杂交稻特殊配合力与杂种优势、亲本间遗传距离的相 关性

杂交水稻育种的实质是配合力育种, 筛选高特殊配合力的杂交水稻组合才能选育出在生产上有实用价值的强优势组合。文章利用SSR标记检测了9个三系杂交稻亲本(5个不育系和4个恢复系)之间的遗传距离, 结合20个杂交稻组合(5×4 NCII)的产量表现, 分析了杂交水稻特殊配合力(Special combining ability, SCA)效应与产量杂种优势、亲本间遗传距离的相关性。结果表明, 特殊配合力效应与对照优势(相关系数r₁=0.5609)、平均优势(相关系数_r2=0.541)之间均呈显著正相关, 而与亲本遗传距离之间相关不显著, 相关系数(r=0.2143)较小。说明本研究所配组合的特殊配合力效应能充分反映杂种优势, 选用的杂交亲本能组配出强优势组合; 而杂交亲本遗传距离的大小并不能反映特殊配合力效应, 分子标记遗传距离与特殊配合力的相关性还有待于进一步的探讨。     

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.     

Parental Selection of Hybrid Breeding Based On Maternal and Paternal Inheritance of Traits in Rapeseed (Brassica napus L.)

Parental selection is crucial for hybrid breeding, but the methods available for such a selection are not very effective. In this study, a 6×6 incomplete diallel cross was designed using 12 rapeseed germplasms, and a total of 36 hybrids together with their parental lines were planted in 4 environments. Four yield-related traits and seed oil content (OC) were evaluated. Genetic distance (GD) was estimated with 359 simple sequence repeats (SSRs) markers. Heterosis levels, general combining ability (GCA) and specific combining ability (SCA) were evaluated. GD was found to have a significant correlation with better-parent heterosis (BPH) of thousand seed weight (TSW), SCA of seeds per silique (SS), TSW, and seed yield per plant (SY), while SCA showed a statistically significant correlation with heterosis levels of all traits at 1% significance level. Statistically significant correlations were also observed between GCA of maternal or paternal parents and heterosis levels of different traits except for SS. Interestingly, maternal (TSW, SS, and OC) and paternal (siliques per plant (SP) and SY) inheritance of traits was detected using contribution ratio of maternal and paternal GCA variance as well as correlations between GCA and heterosis levels. Phenotype and heterosis levels of all the traits except TSW of hybrids were significantly correlated with the average performance of parents. The correlations between SS and SP, SP and OC, and SY and OC were statistically significant in hybrids but not in parents. Potential applications of parental selection in hybrid breeding were discussed.     

苜蓿SSR遗传距离与杂种优势的相关性分析

利用SSR标记技术对40份苜蓿材料（6个雄性不育株系和34个苜蓿品种）的遗传距离进行分析,并利用其中的6个不育株系与14个苜蓿品种测交,进一步对遗传距离(GD)、产量配合力与杂种优势效应进行相关性分析。结果表明,25对SSR引物共扩增出189条谱带,其中多态性条带136条,平均多态性位点百分率为69.23%;40份苜蓿材料的遗传距离为0.1818～0.9091,平均0.4544;各亲本一般配合力及GD均与杂种优势效应存在显著正相关,其中亲本一般配合力与杂种优势的相关性要高于GD.因此,仅以SSR遗传距离还不足以准确的组配强优势组合,需结合各性状配合力的分析,以充分发挥杂种优势效应。     

中国17个优良玉米自交系的分子标记杂合性及其与杂交种性状的关系研究

在玉米单交种育种中 ,鉴定高产杂交种和具有优良特性的自交系是一个重要的问题。研究以 1 7个优良玉米自交系为亲本 ,按照双列杂交配组合 ,利用 RAPD技术分析了 1 7个自交系的多态性以及 RAPD标记与 9个重要农艺性状 (包括产量 )的关系。基于 RAPD标记计算的相似系数聚类将 1 7个自交系分为 5个类群 ,经分析与系谱亲缘关系基本一致。杂交种性状及其特殊配合力与亲本间的遗传距离是高度相关的 ,与聚类前比较 ,聚类后平均遗传距离与平均产量、平均特殊配合力的相关系数显著提高 ,类间平均产量高于类内平均产量。RAPD技术可揭示优良玉米自交系的系谱亲缘关系 ,将自交系划分成不同的类群 ,从而为选择类间自交系杂交 ,进行亲本选配和分子标记辅助育种提供一种方法。     

Heterosis for biomass yield and related traits in five hybrids of Arabidopsis thaliana L. Heynh

Heterosis is of utmost economic importance in plant breeding. However, its underlying molecular causes are still unknown. Given the numerous advantages of Arabidopsis thaliana as a model species in plant genetics and genomics, we assessed the extent of heterosis in this species using five hybrids derived from five ecotypes. Parents, F(1) and F(2), generations in both reciprocal forms were grown in a greenhouse experiment with four replications. Mid-parent heterosis (MPH) and best-parent heterosis (BPH) averaged across hybrids were surprisingly high for biomass yield (MPH: 60.3%; BPH: 32.9%) and rosette diameter (MPH: 49.4%; BPH: 34.8%), but smaller for flowering date (MPH: 27.5%; BPH: 18.5%), seed yield (MPH: 18.9%; BPH: 1.7%), and yield components. Individual hybrids varied considerably in their MPH and BPH values for all traits, one cross displaying 140.1% MPH for biomass yield. MPH was not associated with parental genetic distance determined from molecular markers. Reciprocal effects were significant only in a few cases. With a proper choice of hybrids, our results encourage the use of Arabidopsis as a model species for investigating the molecular causes of heterosis.     

The use of SSRs for predicting the hybrid yield and yield heterosis in 15 key inbred lines of Chinese maize

A challenge to maize breeders is to predict and identify inbred lines that can produce highly heterotic hybrids precisely. In the present study we surveyed the genetic diversity among 15 elite inbred lines of maize in China with SSR markers and assessed the relationship between SSR marker and hybrid yield/yield heterosis in a diallel set of 105 crosses. Forty-three SSR primers selected from all sixty-three primers gave stable profiles amplified in the sample of 15 inbred lines, which could clearly resolve on 4% metaphor agarose gel. The average number of alleles per SSR locus was 4.44 with a range from 2 to 9. The polymorphism information content (PIC) for the SSR loci varied from 0.28 to 0.81 with a mean of 0.6281. Genetic similarity (GS) among 15 lines was estimated with 191 alleles identified as raw data, the Nei's coefficient of GS ranged from 0.492 for 478 vs HZ4 up to 0.745 for E28 to ZH64 with a mean of 0.619. The cluster diagram based upon the SSR data grouped the 15 lines into families consistent with the yield heterotic response of these. Genetic distance (GD) based on SSR data was significantly correlated with hybrid yield/yield heterosis, the correlation coefficient (r) being 0.5432 and 0.4271 in 1999 and 0.4305 and 0.3614 in 1998 field test, respectively, whereas the determination coefficient (r2) was lower. The correlation between GD based on SSR data and hybrid yield/yield heterosis changed alone with the difference of number and pedigree relationship among parents that were used in this study. SSR makers showed high polymorphism and could be used to assess the relationship between inbred lines of maize, but it was difficult to predict the yield heterosis of maize.     

Identification of Indica rice chromosome segments for the improvement of Japonica inbreds and hybrids

Exploitation of heterosis has brought significant advance in plant breeding and agricultural production, although its genetic basis is still poorly understood. In this study, a total of 66 chromosome segment substitution (CSS) lines, derived from a cross between japonica rice inbred line Asominori (as the recurrent parent) and indica rice inbred line IR24 (as the donor parent), were used to investigate the genetic basis of heterosis in indica × japonica inter-subspecific rice hybrids. Each CSS line was crossed with the background parent Asominori, and the heterosis of F(1) hybrids was estimated by comparing the F(1) performance with its two parental lines. Field experiments were carried out across six different environments to evaluate yield and yield-related traits in the 66 CSS lines and their 66 corresponding F(1) hybrids. Quantitative trait loci (QTL) analyses were conducted using a likelihood ratio test based on the stepwise regression. Thirty-six QTL were identified with significant effects in CSSL, 21 with significant effects in hybrids and 13 with significant effects in both. On the basis of average dominance degree, of all the 70 QTL affecting yield-related agronomic traits, 28.6% (20) showed an overdominance, 35.7% (25) a partial dominance and 30% (21) an additive effect, indicating that all effects contribute to trait variation in japonica-indica rice hybrids. Effects of these QTL were examined to identify Indica rice chromosome segments of interest for the improvement of japonica inbred lines and hybrids.     

Relationship of Parental Genetic Distance with Heterosis and Specific Combining Ability in Sesame (<Emphasis Type="Italic">Sesamum indicum</Emphasis> L.) Based on Phenotypic and Molecular Marker Analysis

The genetic distance analysis for selection of suitable parents has been established and effectively used in many crops; however, there is dearth of conclusive report of relationship of genetic distance analysis with heterosis in sesame. In the present study, an attempt was made to estimate the associations of genetic distances using SSR (GDSSR), seed-storage protein profiling (GDSDS) and agro-morphological traits (GDMOR) with hybrid performance. Seven parents were selected from 60 exotic and Indian genotypes based on genetic distance from clustering pattern based on SSR, seed-storage protein, morphological traits and per se performance. For combining ability analysis, 7 parents and 21 crosses generated from 7 × 7 half diallel evaluated at two environments in a replicated field trial during pre-kharif season of 2013. Compared with the average parents yield (12.57 g plant^?1), eight hybrids had a significant (P < 0.01) yield advantage across environments, with averages of 26.94 and 29.99% for better-parent heterosis (BPH) and mid-parent heterosis (MPH), respectively, across environments. Highly significant positive correlation was observed between specific combining ability (SCA) and per se performance (0.97), while positive non-significant correlation of BPH with GDSSR (0.048), and non-significant negative correlations with GDMOR (? 0.01) and GDSDS (? 0.256) were observed. The linear regressions of SCA on MPH, BPH and per se performance of F₁s were significant with R² value of 0.88, 0.84 and 0.95 respectively. The present findings revealed a weak association of GDSSR with F₁’s performance; however, SCA has appeared as an important factor in the determination of heterosis and per se performance of the hybrids. The present findings also indicated that parental divergence in the intermediate group would likely produce high heterotic crosses in sesame.     

Magnitude of hybrid vigor retained in double haploid lines of some heterotic rice hybrids

Sixty one double haploid (DH) lines were evaluated for yield components and yield and compared with the three corresponding hybrids from which they were derived through anther culture. Analysis of a 6×6 diallel experiment led to the selection of these hybrids based on their high heterosis and revealed the nature of gene action of the characters under evaluation. The DH lines along with the hybrids and parents were planted following a simple lattice design with two replications. The results show that in DH lines the values of the characters expressing predominantly additive genetic effects could reach the heterotic level of the hybrids. For characters including yield showing predominantly dominance effects, values in the DH lines were significantly lower than those of the corresponding hybrids. The promising DH lines, however, possess a higher yield potential than the better parents.     

Assessing genetic variation to predict the breeding value of winter triticale cultivars and lines

The objective of this study was to evaluate the usefulness of amplified fragment length polymorphism (AFLP) analyses for selection of the best parents for breeding of hybrid winter triticale. Phenotypic diversity was measured for 8 agronomic traits in 10 parents and 27 F1 hybrids. Genotypic diversity was measured by 91 AFLP markers. Coefficients of correlation of genetic similarity (AFLP-GS) with both Euclidean distances and mean values of the traits were generally not significant. A correction of the preliminary binary matrix into trait-specific derived matrices increased the values of 5 correlation coefficients to a significant level. The correlation of AFLP-GS with mid-parent heterosis of grain weight per plant was low but significant (r = -0.452). Our study confirms the effectiveness of marker preselection for obtaining AFLP-GS better correlated with heterosis. The use of derived matrices is promising for reducing the number of cross combinations tested for specific combining ability.     

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.     

Field evaluation of tetraploid somatic potato hybrids

Tetraploid somatic hybrids, derived from ten different fusion combinations, were grown in the field to study the inheritance of quantitatively-controlled agronomic traits. The data from two seasons showed that the heterosis in yield varied between 70% and 230% compared to the mid-parent value of the dihaploid fusion clones. The relative yield of the hybrids compared to two standard varieties (= 100%) ranged from 47% to 106%. Despite large differences in yield between the single hybrids of the same fusion combination, no significant differences could be detected between the hybrids. The analysis of the yield components revealed that the heterosis was mainly due to a highly increased tuber weight, whereas the tuber number was in general intermediate to that of the parent clones. In three combinations, the starch content of the hybrids was 15–20% higher than the midparent value and in the others it was intermediate. Fusion of clones with long oval tubers and with round or round-oval ones showed the round/round-oval form was dominant. Fusion combinations of tuber flesh colour indicated that in most cases yellow was dominant to white, while red skin colour was dominant over yellow.     

Overdominant epistatic loci are the primary genetic basis of inbreeding depression and heterosis in rice. I. Biomass and grain yield

To understand the genetic basis of inbreeding depression and heterosis in rice, main-effect and epistatic QTL associated with inbreeding depression and heterosis for grain yield and biomass in five related rice mapping populations were investigated using a complete RFLP linkage map of 182 markers, replicated phenotyping experiments, and the mixed model approach. The mapping populations included 254 F(10) recombinant inbred lines derived from a cross between Lemont (japonica) and Teqing (indica) and two BC and two testcross hybrid populations derived from crosses between the RILs and their parents plus two testers (Zhong 413 and IR64). For both BY and GY, there was significant inbreeding depression detected in the RI population and a high level of heterosis in each of the BC and testcross hybrid populations. The mean performance of the BC or testcross hybrids was largely determined by their heterosis measurements. The hybrid breakdown (part of inbreeding depression) values of individual RILs were negatively associated with the heterosis measurements of their BC or testcross hybrids, indicating the partial genetic overlap of genes causing hybrid breakdown and heterosis in rice. A large number of epistatic QTL pairs and a few main-effect QTL were identified, which were responsible for >65% of the phenotypic variation of BY and GY in each of the populations with the former explaining a much greater portion of the variation. Two conclusions concerning the loci associated with inbreeding depression and heterosis in rice were reached from our results. First, most QTL associated with inbreeding depression and heterosis in rice appeared to be involved in epistasis. Second, most ( approximately 90%) QTL contributing to heterosis appeared to be overdominant. These observations tend to implicate epistasis and overdominance, rather than dominance, as the major genetic basis of heterosis in rice. The implications of our results in rice evolution and improvement 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.     

On the genetic control of heterosis for fruit shape in melon (Cucumis melo L.)

The objective of the present work is to study the genetic basis of heterosis for fruit shape (FS) in melon observed in a cross between the Spanish cultivar "Piel de Sapo" (PS) and the Korean accession PI 161375 (Songwang Charmi [SC]) using a set of near-isogenic lines (NILs) with contrasting phenotypes for FS, each carrying a single chromosomal introgression from SC within the genetic background of PS. We investigated the FS of homozygous NILs, hybrids NIL x PS, and all 2-way crosses between NILs to test the main heterosis hypotheses (dominance, overdominance, and epistatic interactions). Gene action of alleles of quantitative trait loci inducing fruit enlargement was dominance, whereas those inducing rounder fruit were additive or recessive. Only minor epistatic interactions were found. Therefore, the most plausible explanation for FS heterosis in this cross is in agreement with the dominance complementation hypothesis. Over 70% of the hybrid heterosis could be achieved by combining just 2 loci, indicating that the genetic control of FS heterosis in this cross is relatively simple. FS is proposed as a reproductive trait in melon because of the high correlation to the number of seeds produced along the fruit longitudinal axis.     

Heterosis breeding in rice (Oryza sativa L.)

Summary Studies conducted at the International Rice Research Institute (IRRI) during 1980 and 1981 have shown up to 73% heterosis, 59% heterobeltiosis and 34% standard heterosis for yield in rice. The latter was estimated in comparison to commercial varieties: IR36 and IR42 (yield 4–5 t/ha in wet season trials and 7–8 t/ha in dry season trials). Generally speaking, absolute yield was lower and extent of standard heterosis was higher in wet season than in dry season with some exception. Yields up to 5.9 t/ha (22% standard heterosis) in the wet season and 10.4 t/ha (34% standard heterosis) in the dry season were obtained. Most of the hybrids performed better in some season while some performed better in both seasons. Hybrids showed better lodging resistance although they were 5–10 cm taller. F₁ hybrids had significant positive correlations with the parental traits viz., yield (r = 0.446), tillering (r = 0.746), height (r = 0.810) and flowering (r = 0.843). Selection of parents among elite breeding lines on the basis of their per se yield performance, diverse origin and resistance to insects and diseases should give heterotic combination. Yield advantage of hybrids was due primarily to increase in number of spikelets per unit area even though tiller number was reduced. Grain weight was either the same or slightly higher. High yielding hybrids also showed significant heterosis and heterobeltiosis for total dry matter and harvest index. For commercial utilization of heterosis in rice, effective male sterility and fertility restoration systems are available and up to 45% natural outcrossing on male sterile lines has been observed. Consequently, F₁ rice hybrid have been successfully developed and used in China. Prospects of developing hybrid rice varieties elsewhere appear bright especially in countries that have organized seed production, certification and distribution programs and where hybrid seed can be produced at a reasonable cost.     

Genetic diversity and its relationship to hybrid performance and heterosis in rice as revealed by PCR-based markers

Ten elite inbred lines (four japonica, six indica), chosen from those widely used in the hybrid rice breeding program at Human Hybrid Rice Research Center in China, were crossed to produce all possible hybrids excluding reciprocals. The 45 F₁ hybrids along with the ten parents were evaluated for eight traits of agronomic importance, including yield potential, in a replicated field trial. The ten parents were analyzed with 100 arbitrary decamer oligonucleotide primers and 22 microsatellite (simple sequence repeats, SSRs) primer sets via polymerase chain reaction (PCR). Out of the 100 random primers used, 74 were informative and amplified 202 non-redundant bands (variants) with a mean of 2.73 bands per polymorphic primer. All 22 microsatellite primer sets representing 23 loci in the rice genome showed polymorphisms among the ten parents and revealed 90 alleles with an average of 3.91 per SSR locus. Cluster analysis based on Nei's genetic distance calculated from the 291 (202 RAPDs, 89 SSRs) non-redundant variants separated the ten parental lines into two major groups that corresponds to indica and japonica subspecies, which is consistent with the pedigree information. Strong heterosis was observed in hybrids for most of the traits examined. For the 43 diallel crosses (excluding 2 crosses not heading), yield potential, its components (including panicles per plant, spikelets per panicle and 1000-grain weight) and their heterosis in F₁ hybrids showed a significant positive correlation with genetic distance. When separate analyses were performed for the three subsets, yield potential and its heterosis showed significant positive correlations with genetic distance for the 15 indica x indica crosses and the 6 japonica x japonica crosses; however, yield potential and its heterosis were not correlated with genetic distance for the 22 indica x japonica crosses. Results indicated that genetic distance measures based on RAPDs and SSRs may be useful for predicting yield potential and heterosis of intra-subspecific hybrids, but not inter-subspecies hybrids.     

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.