应用QTL分析研究籼稻稻米品质和产量性状的关系
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摘要
高产优质是当下水稻育种重要目标，本研究利用抽穗期变异范围相对较小的特青/IRBB组合所衍生的重组自交系群体，结合分子遗传图谱、稻米品质13个评价指标和产量6个相关性状两年表型数据，分析稻米品质与产量之间遗传控制关系，在理论上减小了不同抽穗期的生态环境（例如，温度和光照）对实验结果的影响，可以较为客观地反映稻米品质和产量之间的遗传关系。
在本研究中，我们共计检测到7个与稻米品质和水稻产量均相关的染色体区域，主要分布于水稻第3、5、6、7、8和10染色体，控制稻米品质和产量的QTL在这些染色体区域呈簇分布。分析位于这些区域米质和产量QTL加性效应方向，结果表明，控制二者的QTL增效等位基因往往来自于不同亲本，即，这些区域在提高产量的同时，致使稻米品质降低，这可能是育种实践中，转育优良农艺性状的同时，常伴随着不利性状的转育，即出现遗传累赘现象的重要原因。
第3染色体着丝粒RM15139-RM16区域：在该区域检测到控制千粒重、每穗实粒数、每穗总粒数和结实率4个产量相关性状以及稻米品质包括碾磨品质、外观品质和粒型3个方面的QTL。千粒重与每穗实粒数和每穗总粒数的增效等位基因分别来自于IRBB和特青，即，该区域在增加千粒重同时降低每穗粒数，位于该区域控制糙米率、粒长、长宽比、垩白米率、垩白度和透明度QTL增效等位基因均来自于IRBB，而整精米率和粒宽QTL增效等位基因来自于特青。
第5染色体短臂RM437-RM18189区域：在该区域仅检测到2个产量相关性状千粒重和结实率QTL，增效等位基因分别来自于特青和IRBB，所检测到的控制稻米品质QTL涉及碾磨品质、外观品质和粒型3个方面。该区域在增加千粒重的同时，垩白度、糙米率、精米率、粒宽及垩白米率亦随之增加，而结实率、粒长、长宽比和透明度降低。
第6染色体Wx基因区域：在该区域除了检测到控制稻米蒸煮食味品质3个评价指标的QTL外，同时检测到控制粒宽、千粒重、糙米率、精米率和垩白的QTL，除控制胶稠度的QTL外，其它QTL增效等位基因均来自于特青。
第6染色体ALK基因区域：在该区域检测到控制碱消值、胶稠度、每穗实粒数和每穗总粒数的QTL，除碱消值外增效等位基因均来自于特青。
第7染色体长臂RM70-RM18区域：在该区域检测到控制每穗总粒数、糙米率、透明度和直链淀粉含量QTL，除每穗总粒数QTL来自于特青外，其它均来自于IRBB。
第8染色体短臂RM5647-RM22755区域：在本研究中，第8染染色体短臂为一多功能区域，同时控制每穗实粒数、每穗总粒数、结实率、粒宽、长宽比、糙米率和垩白米率7个性状，除控制长宽比和结实率的QTL来自于IRBB外，其它均来自于特青。
第10染色体长臂近末端RM6100-RM3123区域：该区域检测到控制千粒重、垩白米率、垩白度、糙米率、粒宽和透明度QTL，除透明度外，控制其它性状的QTL均来自于特青。
另外，在本研究中共计检测到29对互作QTL，除第1染色体外，其它11条染色体均参与其中。这些互作遗传效应较小，对性状表型变异贡献率多在1%以下，一个位点往往与多个位点同时产生互作效应，如第2染色体长臂末端控制千粒重主效应QTL qTGW2.2，与多个主效应QTL之间均存在互作。在所检测到29对互作QTL中，主效应QTL之间互作是其主要部分。
Currently，high yield and quality is one of important aims for rice hybrid breeding. A set ofrecombinant inbred line (RIL) population characterized by a relative narrow range of heading date anddeveloped from crosses between Teqing and IRBB lines was utilized to analyze the genetic relationshipbetween rice quality involving13indexes and6grain yield-related traits of indica rice with theassistance of molecular linkage map and phenotypic data. Theoretically, the effect of ecologicalenvironment on assay results due to various heading date among the lines was diminished, as a result,the genetic relationship between rice quality and grain yield was reflected objectively.
In this study, a total of7chromosomal regions related to both rice quality and grain yield weredetected. They were mainly distributed on chromosome3,5,6,7,8and10, on which QTLsconditioning rice quality and grain yield were clustered together. Surveying the additive directions ofthese QTLs, we found that alleles for increasing rice quality and grain yield were often from differentparents. That is, grain yield were improved due to a certain multi-function region followed bydeleterious effects on rice quality. This may be the main cause for cultivating some elite agronomictraits at the expense of deleterious agronomic traits, that is, genetic drag in rice hybrid breeding practice.
RM15139-RM16region on chromosome3: QTLs for4grain yield-related traits including1000-grain weight (TGW), number of grains per panicle (NGP), number of spikelets per panicle (NSP)and spikelet fertility (SF) were detected as well as QTLs for rice quality involving rice milled quality,appearance quality and grain shape in this region. Allele for increasing TGW in this region was fromIRBB while the enhancing alleles of NGP and NSP were from Teqing, which means that this regionincreased TGW with NGP and NSP decreased simultaneously. QTLs conditioning brown rice ratio(BRR), grain length (GL), ratio of grain length to grain width (LWR), percentage of chalky grain (PCG),degree of chalkiness (DC) and the endosperm transparency (ET) were also detected in this region andall QTLs had the enhancing alleles from IRBB, however, QTLs detected in this region for head riceratio (HRR) and grain width (GW) both had the enhancing alleles from Teqing.
RM15139-RM16region on chromosome5: QTLs for TGW and SF were the only two grainyield-related QTLs detected in this region and the enhancing alleles were from Teqing and IRBB,respectively. QTLs detected for rice quality involved3aspects including rice milled quality, appearancequality and grain shape in this region. This region increased TGW, DC, BRR, milled rice ratio (MRR),GW and PCG with SF, GL, LWR and ET decreased simultaneously.
Wx region on chromosome6: In additional to3QTLs conditioning3indexes for cooking andeating quality respectively were detected in this region, QTL for GW, TGW, BRR, MRR and chalkinesswere also detected in this region. All QTLs except QTL for gel consistency (GC) had enhancing allelesfrom Teqing.
Alk region on chromosome6: QTLs conditioning alkali-spread score (ALK), GC, NGP and NSPwere detected in this region and all QTLs except QTL for ALK had enhancing alleles from Teqing.
RM70-RM18region on chromosome7: QTLs conditioning NSP, BRR, ET and amylose content(AC) were detected in this region and all QTLs except QTL for NSP had enhancing alleles from IRBB.
RM5647-RM22755region on chromosome8: In our study, the short arm of chromosome8was amultifunction region and controlled7rice traits including NGP, NSP, SF, GW, LWR, BRR and PCG.All QTLs except QTLs for LWR and SF had enhancing alleles from Teqing.
RM6100-RM3123region on chromosome10: QTLs conditioning TGW, PCG, DC, BRR, GW andET were detected in this region and all QTLs except QTL for ET had enhancing alleles from Teqing.
Additionally, a total of29couple epistatic QTLs involving with all chromosomes exceptchromosome1were detected in this study. These epistatic QTLs had a relative small genetic effect andthe phenotypic variation explained by them was often under1%. One locus usually interacted withmany loci and generated genetic effects, for example, main-effect QTL qTGW2.2located on the distalof long arm of chromosome2interacted with many main-effect QTLs for TGW. Among the29coupleepistatic QTLs, interactions between main-effect QTLs accounted for a most part.
在本研究中，我们共计检测到7个与稻米品质和水稻产量均相关的染色体区域，主要分布于水稻第3、5、6、7、8和10染色体，控制稻米品质和产量的QTL在这些染色体区域呈簇分布。分析位于这些区域米质和产量QTL加性效应方向，结果表明，控制二者的QTL增效等位基因往往来自于不同亲本，即，这些区域在提高产量的同时，致使稻米品质降低，这可能是育种实践中，转育优良农艺性状的同时，常伴随着不利性状的转育，即出现遗传累赘现象的重要原因。
第3染色体着丝粒RM15139-RM16区域：在该区域检测到控制千粒重、每穗实粒数、每穗总粒数和结实率4个产量相关性状以及稻米品质包括碾磨品质、外观品质和粒型3个方面的QTL。千粒重与每穗实粒数和每穗总粒数的增效等位基因分别来自于IRBB和特青，即，该区域在增加千粒重同时降低每穗粒数，位于该区域控制糙米率、粒长、长宽比、垩白米率、垩白度和透明度QTL增效等位基因均来自于IRBB，而整精米率和粒宽QTL增效等位基因来自于特青。
第5染色体短臂RM437-RM18189区域：在该区域仅检测到2个产量相关性状千粒重和结实率QTL，增效等位基因分别来自于特青和IRBB，所检测到的控制稻米品质QTL涉及碾磨品质、外观品质和粒型3个方面。该区域在增加千粒重的同时，垩白度、糙米率、精米率、粒宽及垩白米率亦随之增加，而结实率、粒长、长宽比和透明度降低。
第6染色体Wx基因区域：在该区域除了检测到控制稻米蒸煮食味品质3个评价指标的QTL外，同时检测到控制粒宽、千粒重、糙米率、精米率和垩白的QTL，除控制胶稠度的QTL外，其它QTL增效等位基因均来自于特青。
第6染色体ALK基因区域：在该区域检测到控制碱消值、胶稠度、每穗实粒数和每穗总粒数的QTL，除碱消值外增效等位基因均来自于特青。
第7染色体长臂RM70-RM18区域：在该区域检测到控制每穗总粒数、糙米率、透明度和直链淀粉含量QTL，除每穗总粒数QTL来自于特青外，其它均来自于IRBB。
第8染色体短臂RM5647-RM22755区域：在本研究中，第8染染色体短臂为一多功能区域，同时控制每穗实粒数、每穗总粒数、结实率、粒宽、长宽比、糙米率和垩白米率7个性状，除控制长宽比和结实率的QTL来自于IRBB外，其它均来自于特青。
第10染色体长臂近末端RM6100-RM3123区域：该区域检测到控制千粒重、垩白米率、垩白度、糙米率、粒宽和透明度QTL，除透明度外，控制其它性状的QTL均来自于特青。
另外，在本研究中共计检测到29对互作QTL，除第1染色体外，其它11条染色体均参与其中。这些互作遗传效应较小，对性状表型变异贡献率多在1%以下，一个位点往往与多个位点同时产生互作效应，如第2染色体长臂末端控制千粒重主效应QTL qTGW2.2，与多个主效应QTL之间均存在互作。在所检测到29对互作QTL中，主效应QTL之间互作是其主要部分。
Currently，high yield and quality is one of important aims for rice hybrid breeding. A set ofrecombinant inbred line (RIL) population characterized by a relative narrow range of heading date anddeveloped from crosses between Teqing and IRBB lines was utilized to analyze the genetic relationshipbetween rice quality involving13indexes and6grain yield-related traits of indica rice with theassistance of molecular linkage map and phenotypic data. Theoretically, the effect of ecologicalenvironment on assay results due to various heading date among the lines was diminished, as a result,the genetic relationship between rice quality and grain yield was reflected objectively.
In this study, a total of7chromosomal regions related to both rice quality and grain yield weredetected. They were mainly distributed on chromosome3,5,6,7,8and10, on which QTLsconditioning rice quality and grain yield were clustered together. Surveying the additive directions ofthese QTLs, we found that alleles for increasing rice quality and grain yield were often from differentparents. That is, grain yield were improved due to a certain multi-function region followed bydeleterious effects on rice quality. This may be the main cause for cultivating some elite agronomictraits at the expense of deleterious agronomic traits, that is, genetic drag in rice hybrid breeding practice.
RM15139-RM16region on chromosome3: QTLs for4grain yield-related traits including1000-grain weight (TGW), number of grains per panicle (NGP), number of spikelets per panicle (NSP)and spikelet fertility (SF) were detected as well as QTLs for rice quality involving rice milled quality,appearance quality and grain shape in this region. Allele for increasing TGW in this region was fromIRBB while the enhancing alleles of NGP and NSP were from Teqing, which means that this regionincreased TGW with NGP and NSP decreased simultaneously. QTLs conditioning brown rice ratio(BRR), grain length (GL), ratio of grain length to grain width (LWR), percentage of chalky grain (PCG),degree of chalkiness (DC) and the endosperm transparency (ET) were also detected in this region andall QTLs had the enhancing alleles from IRBB, however, QTLs detected in this region for head riceratio (HRR) and grain width (GW) both had the enhancing alleles from Teqing.
RM15139-RM16region on chromosome5: QTLs for TGW and SF were the only two grainyield-related QTLs detected in this region and the enhancing alleles were from Teqing and IRBB,respectively. QTLs detected for rice quality involved3aspects including rice milled quality, appearancequality and grain shape in this region. This region increased TGW, DC, BRR, milled rice ratio (MRR),GW and PCG with SF, GL, LWR and ET decreased simultaneously.
Wx region on chromosome6: In additional to3QTLs conditioning3indexes for cooking andeating quality respectively were detected in this region, QTL for GW, TGW, BRR, MRR and chalkinesswere also detected in this region. All QTLs except QTL for gel consistency (GC) had enhancing allelesfrom Teqing.
Alk region on chromosome6: QTLs conditioning alkali-spread score (ALK), GC, NGP and NSPwere detected in this region and all QTLs except QTL for ALK had enhancing alleles from Teqing.
RM70-RM18region on chromosome7: QTLs conditioning NSP, BRR, ET and amylose content(AC) were detected in this region and all QTLs except QTL for NSP had enhancing alleles from IRBB.
RM5647-RM22755region on chromosome8: In our study, the short arm of chromosome8was amultifunction region and controlled7rice traits including NGP, NSP, SF, GW, LWR, BRR and PCG.All QTLs except QTLs for LWR and SF had enhancing alleles from Teqing.
RM6100-RM3123region on chromosome10: QTLs conditioning TGW, PCG, DC, BRR, GW andET were detected in this region and all QTLs except QTL for ET had enhancing alleles from Teqing.
Additionally, a total of29couple epistatic QTLs involving with all chromosomes exceptchromosome1were detected in this study. These epistatic QTLs had a relative small genetic effect andthe phenotypic variation explained by them was often under1%. One locus usually interacted withmany loci and generated genetic effects, for example, main-effect QTL qTGW2.2located on the distalof long arm of chromosome2interacted with many main-effect QTLs for TGW. Among the29coupleepistatic QTLs, interactions between main-effect QTLs accounted for a most part.
引文
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