基于混合线性模型的油用向日葵农艺性状及产质量遗传研究
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摘要
本研究利用无血缘关系、具有丰富遗传背景的15个自交系，按照不完全双列杂交（NC designⅡ）方式，构建出包括亲本群体、F1群体和F2群体在内的共计64个遗传群体及交配体系，采用包括环境互作在内的加性-显性遗传模型（AD遗传模型）、加性-显性-母体遗传模型（ADM遗传模型）以及二倍体种子遗传模型，分析了油用向日葵的7个主要农艺性状、产量、含油率、亚油酸含量以及油酸含量的遗传规律、基因效应及特点；并利用多元条件分析方法，分析了农艺性状对产量、含油率的遗传贡献率。旨在为油用向日葵“三系”育种实践和目标性状的间接、直接选择提供依据；为自交系改良及杂种优势利用提供理论指导。本研究结果如下，
1．所观测的7个农艺性状中，株高、茎粗、盘径、百粒重、籽仁率和单盘粒重等6个农艺性状主要受加性效应和显性效应共同控制，但以加性遗传效应为主。在加性效应中，以单盘粒重的最高，籽仁率的最低；在显性效应中，籽仁率的显性效应最强，结实率的最低。结实率的遗传表现主要受控于加性效应和显性×环境互作；性状间的遗传相关性以加性遗传相关为主。
2．单盘粒重、株高和百粒重对产量的遗传贡献大，贡献率在96.0%以上。产量效应的表达主要以显性效应（杂种优势）为主。多数杂交种的单盘粒重对其产量具有高水平的贡献值，单盘粒重的显性效应可作为杂交种产量的首选选择指标；株高、百粒重可作为杂交种显性效应的辅助选择性状。对提高杂交种产量而言，显性效应比加性效应更为重要。
3．所观测的7个农艺性状对籽实含油率的遗传贡献程度有所不同。在加性贡献中，百粒重、盘径、单盘粒重和结实率的加性净遗传效应对籽实含油率的贡献率较大，其中以百粒重的加性贡献率最高为82.32%；在显性贡献中，最有利于通过杂种优势来提高籽实含油率的性状为结实率。在提高籽实含油率方面，农艺性状对含油率的加性贡献比非加性贡献更为重要，农艺性状与单位面积产油量以显性相关为主。
4．油用向日葵在不同发育时期的地上干物质累积量，主要受等位基因的显性效应影响，出苗后40～70d和90～100d是显性效应基因的开启处于相对稳定、高水平表达时期，这两个时间段正好处于现蕾-开花和灌浆-生理成熟时期。因此，该时期是加强田间管理及水肥运筹调控的时期。特殊环境下，苗前期易受环境影响；地上干物质重的杂优表现以群体平均优势为主，也存在一定的超亲优势；在出苗50d内，群体平均优势和群体超亲优势表达明显，在此时段内干物质积累量对产量的显性效应贡献率最高。
5．油用向日葵籽实含油率的遗传表现主要取决于显性遗传效应和母体遗传效应。开花20～25d时段是加性效应表达的高峰期；开花20～30d时段是显性微效多基因被激活及表达的活跃期，即杂优表现突出的时期；母体效应最活跃期为开花后的35～40d时段；在开花20d内（开花初期和中期），基因效应的表达易受到环境条件的影响。
6．含油率、亚油酸含量、油酸含量同时受胚、细胞质及母体植株等3套遗传体系基因的影响。含油率和亚油酸含量主要受控于细胞质效应和母体效应，油酸含量的遗传表现以细胞质互作效应为主。含油率、亚油酸含量的细胞质遗传率大于胚遗传率和母体遗传率；含油率、亚油酸含量和油酸含量之间的相关性以细胞质相关和母体相关为主，其中含油率、亚油酸含量与油酸含量呈负相关，含油率与亚油酸含量呈正相关；控制含油率和亚油酸含量的细胞质基因及母体显性基因存在基因连锁或一因多效。
The research used15inbred lines of the sunflower which had no blood relationship andhad rich genetic background. According to incomplete diallel cross (NC Design II), the64populations were constructed, which included the parental populations, F1populations andF2populations. The genetic models of Additive-Dominance,Additive-Dominance-Maternaland Diploid-Embryo considering Gene-Environment interaction and the method ofmultivariate conditional analysis were employed to evaluate the gene effects andcharacteristics of the main agronomic and quality traits such as yield and oil content. Thepurpose were providing reference for the direct or indirect selection of the target traits andguidance for inbred lines selection and the utilization of the heterosis in oil sunflowerbreeding by Three-Line-Systems. The results were summarized as the followings,
1. Six agronomic characters, such as plant height, stem diameter, disc diameter,weight of100grains, kernel percentage, grain weight per disc were mainly controlled bythe additive effect and dominance effect, but the additive effect was more important. Foradditive effect, it was predominant on the grain weight per disc, while it was slightest onkernel percentage. For dominance effect, it was predominant on the kernel percentage,while it was small fertility. The fertility was mainly controlled by the additive effect anddominance×environment effect. Genetic correlation between traits was controlled by theadditive genes.
2. The main contribution to yield was from grain weight per disc, plant height andweight of100grains, which exceeded96%respectively. The grain per disc of most hybridshad a high level of contribution value to yield and yield expression was mainly controlledby the dominance effect (heterosis). According to enhancing the yield of hybrid, it wastherefore proposed that the dominance effect of the grain weight per disc could used asprimary selecting index for hybrid yield and plant height and weight of100grains assecondary indexes. As to enhance the yield of hybrid, the dominance effect seems to bemore important than the additive effect.
3. The observed seven agronomic characters made different levels of geneticcontribution to oil content of seeds. The great contribution to oil content was made by theadditive net genetic effect of weight of100grains, disc diameter, grain weight per disc andfertile percentage, among them weight of100grains made the maximum with82.32%. As to enhance the oil content of seeds, the additive contribution of agronomic charactersseems to be more important than the non-additive contribution. Among the dominancecontribution, the genetic trends of fertility percentage were favorable. Those weredominant genetic correlation between agronomic characters and oil production per unitarea.
4. The accumulation biomass of aboveground dry matter of oil sunflower in differentdevelopmental stages was mainly affected by the dominance alleles. The phases of40～70d and90～100d after seed emerging were budding-flowering and grouting-physiologicalmaturing respectively, among which the dominant effect gene’s expression was steady,continuous and high. In the two phases it was suggested that field management bereinforced and care about the regulation on water and fertilizer. In specific environment,the seedling was easy to be affected by environments at the early stage of seed emergence.Heterosis of aboveground dry matter was mainly of population herterosis over mid-parentswith some of heterobeltiosis. In50d after seed emergence, population heterosis overmid-parents and over better-parents was most obvious phase. From seed emergence to50d,the accumulation biomass of dry matter made the highest contribution to the dominanteffect of yield.
5. The oil content of seeds was controlled by both dominance effect and maternaleffect. The additive effect had been detected at peak during the days of20～25d afterflowered, and there were highly significant dominant effects during20～30d afterflowering. It was the predominant stage of heterosis performance, while the maternal effectwas active from35to45days after flowering. The expression of gene effect was easy to beaffected by environment in20d after flowering.
6. The oil content(OC), linoleic acid content(LAC) and oleic acid content(OAC) traitswere simultaneously controlled by the genetic effects from embryo genes,cytoplasmicgenes and maternal plant genes.OC and LAC were mainly controlled by cytoplasmic andmaternal genetic effects,while expression of OAC was mainly controlled bycytoplasmic×environment interaction effect.The cytoplasmic heritabilities were larger thanthe embryo and maternal heritabilities for OC and LAC traits,but thecytoplasmic×environment interaction heritabilities were more important than others effectfor OAC.It were more important that the positive cytoplasmic correlations(rC) and maternaldominant correlations(rDm) between the OC and LAC, while OC was negative correlationwith OAC; Moreover,Significant negative correlation were observed between the LAC andOAC, including cytoplasmic effect,maternal additive effect, maternal dominant effect and embryo dominant×environment interaction effect.There were genetic linkage orpleiotropism among cytoplasmic genes of OC and LAC,and including maternal dominantgene. while genetic linkage or pleiotropism were revealed among maternal additive geneand maternal dominant gene for OAC and LAC.
1．所观测的7个农艺性状中，株高、茎粗、盘径、百粒重、籽仁率和单盘粒重等6个农艺性状主要受加性效应和显性效应共同控制，但以加性遗传效应为主。在加性效应中，以单盘粒重的最高，籽仁率的最低；在显性效应中，籽仁率的显性效应最强，结实率的最低。结实率的遗传表现主要受控于加性效应和显性×环境互作；性状间的遗传相关性以加性遗传相关为主。
2．单盘粒重、株高和百粒重对产量的遗传贡献大，贡献率在96.0%以上。产量效应的表达主要以显性效应（杂种优势）为主。多数杂交种的单盘粒重对其产量具有高水平的贡献值，单盘粒重的显性效应可作为杂交种产量的首选选择指标；株高、百粒重可作为杂交种显性效应的辅助选择性状。对提高杂交种产量而言，显性效应比加性效应更为重要。
3．所观测的7个农艺性状对籽实含油率的遗传贡献程度有所不同。在加性贡献中，百粒重、盘径、单盘粒重和结实率的加性净遗传效应对籽实含油率的贡献率较大，其中以百粒重的加性贡献率最高为82.32%；在显性贡献中，最有利于通过杂种优势来提高籽实含油率的性状为结实率。在提高籽实含油率方面，农艺性状对含油率的加性贡献比非加性贡献更为重要，农艺性状与单位面积产油量以显性相关为主。
4．油用向日葵在不同发育时期的地上干物质累积量，主要受等位基因的显性效应影响，出苗后40～70d和90～100d是显性效应基因的开启处于相对稳定、高水平表达时期，这两个时间段正好处于现蕾-开花和灌浆-生理成熟时期。因此，该时期是加强田间管理及水肥运筹调控的时期。特殊环境下，苗前期易受环境影响；地上干物质重的杂优表现以群体平均优势为主，也存在一定的超亲优势；在出苗50d内，群体平均优势和群体超亲优势表达明显，在此时段内干物质积累量对产量的显性效应贡献率最高。
5．油用向日葵籽实含油率的遗传表现主要取决于显性遗传效应和母体遗传效应。开花20～25d时段是加性效应表达的高峰期；开花20～30d时段是显性微效多基因被激活及表达的活跃期，即杂优表现突出的时期；母体效应最活跃期为开花后的35～40d时段；在开花20d内（开花初期和中期），基因效应的表达易受到环境条件的影响。
6．含油率、亚油酸含量、油酸含量同时受胚、细胞质及母体植株等3套遗传体系基因的影响。含油率和亚油酸含量主要受控于细胞质效应和母体效应，油酸含量的遗传表现以细胞质互作效应为主。含油率、亚油酸含量的细胞质遗传率大于胚遗传率和母体遗传率；含油率、亚油酸含量和油酸含量之间的相关性以细胞质相关和母体相关为主，其中含油率、亚油酸含量与油酸含量呈负相关，含油率与亚油酸含量呈正相关；控制含油率和亚油酸含量的细胞质基因及母体显性基因存在基因连锁或一因多效。
The research used15inbred lines of the sunflower which had no blood relationship andhad rich genetic background. According to incomplete diallel cross (NC Design II), the64populations were constructed, which included the parental populations, F1populations andF2populations. The genetic models of Additive-Dominance,Additive-Dominance-Maternaland Diploid-Embryo considering Gene-Environment interaction and the method ofmultivariate conditional analysis were employed to evaluate the gene effects andcharacteristics of the main agronomic and quality traits such as yield and oil content. Thepurpose were providing reference for the direct or indirect selection of the target traits andguidance for inbred lines selection and the utilization of the heterosis in oil sunflowerbreeding by Three-Line-Systems. The results were summarized as the followings,
1. Six agronomic characters, such as plant height, stem diameter, disc diameter,weight of100grains, kernel percentage, grain weight per disc were mainly controlled bythe additive effect and dominance effect, but the additive effect was more important. Foradditive effect, it was predominant on the grain weight per disc, while it was slightest onkernel percentage. For dominance effect, it was predominant on the kernel percentage,while it was small fertility. The fertility was mainly controlled by the additive effect anddominance×environment effect. Genetic correlation between traits was controlled by theadditive genes.
2. The main contribution to yield was from grain weight per disc, plant height andweight of100grains, which exceeded96%respectively. The grain per disc of most hybridshad a high level of contribution value to yield and yield expression was mainly controlledby the dominance effect (heterosis). According to enhancing the yield of hybrid, it wastherefore proposed that the dominance effect of the grain weight per disc could used asprimary selecting index for hybrid yield and plant height and weight of100grains assecondary indexes. As to enhance the yield of hybrid, the dominance effect seems to bemore important than the additive effect.
3. The observed seven agronomic characters made different levels of geneticcontribution to oil content of seeds. The great contribution to oil content was made by theadditive net genetic effect of weight of100grains, disc diameter, grain weight per disc andfertile percentage, among them weight of100grains made the maximum with82.32%. As to enhance the oil content of seeds, the additive contribution of agronomic charactersseems to be more important than the non-additive contribution. Among the dominancecontribution, the genetic trends of fertility percentage were favorable. Those weredominant genetic correlation between agronomic characters and oil production per unitarea.
4. The accumulation biomass of aboveground dry matter of oil sunflower in differentdevelopmental stages was mainly affected by the dominance alleles. The phases of40～70d and90～100d after seed emerging were budding-flowering and grouting-physiologicalmaturing respectively, among which the dominant effect gene’s expression was steady,continuous and high. In the two phases it was suggested that field management bereinforced and care about the regulation on water and fertilizer. In specific environment,the seedling was easy to be affected by environments at the early stage of seed emergence.Heterosis of aboveground dry matter was mainly of population herterosis over mid-parentswith some of heterobeltiosis. In50d after seed emergence, population heterosis overmid-parents and over better-parents was most obvious phase. From seed emergence to50d,the accumulation biomass of dry matter made the highest contribution to the dominanteffect of yield.
5. The oil content of seeds was controlled by both dominance effect and maternaleffect. The additive effect had been detected at peak during the days of20～25d afterflowered, and there were highly significant dominant effects during20～30d afterflowering. It was the predominant stage of heterosis performance, while the maternal effectwas active from35to45days after flowering. The expression of gene effect was easy to beaffected by environment in20d after flowering.
6. The oil content(OC), linoleic acid content(LAC) and oleic acid content(OAC) traitswere simultaneously controlled by the genetic effects from embryo genes,cytoplasmicgenes and maternal plant genes.OC and LAC were mainly controlled by cytoplasmic andmaternal genetic effects,while expression of OAC was mainly controlled bycytoplasmic×environment interaction effect.The cytoplasmic heritabilities were larger thanthe embryo and maternal heritabilities for OC and LAC traits,but thecytoplasmic×environment interaction heritabilities were more important than others effectfor OAC.It were more important that the positive cytoplasmic correlations(rC) and maternaldominant correlations(rDm) between the OC and LAC, while OC was negative correlationwith OAC; Moreover,Significant negative correlation were observed between the LAC andOAC, including cytoplasmic effect,maternal additive effect, maternal dominant effect and embryo dominant×environment interaction effect.There were genetic linkage orpleiotropism among cytoplasmic genes of OC and LAC,and including maternal dominantgene. while genetic linkage or pleiotropism were revealed among maternal additive geneand maternal dominant gene for OAC and LAC.
引文
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