杂交稻骨干亲本“蜀恢527”产量性状及其一般配合力的遗传基础研究
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摘要
骨干亲本在品种培育过程中发挥了至关重要的作用。本研究以籼型杂交稻骨干恢复系蜀恢527为轮回亲本，以高产恢复系ZDZ057、辐恢838和特青为供体，构建3个高代回交产量选择群体。在不同环境下，进行产量及其相关性状表型评价，结合基因型分析，充分发掘供体中优良等位基因。同时，以高代回交群体中的57个产量导入系为父本，与4类核心不育系，即：协青早A（协A，矮败型）、II-32A（IIA，印水型）、冈46A（冈A，冈型）和金23A（金A，野败型）分别测交，剖析产量及其相关性状一般配合力和杂种优势遗传基础。最后，在测交F1组合中，选择与对照组合协A/蜀恢527相比，产量优势表现稳定的极端组合：协A/WD7（低产比较优势杂种）和协A/WD34（高产比较优势杂种）。通过RNA-Seq对这3个杂种及4个亲本进行苗期、分蘖期和幼穗分化期转录组分析，探讨杂种优势形成机理。主要研究结果如下：
1．在两个环境（安徽和海南）下，3个产量选择群体共检测到94个与产量及其相关性状紧密连锁的QTL，这些位点主要集中在12个QTL簇上。在所有位点中，60.1%QTL有利等位基因来自供体，40%QTL能在不同环境下被重复检测到。
2．本研究发现，影响每穗实粒数、每穗总粒数和单株产量性状的位点更多是以两个或多个QTL的非随机关联在高性状值导入系中存在。7个位点在不同群体间存在复等位基因，并且复等位基因间存在效应上的差异。
3．配合力分析发现，产量及其相关性状受基因加性和非加性效应共同影响。而且，除单株产量外，产量相关性状主要受加性效应控制。
4.两个群体共检测到34个与产量及其相关性状一般配合力紧密连锁的QTL。其中，21个（61.8%）QTL有利等位基因来自骨干亲本蜀恢527。高、低产量一般配合力导入系分组比较分析发现，标记RM508和RM587在8个低产一般配合力导入系中被供体ZDZ057或特青等位基因一致替换，同时，在高产一般配合力导入系WD34、WD36和WD43中，标记RM4584和RM224被ZDZ057等位基因一致导入。根据一般配合力定位结果，这些标记与产量及主要构成因子QTL紧密连锁。由此推断，这些位点可能是引起蜀恢527产量一般配合力变化的重要区段。
5．应用比较优势（comparative heterosis, HC）和中亲优势（mid-parent heterosis, HMP），在8个测交群体定位到79个与产量及其相关性状杂种优势连锁的QTL。其中，49.4%QTL（39）是以超显性形式存在，特别是对于单株产量和单株有效穗性状；而加性主要是对于每穗实粒数和每穗总粒数。
6．转录组分析发现，3个时期，杂种协A/WD7与亲本间差异基因最多（3589个），其次是杂种协A/WD34（3146个），最后是协A/蜀恢527（2912个）。除了杂种协A/WD7在分蘖期外，3个杂种在3个时期全部表现为加性表达基因多于非加性，而且无论是加性还是非加性表达基因，都是上调表达基因多于下调表达基因。
7．对照组合协A/蜀恢527与亲本间差异表达基因GO及代谢途径分析发现，参与光合碳同化和碳水化合物代谢过程中淀粉和蔗糖合成关键基因在杂种中表达明显上调，可能是对照组合协A/蜀恢527杂种优势形成的主要原因。
8．相对于对照组合，在低产比较优势杂种协A/WD7中参与光合作用光捕获以及转录调控相关基因表达明显下调，这些基因表达下调与其产量优势下降关系密切。另外，在高产比较优势杂种协A/WD34中，生物钟节律相关基因OsTOC1和OsGI表达上调，OsCCA1表达下调，而且大量碳水化合物代谢相关基因表达上调，这些基因表达变化可能最终导致其产量优势提高。由于OsGI过表达导致导入系WD34和杂种协A/WD34抽穗期延迟。
Key parents play an important role in the processing of hybrid breeding. In this study, threeselected advanced backcross populations were constructed using Shuhui527(SH527) as recurrent parent,a key parent of hybrid rice, and three high-yielding cultivars i.e. ZDZ057, Fuhui838(FH838) andTeqing as donors, respectively. Phenotyping and genotyping for these three populations were conductedfor identifying QTL associated with yield and its related traits from donors under different environments.Secondly,228F1combinations were developed by testcrossing57introgression lines (ILs) fromabove-mentioned populations with four types of male sterile lines i.e. XieqingzaoA (XA), II-32A (IIA),Gang46A (GA) and Jin23A (JA), and evaluated for two years in Anhui for detecting on the geneticbases of general combining ability (GCA) and heterosis affecting yield and its related traits. Lastly,tanscriptomic analysis was conducted for two hybrids with high-and low-yielding comparativeheterosis (HC) selected from228testcrossing combinations combing with check combinations (CC)with their parents at three stages for elucidating the mechanism of heterosis. The main resultssummarized as follows:
1. A total of94QTL controlling yield and its related traits were identified under two environments,and mainly concentrated on12QTL clusters. For these QTL,60.1%QTL had favorable alleles from thedonors and about40%QTL were detected under two environments.
2. The strong and frequent non-random associations between or among QTL affecting filled grainnumber per panicle (GN), spikelets number per panicle (SN) and grain yield per plant (GY) wereobserved in the ILs with high-trait values. Multi-alleles were found in seven QTL and had differenteffects among populations.
3. Additive and non-additive gene effects jointly determined the expression of yield and its relatedtraits. Moreover, addictive effects were more important than non-addictive for all traits except GY.
4. Thirty-four QTL affecting GCA of yield and its related traits were identified in the twopopulations. Of these,61.8%(21) had favorable alleles from key parent, SH527. Markers RM508andRM587, RM4584and RM224were substituted by ZDZ057alleles in eight ILs with Low-yielding GCAand Teqing alleles in the three ILs with high-yielding GCA, respectively. According to the QTLmapping results, these markers tightly associated with yield and key components, which can be inferredthat these loci probably played a key role in controlling yield GCA of SH527.
5. Seventy-nine QTL associated with heterosis of yield and its related traits were identified usingcomparative heterosis (HC) and mid-parent heterosis (HMP) values in the eight testcrossing populations.Of these,49.4%QTL present overdominance, especially for panicle number (PN) and GY, andconversely, additivity for GN and SN.
6. The great numbers of differential expression genes (DGEs) were found between XA/WD7withits parents (3589) at three stages by transcriptomic analysis, second is XA/WD34with XA and WD34(3146), and last is XA/SH527and its parents (2912). Except XA/WD7at tillering stage, three hybrids had more additive expression genes than non-additive at three stages, furthermore, the number ofup-regulated genes were always more than down-regulated whether or not additive and non-additiveexpression genes.
7. In comparison with parents XA and SH527, a lot of up-regulated genes related to carbonassimilation in photosynthesis and starch and sucrose biosynthesis in carbohydrate metabolism wereobserved in hybrid XA/SH527, which probably resulted in that XA/SH527had strong yield vigor thanits parents.
8. Comparing with the check combination, numerous down-regulated genes associated withlight-harvesting in photosynthesis and regulation of transcription exsited in XA/WD7. In addition,expression changes related to circadian rhythm (CCA1, TOC1and GI) and carbohydrate metabolismwere observed in hybrid XA/WD34. These DGEs could play an important role in affecting yieldheterosis in XA/WD7and XA/WD34. Furthermore, Heading dates (HD) of WD34and XA/WD34weredelayed due to the overexpression of OsGI gene.
1．在两个环境（安徽和海南）下，3个产量选择群体共检测到94个与产量及其相关性状紧密连锁的QTL，这些位点主要集中在12个QTL簇上。在所有位点中，60.1%QTL有利等位基因来自供体，40%QTL能在不同环境下被重复检测到。
2．本研究发现，影响每穗实粒数、每穗总粒数和单株产量性状的位点更多是以两个或多个QTL的非随机关联在高性状值导入系中存在。7个位点在不同群体间存在复等位基因，并且复等位基因间存在效应上的差异。
3．配合力分析发现，产量及其相关性状受基因加性和非加性效应共同影响。而且，除单株产量外，产量相关性状主要受加性效应控制。
4.两个群体共检测到34个与产量及其相关性状一般配合力紧密连锁的QTL。其中，21个（61.8%）QTL有利等位基因来自骨干亲本蜀恢527。高、低产量一般配合力导入系分组比较分析发现，标记RM508和RM587在8个低产一般配合力导入系中被供体ZDZ057或特青等位基因一致替换，同时，在高产一般配合力导入系WD34、WD36和WD43中，标记RM4584和RM224被ZDZ057等位基因一致导入。根据一般配合力定位结果，这些标记与产量及主要构成因子QTL紧密连锁。由此推断，这些位点可能是引起蜀恢527产量一般配合力变化的重要区段。
5．应用比较优势（comparative heterosis, HC）和中亲优势（mid-parent heterosis, HMP），在8个测交群体定位到79个与产量及其相关性状杂种优势连锁的QTL。其中，49.4%QTL（39）是以超显性形式存在，特别是对于单株产量和单株有效穗性状；而加性主要是对于每穗实粒数和每穗总粒数。
6．转录组分析发现，3个时期，杂种协A/WD7与亲本间差异基因最多（3589个），其次是杂种协A/WD34（3146个），最后是协A/蜀恢527（2912个）。除了杂种协A/WD7在分蘖期外，3个杂种在3个时期全部表现为加性表达基因多于非加性，而且无论是加性还是非加性表达基因，都是上调表达基因多于下调表达基因。
7．对照组合协A/蜀恢527与亲本间差异表达基因GO及代谢途径分析发现，参与光合碳同化和碳水化合物代谢过程中淀粉和蔗糖合成关键基因在杂种中表达明显上调，可能是对照组合协A/蜀恢527杂种优势形成的主要原因。
8．相对于对照组合，在低产比较优势杂种协A/WD7中参与光合作用光捕获以及转录调控相关基因表达明显下调，这些基因表达下调与其产量优势下降关系密切。另外，在高产比较优势杂种协A/WD34中，生物钟节律相关基因OsTOC1和OsGI表达上调，OsCCA1表达下调，而且大量碳水化合物代谢相关基因表达上调，这些基因表达变化可能最终导致其产量优势提高。由于OsGI过表达导致导入系WD34和杂种协A/WD34抽穗期延迟。
Key parents play an important role in the processing of hybrid breeding. In this study, threeselected advanced backcross populations were constructed using Shuhui527(SH527) as recurrent parent,a key parent of hybrid rice, and three high-yielding cultivars i.e. ZDZ057, Fuhui838(FH838) andTeqing as donors, respectively. Phenotyping and genotyping for these three populations were conductedfor identifying QTL associated with yield and its related traits from donors under different environments.Secondly,228F1combinations were developed by testcrossing57introgression lines (ILs) fromabove-mentioned populations with four types of male sterile lines i.e. XieqingzaoA (XA), II-32A (IIA),Gang46A (GA) and Jin23A (JA), and evaluated for two years in Anhui for detecting on the geneticbases of general combining ability (GCA) and heterosis affecting yield and its related traits. Lastly,tanscriptomic analysis was conducted for two hybrids with high-and low-yielding comparativeheterosis (HC) selected from228testcrossing combinations combing with check combinations (CC)with their parents at three stages for elucidating the mechanism of heterosis. The main resultssummarized as follows:
1. A total of94QTL controlling yield and its related traits were identified under two environments,and mainly concentrated on12QTL clusters. For these QTL,60.1%QTL had favorable alleles from thedonors and about40%QTL were detected under two environments.
2. The strong and frequent non-random associations between or among QTL affecting filled grainnumber per panicle (GN), spikelets number per panicle (SN) and grain yield per plant (GY) wereobserved in the ILs with high-trait values. Multi-alleles were found in seven QTL and had differenteffects among populations.
3. Additive and non-additive gene effects jointly determined the expression of yield and its relatedtraits. Moreover, addictive effects were more important than non-addictive for all traits except GY.
4. Thirty-four QTL affecting GCA of yield and its related traits were identified in the twopopulations. Of these,61.8%(21) had favorable alleles from key parent, SH527. Markers RM508andRM587, RM4584and RM224were substituted by ZDZ057alleles in eight ILs with Low-yielding GCAand Teqing alleles in the three ILs with high-yielding GCA, respectively. According to the QTLmapping results, these markers tightly associated with yield and key components, which can be inferredthat these loci probably played a key role in controlling yield GCA of SH527.
5. Seventy-nine QTL associated with heterosis of yield and its related traits were identified usingcomparative heterosis (HC) and mid-parent heterosis (HMP) values in the eight testcrossing populations.Of these,49.4%QTL present overdominance, especially for panicle number (PN) and GY, andconversely, additivity for GN and SN.
6. The great numbers of differential expression genes (DGEs) were found between XA/WD7withits parents (3589) at three stages by transcriptomic analysis, second is XA/WD34with XA and WD34(3146), and last is XA/SH527and its parents (2912). Except XA/WD7at tillering stage, three hybrids had more additive expression genes than non-additive at three stages, furthermore, the number ofup-regulated genes were always more than down-regulated whether or not additive and non-additiveexpression genes.
7. In comparison with parents XA and SH527, a lot of up-regulated genes related to carbonassimilation in photosynthesis and starch and sucrose biosynthesis in carbohydrate metabolism wereobserved in hybrid XA/SH527, which probably resulted in that XA/SH527had strong yield vigor thanits parents.
8. Comparing with the check combination, numerous down-regulated genes associated withlight-harvesting in photosynthesis and regulation of transcription exsited in XA/WD7. In addition,expression changes related to circadian rhythm (CCA1, TOC1and GI) and carbohydrate metabolismwere observed in hybrid XA/WD34. These DGEs could play an important role in affecting yieldheterosis in XA/WD7and XA/WD34. Furthermore, Heading dates (HD) of WD34and XA/WD34weredelayed due to the overexpression of OsGI gene.
引文
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