Extensive heterosis in growth of yeast hybrids is explained by a combination of genetic models

Heterosis, also known as hybrid vigor, is the superior performance of a heterozygous hybrid relative to its homozygous parents. Despite the scientific curiosity of this phenotypic phenomenon and its significance for food production in agriculture, its genetic basis is insufficiently understood. Studying heterosis in yeast can potentially yield insights into its genetic basis, can allow one to test the different hypotheses that have been proposed to explain the phenomenon and allows better understanding of how to take advantage of this phenomenon to enhance food production. We therefore crossed 16 parental yeast strains to form 120 yeast hybrids, and measured their growth rates under five environmental conditions. A considerable amount of dominant genetic variation was found in growth performance, and heterosis was measured in 35% of the hybrid–condition combinations. Despite previous reports of correlations between heterosis and measures of sequence divergence between parents, we detected no such relationship. We used several analyses to examine which genetic model might explain heterosis. We found that dominance complementation of recessive alleles, overdominant interactions within loci and epistatic interactions among loci each contribute to heterosis. We concluded that in yeast heterosis is a complex phenotype created by the combined contribution of different genetic interactions.     

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。     

Optimal allocation in designs for assessing heterosis from cDNA gene expression data

Heterosis is defined as the superiority of a hybrid cross over its two parents. Plant and animals breeders have long been exploiting heterosis, but the causes of this phenomenon are as yet only partly understood. Recently, chip technology has opened up the opportunity to study heterosis at the gene expression level. This article considers the cDNA chip technology, which allows assaying two genotypes simultaneously on the same chip. Heterosis involves the response of at least three genotypes (two parents and their hybrid), so a chip or microarray constitutes an incomplete block, which raises a design problem specific to heterosis studies. The question to be answered is how genotype pairs should be allocated to chips. We address this design problem for two types of heterosis: midparent heterosis and better-parent heterosis. The general picture emerging from our results is that most of the resources should be allocated to parent-hybrid pairs, while chips with parent-parent pairs or hybrid-reciprocal pairs should be used sparingly or not at all.     

分子标记在油菜杂种优势利用中的研究进展

杂种优势是一种普遍存在的生物学现象, 是提高作物产量的重要途径, 因而受到越来越广泛的重视。分子标记是近年兴起的一种新型的生物技术, 国内外已利用RFLP、RAPD、AFLP、SSR、ISSR和SRAP等多种分子标记技术对油菜的杂种 优势进行了广泛的研究, 取得了很多有价值的研究成果。本文综述了分子标记在油菜授粉控制系统中关键基因标记、亲本遗传多样性分析、杂种优势群划分、杂种纯度鉴定和杂种优势预测中的研究进展。通过对不同作物间的比较, 探讨了分子标记在油菜杂种优势利用中的研究深度, 并对分子标记在油菜杂种优势预测中应注意的问题进行了讨论。     

分子标记在油菜杂种优势利用中的研究进展

杂种优势是一种普遍存在的生物学现象,是提高作物产量的重要途径,因而受到越来越广泛的重视。分子标记是近年兴起的一种新型的生物技术,国内外已利用RFLP、RAPD、AFLP、SSR、ISSR和SRAP等多种分子标记技术对油菜的杂种优势进行了广泛的研究,取得了很多有价值的研究成果。本文综述了分子标记在油菜授粉控制系统中关键基因标记、亲本遗传多样性分析、杂种优势群划分、杂种纯度鉴定和杂种优势预测中的研究进展。通过对不同作物间的比较,探讨了分子标记在油菜杂种优势利用中的研究深度,并对分子标记在油菜杂种优势预测中应注意的问题进行了讨论。     

Epigenetic Variations in Plant Hybrids and Their Potential Roles in Heterosis

Heterosis,one of the most important biological phenomena,refers to the phenotypic superiority of a hybrid over its genetically diverse parents with respect to many traits such as biomass,growth rate and yield.Despite its successful application in breeding and agronomic production of many crop and animal varieties,the molecular basis of heterosis remains elusive.The classic genetic explanations for heterosis centered on three hypotheses:dominance (Davenport,1908;Bruce,1910;Keeble and Pellew,1910;Jones,1917),overdominance (East,1908;Shull,1908) and epistasis (Powers,1944;Yu et al.,1997).However,these hypotheses are largely conceptual and not connected to molecular principles,and are therefore insufficient to explain the molecular basis of heterosis (Birchler et al.,2003).Recently,many studies have explored the molecular mechanism of heterosis in plants at a genome-wide level.These studies suggest that global differential gene expression between hybrids and parental lines potentially contributes to heterosis in plants (e.g.,Swanson-Wagner et al.,2006;Zhang et al.,2008;Wei et al.,2009;Song et al.,2010).Research suggests that genetic components,including cis-acting elements and trans-acting factors,are critical regulators of differential gene expression in hybrids (Hochholdinger and Hoecker,2007;Springer and Stupar,2007;Zhang et al.,2008).However,other research indicates that epigenetic components,the regulators of chromatin states and genome activity,also have the potential to impact heterosis (e.g.,Ha et al.,2009;He et al.,2010;Groszmann et al.,2011;Barber et al.,2012;Chodavarapu et al.,2012;Greaves et al.,2012a;Shen et al.,2012).     

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.     

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.     

杂交水稻育种将迎来新时代

杂种优势是生物界一种普遍的现象,在许多作物中得到应用．杂交水稻自20世纪70年代开始在中国大规模种植,对于粮食生产具有举足轻重的作用．现有的“三系法”和“两系法”杂交育种技术对粮食增产贡献巨大,但它们的技术缺陷也非常明显．本文在总结已有杂交育种技术的操作流程及优缺点的基础上,阐述了利用智能不育杂交育种技术,实现隐性雄性核不育材料在杂交水稻中应用的技术流程．这种飞跃性技术的运用将推动杂交水稻的生产进入一个新的时代．     

Predictive potential of DNA markers in heterosis breeding of maize

PCR-analysis of maize inbred lines has been carried out. Genetic distances between the lines have been calculated, allelic composition and heterosis level of F-hybrids have been determined. Heterosis level of hybrid seed yield rised according to increasing of genetic distances between initial lines. Correlation of allelic composition of inbred line microsatellite loci and heterosis level of the respective hybrids has been revealed.     

Enriched partial correlations in genome-wide gene expression profiles of hybrids (A. thaliana): a systems biological approach towards the molecular basis of heterosis

Heterosis is a well-known phenomenon but the underlying molecular mechanisms are not yet established. To contribute to the understanding of heterosis at the molecular level, we analyzed genome-wide gene expression profile data of Arabidopsis thaliana in a systems biological approach. We used partial correlations to estimate the global interaction structure of regulatory networks. Our hypothesis states that heterosis comes with an increased number of partial correlations which we interpret as increased numbers of regulatory interactions leading to enlarged adaptability of the hybrids. This hypothesis is true for mid-parent heterosis for our dataset of gene expression in two homozygous parental lines and their reciprocal crosses. For the case of best-parent heterosis just one hybrid is significant regarding our hypothesis based on a resampling analysis. Summarizing, both metabolome and gene expression level of our illustrative dataset support our proposal of a systems biological approach towards a molecular basis of heterosis.     

Comparative gene expression profiles between heterotic and non-heterotic hybrids of tetraploid Medicago sativa

Background  

Heterosis, the superior performance of hybrids relative to parents, has clear agricultural value, but its genetic control is unknown. Our objective was to test the hypotheses that hybrids expressing heterosis for biomass yield would show more gene expression levels that were different from midparental values and outside the range of parental values than hybrids that do not exhibit heterosis.     

Heterosis of biomass production in Arabidopsis. Establishment during early development

Heterosis has been widely used in agriculture to increase yield and to broaden adaptability of hybrid varieties and is applied to an increasing number of crop species. We performed a systematic survey of the extent and degree of heterosis for dry biomass in 63 Arabidopsis accessions crossed to three reference lines (Col-0, C24, and Nd). We detected a high heritability (69%) for biomass production in Arabidopsis. Among the 169 crosses analyzed, 29 exhibited significant mid-parent-heterosis for shoot biomass. Furthermore, we analyzed two divergent accessions, C24 and Col-0, the F(1) hybrids of which were shown to exhibit hybrid vigor, in more detail. In the combination Col-0/C24, heterosis for biomass was enhanced at higher light intensities; we found 51% to 66% mid-parent-heterosis at low and intermediate light intensities (60 and 120 micromol m(-2) s(-1)), and 161% at high light intensity (240 micromol m(-2) s(-1)). While at the low and intermediate light intensities relative growth rates of the hybrids were higher only in the early developmental phase (0-15 d after sowing [DAS]), at high light intensity the hybrids showed increased relative growth rates over the entire vegetative phase (until 25 DAS). An important finding was the early onset of heterosis for biomass; in the cross Col-0/C24, differences between parental and hybrid lines in leaf size and dry shoot mass could be detected as early as 10 DAS. The widespread occurrence of heterosis in the model plant Arabidopsis opens the possibility to investigate the genetic basis of this phenomenon using the tools of genetical genomics.     

Biochemical and physiological basis of heterosis

The phenomenon of heterosis or hybrid vigor has long been recognized to have a genetic basis. Heterosis expressed in hybrids often has been correlated with the heterozygosity inherent to a hybrid produced from the cross of parents of different genetic backgrounds. Our understanding of the molecular mechanisms that elicit heterosis remains incomplete; however, a number of critical experiments have contributed to our understanding of the physiological processes and biochemical mechanisms operative in the expression of heterosis. These experiments and the genetic mechanisms thought to underlie them are considered here. Based upon these findings, heterosis does not appear to have a single cause, but rather may result by reason of the action of either single or multiple genes’ in the nucleus or the cytoplasm or both. Pertinent examples of gene action which potentiates heterotic effects are discussed.     

Genome‐wide identification and analysis of heterotic loci in three maize hybrids

Heterosis, or hybrid vigour, is a predominant phenomenon in plant genetics, serving as the basis of crop hybrid breeding, but the causative loci and genes underlying heterosis remain unclear in many crops. Here, we present a large‐scale genetic analysis using 5360 offsprings from three elite maize hybrids, which identifies 628 loci underlying 19 yield‐related traits with relatively high mapping resolutions. Heterotic pattern investigations of the 628 loci show that numerous loci, mostly with complete–incomplete dominance (the major one) or overdominance effects (the secondary one) for heterozygous genotypes and nearly equal proportion of advantageous alleles from both parental lines, are the major causes of strong heterosis in these hybrids. Follow‐up studies for 17 heterotic loci in an independent experiment using 2225 F₂ individuals suggest most heterotic effects are roughly stable between environments with a small variation. Candidate gene analysis for one major heterotic locus (ub3) in maize implies that there may exist some common genes contributing to crop heterosis. These results provide a community resource for genetics studies in maize and new implications for heterosis in plants.     

Understanding the genetic and molecular constitutions of heterosis for developing hybrid rice

The wide adoption of hybrid rice has greatly increased rice yield in the last several decades. The utilization of heterosis facilitated by male sterility has been a common strategy for hybrid rice development. Here, we summarize our efforts in the genetic and molecular understanding of heterosis and male sterility together with the related progress from other research groups. Analyses of F1 diallel crosses show that strong heterosis widely exists in hybrids of diverse germplasms, and inter-subsp...