Genetic variation in testcrosses of European maize inbreds for resistance to the European corn borer and relations to line per se performance

The European com borer (ECB) Ostrinia nubilalis H. is a major pest in World maize Zea mays L. production. Objectives of this study were to (1) investigate the genotypic variance and covariance in testcrosses of European flint and dent inbreds for ECB resistance and agronomic traits, and (2) estimate the correlation between line per se and testcross performance for ECB resistance traits. A total of 16 flint and 24 dent inbreds and their testcrosses with two testers from the opposite germplasm pool were evaluated in four and three German environments, respectively. Using artificial infestation with ECB larvae, resistance was assessed by damage rating of stalks, tunnel length in dissected stalks, and relative yield of infested plots compared with protected plots. Yield losses due to ECB damage in testcross hybrids amounted to 40%. Significant genotypic variances between flint and dent lines and high heritabilities were found for damage rating of stalks for both line per se and testcross performance. Heritabilities were low or intermediate for tunnel length and relative grain yield. Correlations between line per se and testcross performance were tight for the damage rating of stalks and moderate for tunnel length and relative yield in both flint and dent germplasm. For damage rating of stalks, per se performance of lines tested in a few environments can be used to predict their testcross performance. In contrast, assessment of testcross performance for tunnel length and relative yield requires evaluating testcrosses with several testers in multi-environment trials.     

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.     

Variation and covariation for quantitative resistance to head blight (Fusarium culmorum) in two testcross series of S2 lines in winter rye

The amount of genetic variation among inbred lines and testcrosses, and covariation between both genetic materials, are of crucial importance for selection efficiency in hybrid breeding. To estimate these quantitative genetic parameters for resistance of winter rye (Secale cereale) to head blight caused by Fusarium culmorum, 88 three-way cross hybrids, produced by crossing each of 44 S₂ Carsten inbred lines with two unrelated Petkus single-cross testers, were evaluated along with the parental lines over 2 years. Resistance traits were head-blight rating and grain weight per spike relative to the non-inoculated control. Significant genotypic variation occurred among lines and in both testcross series. S₂ lines displayed considerably more variation than testcross series. Genotype × environment interaction was more marked among the inbred lines, while estimates of heritability were similar for both genetic materials. Testcrosses showed heterosis for head-blight resistance. No relationship existed between S₂ lines and the two testcross series for any resistance trait. This might be caused by an association between inbreeding and Fusarium-head-blight susceptibility and different inbreeding depression among the S₂ population. The phenotypic correlations between the testcross series were moderate for both traits (r = 0.58, P < 0.01). In conclusion, Fusarium-head-blight resistance has to be selected at the respective heterozygosity levels.     

Relationships between line per se and testcross performance for agronomic traits in two broad-based populations of maize

Degree of the association between line per se performance (LP) and testcross performance (TP) is important in breeding programs and simultaneous improvement of commercial hybrids and their parental lines. This experiment was designed to study genetic variability and genetic correlation for several agronomic traits in two maize (Zea mays L.) broad-based populations (NS12-SG and NS14-SG). Independent trials with 80 entries of S₁ progenies as well as their testcrosses were arranged according to an incomplete block design with replicates in sets. Grain yield, stay green, anthesis-silking interval, stalk water content and grain moisture were evaluated in four environments. The anthesis-silking interval had the highest genetic variation, followed by stay green. High heritability estimates (>0.50) for all traits, pointed out that further selection would be possible. Genetic correlations between line per se and testcross performance were lowest for grain yield (0.396** and 0.592**, for NS12-SG and NS14-SG, respectively), and highest for grain moisture (0.937** and 0.821**, respectively). High correlations between line per se and their testcrosses for stay green, anthesis-silking interval, stalk water content and grain moisture indicated that additive gene action might be more important than dominance in controlling the expression of these traits.     

Quantitative genetic studies on breeding maize for adaptation to organic farming

Organic farming has gained in importance in Germany during recent years. Therefore an increasing demand exists for varieties with specific adaptation to this farming system. In the present study we therefore conducted comparative field experiments with modern maize breeding materials under organic versus conventional farming conditions (ORG and CON, respectively) to estimate quantitative genetic parameters needed for developing optimal breeding strategies and to investigate the perspectives of selection for specific adaptation to ORG. Starting from two broad samples of elite germplasm, consisting of 178 flint and dent lines, respectively, fractions of 11 flint and 11 dent lines were selected based on their testcross performance under ORG. A corresponding set of lines was selected under CON. Testcross performance was evaluated in three regions of Germany and selection of superior lines was practiced across two stages in 2004 and 2005, respectively. The specifically selected lines were crossed in a factorial manner for production of experimental inter-pool single-cross hybrids which were field-tested under ORG and CON in two regions in 2006. Average grain yields were about 16% lower under ORG than under CON. Variance components and entry-mean heritability coefficients under ORG largely resembled those obtained under CON. Phenotypic correlations between ORG and CON were moderate for grain yield and strong for grain dry matter content. No consistent estimates were obtained for the corresponding genotypic correlation for grain yield. At the first stage of testcross selection no evidence of specific adaptation to ORG or CON was observed whereas the factorial crosses tested in 2006 displayed distinct, yet non-significant, advantages when evaluated under the respective target farming system. A small top fraction of hybrids showed outstanding performance under both ORG and CON. The chances of detecting such broadly adapted genotypes are increased if ORG test sites are included in the regular testing system.     

Vernalization response and earliness per se in cultivars representing different eras of wheat breeding in Argentina

Argentine wheat cultivars are assumed to be essentially vernalization insensitive or very slightly sensitive. However, there are only speculations on this lack of vernalization requirement and a greater unawareness on the variation in earliness per se. The aims of this research have been to determine the extent of variability in vernalization requirement and earliness per se, and how the variability in both traits was produced by breeding programs, through the release of wheat cultivars from the 1930's to the 1990's in Argentina. Sixty-eight cultivars, selected among those of highest performance in each era, were evaluated under field and glasshouse conditions for their vernalization response and earliness per se. Forty per cent of the cultivars showed some vernalization response. There was a decrease in this requirement along the first decades of the analysed breeding period, likely in response to the considerable introgression of CIMMYT germplasm. This initial trend to release earlier cultivars was also evidenced in a clear decrease in earliness per se. As this tendency in both characteristics was reverted during the last two decades, it may denote that certain vernalization response and not extreme earliness per se, may contribute to achieve higher yield cultivars in our area. This revised version was published online in July 2006 with corrections to the Cover Date.     

Towards Enhancement of Early‐Stage Chilling Tolerance and Root Development in Sorghum F1 Hybrids

Sorghum (Sorghum bicolor L. Moench) is regarded a drought‐tolerant alternative to maize as a bioenergy and fodder crop, but its early‐stage chilling sensitivity is obstructing a successful implementation in temperate areas. While several studies have identified quantitative trait loci (QTL) underlying chilling tolerance‐related traits in sorghum lines, little is known about the inheritance of these traits in F₁ hybrids. We have conducted a comprehensive approach to analyse heterosis, combining ability and the relation between line per se and hybrid performance for emergence and early shoot and root development comprising both field trials and controlled environment experiments including chilling tests. To our best knowledge, this is the first study analysing heterosis for sorghum root parameters under chilling. Our results show that most traits are heterotic and that the mid‐parent values are rather poor predictors of hybrid performance. Hybrid breeding programmes should focus on efficient GCA tests and the establishment of genetically diverse pools to maximise heterosis rather than on a too strict selection among lines based on their per se performance. The medium‐to‐high heritabilities estimated for seedling emergence and juvenile biomass suggest that a robust breeding progress for these complex traits is feasible.     

Construction of a crop—wild hybrid population for broadening genetic diversity in cultivated sunflower and first evaluation of its combining ability: the concept of neodomestication

The genetic base of sunflower elite lines is very narrow, due to many years of selection and breeding. To broaden the genetic diversity of the cultivated sunflower, in 1995 73 wild sunflower populations were crossed with 3 cultivated lines (Testers), and 219 hybrid offspring’s were evaluated in the field. GCA and SCA effects were computed suggesting for all traits a genetic potential for improvement through selection. Study of the hybrids revealed that the wild accessions bear different genetic abilities to combine with the testers for traits of morphological architecture, phenology and yield (seed weight and seed oil). The variance due to GCA and SCA showed that gene action was additive for days to flowering, branching and plant height. Genotypes derived from the same geographic origin may have either good or poor general combing ability. The correlation between GCA and per se genotype performance was positive for all traits except for seed oil content. This was the first attempt to evaluate wild-cultivated hybrids in sunflower on a large scale and will be the starting point for the management of hybrid Helianthus annuus populations for breeding. GCA and SCA estimations will facilitate the definition of strategies to manage and exploit the natural diversity for this crop.     

Adaptation of tropical maize germplasm to temperate environments

Maize (Zea mays L.) is one of few crops that can offer significant genetic gains with the utilization of genetic diversity. Genetically broad-based germplasm has the potential to contribute useful and unique alleles to U.S. Corn Belt breeding programs not present in current U.S. genome sequences (e.g. B73, NAM, etc.). Our objectives were to determine if unique tropical genetic materials have been effectively adapted to temperate environments and how their agronomic performance was relative to adapted populations. An important long-term objective of the Iowa and North Dakota maize breeding programs has been, in addition to the typical elite by elite line pedigree selection cultivar development process, to adapt exotic and unique germplasm, maximize their genetic improvement, and develop unique products for breeding and commercial uses. Stratified mass selection methodology for earliness has been utilized for the adaptation of tropical and temperate populations to Iowa and North Dakota environments. This method has allowed screening of up to 25,000 genotypes per population cycle at a rate of one cycle per year. In addition, the estimated cost per year our programs had for the adaptation of each population was less than $2,000 which could successfully be applied in any breeding program across the globe. This cost has been less than 1 % of the total cost for finding minor genes on the same trait. Our results showed the successful adaptation of exotic populations was independent from genetic background. We can speculate there are a few major genes responsible for most of flowering date expression. We encourage the use of technology to target traits according to their genetic complexity. Stratified mass selection at the phenotypic level has been successful. Each of the populations with either 25 of 100 % tropical germplasm are available for anyone who may desire to expand the germplasm base of their breeding programs with tropical germplasm adapted to temperate mid- and short-season U.S. Corn Belt environments.     

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.     

Doubled haploids in tropical maize: <Emphasis Type="SmallCaps">II</Emphasis>. Quantitative genetic parameters for testcross performance

Single crosses (SC) of elite inbreds and open-pollinated populations (OP) are suitable source germplasm for doubled haploid (DH) line development in hybrid maize breeding, given that they combine a high population mean ([`(x)] \overline{x} ) for testcross performance with adequate response to selection ( \Updelta G \Updelta G ). This is the first study reporting testcross grain yield (TCGY) and dry matter content (TCDMC) evaluations of 131 DH lines developed from ten tropical source germplasm comprising five OP (OP1–OP5) and five SC (SC1–SC5). Gene diversity (d) and the average number of alleles (a <sub>r</sub> ) per locus was estimated for DH lines based on 24 simple sequence repeat markers. Analysis across three environments revealed no significant differences between [`(x)] \overline{x} of OP- and SC-derived DH lines for TCGY and TCDMC. Significant genetic variance for both traits was only detected among OP-derived DH lines which may be explained by a larger number of segregating quantitative trait loci (QTL) as suggested by higher d and a _r values than in SC-derived DH lines. The usefulness criterion ( U = [`(x)] + \Updelta G U = \overline{x} + \Updelta G ) was higher for OP-derived DH lines for TCDMC, but higher for SC-derived DH lines for TCGY. DH lines from OP1, OP2, and OP3 showed high TCGY, suggesting that they may be useful in tropical hybrid breeding. We conclude that tropical OP represent a valuable source of untapped genetic variation that can efficiently be exploited with DH technology for hybrid maize breeding.     

Optimization of DH-line based recurrent selection procedures in maize under a restricted annual loss of genetic variance

Doubled haploid (DH) lines are increasingly being used in commercial hybrid maize (Zea mays L.) breeding. They allow for various quantitative genetic and logistic advantages provided that they are implemented in efficient and optimally allocated breeding procedures. In the present study, a new software was applied to optimize two recurrent selection (RS) schemes for hybrid maize breeding based on DH lines under a restricted annual budget and an upper limit for the relative annual loss of genetic variance. This software maximizes the expected gain from selection in general combining ability by means of quantitative genetic model calculations. Optimization results are compared for one, two, and three stages of testcross selection under different assumptions regarding the evaluation of lines per se and the annual budget. Results show that the optimum allocation of technical and budget resources to the individual steps of an RS program and the efficiency of alternative RS procedures are decisively determined by the number of selection stages. Under standard assumptions, one-stage selection was superior to two- and three-stage selection. Thus, shortening the length of an RS scheme considerably increases its efficiency. By intercrossing a reduced number of selected lines for starting a new RS cycle, the short-term response to selection may be increased, but the population size and, thus, the selection limits in the long run are diminished. Therefore, fair comparisons of alternative RS procedures require to define the intended time span for maximizing the genetic gain from RS and to restrict the relative annual loss of genetic variance accordingly.     

Identification of genomic regions carrying QTL for agronomic and quality traits in rye (Secale cereale) introgression libraries

Introgression libraries can be used to localize genomic regions carrying quantitative trait loci (QTL). We used this approach to detect QTL regions affecting the per se performance of agronomic and quality traits with two rye (Secale cereale L.) introgression libraries. Our objectives were to detect candidate introgression lines (pre‐ILs) that have a different per se performance than the recurrent parent and to identify the underlying QTL regions. The introgression libraries containing 40 BC₂S₃ lines each were established with marker‐assisted backcrossing from crosses of the heterozygous Iranian primitive rye accession Altevogt 14160 and the elite inbred line L2053‐N. To assess the phenotypic effect of the donor chromosome segments (DCS) the pre‐ILs were evaluated for grain yield, plant height, thousand‐kernel weight, test weight, falling number and protein content in replicated field trials at five locations in Germany over 2 years. In total, 58 significant (P < 0.05) differences between pre‐ILs and L2053‐N were observed in each introgression library. The DCS in pre‐ILs differing from the recurrent parent possess most likely the responsible QTL. Genomic regions carrying favourable QTL alleles were detected for test weight, thousand‐kernel weight and protein content. We conclude that Altevogt 14160 can not only be used to enrich the genetic variation of the restricted hybrid rye gene pools but will also allow the breeder to efficiently detect favourable QTL for marker‐assisted selection.     

Farmers’ participation and breeding for durable disease resistance in the Andean region

In the Andean region, the Preduza project and its partners combined breeding for durable disease resistance using locally adapted cultivars and farmer participatory methods. The approach taken resembles participatory variety selection (PVS). Farmers participated in the selection of advanced materials, rather than finished cultivars. This paper describes this approach and reports experiences with farmers–breeders collaboration. As breeders involved farmers as participants, they learned more about the most important criteria of male and female farmers for preferred cultivars in the marginal environments of Andean cropping systems. This approach encouraged the use of locally adapted cultivars (often landraces), made the breeders less dependent on foreign materials, and has resulted in selection and development of new wheat, barley, common bean, quinoa, potato and maize cultivars. Breeding programmes based on crossing locally adapted cultivars followed by selection by the breeders in the early phases of the breeding programmes and by participatory selection with the farmers in the more advanced stages of the breeding programmes appeared successful. It became clear that breeders must be well acquainted with the farmer preferences such as the requirements for specific agronomic, storage, processing and marketing traits. Over a period of five years the centralized formal breeding approach predominantly based on material produced by the international institutes was replaced by decentralized breeding approaches based largely on local germplasm with extensive farmer participation.     

花期玉米(Zea mays)穗三叶叶形结构的遗传效应及配合力分析

为了剖析玉米叶形结构的遗传规律,进而拓宽优良株型自交系的遗传基础。本研究以12份不同叶形结构玉米自交系组配的32份F1杂交种为试材,在2个生态环境下,花期采用加性-显性-母体遗传模型(ADM)对其穗三叶叶长、叶宽、叶夹角、叶向值及叶面积进行遗传效应和配合力分析。结果表明:(1)玉米穗三叶叶长、叶宽及叶向值主要受基因的加性效应调控,其次是显性效应,同时还兼受加性×环境互作效应或母体×环境互作效应等遗传体系的调控,育种改良中宜在早代对其进行选择;叶夹角主要受基因的母体效应调控,其次是加性效应,另外还受加性×环境互作效应及母体×环境互作效应影响,育种中宜选择叶夹角较小的材料作为母本进行改良;叶面积只受基因的显性效应及显性×环境互作效应的调控,其应从较晚世代中进行遗传选择。(2)除父本叶长的一般配合力差异不显著外,父本其余性状的一般配合力和母本全部性状的一般配合力间差异均显著或极显著,且这5个叶形相关性状的全部特殊配合力间差异极显著。(3)从相应自交系各性状的一般配合力相对效应值分析发现,‘锋1999马’和‘锋1913硬’的综合性状表现优良,有利于组配出叶片大小适中及株型紧凑的优良耐密高产F1杂交组合。研究结果表明,玉米穗三叶5个叶形结构的遗传效应不尽相同,相应亲本5个叶形结构的一般配合和特殊配合力间存在明显差异,因此在玉米叶形结构遗传改良上应按照相应性状的遗传特征选择对应改良策略进行改良,并根据综合性状表现优良的亲本有目的地组配杂交组合,提高玉米理想株型育种效率。本研究为进一步剖析玉米叶形结构的遗传机理及玉米理想株型育种提供参考。     

Lessons learnt from forty years of international spring bread wheat trials

Yield potential is the maximum attainable yield within the limits imposed by the production environment. Better understanding of these constraints and the underlying causes of genotype × environment interaction will improve productivity regionally and globally. For 40 years, the International Maize and Wheat Improvement Center (CIMMYT) has distributed wheat yield trials, and collaborators from across the world have provided yield, disease, and agronomic data. Various analyzes of these data have been conducted over the years to assess the effectiveness of CIMMYT’s Mexican based breeding program, and to identify key selection environments and genotypes with broad adaptation. Analysis of these data confirmed the value of shuttle breeding in Mexico. Well-watered and terminal heat stress selection environments generated in Mexico associate well with their global target areas. Germplasm targeting dry areas is selected and screened for drought tolerance in Mexico using limited irrigation. This type of screening correlated well with environments in South Asia, but less so with sites in West Asia and South America. On the basis of these findings, drought screening in Mexico has been modified to better reflect a wider array of drier environments. In contrast, CIMMYT’s high rainfall environment at Toluca, the other arm of the Mexico-based shuttle, was found to be a poor predictor of global high rainfall areas. The analysis of these data led to the identification of key locations in countries outside Mexico. Integrating information from these sites with that obtained in Mexico has helped improve the efficiency of CIMMYT’s global wheat breeding effort.     

Efficiency of Recurrent Selection Methods to Improve the Line Value of a Population

Five breedings methods have been studied for their efficiency in improving the line value of a population, i.e. the value of all lines that can be derived from the population. These methods are: half-sib family recurrent selection (HSF-RS), full-sib family recurrent selection (FSF-RS), half-sib progeny recurrent selection (HSP-RS), recurrent selection (S₁-RS) and the single-doubled-haploid descent recurrent selection (SDH-RS). All breeding methods are compared on the basis of the genetic advance per unit of time, with the constraint of the same effective size after the first cycle of selection. Two modalities of SDH-RS are considered for an annual plant without an off-season generation: one with a cycle in four years (SDH₁-RS) and the other with a cycle in five years (SDH₂-RS). For all methods the possibility of one off-season generation for intercrossing is considered. Fields of efficiency of the best methods are determined considering five main parameters, the heritability, the importance of additive variance, the correlation QL between the additive effect for line value and the additive effect for the per se value, the length of the cycle for SDH-RS and the selection intensities. SDH-RS is the best method when the heritability is low, with a low proportion of additive variance and with a relatively low correlation QL between additive effects for per se and line values. Without off-season generation, S₁-RS is the best method among the remaining realistic situations; with one off-season generation for intercrossing it is FSF-RS. The use of an off-season generation reduces the domain of efficiency of SDH-RS while an increase in the selection intensity increases this domain. With a selection intensity for SDH-RS equal to that for S₁-RS, SDH-RS is the best in a large range of realistic situations except for SDH₂ with one offseason generation and medium to high correlation QL.     

Proof of concept to unmask the breeding value of genetic resources of barley (Hordeum vulgare) with a hybrid strategy

Broadening the genetic base of elite breeding programmes is crucial for further breeding success. The absence of major adaption genes, however, often masks the grain yield breeding value of genetic resources. We assessed the ability of a hybrid strategy to provide unbiased performance estimates of 21 barley genetic resources. By crossing them to elite tester lines, 25 three-way hybrids were produced and evaluated together with a part of their parents and eight elite hybrids for important agronomic traits in replicated field trials in four environments. The phenotypic data analyses revealed that the hybrid strategy facilitated to identify promising resources by substantially improving lodging resistance. Combining genotypic data for 5,562 SNPs with the phenotypic data highlighted the potential to boost the diversity of the elite breeding pool via targeted introgression of genetic resources into the male and female heterotic pools. We propose an application of the hybrid strategy for genetic resources of entire genebank collections and to use genome-wide predictions to support a targeted choice of accessions with high value for barley breeding.     

辣木优良单株筛选比较

为筛选出优良的辣木果用种源,对30个辣木家系(每个家系15～30株)进行连续3年的跟踪观测及综合评价,以辣木种子产量为主要指标,同时兼顾株高、冠幅、地径以及果荚的相关性状指标(包括单果重、果荚长、果荚宽、单荚种子数等),经初选、复选和决选,从30个家系中最终筛选出丰产、果型好、种子饱满的果用优良单株30株。研究结果可为后期辣木良种的选育提供基础材料,同时为干热区辣木树种的选育提供理论参考和技术方法。     

Direct and indirect selection for yield in chickpea

Summary The efficiency of indirect selection for seed yield was compared with direct selection for yield per se in chickpea. A total of 2500 single F₂ plants, derived from 50 crosses with 50 plants from each cross, were divided into five sub-populations (SP1 to SP5) of 500 plants each by including 10 plants from each of the 50 crosses. The five sub-populations were advanced upto F₆ by exercising 10% selection intensity for four successive generations for number of pods per plant in SP1, number of seeds per pod in SP2, seed weight in SP3, seed yield in SP4 and random selection in SP5. The efficiency of direct and indirect selection for yield was evaluated by comparing groups of 50 F₆ lines from each sub-population. SP1 and SP3 F₆ lines showed higher mean grain yield than the other three methods. SP1 and SP3 were found to be almost equally efficient in developing F₆ lines which were significantly superior to the check. This suggests that indirect selection for yield via pod number and seed weight is more efficient than direct selection for yield.