted at different developmental stages under field conditions at two locations (Dongbeiwang in 2007 and Shangzhuang in 2008) for five quantitative root traits. The aims were to (1) understand the genetic basis of root growth in the field; (2) investigate the contribution of root traits to grain yield (GY); and (3) detect QTLs controlling root traits at the seedling (I), silking (II) and maturation (III) stages. Axial root (AR)-related traits showed higher heritability than lateral root (LR)-related traits, which indicated stronger environmental effects on LR growth. Among the three developmental stages, root establishment at stage I showed the closest relationship with GY (r = 0.33-0.43, P 0.001). Thirty QTLs for RSA were detected in the BC(4)F(3) population and only 13.3 % of the QTLs were detected at stage III. Most important QTLs for root traits were located on chromosome 6 near the locus umc1257 (bin 6.02-6.04) at stage I, and chromosome 10 near the locus umc2003 (bin 10.04) for number of AR across all three developmental stages. The regions of chromosome 7 near the locus bnlg339 (bin 7.03) and chromosome 1 near the locus bnlg1556 (bin 1.07) harbored QTLs for both GY- and LR-related traits at stages I and II, respectively. These results help to understand the genetic basis of root development under field conditions and their contribution to grain yield.

Joint linkage association mapping (JLAM) combines the advantages of linkage mapping and association mapping, and is a powerful tool to dissect the genetic architecture of complex traits. The main goal of this study was to use a cross-validation strategy, resample model averaging and empirical data analyses to compare seven different biometrical models for JLAM with regard to the correction for population structure and the quantitative trait loci (QTL) detection power. Three linear models and four linear mixed models with different approaches to control for population stratification were evaluated. Models A, B and C were linear models with either cofactors (Model-A), or cofactors and a population effect (Model-B), or a model in which the cofactors and the single-nucleotide polymorphism effect were modeled as nested within population (Model-C). The mixed models, D, E, F and G, included a random population effect (Model-D), or a random population effect with defined variance structure (Model-E), a kinship matrix defining the degree of relatedness among the genotypes (Model-F), or a kinship matrix and principal coordinates (Model-G). The tested models were conceptually different and were also found to differ in terms of power to detect QTL. Model-B with the cofactors and a population effect, effectively controlled population structure and possessed a high predictive power. The varying allele substitution effects in different populations suggest as a promising strategy for JLAM to use Model-B for the detection of QTL and then to estimate their effects by applying Model-C.

Winter triticale, an abundant biomass source for producing biogas in Central Europe, has been bred until now for increased grain yield but not for early biomass yield. Therefore, our objectives were to assess the feasibility of breeding triticale for simultaneous improvement of early biomass and grain yield as well as to predict the early biomass yield using an index of traits measured at maturity stage. A representative sample of 100 triticale lines was evaluated in Experiment I at four locations in Germany. Traits were measured at two growth stages (early dough and full maturity). Genetic variances were significantly different from zero for all traits. The correlated response for early biomass yield estimated from grain yield (0.53) was high, which points to the efficiency of indirect selection. A multiple regression model was developed to predict early biomass yield. The reliability of the model was confirmed in an independent Experiment II with 49 lines. The calibration model explained 55% of total variation for early biomass yield. This clearly underlines that at least during earlier stages of selection, field testing can be carried out for grain yield and related traits to select potential genotypes with high biomass yield based on the developed regression model.

Hybrid breeding facilitates a maximum exploitation of heterosis. The objectives of our research were to (i) examine the magnitude of heterosis over the mid- and better parent for yield, yield components, and quality traits in Central European spring durum wheat and (ii) investigate the efficiency of parental selection based on mid-parent value or general combining ability effects. Sixteen inbred lines and 40 incomplete factorial crosses were field-evaluated for eight agronomic traits at 10 environments in Germany. For grain yield, the hybrids yielded on average 10% higher than the mid-parent performance, and the maximum superiority was 22%. The significantly positive contrast between inbreeding depression and mid-parent heterosis for grain yield indicated the presence of positive additive x additive epistatic interactions. Furthermore, the best hybrid outperformed the best line variety by 1.01 Mg ha ―1 . We conclude that there is a potential for hybrid breeding in durum wheat.

Resistance to Fusarium head blight (FHB) is of great importance in wheat breeding programs in the northern hemisphere. In Europe, breeders prefer adapted germplasm as resistance donor because of high grain yield and quality demands. Our objective was to identify chromosomal regions affecting FHB resistance among 455 European soft winter wheat (Triticum aestivum L.) lines using a genome-wide association mapping approach and to analyze the importance of epistatic interactions. All entries were evaluated for FHB resistance by inoculation in two environments and several ratings. Wheat was genotyped by 115 simple sequence repeat markers randomly distributed across the genome and two allele-specific markers for Rht-B1 and Rht-D1 genes. The genome-wide scan revealed nine significant (P < 0.05) marker–phenotype associations on seven chromosomes including dwarfing gene Rht-D1. Using a Bonferroni–Holm correction, three significant associations remained on chromosomes 1B, 1D, and 2D. The proportion of the genotypic variance explained simultaneously by individual markers was 36% and increased to 50% when two digenic epistatic interactions were considered, one of them associated with Rht-B1. In conclusion, new genomic regions on chromosomes 1D and 3A could be found for FHB resistance in European wheat and the effect of epistatic interactions was substantial.

Heterotic groups are an essential prerequisite for effi cient hybrid breeding. The present study in winter triticale (×Triticosecale Wittmack) was performed to (i) evaluate the magnitude of heterosis, (ii) investigate the effi ciency of the prediction of hybrid performance based on midparent value or general combining ability (GCA) effects, and (iii) identify two heterotic groups by employing principle coordinate analysis and an enumeration algorithm for maximizing F 1 performance, midparent heterosis, and σ 2 GCA / σ 2 SCA ratio for grain yield. Twenty-one inbred lines and their 210 diallel crosses were fi eld- evaluated for grain yield at fi ve agroecologically diverse locations in Germany. On average, the hybrids yielded 8.6% higher than the midparent performance, and the maximum superiority was 12.4%. Hybrid performance was predicted more reliably with GCA effects rather than with midparent performance. An enumeration algorithm based on optimum allocation of parents to heterotic groups, improved all three criteria. F 1 performance, midparent heterosis, and σ 2 GCA /σ 2 SCA ratio increased by about 3, 24, and 70%, respectively, as compared with the base situation of having no heterotic groups. These improvements substantiate the potential advantages, when heterotic groups are developed in hybrid breeding.

The objective of our study was to examine the influence of genetic divergence on the ratio of the components of variance for specific (σ 2 SCA ) and general (σ 2 GCA ) combining ability using experimental data in triticale (xTriticosecale Wittm.). In total, 21 lines and their 210 crosses were evaluated for grain yield in field trials. Published molecular data were reanalyzed, indicating an optimum of two subgroups. The estimates of σ 2 SCA and σ 2 GCA were determined for the total diallel and between the two subgroups. The ratio of σ 2 SCA VS. σ 2 GCA tended to be lower for crosses between than within genetically distinct groups. Our experimental findings can be interpreted as an indicator of a more favorable ratio of σ 2 SCA vs. σ 2 GCA in situations with two genetically distinct populations than in situations with populations that are not genetically distinct.