Heterosis Is Prevalent for Multiple Traits in Diverse Maize Germplasm

Background

Heterosis describes the superior phenotypes observed in hybrids relative to their inbred parents. Maize is a model system for studying heterosis due to the high levels of yield heterosis and commercial use of hybrids.

Methods

The inbred lines from an association mapping panel were crossed to a common inbred line, B73, to generate nearly 300 hybrid genotypes. Heterosis was evaluated for seventeen phenotypic traits in multiple environments. The majority of hybrids exhibit better-parent heterosis in most of the hybrids measured. Correlations between the levels of heterosis for different traits were generally weak, suggesting that the genetic basis of heterosis is trait-dependent.

Conclusions

The ability to predict heterosis levels using inbred phenotype or genetic distance between the parents varied for the different traits. For some traits it is possible to explain a significant proportion of the heterosis variation using linear modeling while other traits are more difficult to predict.     

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．     

Comparative analysis of inbred and hybrid maize at the diploid and tetraploid levels

Heterosis often occurs in offspring derived from a cross between inbred or divergent parents and can be observed as the superior performance of these hybrids for a wide variety of characters. Heterosis was compared in maize lines at two ploidy levels, diploid and tetraploid, to gain a better understanding of the interaction of heterosis and ploidy level. Employing genetically identical diploid and tetraploid maize derived from four different inbred lines, we investigated heterosis for 11 morphological traits, including several plant height measures, as well as flowering time for both silks and anthers. We find that the heterotic response of a certain hybrid differs between diploid and tetraploid lines, and that the response at one ploidy cannot serve as a predictor for the other. Also, progressive heterosis was found for several of the characters in the tetraploid double-cross hybrid, which can have four different alleles at one locus, compared to the double-cross diploid hybrids, which can only possess two alleles per locus. Overall, the results indicate that the heterotic response of tetraploid maize lines differs significantly from that of the diploid. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

冬小麦植物铁载体分泌的杂种效应

缺铁是石灰性土壤常见的植物营养问题之一.禾本科植物种或基因型的植物铁载体分泌能力与耐缺铁有关,提高植物铁载体分泌能力是改良缺铁的土壤上植物铁aestivum L.) 3个杂交种及其4个亲本在缺铁营养液中植物铁载体的分泌及杂种的效应.植物铁载体的分泌率通过根分泌物对新形成的Fe(OH)3的活化能力进行测定, 在缺铁症出现时每隔2、3天测定1次.在缺铁条件下,所有基因型都分泌较多的植物铁载体,并且随缺铁症状的发展分泌量增加.杂交种具有对缺铁更敏感的反馈系统,在缺铁条件下,杂交种比亲本分泌铁载体的速度更快、量更高.通过分析杂交种和亲本的关系,认为可以通过对亲本分泌植物铁载体能力和配合力的选择,利用杂种优势来提高小麦铁的利用效率.     

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。     

Do rice hybrids have heterosis for insect resistance? A study with Nilaparvata lugens (Hemiptera: Delphacidae) and Marasmia patnalis (Lepidoptera: Pyralidae)

Antibiosis-based resistance to two insect pests of rice, Nilaparvata lugens (St?l) (Hemiptera: Delphacidae) and Marasmia patnalis Bradley (Lepidoptera: Pyralidae), was compared in 11 F1 hybrids and their parental lines. Our objective was to determine whether hybrids show heterosis (hybrid vigor) for insect resistance or susceptibility. Heterosis is defined as the amount by which a hybrid exceeds its midparent value or its better parent. Overall, we did not find evidence of heterosis or heterobeltiosis (a type of heterosis in which a hybrid exceeds its better parent) for antibiosis-based resistance or susceptibility to either of the insects. One hybrid, IR64616H, seemed more resistant to M. patnalis than its better parent but none of the other hybrids showed heterobeltiosis for resistance or susceptibility to either insect. Three hybrids had resistance to N. lugens that exceeded their midparent value, possibly due to dominant resistance in one of the parents. The increased frequency and severity of insect outbreaks on hybrid rice that have been reported in China may be attributable to factors other than diminished antibiosis in hybrids, such as greater attractiveness of hybrids to migrating or dispersing insects or differences in agronomic practices applied to hybrids and inbred rice cultivars.     

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.     

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.     

Gene expression analyses in maize inbreds and hybrids with varying levels of heterosis

Background  

Heterosis is the superior performance of F₁ hybrid progeny relative to the parental phenotypes. Maize exhibits heterosis for a wide range of traits, however the magnitude of heterosis is highly variable depending on the choice of parents and the trait(s) measured. We have used expression profiling to determine whether the level, or types, of non-additive gene expression vary in maize hybrids with different levels of genetic diversity or heterosis.     

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.     

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.     

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.     

Molecular marker genotypes, heterozygosity and genetic interactions explain heterosis in Arabidopsis thaliana

The underlying mechanisms for hybrid vigor or heterosis are elusive. Here we report a population of recombinant inbred lines (RILs), derived from the two ecotypes, Col and Ler, which can serve as a permanent resource for studying the molecular basis of hybrid vigor in Arabidopsis. Using a North Carolina mating design III (NCIII), we determined the additive and dominant nature of gene action in this population. We detected heterosis among crosses of RILs with one of the two parents (Col and Ler) and analyzed genotypes and heterozygosities for RILs and test cross families (RILs crossed to Col and Ler) using a total of 446 published molecular markers. The performance of RILs and additive and dominant components in the test cross families were used to analyze QTLs for 16 traits, using QTL cartographer and composite interval mapping with 1,000 permutations for each trait. Our data suggest that locus-specific and/or genome-wide differential heterozygosity, including epistasis, plays an important role in the generation of the observed heterosis. Furthermore, the hybrid vigor occurred between two closely related ecotypes, and provides a general mechanism for novel variation generated between genetically similar materials.     

Spatial and temporal expression modes of MicroRNAs in an elite rice hybrid and its parental lines

Heterosis is a commonly observed phenomenon in nature and refers to the superior performance of hybrids relative to both parents. The molecular mechanisms of heterosis are mostly unknown. Quantitative trait locus (QTL) mapping has been used to explain the genetic basis of heterosis, and large amounts of QTLs have been mapped for various agronomic traits, but the nature of QTL contributing to heterosis is still enigmatic. MicroRNAs (miRNAs) are master regulators in the processes of plant development and trait performance, and many miRNAs are predicted to reside in QTL intervals. We analyzed the expression modes of miRNAs, which were picked up from miRNA databases and chosen from those predicted from QTL intervals by bioinformatic approaches, in different organs at developmental stages of an elite rice hybrid and its parents. All possible modes of action for miRNA expression were detected, but most miRNAs showed nonadditive mode, and different stages and distinct organs displayed different patterns of miRNA expression. A large proportion of miRNAs were not detected for expression in leaves but expressed in the culms and roots of the hybrid at tillering stage. MiRNAs from grain-weight QTL intervals have multiple effects on grain development. Together, our results reveal that miRNAs, especially those from QTL intervals, play roles in heterotic performance in this elite rice hybrid, our results also shade new light on understanding the molecular mechanisms of heterosis.     

盐度对长牡蛎和近江牡蛎及其杂交稚贝生长和存活的影响

2011年8月以长牡蛎自繁组GG(Crassostrea gigas♀×C.gigas♂),近江牡蛎自繁组AA(C.ariakensis♀×C.ariakensis♂)、正交组GA(Crassostrea gigas♀×C.ariakensis♂)、反交组AG(C.ariakensis♀×C.gigas♂)为实验材料,开展了稚贝对盐度的适应性研究。结果发现长牡蛎的最适生存盐度为15—35,最适生长盐度为25—35;近江牡蛎的最适生存盐度为10—25,最适生长盐度为20—25;GA的最适生存盐度为15—30,最适生长盐度为15—30,AG的最适生存盐度为20—30,最适生长盐度为20—25。GG对低盐度敏感,AA对高盐度敏感,AG具有高盐度存活的杂种优势,在盐度30时,中亲杂种优势HG×A为13.32,单亲杂种优势HGA和HAG分别为1.89和27.88,在盐度40时,HAG上升到400,GA和AG都不具有生长优势。杂种稚贝对盐度适应介于双亲之间,且表现出一定程度的父系遗传特点。