利用染色体片段置换系剖析水稻籼粳亚种间产量相关农艺性状杂种优势的遗传基础
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摘要
水稻(Oryza sativa)是世界主要粮食作物之一,在世界上一百二十个国家和地区广泛栽培种植,全球一半以上的人口以稻米为主食,而在中国有60%以上的人口以稻米为日常生活的主食。随着人口的不断增长以及可利用耕地面积的逐年减少,粮食安全问题日益严峻,因此提高水稻产量显得尤为重要。其中,水稻杂种优势利用是增加稻谷产量的重要途径之一。
近百年来,杂种优势遗传机理的研究水平一直落后于对其在生产上的利用程度,这种杂种优势理论研究滞后于生产实践的局面势必影响杂种优势在现实中的进一步大规模利用。研究水稻的杂种优势形成机理,不仅对水稻杂交种的选育具有重要的指导意义,同时,水稻作为单子叶植物基因组研究的模式植物,将为研究禾谷类作物杂种优势理论提供模式借鉴。因此,广泛开展水稻杂种优势遗传基础的研究和探讨具有十分重要的理论价值和现实意义。
水稻籼／粳亚种间的杂种优势强于籼/籼或粳/粳亚种内的杂种优势,利用籼粳亚种间杂种优势是进一步提高水稻单产的重要技术策略。本研究利用来自日本的典型粳稻品种Asominori和国际水稻所(IRRI)育成的典型籼稻品种IR24为双亲构建的互为受体(遗传背景)和供体(置换片段)的两套染色体片段置换系群体及其与各自对应背景亲本杂交构建的两套杂种F1群体共四套材料,在两年两点四个环境(2007、2008年南京和南昌)对水稻籼粳亚种间组合产量构成性状和穗形相关性状等与产量密切相关的农艺性状进行了杂种优势的研究,利用SSR标记重新构建的染色体片段置换系基因型图谱检测产量相关农艺性状增(减)效杂种优势位点(Heterotic loci, HL),并通过利用染色体片段置换系群体检测到的具有加性效应的QTL和对应杂种F1群体检测到的具有显性效应的QTL之间的加显性效应剖析了产量相关农艺性状杂种优势形成的遗传基础。主要研究结果如下：
1、利用SSR标记重新鉴定了以Asominori为受体亲本,IR24为供体染色体片段置换系(简称AIS)和以IR24为受体亲本,Asominori为供体的染色体片段置换系(简称IAS)的基因型。利用全基因组筛选到的137个SSR多态性标记构建了AIS置换系基因型图谱和132个SSR多态性标记构建了IAS置换系基因型图谱,获得了QTL分析所需要的室内分子标记数据。
2、两套染色体片段置换系大部分农艺性状表现出明显的超亲分离现象；各农艺性状杂种优势均表现出变幅极大的分离,但不同农艺性状杂种优势的表现程度不一；不同农艺性状年度间均表现出显著差异,部分性状基因型与环境显著互作。
3、无论是高值亲本,还是低值亲本,都包含有对性状表现型起增效或者起减效作用的等位基因；这些数量性状位点分散在双亲的基因组中,个别区段有连锁。
4、采用基于逐步回归和极大似然比估计相结合的完备区间作图法,利用QTL IciMapping软件为检测平台,以经验阈值LOD≥3.0作为判断产量相关农艺性状杂种优势QTL存在与否的标准,对8个产量相关性状杂种优势效应进行了QTL定位。在四个环境中利用AIS和IAS对应杂种F1群体,分别定位了83个和53个与产量构成性状相关的增(减)效杂种优势标记位点。其中AIS对应杂种F1群体中发现13个标记位点能在多个环境重复检测到,有些位点能同时影响多个性状的表现,位于第5和7染色体上的RM267和RM82标记位点杂合状态下减少杂种F1代单株产量、每穗实粒数和结实率RM82标记位点还同时增加杂种F1代单株库容和株高。来自第2染色体上的RM5390标记位点杂合状态下能同时增加单株产量、每穗实粒数和结实率的表现。来自第4和6染色体上的RM252和RM217标记位点杂合状态下降低杂种F1的株高。IAS对应杂种F1群体中只发现1个标记位点能在多个环境重复检测到,来自第1染色体上的RM488标记位点杂合状态下增加杂种F1的株高。从HL效应的方向来看,AIS对应F1群体中检测到39个增效位点,占总位点数的46.99%；IAS对应F1群体中检测到22个增效位点,占总位点数的41.51%；两套F1群体还分别检测到44个和31个减效位点,这些减效位点主要集中在单株产量、每穗实粒数和结实率等与籼粳不育基因密切相关的性状,通过广亲和材料的应用或者染色体片段置换技术可以消除减效位点的负向效应。
5、利用AIS和IAS对应杂种Fl群体四个环境中的田间表型数据结合QTL分析软件,分别定位了40个和22个与穗形性状相关的增(减)效杂种优势标记位点。其中AIS对应杂种F1群体中发现2个增(减)效杂种优势标记位点能在多个环境重复检测到,来自第8染色体上的RM284标记位点能增加杂种F1的穗长,来自第8染色体上的标记位点RM331能增加杂种F1的一次枝梗数；IAS对应杂种F1群体中未发现能在多个环境重复检测到的增(减)效杂种优势标记位点。两套群体在第8染色体上检测到相同的杂种优势标记位点RM284,杂合状态下都表现增加杂种F1的穗长性状。从HL效应的方向来看,AIS对应F1群体中有27个增效位点,占总位点数的67.50%；IAS对应F1群体中有12个增效位点,占总位点数的54.55%；两套F1群体还分别检测到13个和10个减效位点。育种过程中可根据不同育种目标灵活利用穗形相关性状增(减)杂种优势位点来改良穗形性状。
6、利用AIS及其对应F1群体在四个环境分别检测到99个和79个与产量性状相关的加性与显性效应的QTL,其中具有加性效应的QTL分布在全部12条染色体上具有显性效应的QTL分布在除了Chr.9和Chr.11外的10条染色体上。而利用IAS及其对应F1群体在四个环境分别检测到116个和125个与产量性状相关的加性与显性效应的QTL,其中加性与显性效应的QTL都分布在全部12条染色体上。通过产量构成性状QTL的显性效应值D和加性效应值A的比值(平均显性度,|D/A|)分析,发现粳稻背景下共有52个QTL表现超显性效应,约占全部QTL的34.44%；61个QTL表现部分显性效应,约占全部QTL的40.40%；22个QTL表现显性效应,约占全部QTL的14.57%；16个QTL表现加性效应,约占全部QTL的10.60%；籼稻背景下,共有61个QTL表现超显性效应,约占全部QTL的33.70%；69个QTL表现部分显性效应,约占全部QTL的38.12%；32个QTL表现完全显性效应,约占全部QTL的17.68%；19个QTL表现加性效应,约占全部QTL的10.50%。因此,可以看出,QTL的超显性效应和显性效应(主要是部分显性)可能是水稻产量构成性状杂种优势形成的重要原因。
7、利用AIS及其对应F1群体在不同环境中分别检测到52个和55个与穗形性状相关的加性与显性效应的QTL,其中具有加性效应的QTL分布在除了第9和11染色体外的10条染色体上,具有显性效应的QTL分布在除了第5,9和11染色体外的9条染色体上。而利用IAS及其对应F1群体在不同环境中分别检测到75个和62个与穗形性状相关的加性与显性效应的QTL,都分布在于12条染色体上。通过穗形相关性状QTL平均显性度(|D/A|)的比较分析,发现粳稻背景下共有18个QTL表现超显性效应,约占全部QTL的21.95%;45个QTL表现部分显性效应,约占全部QTL的54.88%；11个QTL表现显性效应,约占全部QTL的13.41%；8个QTL表现加性效应,约占全部QTL的9.76%；籼稻背景下,共有26个QTL表现超显性效应,约占全部QTL的24.76%；47个QTL表现部分显性效应,约占全部QTL的44.76%；18个QTL表现完全显性效应,约占全部QTL的17.14%；17个QTL表现加性效应,约占全部QTL的16.19%。因此,可以看出,QTL的显性效应(主要是部分显性)可能是水稻穗形相关性状杂种优势形成的重要原因。
8、从产量相关农艺性状杂种优势位点以及加性和显性效应的QTL在染色体上的分布来看,发现在第1,2,3,4,6,7,8,9和12染色体上存在QTL的“簇状”分布现象,并对QTL簇与产量相关农艺性状的遗传关系进行了分析,发现各相关农艺性状普遍存在QTL的多效性,因此认为QTL的多效性可能是产量相关农艺性状以及杂种优势的遗传基础之一
With the development of social economy, greater demand was put on world food production due to the decreasing arable lands and increasing global population. The yield increase of rice, a main staple for a large fragment of the world population (amounting to half global population and 60%of population in our country), would contribute significantly to cope with the increasing global food crisis. Heterosis application was the principal way to improve rice grain yield.
However, since the advancement of research on the genetic basis of heterosis was lagged behind the utilization in the field production in last century, which limited a larger scale of heterosis utilization in crop production. The research of heterosis in rice is not only meaningful to guide the rice hybrid breeding, but also provide a theoretical model for the research of heterosis in the other cereal crops because rice is a model plant in monocotyledonous genomic research. Therefore, it is very important to conduct the research of heterosis and dissect its genetic basis for the theoretical value and realistic meaning in hybrid breeding of rice.
The extent of heterosis expressed in inter-subspecific crosses follows the general trend: indica/japonica> indicalindica> japonica/japonica. The extensive exploitation of inter-subspecific heterosis derive from indica/japonica is a very important strategy to enhance the yield level in rice production. The inter-subspecific cross between a japonica cv. Asominori and an indica cv. IR24 demonstrated huge heterosis in F1 hybrid. In our study, we conducted our experiments across four environments (2007,2008 in Nanjing; 2007, 2008 in Nanchang), mainly focus on the dissection and discussion of the genetic basis of heterosis about yield-related agronomic traits using two sets of chromosome segment substitution lines (CSSLs) derived from Asominori (a japonica cultivar) and IR24 (an indica cultivar) as reciprocal donor parent and recipient parent (background parent) and F1 derivatives from these two set of CSSLs population. QTL were detected by statistical software QTL IciMapping using two new different SSR map. The main conclusions are as follows:
1. Genotypes of these two sets of 66 CSS lines (one is indica c.v. IR24 chromosome segment substitution lines with Japonica c.v. Asominori genetic background; the other is japonica c.v. Asominori chromosome segment substitution lines with indica c.v. IR24 genetic background) were rescreened by 137 and 132 good polymorphism SSR markers, replacing 82 original RFLP markers. The purpose of replacing with SSR markers was to simplify the procedure of experiments by using RFLP markers and to enhance the efficiency of QTL detection using new maps with higher density of SSR markers.
2. Most of the agronomic traits showed transgressive segregation phenomenon in two CSSLs population; Heterosis of each agronomic trait exhibited a huge range of segregation, however different traits showed different level of heterosis; The studied traits also showed significant difference between years and genotypes, the interaction between genotypes and environments also exhibited a significance level.
3. Not only the high value parent but also the low value parent has detected many alleles at QTL with positive or negative effects in the whole genome, some chromosome segments were tightly linked.
4. We have detected 83 and 53 heterotic loci (HL) affecting yield component traits in Fi derivatives of AIS and IAS population, respectively. Of all the HL, only 13 HL could be detected repeatedly in two environments at least which reflected heterosis of yield component traits were very sensitive to environmental influence. Chances were the effect of pleiotropism, chromosome segments RM267 and RM82 on Chr.5 and Chr.7 could negatively affect grain weight plant-1 (GWP), grains per panicle (GPP) and seed-setting rate (SSR), simultaneously. RM82 could enhance the performance of sink capacity plant-1(SCP) and plant height (PH) as well. Chromosome segment RM5390 as an enhancing heterotic locus on Chr.2 could positively affect GWP, GPP and SSR traits simultaneously. RM252 and RM217 on Chr.4 and Chr.6 could decrease PH of F1 hybrids. Considering the effect of heterotic loci,39 with positive heterosis (amounting to 46.99%) and 44 (53.01%) with negative heterosis in AIS F1 derivative; 22 HL (41.51%) with positive and 31 (58.49%) with negative effect of heterosis in IAS F1 derivative. Possibly, we could create high-yield cultivar through pyramiding HL with positive effect and deleting HL negative effect by molecular tools such as SSR polymorphism markers.
5. We have detected 40 and 22 HL affecting panicle morphological traits in F1 derivatives of AIS and IAS population, respectively. Of all the HL were detected,27 with positive heterosis (67.50%) and 13 (32.50%) with negative heterosis in AIS F] derivative; 12 HL (54.55%) with positive and 10 (45.45%) with negative effect of heterosis in IAS F1 derivatives. Only 2 HL in AIS F1 derivative could be detected repeatedly in two more environments which reflected heterosis of panicle morphological traits were very sensitive to environment, too. Chromosome segments RM284 and RM331 on Chr.8 could increase the performance of panicle length (PL) and primary branches number (PBN), respectively. None HL could be detected in IAS F1 derivative. But we found that RM284 on Chr.8 could be detected in two sets of F1 derivatives, simultaneously.
6. Through mapping additive QTL and dominance QTL affecting yield component traits, we could dissect the genetic basis of heterosis. Eighty-nine additive QTL and 79 dominance QTL affecting eight yield component traits (grains weight per plant, GWP; sink capacity per plant, SCP; spikelets per panicle, SPP; grains per panicle, GPP; seed-setting rate, SSR; thousand-grain weight, TGW; panicles per plant, PPP; plant height, PH) were detected across four environments in AIS and F1 derivative population; 117 additive QTL and 125 dominance QTL located on all 12 chromosomes affecting eight yield-related traits were detected across the four same environments in IAS and F1 derivative population. Through comparing dominance effects and additive effects of those 151 QTL totally affecting eight yield-related traits in AIS and its corresponding F1 population, we found 52 QTL with over-dominance effect (amounting to 34.44%),61 QTL with partial dominance effect (amounting to 40.40%),22 QTL with complete dominance (amounting to 14.57%), 16 QTL with additive effect (amounting to 10.60%); we also detected 181 QTL totally affecting five panicle traits in IAS and its corresponding F1 population, of all the QTL with additive and dominance effect,61 QTL were identified with over-dominance effect (amounting to 33.70%),69 QTL with partial dominance effect (amounting to 38.12%),32 QTL with complete dominance effect (amounting to 17.68%),19 QTL with additive effect (amounting to 10.50%). Results indicated that both dominance (mainly were partial dominance) and over-dominance effects of QTL may be as the major reason in underlying the genetic basis of heterosis of yield component traits in rice hybrids.
7. We detected 52 QTL with additive effects and 55 QTL with dominance effects affecting five panicle morphological structural traits (panicle length, PL; density of spikelets per panicle, DSP; primary branches number, PBN; secondary branches number, SBN; density of secondary branches number, DSBN) across different environments in AIS and F1 derivative population. We found 72 additive QTL and 62 dominance QTL affecting five panicle morphological structural traits were detected across three same environments in IAS and F1 population. Comparing dominance effect and additive effect of those 82 QTL affecting five panicle morphological structural traits in AIS and its corresponding F1 population, we found 18 QTL with over-dominance effect (amounting to 21.95%),45 QTL with partial dominance effect (amounting to 54.88%),11 QTL with complete dominance (amounting to 13.41%),8 QTL with additive effect (9.76%); we also detected 108 QTL affecting five panicle traits in IAS and its corresponding F1 population, of all the QTL with additive and dominance effect,26 QTL were identified with over-dominance effect (amounting to 24.76%),47 QTL with partial dominance effect (amounting to 44.76%),18 QTL with complete dominance effect (amounting to 17.14%),17 QTL with additive effect (amounting to 16.19%). Results indicated that dominance (mainly were partial dominance) of QTL plays an important role in underlying the genetic basis of heterosis of panicle traits in rice hybrids.
8. We have detected many QTL clusters affecting agronomic traits located on chromosome 1,2,3,4,6,7,8,9 and 12 maybe because of pleiotropic effects of QTL or close linkage of some markers. We also analyzed the genetic relationships between QTL clusters and yield-related agronomic traits and found pleiotropic QTL was a common phenomenon which exhibited the pleiotropic phenomenon might be one of the genetic bases of yield-related traits and heterosis.
近百年来,杂种优势遗传机理的研究水平一直落后于对其在生产上的利用程度,这种杂种优势理论研究滞后于生产实践的局面势必影响杂种优势在现实中的进一步大规模利用。研究水稻的杂种优势形成机理,不仅对水稻杂交种的选育具有重要的指导意义,同时,水稻作为单子叶植物基因组研究的模式植物,将为研究禾谷类作物杂种优势理论提供模式借鉴。因此,广泛开展水稻杂种优势遗传基础的研究和探讨具有十分重要的理论价值和现实意义。
水稻籼／粳亚种间的杂种优势强于籼/籼或粳/粳亚种内的杂种优势,利用籼粳亚种间杂种优势是进一步提高水稻单产的重要技术策略。本研究利用来自日本的典型粳稻品种Asominori和国际水稻所(IRRI)育成的典型籼稻品种IR24为双亲构建的互为受体(遗传背景)和供体(置换片段)的两套染色体片段置换系群体及其与各自对应背景亲本杂交构建的两套杂种F1群体共四套材料,在两年两点四个环境(2007、2008年南京和南昌)对水稻籼粳亚种间组合产量构成性状和穗形相关性状等与产量密切相关的农艺性状进行了杂种优势的研究,利用SSR标记重新构建的染色体片段置换系基因型图谱检测产量相关农艺性状增(减)效杂种优势位点(Heterotic loci, HL),并通过利用染色体片段置换系群体检测到的具有加性效应的QTL和对应杂种F1群体检测到的具有显性效应的QTL之间的加显性效应剖析了产量相关农艺性状杂种优势形成的遗传基础。主要研究结果如下：
1、利用SSR标记重新鉴定了以Asominori为受体亲本,IR24为供体染色体片段置换系(简称AIS)和以IR24为受体亲本,Asominori为供体的染色体片段置换系(简称IAS)的基因型。利用全基因组筛选到的137个SSR多态性标记构建了AIS置换系基因型图谱和132个SSR多态性标记构建了IAS置换系基因型图谱,获得了QTL分析所需要的室内分子标记数据。
2、两套染色体片段置换系大部分农艺性状表现出明显的超亲分离现象；各农艺性状杂种优势均表现出变幅极大的分离,但不同农艺性状杂种优势的表现程度不一；不同农艺性状年度间均表现出显著差异,部分性状基因型与环境显著互作。
3、无论是高值亲本,还是低值亲本,都包含有对性状表现型起增效或者起减效作用的等位基因；这些数量性状位点分散在双亲的基因组中,个别区段有连锁。
4、采用基于逐步回归和极大似然比估计相结合的完备区间作图法,利用QTL IciMapping软件为检测平台,以经验阈值LOD≥3.0作为判断产量相关农艺性状杂种优势QTL存在与否的标准,对8个产量相关性状杂种优势效应进行了QTL定位。在四个环境中利用AIS和IAS对应杂种F1群体,分别定位了83个和53个与产量构成性状相关的增(减)效杂种优势标记位点。其中AIS对应杂种F1群体中发现13个标记位点能在多个环境重复检测到,有些位点能同时影响多个性状的表现,位于第5和7染色体上的RM267和RM82标记位点杂合状态下减少杂种F1代单株产量、每穗实粒数和结实率RM82标记位点还同时增加杂种F1代单株库容和株高。来自第2染色体上的RM5390标记位点杂合状态下能同时增加单株产量、每穗实粒数和结实率的表现。来自第4和6染色体上的RM252和RM217标记位点杂合状态下降低杂种F1的株高。IAS对应杂种F1群体中只发现1个标记位点能在多个环境重复检测到,来自第1染色体上的RM488标记位点杂合状态下增加杂种F1的株高。从HL效应的方向来看,AIS对应F1群体中检测到39个增效位点,占总位点数的46.99%；IAS对应F1群体中检测到22个增效位点,占总位点数的41.51%；两套F1群体还分别检测到44个和31个减效位点,这些减效位点主要集中在单株产量、每穗实粒数和结实率等与籼粳不育基因密切相关的性状,通过广亲和材料的应用或者染色体片段置换技术可以消除减效位点的负向效应。
5、利用AIS和IAS对应杂种Fl群体四个环境中的田间表型数据结合QTL分析软件,分别定位了40个和22个与穗形性状相关的增(减)效杂种优势标记位点。其中AIS对应杂种F1群体中发现2个增(减)效杂种优势标记位点能在多个环境重复检测到,来自第8染色体上的RM284标记位点能增加杂种F1的穗长,来自第8染色体上的标记位点RM331能增加杂种F1的一次枝梗数；IAS对应杂种F1群体中未发现能在多个环境重复检测到的增(减)效杂种优势标记位点。两套群体在第8染色体上检测到相同的杂种优势标记位点RM284,杂合状态下都表现增加杂种F1的穗长性状。从HL效应的方向来看,AIS对应F1群体中有27个增效位点,占总位点数的67.50%；IAS对应F1群体中有12个增效位点,占总位点数的54.55%；两套F1群体还分别检测到13个和10个减效位点。育种过程中可根据不同育种目标灵活利用穗形相关性状增(减)杂种优势位点来改良穗形性状。
6、利用AIS及其对应F1群体在四个环境分别检测到99个和79个与产量性状相关的加性与显性效应的QTL,其中具有加性效应的QTL分布在全部12条染色体上具有显性效应的QTL分布在除了Chr.9和Chr.11外的10条染色体上。而利用IAS及其对应F1群体在四个环境分别检测到116个和125个与产量性状相关的加性与显性效应的QTL,其中加性与显性效应的QTL都分布在全部12条染色体上。通过产量构成性状QTL的显性效应值D和加性效应值A的比值(平均显性度,|D/A|)分析,发现粳稻背景下共有52个QTL表现超显性效应,约占全部QTL的34.44%；61个QTL表现部分显性效应,约占全部QTL的40.40%；22个QTL表现显性效应,约占全部QTL的14.57%；16个QTL表现加性效应,约占全部QTL的10.60%；籼稻背景下,共有61个QTL表现超显性效应,约占全部QTL的33.70%；69个QTL表现部分显性效应,约占全部QTL的38.12%；32个QTL表现完全显性效应,约占全部QTL的17.68%；19个QTL表现加性效应,约占全部QTL的10.50%。因此,可以看出,QTL的超显性效应和显性效应(主要是部分显性)可能是水稻产量构成性状杂种优势形成的重要原因。
7、利用AIS及其对应F1群体在不同环境中分别检测到52个和55个与穗形性状相关的加性与显性效应的QTL,其中具有加性效应的QTL分布在除了第9和11染色体外的10条染色体上,具有显性效应的QTL分布在除了第5,9和11染色体外的9条染色体上。而利用IAS及其对应F1群体在不同环境中分别检测到75个和62个与穗形性状相关的加性与显性效应的QTL,都分布在于12条染色体上。通过穗形相关性状QTL平均显性度(|D/A|)的比较分析,发现粳稻背景下共有18个QTL表现超显性效应,约占全部QTL的21.95%;45个QTL表现部分显性效应,约占全部QTL的54.88%；11个QTL表现显性效应,约占全部QTL的13.41%；8个QTL表现加性效应,约占全部QTL的9.76%；籼稻背景下,共有26个QTL表现超显性效应,约占全部QTL的24.76%；47个QTL表现部分显性效应,约占全部QTL的44.76%；18个QTL表现完全显性效应,约占全部QTL的17.14%；17个QTL表现加性效应,约占全部QTL的16.19%。因此,可以看出,QTL的显性效应(主要是部分显性)可能是水稻穗形相关性状杂种优势形成的重要原因。
8、从产量相关农艺性状杂种优势位点以及加性和显性效应的QTL在染色体上的分布来看,发现在第1,2,3,4,6,7,8,9和12染色体上存在QTL的“簇状”分布现象,并对QTL簇与产量相关农艺性状的遗传关系进行了分析,发现各相关农艺性状普遍存在QTL的多效性,因此认为QTL的多效性可能是产量相关农艺性状以及杂种优势的遗传基础之一
With the development of social economy, greater demand was put on world food production due to the decreasing arable lands and increasing global population. The yield increase of rice, a main staple for a large fragment of the world population (amounting to half global population and 60%of population in our country), would contribute significantly to cope with the increasing global food crisis. Heterosis application was the principal way to improve rice grain yield.
However, since the advancement of research on the genetic basis of heterosis was lagged behind the utilization in the field production in last century, which limited a larger scale of heterosis utilization in crop production. The research of heterosis in rice is not only meaningful to guide the rice hybrid breeding, but also provide a theoretical model for the research of heterosis in the other cereal crops because rice is a model plant in monocotyledonous genomic research. Therefore, it is very important to conduct the research of heterosis and dissect its genetic basis for the theoretical value and realistic meaning in hybrid breeding of rice.
The extent of heterosis expressed in inter-subspecific crosses follows the general trend: indica/japonica> indicalindica> japonica/japonica. The extensive exploitation of inter-subspecific heterosis derive from indica/japonica is a very important strategy to enhance the yield level in rice production. The inter-subspecific cross between a japonica cv. Asominori and an indica cv. IR24 demonstrated huge heterosis in F1 hybrid. In our study, we conducted our experiments across four environments (2007,2008 in Nanjing; 2007, 2008 in Nanchang), mainly focus on the dissection and discussion of the genetic basis of heterosis about yield-related agronomic traits using two sets of chromosome segment substitution lines (CSSLs) derived from Asominori (a japonica cultivar) and IR24 (an indica cultivar) as reciprocal donor parent and recipient parent (background parent) and F1 derivatives from these two set of CSSLs population. QTL were detected by statistical software QTL IciMapping using two new different SSR map. The main conclusions are as follows:
1. Genotypes of these two sets of 66 CSS lines (one is indica c.v. IR24 chromosome segment substitution lines with Japonica c.v. Asominori genetic background; the other is japonica c.v. Asominori chromosome segment substitution lines with indica c.v. IR24 genetic background) were rescreened by 137 and 132 good polymorphism SSR markers, replacing 82 original RFLP markers. The purpose of replacing with SSR markers was to simplify the procedure of experiments by using RFLP markers and to enhance the efficiency of QTL detection using new maps with higher density of SSR markers.
2. Most of the agronomic traits showed transgressive segregation phenomenon in two CSSLs population; Heterosis of each agronomic trait exhibited a huge range of segregation, however different traits showed different level of heterosis; The studied traits also showed significant difference between years and genotypes, the interaction between genotypes and environments also exhibited a significance level.
3. Not only the high value parent but also the low value parent has detected many alleles at QTL with positive or negative effects in the whole genome, some chromosome segments were tightly linked.
4. We have detected 83 and 53 heterotic loci (HL) affecting yield component traits in Fi derivatives of AIS and IAS population, respectively. Of all the HL, only 13 HL could be detected repeatedly in two environments at least which reflected heterosis of yield component traits were very sensitive to environmental influence. Chances were the effect of pleiotropism, chromosome segments RM267 and RM82 on Chr.5 and Chr.7 could negatively affect grain weight plant-1 (GWP), grains per panicle (GPP) and seed-setting rate (SSR), simultaneously. RM82 could enhance the performance of sink capacity plant-1(SCP) and plant height (PH) as well. Chromosome segment RM5390 as an enhancing heterotic locus on Chr.2 could positively affect GWP, GPP and SSR traits simultaneously. RM252 and RM217 on Chr.4 and Chr.6 could decrease PH of F1 hybrids. Considering the effect of heterotic loci,39 with positive heterosis (amounting to 46.99%) and 44 (53.01%) with negative heterosis in AIS F1 derivative; 22 HL (41.51%) with positive and 31 (58.49%) with negative effect of heterosis in IAS F1 derivative. Possibly, we could create high-yield cultivar through pyramiding HL with positive effect and deleting HL negative effect by molecular tools such as SSR polymorphism markers.
5. We have detected 40 and 22 HL affecting panicle morphological traits in F1 derivatives of AIS and IAS population, respectively. Of all the HL were detected,27 with positive heterosis (67.50%) and 13 (32.50%) with negative heterosis in AIS F] derivative; 12 HL (54.55%) with positive and 10 (45.45%) with negative effect of heterosis in IAS F1 derivatives. Only 2 HL in AIS F1 derivative could be detected repeatedly in two more environments which reflected heterosis of panicle morphological traits were very sensitive to environment, too. Chromosome segments RM284 and RM331 on Chr.8 could increase the performance of panicle length (PL) and primary branches number (PBN), respectively. None HL could be detected in IAS F1 derivative. But we found that RM284 on Chr.8 could be detected in two sets of F1 derivatives, simultaneously.
6. Through mapping additive QTL and dominance QTL affecting yield component traits, we could dissect the genetic basis of heterosis. Eighty-nine additive QTL and 79 dominance QTL affecting eight yield component traits (grains weight per plant, GWP; sink capacity per plant, SCP; spikelets per panicle, SPP; grains per panicle, GPP; seed-setting rate, SSR; thousand-grain weight, TGW; panicles per plant, PPP; plant height, PH) were detected across four environments in AIS and F1 derivative population; 117 additive QTL and 125 dominance QTL located on all 12 chromosomes affecting eight yield-related traits were detected across the four same environments in IAS and F1 derivative population. Through comparing dominance effects and additive effects of those 151 QTL totally affecting eight yield-related traits in AIS and its corresponding F1 population, we found 52 QTL with over-dominance effect (amounting to 34.44%),61 QTL with partial dominance effect (amounting to 40.40%),22 QTL with complete dominance (amounting to 14.57%), 16 QTL with additive effect (amounting to 10.60%); we also detected 181 QTL totally affecting five panicle traits in IAS and its corresponding F1 population, of all the QTL with additive and dominance effect,61 QTL were identified with over-dominance effect (amounting to 33.70%),69 QTL with partial dominance effect (amounting to 38.12%),32 QTL with complete dominance effect (amounting to 17.68%),19 QTL with additive effect (amounting to 10.50%). Results indicated that both dominance (mainly were partial dominance) and over-dominance effects of QTL may be as the major reason in underlying the genetic basis of heterosis of yield component traits in rice hybrids.
7. We detected 52 QTL with additive effects and 55 QTL with dominance effects affecting five panicle morphological structural traits (panicle length, PL; density of spikelets per panicle, DSP; primary branches number, PBN; secondary branches number, SBN; density of secondary branches number, DSBN) across different environments in AIS and F1 derivative population. We found 72 additive QTL and 62 dominance QTL affecting five panicle morphological structural traits were detected across three same environments in IAS and F1 population. Comparing dominance effect and additive effect of those 82 QTL affecting five panicle morphological structural traits in AIS and its corresponding F1 population, we found 18 QTL with over-dominance effect (amounting to 21.95%),45 QTL with partial dominance effect (amounting to 54.88%),11 QTL with complete dominance (amounting to 13.41%),8 QTL with additive effect (9.76%); we also detected 108 QTL affecting five panicle traits in IAS and its corresponding F1 population, of all the QTL with additive and dominance effect,26 QTL were identified with over-dominance effect (amounting to 24.76%),47 QTL with partial dominance effect (amounting to 44.76%),18 QTL with complete dominance effect (amounting to 17.14%),17 QTL with additive effect (amounting to 16.19%). Results indicated that dominance (mainly were partial dominance) of QTL plays an important role in underlying the genetic basis of heterosis of panicle traits in rice hybrids.
8. We have detected many QTL clusters affecting agronomic traits located on chromosome 1,2,3,4,6,7,8,9 and 12 maybe because of pleiotropic effects of QTL or close linkage of some markers. We also analyzed the genetic relationships between QTL clusters and yield-related agronomic traits and found pleiotropic QTL was a common phenomenon which exhibited the pleiotropic phenomenon might be one of the genetic bases of yield-related traits and heterosis.
引文
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