Identification and validation of a yield-enhancing QTL cluster in rice (Oryza sativa L.)

Improvement of rice grain yield (YD) is an important goal in rice breeding. YD is determined by its related traits such as spikelet fertility (SF), 1,000-grain weight (TGW), and the number of spikelets per panicle (SPP). We previously mapped quantitative trait loci (QTLs) for SPP and TGW using the recombinant inbred lines (RILs) derived from the crosses between Minghui 63 and Teqing. In this study, four QTLs for SF and four QTLs for YD were detected in the RILs. Comparison of the locations of QTLs for these three yield-related traits identified one QTL cluster in the interval between RM3400 and RM3646 on chromosome 3. The QTL cluster contained three QTLs, SPP3a, SF3 and TGW3a, but no YD QTL was located there. To validate the QTL cluster, a BC₄F₂ population was obtained, in which SPP3a, SF3 and TGW3a were simultaneously mapped to the same region. SPP3a, SF3 and TGW3a explained 36.3, 29.5 and 59.0 % of phenotype variance with additive effect of 16.4 spikelets, 6 % SF and 1.8 g grain weight, respectively. In the BC₄F₂ population, though the region has opposite effects on TGW and SPP/SF, a YD QTL YD3 identified in this cluster region can increase 4.6 g grains per plant, which suggests this QTL cluster is a yield-enhancing QTL cluster and can be targeted to improve rice yield by marker aided selection.     

水稻种子衰老相关基因定位

利用ZS97×MH63组合衍生的160份重组自交系进行种子衰老遗传分析。每份自交系3批种子,即在武汉室温条件下存放3年的自然衰老种子,收获3个月后的种子,收获3个月后再经过高温、高湿加快衰老处理的种子,供25℃条件下考查发芽率和发芽势来衡量种子衰老状况。复合区间作图方法进行了2批衰老种子的发芽势与发芽率QTL定位。自然     

利用Ⅱ-32B/A7444组合CSSL群体定位水稻7个穗部性状QTL

为发掘水稻穗部性状有利等位变异,构建了以籼稻保持系II-32B为遗传背景的A7444染色体片段置换系群体;利用QTL Ici Mapping 4.1软件对该群体7个穗部性状进行了QTL定位。结果 2年共检测到26个QTL。2年均检测到的13个QTL中,控制一次枝梗数的4个QTL位于第1、第6、第8和第9染色体,平均贡献率分别为15.16%、13.10%、29.74%和11.21%,平均加性效应分别为-1.40、1.01、1.11和0.77。控制二次枝梗数的2个QTL位于第6和第8染色体,平均贡献率分别为10.97%和21.39%,平均加性效应分别为5.45和6.36。控制每穗总粒数的3个QTL位于第2、第6和第8染色体,平均贡献率分别为8.65%、12.52%和31.22%,平均加性效应分别为-18.61、22.23和31.87。控制每穗实粒数的1个QTL位于第8染色体,平均贡献率为28.06%,平均加性效应30.85。控制千粒重的2个QTL位于第2染色体,平均贡献率分别为44.65%和17.51%,平均加性效应分别为2.88和-2.51。控制粒宽的1个QTL位于第10染色体,平均贡献率为21.96%,平均加性效应为0.11。第2、第6和第8染色体分别存在同时控制二次枝梗数、每穗总粒数和每穗实粒数QTL的区段。qSBN6和qSBN8所在区间与Hd1和DTH8的相同,但分别存在16处和1处碱基差异,推测为Hd1和DTH8的不同等位基因。qSBN2为新检测到的控制二次枝梗数位点。研究结果为实施分子标记聚合育种提供了有用信息。     

QTL analysis of cooked rice grain elongation, volume expansion, and water absorption using a recombinant inbred population

The traits of elongation, volume expansion, and water absorption are very important in determining the quality of cooked rice grains. In this study, quantitative trait loci (QTL) analysis of these traits was performed using a recombinant inbred population derived from a cross between two indica cultivars, ‘Zhenshan 97’ and ‘Minghui 63 ,’ which are the parents of the most widely grown hybrid rice in China. Using a linkage map based on 221 molecular marker loci covering a total of 1796 cM, a total of 33 QTLs were identified for the nine traits tested. QTLs were detected on chromosomes 1– 3 , 5– 9 , and 11 , respectively. The QTLs identified included three for cooked rice grain length elongation (chromosomes 2 , 6 , and 11), six for width expansion (chromosomes 1‐ 3 , 6 , 9 , and 11) and two for water absorption (chromosomes 2 and 6). Interestingly, a single QTL located near the wx gene on chromosome 6 seemed to influence all the traits tested for the cooked rice quality.     

水稻千粒重和垩白粒率的QTL及其互作分析

产量因子千粒重和稻米品质指标垩白粒率密切相关。本研究以越光/Kasalath//越光BIL群体为材料,分析千粒重和垩白粒率的相关性、QTL、上位性互作及其环境的互作效应。相关分析表明,群体千粒重和垩白粒率在2005年和2006年均呈极显著正相关,相关系数分别为0.42和0.35 (P<0.001)。2年共检测到千粒重QTL 11个,其中5个在2年重复检测到,5个具有环境互作效应;千粒重上位性互作8对,7对与环境存在互作。垩白粒率QTL 6个,3个具有环境互作效应;上位性互作9对,其中4对具有上位性环境互作效应。比较分析发现3个主效QTL同时控制千粒重和垩白粒率的表现,千粒重和垩白粒率的增效等位基因来自同一亲本;1对上位性互作同时对千粒重和垩白粒率有相同的影响。一些与垩白粒率不相关的千粒重主效QTL,如qTGW-3c、qTGW-4a和qTGW-6b,可为育种所利用。对利用QTL定位结果进行千粒重和垩白粒率分子辅助选择育种进行了探讨。     

发掘人工合成小麦中千粒重QTL的有利等位基因

以人工合成小麦Am3为供体亲本,普通小麦莱州953为轮回亲本,经5次回交然后自交,培育出含85个株系的F_2:3群体。以该群体为材料,用348对多态性SSR标记,进行全基因组扫描,发掘人工合成小麦中千粒重QTL的有利等位基因。利用复合区间作图法检测到3个千粒重QTL,其对表型变异的贡献率为10.9%~33.79%。其中,Am3的等位基因能够增加千粒重2.3~4.8 g。相关分析表明,该导入系群体的千粒重与穗粒数、穗数和株高无显著相关性。千粒重QTL与穗粒数、穗数性状的QTL不在同一位置,这有利于高千粒重基因与其他产量性状基因的聚合。采用混合线性模型作图法检测到1个千粒重QTL(QGw.caas-3D),该QTL与环境互作效应小,而且与复合区间作图法在3个环境中都检测到的QTL相同,表明QGw.caas-3D是一个稳定的主效QTL。     

利用品质性状的回交选择导入系挖掘水稻抗纹枯病QTL

将优质、抗纹枯病的高秆供体Tarom Molaii和Binam导入半矮秆IR64和特青背景,培育品质性状回交选择构建的4个导入系群体IR64/Tarom Molaii、特青/Tarom Molaii、IR64/Binam和特青/Binam,定位了影响水稻抗纹枯病病级(disease scale, DS)、相对病斑高度(relative lesion height, RH)和株高(plant height, PH)的QTL。结果表明,4个导入系群体的DS与RH高度相关,两者与PH呈显著负相关。导入系后代各性状均呈现超亲分离,出现抗性明显优于双亲的抗病个体,其中40%左右属半矮秆抗病类型。采用单向方差分析,在这4个群体中分别定位到10、8、8和6个影响3个性状的QTL,多数基因座上降低DS和RH即增强抗病性同时增加株高的等位基因均来自两个供体。未在同一供体两个不同背景下检测到影响3个性状的相同QTL,表明抗纹枯病QTL表达有明显的遗传背景效应。PH与DS及PH与RH被定位在同一个显著标记位点的QTL数分别占两个性状QTL总数的38%和52%,表明水稻纹枯病抗性与株高关系密切,两者存在许多连锁位点。与以往相同群体品质性状QTL的定位结果相比,发现品质性状QTL与抗纹枯病QTL大多分布在染色体的不同区域,彼此独立遗传。对利用目标性状选择导入系定位非目标性状QTL的效果、影响因素及育种应用进行了探讨,强调了目标性状选择导入系对非目标性状QTL发掘及育种应用的重要性。     

Dissection of component QTL expression in yield formation in rice

Yield and its components were investigated by using a population of 241 recombinant inbred lines (F₉ RILs) derived from an elite hybrid rice cross of ‘Zhenshan 97’בMinghui 63′. Quantitative trait loci (QTLs) for causal analysing of yield traits were detected at different yield component (YC) influences by conditional and unconditional QTL mapping methods. The number of QTLs significantly affecting yield was different at component‐special influence. Some QTLs controlling yield identified in one component influence were undetectable at the others. More QTLs for yield could be detected at different YC influences. It is possible to reveal that causal gene expression for yield could be different at different YC influences. Mapping QTLs for component effects of yield could help us in understanding the nature of cause‐effect traits for the formation of grain yield.     

水稻黑条矮缩病抗性QTL分析

利用珍汕97B/明恢63的重组自交系群体,采用自然发病鉴定的方法,以穴发病率为表型值,对各株系进行黑条矮缩病抗性鉴定。群体的穴发病率偏向于抗病亲本,且呈连续性分布,表明水稻黑条矮缩病抗性受数量性状基因控制。利用WinQTLcart 2.5软件对黑条矮缩病抗性QTL进行分析,共检测到6个QTL,其中第6、11染色体上各有2个,第7、9染色体各有1个。第6、7、9染色体上的4个QTL在两个地点都能检测到,是稳定表达的QTL,尤其是第6染色体上的2个QTL,LOD值分别为12.09和9.77,贡献率分别为20.20%和18.68%,是主效QTL,能在分子标记辅助选择育种中加以利用。     

影响水稻穗部性状及籽粒碾磨品质的QTL及其环境互作分析

利用优质恢复系测258为轮回亲本与粳型糯稻新品系IR75862杂交创制的BC₁F₇回交导入系群体,在广西南宁和海南三亚定位了产量相关性状(二次枝梗数、穗总粒数、穗实粒数、粒重和穗重)、粒型(粒长、宽、厚)和碾磨品质(糙米率、精米率和整精米率)的主效QTL并剖析其环境互作效应。双亲在穗实粒数、千粒重、粒长和粒宽及整精米率等性状上存在显著差异。各产量相关性状间呈极显著正相关,而与千粒重和粒长呈极显著负相关。多数产量及粒型相关性状与3种碾磨品质相关不显著。在南宁和三亚环境下检测到影响产量相关性状、粒型及碾磨品质的主效QTL共计57个,包括二次枝梗数的6个,穗实粒数4个,穗总粒数、粒重和穗重各5个,粒长9个,粒宽7个,粒厚1个,糙米率4个,精米率5个和整精米率6个,分布在除第11染色体外的所有染色体上。多数影响枝梗数、穗粒数和粒重的QTL成簇分布,而且与影响BR、MR和HR的QTL分布在不同染色体区域。在第2、第3、第4、第5和第6染色体上鉴定出影响穗粒数、粒重、粒型及碾磨品质的重要QTL,这些QTL在以往不同遗传背景和环境下被多次检测到。在第8染色体RM152~RM310区间鉴定到1个影响粒长和粒宽的新的QTL,能同步增加粒宽和粒长。鉴定出的这些稳定表达的QTL具有标记辅助选择育种的应用价值。整精米率是受环境影响最大的性状,其QTL的环境互作效应明显。对QTL的环境互作效应特点及其在品种标记辅助改良中的作用进行了深入探讨。     

利用双向导入群体检测遗传背景对耐盐QTL定位的影响

以优质粳稻品种Lemont与高产籼稻品种特青为亲本培育的高代双向回交导入系为材料,在温室140 mmol L^-1 NaCl胁迫条件下定位影响苗期叶片盐害级别(SST)、幼苗存活天数(SDS)、地上部K⁺浓度(SKC)和地上部Na⁺浓度(SNC)及人工气候室条件下影响地上部K⁺、Na⁺浓度的QTL。双向导入系的大部分遗传背景与各自的受体亲本相同,其中Lemont背景导入系中轮回亲本Lemont的基因组平均占83.8%,特青背景导入系中轮回亲本特青基因组平均占88.9%。各耐盐相关性状在两个背景群体中均出现超亲分离,多数性状的频率分布呈相互重叠状态,表明双亲作为供体相互导入各耐盐性状基因的效应大致相当。两个背景导入系群体中分别检测到影响上述耐盐相关性状的QTL各18个,同一性状在两个背景导入系中未能检测到任何相同表达的QTL,表明耐盐QTL表达具有很强的遗传背景效应,同时也说明这些耐盐QTL的效应可能较小。温室和人工气候室两种环境下仅在特青背景导入系中检测到1个影响SKC的相同QTL,表明耐盐QTL与环境的互作非常明显。虽然双亲均表现中等感盐,但QTL定位结果表明双亲中都存在一些提高耐盐相关性状的有利等位基因。研究认为,利用分子标记技术挖掘“隐蔽”于育成品种中的耐盐基因,进一步利用分子标记辅助选择技术对这些非等位耐盐基因进行聚合,完全有可能提高育成品种的耐盐水平。     

水稻抽穗期数量性状基因的定位及遗传效应分析

本研究利用特早抽穗粳稻品种石狩白毛和籼稻品种明恢63杂交的F2分离群体共116株,构建了含88个共显性分子标记的连锁图谱,对水稻(Oryza sativA L．)抽穗期进行基因定位。利用2种分析软件MAPMAKER／QTL和QTLMapper进行分析,共检测到3个抽穗期的数量性状基因座(QTLs)。2个软件共同发现第7染色体上RM214与A5106标记区间内存在1个主效QTL Hd7a（Hd7c),来自明恢63的这个位点的等位基因可使抽穗期延迟。其余2个QTLs分别位于第7、9染色体上。同时检测到有5对位点间存在上位性作用,但相对贡献率较小,表明上位性效应也是影响抽穗期的遗传基础。     

用(培矮64s/Nipponbare)F2群体对水稻产量构成性状的QTL定位分析

用水稻测序品种培矮64s和Nipponbare为亲本构建的含137个SSRs标记的连锁遗传图谱和(培矮64s/Nipponbare)F2群体的180个单株,对水稻的单株有效穗数、穗粒数、穗实粒数、结实率、穗着粒密度、千粒重等6个产量构成性状进行了QTL定位分析.共检测到6个性状的22个QTLs,分布在第1、2、4、5、6、9、10、11、12等9条染色体的14个区域,表型贡献率5.0%～19.3%;相关性较强的性状之间具有较多共同或紧密连锁的QTLs;集中分布的QTLs之间既有同向连锁,也有反向连锁.对不同水稻群体定位的同源QTL进行了比较,对QTL在染色体上的集中分布,以及用QTL定位结果和生物信息学方法相结合预测基因的功能等进行了探讨.     

QTLs identification of crude fat content in brown rice and its genetic basis analysis using DH and two backcross populations

Fat content is a concern for the enhancement of rice for eating, cooking, and storage qualities. To clarify its genetic mechanism, a double haploid (DH) population derived from anther hybrid F₁ of Zhenshan 97B (indica) and Wuyujing 2 (japonica) and two backcross F₁ (BCF₁) populations, which came from the DH lines backcrossing to two parents, were used to scan quantitative trait loci (QTLs) and dissect gene effects for the crude fat content (CFC) in brown rice. Fourteen QTLs were resolved, distributing on chromosomes 1, 3, and 5–9. Three loci were detected repeatedly in two populations, DH or BCF₁. Among these loci, a major QTL, qCFC5, flanking markers RM87 and RM334, was located on chromosome 5, which was detected simultaneously among three populations. The main QTLs had a major role in controlling CFC in brown rice and were modified by several mini-effect QTLs and epistatic affection. Wenjun Liu and Jing Zeng are contributed equally to this paper.     

不同环境条件下稻米组氨酸和精氨酸的胚乳和母体植株QTL分析

利用汕优63重组自交系与双亲回交产生的BC1F1和BC2F1群体,采用新发展的包括环境互作效应在内的多遗传体系QTL作图方法和基因定位软件,对稻米两种半必需氨基酸（组氨酸和精氨酸）进行三倍体胚乳和二倍体母体植株等不同遗传体系的QTL定位分析。共检测到10个控制组氨酸含量的QTL以及8个控制精氨酸含量的QTL。全部QTL均具有极显著的三倍体胚乳和二倍体母体植株基因的加性主效应,其中4个QTL具有显著或极显著的三倍体胚乳显性主效应,7个QTL还具有明显的环境互作效应。     

Validating a segment on the short arm of chromosome 6 responsible for genetic variation in the hull silicon content and yield traits of rice

Rice is a typical silicon-accumulating plant and the beneficial effect of silicon on rice has long been recognized. In a previous study using 244 recombinant inbred lines (RILs) of an indica rice cross, Zhenshan 97B/Milyang 46 grown in 2003, four QTLs were detected for hull silicon content. QTL qHUS-6 had the largest effect among these, and the same interval also had significant effects on yield traits in the same population. The primary objective of this study was to validate the QTL effect in this region on HUS and yield traits. The same RIL population and another RIL population of lower heterogeneity were grown in 2004. QTL qHUS-6 was found to have significant additive effects on hull silicon content with a consistent direction in the two populations. From a residual heterozygous line selected from RILs of the same cross, 15 F_2:3 lines that differed only in a 2.15-Mb segment extending from RM587 to RM6119 on the short arm of chromosome 6 were derived. In these lines, qHUS-6 displayed a major effect, so did QTLs for yield traits previously detected in the same region. Two more QTLs for HUS detected in 2003, qHUS-1-1 and qHUS-1-2, also had consistent effects in the Zhenshan 97B/Milyang 46 RIL population in 2004. Thus this study verified three candidate regions for fine mapping HUS QTLs and determining the genetic relationship between silicon content and yield traits in rice.     

水稻抗纹枯病QTL表达的遗传背景及环境效应

利用水稻纹枯病菌强致病菌系RH-9人工接种Lemont导入到特青背景的213个近等基因导入系(TQ-ILs)群体和特青导入到Lemont背景的195个近等基因导入系(LT-ILs)群体,定位和分析了水稻抗纹枯病数量性状座位(quantitative trait loci, QTL)及其表达的环境与遗传背景效应。亲本Lemont对RH-9表现为高度感病,特青表现为中等抗病。人工接种后TQ-ILs群体的相对病斑高度(病斑高度与株高比)呈连续正态分布,LT-IL群体则明显偏向感病亲本Lemont。在不同年份和遗传背景下检测到影响纹枯病相对病斑高度的主效QTL 10个和互作QTL 13个,其中2006年在TQ-IL群体定位到的6个主效QTL在2007年均得到验证,表明这些QTL具有较好年度间的重复性。QSh4是唯一在双向导入系背景下表达的QTL,该位点特青等位基因降低相对病斑高度,提高抗性水平。在TQ-ILs群体中定位到位于第10染色体RM216~RM311区间的QSb10a与在LT-IL群体中定位到的位于相邻区间RM222~RM216的QSb10b的基因作用方向不同,推断这两个QTL存在紧密连锁关系。绝大多数在TQ-IL群体中表达的主效及互作QTL在LT-ILs群体中不表达,表明水稻抗纹枯病QTL具有明显的遗传背景效应。通过比较作图,本研究定位到的其中8个QTL在以往不同群体中同样被检测到,这些主效QTL对通过分子标记辅助选择(marker-assisted selection, MAS)培育水稻抗纹枯病育种可能具有应用价值。研究指出,标记辅助选择在不同遗传背景中能稳定表达的QTL或通过聚合不同抗病QTL是进一步提高水稻纹枯病抗性水平的一个有效途径。     

利用BC_2F_3产量选择导入系定位水稻耐盐QTL

本研究以生产上大而秋推广的籼稻恢复系蜀恢527和明恢86为轮同亲本,以ZDZ057和特青为供体亲本,培育选择了蜀恢527/ZDZ057、明恢86/ZDZ057、蜀恢527/特青和明恢86/特青4个BC2F3产量选择导入系群体,分别选取65个、75个、73个和60个均匀分布在12条染色体上的SSR标记用于以上4个群体的基因型分析.对4个BC2F3群体的幼苗耐盐等级和幼苗存活天数两个性状进行了表型鉴定.采用性状-标记间的单向方差分析对上述性状进行了QTL定位.在群体蜀恢527/ZDZ057、明恢86/ZDZ057、蜀恢527/特青和明恢86/特青中分别检测到了11、15、11和6个控制幼苗耐盐等级和幼苗存活天数的QTL,其中群体明恢86/ZDZ057检测到的15个QTL中有3个QTL具有多效性.其中有2个QTL能存蜀恢527/ZDZ057和明恢86/ZDZ057群体分别检测到,1个QTL可以在蜀恢527/ZDZ057和蜀恢527/特青群体分别检测剑,1个QTL在明恢86/ZDZ057和明恢86/特青群体中分别检测到.QSst2c、QSst10b、QSst12、QSds1b、QSst4b、QSst2d、QSst2e、QSds10、QSst1b、QSst10c、QSst10d和QSds4b能解释的表型变异分别为20.36%、24.06%、22.50%、28.45%、21.39%、23.31%、34.92%、42.18%、20.29%、20.50%、21.3l%和26.27%,位点QSst2c、QSds1b、QSst9a和QSst2e与前人定位在同一区域.QSst12表现为负向超显性,QSst4b和QSds10表现为正向超显性,QSst10b和QSst2d分别表现为负向和正向部分显性.本研究结果将为水稻高产耐盐育种提供有益参考.     

灌溉与自然降雨条件下水稻高代回交导入系产量QTL的定位

利用254个Lemont导入到特青背景的高代回交导入系定位了灌溉（对照）与自然降雨（干旱胁迫）环境下影响单株籽粒产量及其穗部相关性状的QTL。在两种环境下共检测到32个影响单株粒籽产量、千粒重和每穗实粒数的主效QTL,根据不同环境下表达的情况将其分成3类,第1类10个QTL,在两种环境下均被检测到;第2类14个QTL,只在对照条件下检测到;第3类8个QTL,受干旱胁迫诱导,只在胁迫条件下被检测到。此外还检测到9个影响胁迫与对照条件下性状差值的QTL。认为在两种条件下均检测到的相对稳定的3个QTL（QGn11b、QGn12和QGn11b）及影响两种条件下性状差值（即性状稳定性）的9个QTL可能对耐旱性有直接贡献。在所有12个耐旱QTL中,除在QGn5和QGy1的Lemont等位基因减小性状差值（即增强耐旱性）外,其余位点上增强耐旱性的等位基因均来自特青。另外通过与源自相同亲本的不同定位群体在不同环境下定位结果的比较,鉴别出一些受遗传背景和环境影响较小的QTL如QGn3b、QGw1、QGw5、QGy1、QGy5、QGy8和QGy10。对应用QTL定位结果进行标记辅助选择培育耐旱品种进行了探讨。     

QTL analysis on rice grain appearance quality,as exemplifying the typical events of transgenic or backcrossing breeding

Rice grain shape and yield are usually controlled by multiple quantitative trait loci (QTL). This study used a set of F_9–10 recombinant inbred lines (RILs) derived from a cross of Huahui 3 (Bt/Xa21) and Zhongguoxiangdao, and detected 27 QTLs on ten rice chromosomes. Among them, twelve QTLs responsive for grain shape/ or yield were mostly reproducibly detected and had not yet been reported before. Interestingly, the two known genes involved in the materials, with one insect-resistant Bt gene, and the other disease-resistant Xa21 gene, were found to closely link the QTLs responsive for grain shape and weight. The Bt fragment insertion was firstly mapped on the chromosome 10 in Huahui 3 and may disrupt grain-related QTLs resulting in weaker yield performance in transgenic plants. The introgression of Xa21 gene by backcrossing from donor material into receptor Minghui 63 may also contain a donor linkage drag which included minor-effect QTL alleles positively affecting grain shape and yield. The QTL analysis on rice grain appearance quality exemplified the typical events of transgenic or backcrossing breeding. The QTL findings in this study will in the future facilitate the gene isolation and breeding application for improvement of rice grain shape and yield.