A systematic dissection in oilseed rape provides insights into the genetic architecture and molecular mechanism of yield heterosis

Heterosis refers to the better performance of cross progeny compared with inbred parents, and its utilization contributes greatly to agricultural production. Several hypotheses have been proposed to explain heterosis mainly including dominance, over-dominance (or pseudo-overdominance) and epistasis. However, systematic dissection and verification of these hypotheses are rarely documented. Here, comparison of heterosis level across different traits showed that the strong heterosis of composite traits (such as yield) could be attributed to the multiplicative effects of moderate heterosis of component traits, whether at the genome or locus level. Yield heterosis was regulated by a complex trait-QTL network that was characterized by obvious centre-periphery structure, hub QTL, complex up/downstream and positive/negative feedback relationships. More importantly, we showed that better-parent heterosis on yield could be produced in a cross of two near-isogenic lines by the pyramiding and complementation of two major heterotic QTL showing partial-dominance on yield components. The causal gene (BnaA9.CYP78A9) of QC14 was identified, and its heterotic effect results from the heterozygous status of a CACTA-like transposable element in its upstream regulatory region, which led to partial dominance at expression and auxin levels, thus resulting in non-additive expression of downstream responsive genes involved in cell cycle and proliferation, eventually leading to the heterosis of cell number. Taken together, the results at the phenotypic, genetic and molecular levels were highly consistent, which demonstrated that the pyramiding effect of heterotic QTL and the multiplicative effect of individual component traits could well explain substantial parts of yield heterosis in oilseed rape. These results provide in-depth insights into the genetic architecture and molecular mechanism of yield heterosis.     

Molecular marker heterozygosity and hybrid performance in indica and japonica rice

An essential assumption underlying markerbased prediction of hybrid performance is a strong linear correlation between molecular marker heterozygosity and hybrid performance or heterosis. This study was intended to investigate the extent of the correlations between molecular marker heterozygosity and hybrid performance in crosses involving two sets of rice materials, 9 indica and 11 japonica varieties. These materials represent a broad spectrum of the cultivated rice gene pool including landraces, primitive cultivars, historically important cultivars, modern elite cultivars and parents of superior hybrids. Varieties within each set were intermated in all possible nonreciprocal pairs resulting in 36 crosses in the indica set and 55 in the japonica set. The F₁s and their parents, 111 entries in total, were examined for performance of seven traits in a replicated field trial. The parents were surveyed for polymorphisms using 96 RFLP and ten SSR markers selected at regular intervals from a published molecular marker linkage map. Molecular marker genotypes of the F₁ hybrids were deduced from the parental genotypes. The analysis showed that, with very few exceptions, correlations in the indica dataset were higher than in that of their japonica counterparts. Among the seven traits analyzed, plant height showed the highest correlation between heterozygosity and hybrid performance and heteorsis in both indica and japonica datasets. Correlations were low to intermediate between hybrid performance and heterozygosity (both general and specific) in yield and yield component traits in both indica and japonica sets, and also low to intermediate between specific heterozygosity and heterosis in the indica set, whereas very little correlation was detected between heterosis and heterozygosity (either general or specific) in the japonica set. In comparison to the results from our previous studies, we concluded that the relationship between molecular marker heterozygosity and heterosis is variable, depending on the genetic materials used in the study, the diversity of rice germplasms and the complexity of the genetic basis of heterosis.     

Extending the rapeseed gene pool with resynthesized <Emphasis Type="Italic">Brassica napus</Emphasis> II: Heterosis

Hybrid breeding relies on the combination of parents from two differing heterotic groups. However, the genetic diversity in adapted oilseed rape breeding material is rather limited. Therefore, the use of resynthesized Brassica napus as a distant gene pool was investigated. Hybrids were derived from crosses between 44 resynthesized lines with a diverse genetic background and two male sterile winter oilseed rape tester lines. The hybrids were evaluated together with their parents and check cultivars in 2 years and five locations in Germany. Yield, plant height, seed oil, and protein content were monitored, and genetic distances were estimated with molecular markers (127 polymorphic RFLP fragments). Resynthesized lines varied in yield between 40.9 dt/ha and 21.5 dt/ha, or between 85.1 and 44.6% of check cultivar yields. Relative to check cultivars, hybrids varied from 91.6 to 116.6% in yield and from 94.5 to 103.3% in seed oil content. Mid-parent heterosis varied from −3.5 to 47.2% for yield. The genetic distance of parental lines was not significantly correlated with heterosis or hybrid yield. Although resynthesized lines do not meet the elite rapeseed standards, they are a valuable source for hybrid breeding due to their large distance from present breeding material and their high heterosis when combined with European winter oilseed rape.     

Heterotic patterns in rapeseed (<Emphasis Type="Italic">Brassica napus</Emphasis> L.): I. Crosses between spring and Chinese semi-winter lines

Chinese semi-winter rapeseed is genetically diverse from Canadian and European spring rapeseed. This study was conducted to evaluate the potential of semi-winter rapeseed for spring rapeseed hybrid breeding, to assess the genetic effects involved, and to estimate the correlation of parental genetic distance (GD) with hybrid performance, heterosis, general combining ability (GCA) and specific combining ability (SCA) in crosses between spring and semi-winter rapeseed lines. Four spring male sterile lines from Germany and Canada as testers were crossed with 13 Chinese semi-winter rapeseed lines to develop 52 hybrids, which were evaluated together with their parents and commercial hybrids for seed yield and oil content in three sets of field trials with 8 environments in Canada and Europe. The Chinese parental lines were not adapted to local environmental conditions as demonstrated by poor seed yields per se. However, the hybrids between the Chinese parents and the adapted spring rapeseed lines exhibited high heterosis for seed yield. The average mid-parent heterosis was 15% and ca. 50% of the hybrids were superior to the respective hybrid control across three sets of field trials. Additive gene effects mainly contributed to hybrid performance since the mean squares of GCA were higher as compared to SCA. The correlation between parental GD and hybrid performance and heterosis was found to be low whereas the correlation between GCA_{(f + m)} and hybrid performance was high and significant in each set of field trials, with an average of r = 0.87 for seed yield and r = 0.89 for oil content, indicating that hybrid performance can be predicted by GCA_{(f + m)}. These results demonstrate that Chinese semi-winter rapeseed germplasm has a great potential to increase seed yield in spring rapeseed hybrid breeding programs in Canada and Europe.     

Selection and stability of synthetic varieties of Lolium perenne

Summary The performance of three experimental cultivars of Lolium perenne selected for yield or water soluble carbohydrate content was monitored over four generations of seed multiplication under relaxed selection. In each variety the selected trait regressed towards that of the base population from which the selection line derived. This could be accounted for by residual genetic variation within the lines for the selected trait, and in some instances, by association of this variation with the fitness character, seed numbers produced. These results emphasize the need for practical breeding programmes to consider the nature of the gene action controlling the selected trait, if additive, directional selection should be effective in increasing the expression of the character. Where ambidirectional dominance and epistasis are important, consideration should be given to means of achieving reassortment of the controlling genes prior to selection.     

A diallel analysis of heterosis in elite hybrid rice based on RFLPs and microsatellites

Hybrid rice has contributed significantly to the dramatic increase of rice production in the world. Despite this, little attention has been given to studying the genetic basis of heterosis in rice. In this paper, we report a diallel analysis of heterosis using two classes of molecular markers: restriction fragment length polymorphisms, (RFLPs) and microsatellites. Eight lines, which represent a significant portion of hybrid rice germ plasm, were crossed in all possible pairs, and the F₁s were evaluated for yield and yield component traits in a replicated field trial. The parental lines were surveyed for polymorphisms with 117 RFLP probes and ten microsatellites, resulting in a total of 76 polymorphic markers well-spaced in the rice RFLP map. The results indicated that high level heterosis is common among these crosses: more than 100% midparent and 40% better-parent heterosis were observed in many F₁s, including some crosses between maintainer lines. Heterosis was found to be much higher for yield than for yield component traits, which fits a multiplicative model almost perfectly. Between 16 and 30 marker loci (positive markers) detected highly significant effects on yield or its component traits. Heterozygosity was significantly correlated with several attributes of performance and heterosis. Correlations based on positive markers (specific heterozygosity) were large for midparent heterosis of yield and seeds/panicle and also for F₁ kernel weight. These large correlations may have practical utility for predicting heterosis.     

Early changes of gene activity in developing seedlings of Arabidopsis hybrids relative to parents may contribute to hybrid vigour

Hybrid vigour (heterosis) has been used for decades in crop industries, especially in the production of maize and rice. Hybrid varieties usually exceed their parents in plant biomass and seed yield. But the molecular basis of hybrid vigour is not fully understood. In this project, we studied heterosis at early stages of seedling development in Arabidopsis hybrids derived from crossing Ler and C24 accessions. We found that early heterosis is associated with non‐additive gene expression that resulted from earlier changes in gene expression in the hybrids relative to the parents. The non‐additively expressed genes are involved in metabolic pathways, including photosynthesis, critical for plant growth. The early increased expression levels of genes involved in energy production in hybrids is associated with heterosis in the young seedlings that could be essential for biomass heterosis at later developmental stages of the plant.     

超甜玉米种子萌发物质利用性状杂种优势及亲子回归关系分析

为阐明超甜玉米(Zea mays L.var.saccharata Bailey)亲本对F_1种子物质利用性状遗传效应,研究了19份自交系和其测配的10个杂交组合的杂种优势及亲子回归关系。结果表明,超甜玉米自交系及F_1种子的淀粉含量、蛋白质含量、脂肪含量、百粒重、种子物质动用量和种子物质利用率差异较大,10个杂交组合中亲本和F_1种子的淀粉含量、脂肪含量、百粒重均存在显著差异。F_1种子淀粉含量和百粒重均表现出正向超亲优势,即近高亲本遗传;而F_1种子的蛋白质含量、脂肪含量、种子物质动用量和种子物质利用率为近低亲本遗传。聚类分析和杂种优势分析表明,性状差异较大的FH14、Q26、GT22、GT2等亲本测配的杂交组合在种子百粒重或种子物质利用率等性状上具有较强的超亲优势。回归分析表明,母本对F_1种子的淀粉含量、百粒重有负效应,对种子物质动用量和种子物质利用率有正效应;父本对种子淀粉含量有负效应,对种子物质利用率有正效应。     

Associations between quantitative traits and enzyme loci in the F2 population of a maize hybrid

Summary Univariate and multivariate analyses were used to identify associations between eight enzyme marker loci and 11 quantitative traits of maize (Zea mays L.). The material analyzed included inbred lines Wf9 and Pa405, single-cross hybrid Wf9 X Pa405, and the F2 generation of the selfed single-cross hybrid. Each enzyme locus assayed was associated with at least one quantitative trait, and all quantitative traits were associated with genotypes at particular enzyme loci. Significant associations also were found between the level of heterozygosity per individual and nine of 11 quantitative traits. The total contribution to heterosis, for seed yield per plant, of genes linked with the eight enzyme loci, was 27% of the F2 mean and 18% of the difference in mean between the F1 hybrid and the inbred parents. Genes linked with Glu1 accounted for nearly one third of the total dominance effect detected by the eight enzyme loci. The chromosome segments marked by loci with significant effects on seed yield were markedly overdominant. The large heterotic effects of chromosome segments marked by particular loci suggest that enzyme loci could be used to help transfer genes responsible for heterosis to inbred lines. We conclude that analyses of additional inbred lines, F1 hybrids, and F2 populations in more environments will halp identify specific associations between enzyme loci, or chromosome segments which they mark, and important agronomic traits.Cooperative investigations of the USDA, ARS and Dept. of Plant Sciences, South Dakota State Univ. (SDSU), Brookings, Journal Series No. 2039; and the Institute of Animal Resource Ecology, Univ. of British Columbia, Vancouver, B.C. V6T 1W5, Canada     

Relationship between genetic distance and heterosis for yield and morphological traits in winter canola (Brassica napus L.)

Genetic distance among canola cultivars was estimated through multivariate analysis. Thirty cultivars from various sources were analyzed and clustered based upon five morphological characteristics and yield components-crown diameter, number of branches plant^-1, number of pods plant^-1, number of seeds pod^-1 and yield plant^-1 -and placed in three distinct clusters. Two cultivars from each cluster were selected as parents and 15 partial-diallel inter- and intra-cluster crosses were made between the six selected parents and evaluated at two locations in Michigan in 1990/1991. The association between genetic distance and mid-parent heterosis was investigated. The correlation between genetic distance and heterosis was positive and highly significant for seed yield, number of pods plant^-1, and number of seeds pod^-1. Clustering, based on yield and yield-component traits, demonstrated that inter-cluster heterosis was greater than intra-cluster heterosis in the majority of cases.     

Relationship between seed yield heterosis and molecular marker heterozygosity in soybean

 In soybean [Glycine max (L.) Merr.] heterosis has been reported for seed yield. Molecular markers may be useful to select diverse parents for the expression of heterosis and yield improvement. The objective of this study was to determine if molecular markers could be used to predict yield heterosis in soybean. From each Maturity Group (MG) II and III, 21 genotypes were selected on the basis of high yield (HY), different geographic origin (GO), and isozyme loci (ISO) and for diversity in restriction fragment length polymorphisms (RFLP), and crosses were made within MGs and selection criteria groups to obtain 6 F₁ hybrids per group. The 21 parents and the 24 F₁ hybrids of each MG were evaluated for yield in replicated tests at two locations in 2 years, and midparent heterosis (MPH) and high-parent heterosis (HPH) estimates were calculated. On the basis of hybrid performance during the first year, 12 parents (3 per selection criteria group) were chosen in each MG to conduct a second RFLP analysis using 129 probes. Genetic distances (GD_M) for pairs of the 12 genotypes were calculated with this RFLP information and correlated with MPH and HPH estimates. Significant MPH averages for seed yield were observed in the combined analysis of variance in each of the four selection criteria groups of MG II, and in the HY, ISO, and GO of MG III. Significant HPH averages were observed only in the ISO and GO groups of MG II. The greatest frequency of F₁ hybrids with significant MPH was observed in the ISO and GO groups of both MGs. For HPH, the greatest frequency was observed in the ISO group of both MGs. In both MGs, the ISO group had the largest absolute MPH value; the RFLP group had generally the smallest. The observations indicated that the expression of heterosis in seed yield might be associated with diversity in the isozyme loci present in the parents. For the genotypes included in the second RFLP analysis, correlations of GD_Ms with MPH and HPH values on an entry-mean basis were low and not significant, indicating that heterosis in yield may not be associated with genetic diversity at the molecular level as determined by RFLPs. The results suggest that in soybean, parent selection on the basis of RFLPs and isozyme loci to exploit heterosis in seed yield may not be feasible. There was no association between genetic distance estimated by the RFLP analysis and seed yield heterosis, and in spite of the observed relationship between isozyme loci and heterosis for yield, the practicality of using the isozyme markers to select parents may be limited because of the reduced number of assayable isozyme loci in soybean. Received: 8 March 1996 / Accepted: 21 February 1997     

Comparison of phenotypic and molecular distances to predict heterosis and F1 performance in Ethiopian mustard (Brassica carinata A. Braun)

Predicting heterosis and F₁ performance from the parental generation could largely enhance the efficiency of breeding hybrid or synthetic cultivars. This study was undertaken to determine the relationship between parental distances estimated from phenotypic traits or molecular markers with heterosis, F₁ performance and general combining ability (GCA) in Ethiopian mustard (Brassica carinata). Nine inbred lines representing seven different geographic regions of Ethiopia were crossed in half-diallel. The nine parents along with their 36 F₁s were evaluated in a replicated field trail at three locations in Ethiopia. Distances among the parents were calculated from 14 phenotypic traits (Euclidean distance, ED) and 182 random amplified polymorphic DNA (RAPD) markers (Jaccard’s distances, JD), and correlated with heterosis, F₁ performance and GCA sum of parents (GCA_sum). The correlation between phenotypic and molecular distances was low (r=0.34, P≤0.05). Parents with low molecular distance also had low phenotypic distance, but parents with high molecular distance had either high, intermediate or low phenotypic distance. Phenotypic distance was highly significantly correlated with mid-parent heterosis (r=0.53), F₁ performance (r=0.61) and GCA (r=0.79) for seed yield. Phenotypic distance was also positively correlated with (1) heterosis, F₁ performance and GCA for plant height and seeds plant⁻¹, (2) heterosis for number of pods plant⁻¹, and (3) F₁ performance for 1,000 seed weight. Molecular distance was correlated with GCA_sum (r=0.36, P≤0.05) but not significantly with heterosis and F₁ performance for seed yield. For each parent a mean distance was calculated by averaging the distances to the eight other parents. Likewise, mean heterosis was estimated by averaging the heterosis obtained when each parent is crossed with the other eight. For seed yield, both mean ED and JD were significantly correlated with GCA (r=0.90, P≤0.01 for ED and r=0.68, P≤0.05 for JD) and mean heterosis (r=0.79, P≤0.05 for ED and r=0.77, P≤0.05 for JD). In conclusion, parental distances estimated from phenotypic traits better predicted heterosis, F₁ performance and GCA than distances estimated from RAPD markers. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

对生玉米产量性状的杂种优势和配合力分析

本文利用对生基因转育获得的对生与互生自交系及其杂交组合,研究了对生性状对产量性状杂种优势和配合力的遗传效应的影响。结果表明:对生玉米同互生玉米一样具有普遍的杂种优势,在产量性状上F1对生株优势互×互组合大于对×对组合和对×互(互×对)组合;对生性状的转育对产量性状的一般配合力有降低效应,而含有对生基因的互生玉米具有较高的一般配合力效应,因此,在对生玉米育种中可以利用含有不同对生基因的互生自交系作亲本来组配杂交组合(互×互),从而获得较高的制种产量和高产的对生杂交种。     

组学技术揭示水稻杂种优势遗传机制

杂种优势是杂交后代在生长或生殖性状上表现出优于亲本的现象。虽然杂种优势在农业生产上已广为应用,但其分子机理仍不清楚。最近,中国科学家通过分析17个代表性杂交稻(Oryza sativa)品种,共10 074个F2个体的全基因组序列和表型,对水稻产量杂种优势相关位点进行了系统定位和解析。此外,中国另一个科研小组通过整合杂交稻亲本和杂交种的表型组、转录组及基因组等多层次数据,深入研究了超级杂交稻两优培九产量的杂种优势基础。这些研究不仅为杂种优势理论的建立提供了新数据,也为水稻育种实践提供了有益的指导。     

The extent of parental genotypic divergence determines maximal heterosis by increasing fertility in inter-subspecific hybrids of rice (Oryza sativa L.)

The magnitude of heterosis in F₁ hybrids is related not only to the performance of parents per se but also to the genetic diversity between two parents. The extent of genotypic divergence between hybrid rice parents was investigated at the molecular level, using two subsets of rice materials: a subset of doubled haploid (DH) lines derived from an Indica × Japonica cross (Gui630/02428) and another subset of Indica or Japonica lines representative of a broad spectrum of the Asian cultivated rice gene pool, including landraces, primitive cultivars, historically important cultivars, modern elite cultivars, super rice and parents of superior hybrids. 57 entries deliberately selected from the 81-DH lines (in total) were testcrossed to two widely used rice lines in China, photoperiod-sensitive genic male sterile (PGMS) N422s and thermo-sensitive genic male sterile (TGMS) Peiai64s. Results of the two sets of test-cross F₁ populations showed congruently that parental genotypic divergence has a relatively low impact on heterosis for the two yield components, i.e., panicle number and 1000-grain weight, but it has a great bearing on fertility parameters, i.e., filled grains per plant and seedset. Heterosis for grain yield in the two test-cross populations exhibited a sharp maximum when the proportion of Japonica alleles in the male parent was between 50 and 60%, so was the heterosis for fertility parameters correspondingly. Thus fertility parameters were the most sensitive and important factors which were influenced by the extent of parental genotypic divergence. Moreover, our results showed that parents with moderate extent of genotypic divergence played an important role in the use of inter-subspecific rice heterosis.     

利用微卫星标记鉴定杂交水稻冈优22种子纯度的研究

水稻是重要的粮食作物之一,早在70年代我国就已实行了杂交稻的三系配套,目前杂交水稻在我国已大面积种植。由于杂交稻种子的真伪无法从种子形态上鉴别,这给造假者以可乘之机。在正常的情况下,种子的纯度也会因质量控制不严而给农业生产造成大量损失。这些都迫切需要有一种…     

杂交小麦制种研究现状及展望

利用杂种优势对提高小麦单产具有重要作用,杂交小麦制种是小麦种业未来发展的重要方向之一。目前,小麦在强优势杂交组合选配方面取得较大进步。由于缺乏优异高异交结实率的制种亲本资源和高产高效杂交制种体系,导致大面积制种效率低。因此,探究杂交制种技术体系进展、亲本种质资源改良和生产流程标准化等方面的研究现状,将有利于对杂交制种发展瓶颈和未来发展方向等问题的整体把握。综述杂交小麦制种技术的影响因素,并通过商业化大田杂交作物与新型杂交小麦比较,对杂交小麦快速商业化等方面提出建议,对其发展趋势进行展望。     

Combining ability analysis over F1-F5 generations in diallel crosses of bread wheat

Summary Combining ability studies for grain yield and its primary component traits in diallel crosses involving seven diverse wheat cultivars of bread wheat (Triticum aestivum L.) over generations F₁-F₅ are reported. The general and specific combining ability variances were significant in all generations for all the traits except specific combining ability variance for number of spikes per plant in the F₅. The ratio of general to specific combining ability variances was significant for all the traits except grain yield in all the generations. This indicated an equal role of additive and non-additive gene effects in the inheritance of grain yield, and the predominance of the former for its component traits. The presence of significant specific combining ability variances in even the advanced generations may be the result of an additive x additive type of epistasis or evolutionary divergence among progenies in the same parental array. The relative breeding values of the parental varieties, as indicated by their general combining ability effects, did not vary much over the generations. The cheap and reliable procedure observed for making the choice of parents, selecting hybrids and predicting advanced generation (F₅) bulk hybrid performance was the determination of breeding values of the parents on the relative performance of their F₂ progeny bulks.     

Use of a Controlled-Nutrient Experiment to Test Heterosis Hypotheses

A controlled-nutrient (CN) experiment was conducted to test three heterosis hypotheses with reference to tomato yield, and its components, for a set of two inbred lines and their hybrid that had previously exhibited considerable heterosis under field conditions. The CN treatments consisted of periodic applications of differential doses of nutrient solution to plants reared individually in containers filled with vermiculite. Ripe fruit were harvested, counted and weighed over a period of 340 days. The data permitted the partitioning of yield into a closed system of five component variables. Heterosis was not exhibited by yield, nor yield components, at any of the four nutrient levels. Hence the total heterosis phenomenon was classified as nutrient-dependent: heterosis occurring under field conditions, but not under the nutritional restrictions of the CN experiment. Three heterosis hypotheses were examined for their ability to explain all of the nutrient-dependent aspects of the heterosis phenomenon. Hypothesis 1: Heterosis is a consequence of a more efficient hybrid metabolic system in that it can produce more product with equal input. Hypothesis 2: Heterosis is a consequence of the somatic multiplication of additive component traits. Hypothesis 3: Heterosis is a consequence of a faster hybrid growth rate. Although none of the hypotheses are rejected by the field data, the first two are rejected by the CN experimental results. The third hypothesis fits all aspects of the nutrient-dependent heterosis phenomenon remarkably well. It is speculated that the indeterminate pattern of plant development responsible for yield and its components is due to two major gene systems: genes that determine morphogenetic, and genes that determine growth rate manifestations of growth. Under this hypothesis, the CN technique permits separation of the responses due to these two gene systems.