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.     

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.     

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     

The Physiology of Heterosis in Sorghum with Respect to Environmental Stress

The hypothesis that heterosis in biomass production of sorghum(Sorghum bicolor L. Moench) may be ascribed to stability incarbon exchange rate (CER) over a wide range of environmentalconditions was evaluated. This hypothesis was based on previousresults from detached leaves that hybrids sustained greaterCER over a wider temperture range than their parents. Two grain sorghum hybrids (ATx378/RTx430 and ATx378/RTx434)and their parental lines were grown in the greenhouse in a gradientof ambient temperatures under two water regimes (well-irrigatedand drought up to heading). Plant water-use (estimated by weighingpots), leaf area, leaf gas exchange, grain yield, and above-groundbiomass were determined. Significant heterosis was found for biomass, grain yield perplant, and grain number per panicle. No heterosis occurred forharvest index, indicating that heterosis in grain yield wasdue to heterosis in biomass. Neither growth duration nor leafarea could explain heterosis in biomass. CER and stomatal conductancefor hybrid ATx378/RTx430 in the controls were greater than forboth its parents at leaf temperatures above 38 °C. This,however, was not observed in the other hybrid which was lessheterotic for biomass and grain yield in the controls. WhenCER data were subjected to a stability analysis by joint linearregression, the two hybrids had greater CER than their respectiveparents especially under conditions favouring high CER. Whenextreme stress conditions developed, the hybrid's performancedepended on its genetic background more than on heterosis. Sorghum, Sorghum bicolor L. Moench., heterosis, hybrids, photosynthesis, transpiration, stomata, drought, heat, temperature     

SSR heterogenic patterns of parents for marking and predicting heterosis in rice breeding

The most important concerns of hybrid rice breeders are selection of donors to improve parental lines and prediction of hybrid performance. In this study, SSR molecular marker technology and a half-diallel method were used to address these related hybrid production issues. The results show that genetic diversity among the parental lines is certainly related to heterosis. The heterozygosity of each parental pair is significantly associated with the general combining ability, not with the specific combining ability. However, neither genetic diversity nor heterozygosity is a good indicator for predicting heterosis. From these results, it is suggested that donors for improving parents of hybrids be selected from the improved inbred lines by conventional breeding programs. In this investigation, we also discovered that four favorable alleles and six favorable heterogenic patterns on the parental lines significantly contribute to the heterosis of their hybrids in grain yield, whereas six unfavorable alleles and six unfavorable heterogenic patterns significantly reduce heterosis. These noticeable findings could be, in practice, useful for hybrid rice breeding programs with SSR marker-assisted selection. It is suggested that the optimal combinations with the superior grain yield could be bred out by assembling those favorable alleles into their parental lines and by removing the unfavorable alleles from the parental lines. This study also indicates that there is still a great heterosis potential to be exploited in indica/indica hybrids by the same strategy. In indica/japonica hybrid breeding programs, it may also be important to remove unfavorable alleles rather than broaden genetic diversity or heterozygosity of the parents.     

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.     

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。     

Temporal Regulation of the Metabolome and Proteome in Photosynthetic and Photorespiratory Pathways Contributes to Maize Heterosis

Heterosis or hybrid vigor is widespread in plants and animals. Although the molecular basis for heterosis has been extensively studied, metabolic and proteomic contributions to heterosis remain elusive. Here we report an integrative analysis of time-series metabolome and proteome data in maize (Zea mays) hybrids and their inbred parents. Many maize metabolites and proteins are diurnally regulated, and many of these show nonadditive abundance in the hybrids, including key enzymes and metabolites involved in carbon assimilation. Compared with robust trait heterosis, metabolic heterosis is relatively mild. Interestingly, most amino acids display negative mid-parent heterosis (MPH), i.e., having lower values than the average of the parents, while sugars, alcohols, and nucleoside metabolites show positive MPH. From the network perspective, metabolites in the photosynthetic pathway show positive MPH, whereas metabolites in the photorespiratory pathway show negative MPH, which corresponds to nonadditive protein abundance and enzyme activities of key enzymes in the respective pathways in the hybrids. Moreover, diurnally expressed proteins that are upregulated in the hybrids are enriched in photosynthesis-related gene-ontology terms. Hybrids may more effectively remove toxic metabolites generated during photorespiration, and thus maintain higher photosynthetic efficiency. These metabolic and proteomic resources provide unique insight into heterosis and its utilization for high yielding maize and other crop plants.     

Progressive heterosis in genetically defined tetraploid maize

Progressive heterosis, i.e., the additional hybrid vigor in double-cross tetraploid hybrids not found in their single-cross tetraploid parents, has been documented in a number of species including alfalfa,potato, and maize. In this study, four artificially induced maize tetraploids, directly derived from standard inbred lines, were crossed in pairs to create two single-cross hybrids. These hybrids were then crossed to create double-cross hybrids containing genetic material from all four original lines. Replicated fieldbased phenotyping of the materials over four years indicated a strong progressive heterosis phenotype in tetraploids but not in their diploid counterparts. In particular, the above ground dry weight phenotype of double-cross tetraploid hybrids was on average 34% and 56% heavier than that of the single-cross tetraploid hybrids and the double-cross diploid counterparts, respectively. Additionally,whole-genome resequencing of the original inbred lines and further analysis of these data did not show the expected spectrum of alleles to explain tetraploid progressive heterosis under the complementation of complete recessive model. These results underscore the reality of the progressive heterosis phenotype,its potential utility for increasing crop biomass production, and the need for exploring alternative hypothesis to explain it at a molecular level.     

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.     

抗除草剂杂交籼稻亲本的配合力分析

本试验以6份新育成的抗除草剂籼型恢复系为父本,5份生产上广泛应用的不育系为母本,采用不完全双列杂交设计配制了30份杂交组合,对其苗期除草剂抗性和主要农艺性状配合力进行了分析。除草剂抗性鉴定表明,亲本恢复系及三系杂交组合抗性接近完全,两系杂交组合抗性达90%以上。配合力分析表明,不育系除单株有效穗数外其他农艺性状的一般配合力均达到极显著差异;恢复系间一般配合力在所有性状中均达到显著或极显著差异;杂交组合间特殊配合力方差仅在单株产量、结实率、播始历期和千粒重4个性状中达到显著或极显著差异。不育系中,金科1A在单株产量、结实率等7个性状上的一般配合力均为最高,但其特殊配合力方差最小;广占63-4S在千粒重性状上具有最高的一般配合力、最大的特殊配合力方差,在播始历期上具有最高的一般配合力负效应;C815S在株高上的一般配合力负效应最大,同时特殊配合力方差较高。恢复系中,华抗恢101在单株有效穗数上具有最高的一般配合力和特殊配合力方差;华抗恢104在穗长上具有最高的一般配合力,在播始历期上具有最高的一般配合力负效应;华抗恢105在单株产量、结实率等性状上具有最高的一般配合力,在株高上具有最高的一般配合力负效应;华抗恢106在千粒重性状上具有最高的一般配合力和特殊配合力效应方差。利用抗除草剂恢复系配制杂交组合,不仅可以改良其除草剂抗性,也可以通过广泛测配,选择一般配合力强、特殊配合力方差大的亲本配组育成强优势组合。     

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.     

玉米雌穗发育期基因差异表达与杂种优势的研究

杂种优势在提高粮食作物特别是玉米的产量方面具有重要的作用。然而,杂种优势的原理却仍然是一个世界性的难题。用12个玉米自交系及其按不完全双列杂交组配的33个杂交种为材料,分4个不同发育时期取杂交种及其亲本的雌穗组织,利用差异显示技术,分析杂种与亲本的基因差异表达类型及其与7个主要农艺性状的杂种表现和杂种优势的相关关系。发现1):在5种表达类型中单态表达(基因在杂交种和双亲中同时表达的类型)的数量最大,这说明杂种优势的形成不仅与基因的表达与否相关,还与大量基因的上调或下调表达相关;2):在玉米雌幼穗的发育初期杂交种与双亲的基因表达差异最大,这可能与雌穗发育初期器官的形成和发育相关,因此这一时期差异表达(在质的方面)的基因对产量性状和杂种优势的形成具有密切关系;3):综合各种基因表达类型与产量性状和杂种优势的关系,发现某些基因在杂种中的沉默表达可以促进籽粒的发育和抑制幼穗中小花的发育。     

Inheritance of resistance against northern leaf blight of maize using conventional breeding methods

Maize (Zea mays L.) is one of the important cereal crops along with wheat and rice worldwide. The purpose of this study was to use classical genetic approaches to assess the resistance of various maize parents and hybrids to the northern corn leaf blight (NCLB) disease in two different locations in Egypt. Eight parents, 28 F1, and 2 check hybrids were evaluated. The analysis of variance showed high significant variations between maize parents and their hybrids for the studied parameters and NCLB disease, besides there are significant variations between both locations. Results of maize parents showed that Sids 63, Giza 602, and Giza 628 cultivars exhibited the highest values and were resistant to NCLB in both locations comparing with Nubaria 39 and Gemmiza 18 that were susceptible to NCLB disease. Concerning the maize hybrids, analysis of variance and mean squares of growth characters in both locations indicated high significant variations between the maize hybrids including the check hybrids. When combined between the two locations for current parameters against NCLB, the data pointed that the Sakha location values for maize hybrids were much closed to the combining data in parents and the hybrids detected high resistance to this disease comparing with Nubaria location. All tested maize lines (38 lines), including parents and hybrids were classified as follows, two lines were rated as 1 (highly resistant), three were rated as 2 (resistant), sixteen were rated as 3 (moderate resistant), eight were rated 4 (moderately susceptible) and nine were rated 5 (susceptible). The data explaining that the crossing between high resistant maize cultivars produced high levels of resistance to NCLB disease. Therefore, our results verified that classical breeding could efficiently increase the resistance levels of maize germplasm against NCLB disease by developing new cultivars with superior performance in terms of grain yield, disease resistance and grain quality.     

Differential gene expression in liver of inbred chickens and their hybrid offspring

Differential display of mRNA was used to analyse the differences of gene expression in liver between chicken hybrids and their parents in a 4 x 4 diallel crosses in order to study the molecular basis of heterosis in chickens. The results indicated that patterns of gene expression in hybrids differ significantly from their parents. Four patterns of differential gene expression were revealed, which included: (i) bands only detected in the hybrid F1s (UNF1); (ii) bands only absent in the hybrid F1s (ABF1); (iii) bands only detected in the parental P1 or P2 lines (UNP1 and UNP2) and (iv) bands absent in the parental P1 or P2 lines (ABP1 and ABP2). In addition, correlations between patterns of gene expression and heterosis percentages of nine carcass traits of 8-week-old chickens were evaluated. Statistical results showed that negative correlations between heterosis percentages and the percentage of F1-specific bands (UNF1) were significant at P < 0.01 for breast muscle yield, leg muscle yield, wing weight, eviscerated weight and eviscerated weight with giblet of 8-week-old chickens, and at P < 0.05 for intermuscular fat width. Heterosis percentage was negatively correlated with ABP (bands present in the hybrid F1s and one parental line but absent in the other parental line, ABP1 and ABP2) for breast muscle yield, leg muscle yield, wing weight, eviscerated weight and eviscerated weight with giblet of 8-week-old chickens (P < 0.01). Bands detected only in the hybrid F1s but not in either of the parental lines (UNF1) and bands absent in parental P1 or P2 lines (which includes ABP1 and ABP2) may play important roles in chicken heterosis.     

Heterotic patterns of sugar and amino acid components in developing maize kernels

Heterosis is the superior performance of hybrids over their inbred parents. Despite its importance, little is known about the genetic and molecular basis of this phenomenon. Heterosis has been extensively exploited in plant breeding, particularly in maize (Zea mays, L.), and is well documented in the B73 and Mo17 maize inbred lines and their F1 hybrids. In this study, we determined the dry matter, the levels of starch and protein components and a total of 24 low-molecular weight metabolites including sugars, sugar-phosphates, and free amino acids, in developing maize kernels between 8 and 30 days post-pollination (DPP) of the hybrid B73 × Mo17 and its parental lines. The tissue specificity of amino acid and protein content was investigated between 16 and 30 DPP. Key observations include: (1) most of the significant differences in the investigated tissue types occurred between Mo17 and the other two genotypes; (2) heterosis of dry matter and metabolite content was detectable from the early phase of kernel development onwards; (3) the majority of metabolites exhibited an additive pattern. Nearly 10% of the metabolites exhibited nonadditive effects such as overdominance, underdominance, and high-parent and low-parent dominance; (4) The metabolite composition was remarkably dependent on kernel age, and this large developmental effect could possibly mask genotypic differences; (5) the metabolite profiles and the heterotic patterns are specific for endosperm and embryo. Our findings illustrate the power of metabolomics to characterize heterotic maize lines and suggest that the metabolite composition is a potential marker in the context of heterosis research.     

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.     

Corn hybrids display lower metabolite variability and complex metabolite inheritance patterns

We conducted a comparative analysis of the root metabolome of six parental maize inbred lines and their 14 corresponding hybrids showing fresh weight heterosis. We demonstrated that the metabolic profiles not only exhibit distinct features for each hybrid line compared with its parental lines, but also separate reciprocal hybrids. Reconstructed metabolic networks, based on robust correlations between metabolic profiles, display a higher network density in most hybrids as compared with the corresponding inbred lines. With respect to metabolite level inheritance, additive, dominant and overdominant patterns are observed with no specific overrepresentation. Despite the observed complexity of the inheritance pattern, for the majority of metabolites the variance observed in all 14 hybrids is lower compared with inbred lines. Deviations of metabolite levels from the average levels of the hybrids correlate negatively with biomass, which could be applied for developing predictors of hybrid performance based on characteristics of metabolite patterns.     

不同类型杂交水稻组合及其亲本苗期同工酶位点分析

对6个杂交水稻组合及其亲本苗期14种酶的同工酶进行检测,并对其中较为清晰稳定的酶带进行了位点分析,以探讨杂种优势遗传基础。共获得了28个位点的相关资料,包括每个位点的性质、表型多态性和每个位点所编码的酶活性及酶带平均迁移率。所有供试材料在其中25个位点上均未表现出多态性,只有部分组合与其亲本在Est-4、Amy-1、Aat-2等3个位点上存在多态性,其中有81．8％的差异组合其酶带表现出共显性特征。对共显性与杂种优势的关系进行了探讨。