整合分析甘蓝型油菜中控制种子含油量变异的数量性状位点
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摘要
甘蓝型油菜(Brassica napus L.)是世界范围内四大油料作物之一,其菜籽产油除了满足日常食用所需,也可以被加工成工业用油；随着不可再生的原油资源日益告急,提高植物油的产量和品质对国计民生有着重大的战略意义。在过去的数十年间,甘蓝型油菜的菜籽单产量得到了翻番,菜籽油的脂肪酸含量比例可以实现满足不同用途的优化,但是菜籽的含油量提升幅度较小。由于种子含油量是一个典型的数量性状,因此通过数量性状位点遗传定位(QTL mapping)的方法来发掘其调控位点、研究其变异的机理、并应用于育种改良,是切实可行的策略。之前已有许多关于甘蓝型油菜种子含油量QTL的定位研究,但是其中存在一些问题,以致这些研究结果对育种改良实践的促进效果并不明显,如,单项研究中定位得到的QTL数目非常有限、不超过20个,单个QTL对表型变异的贡献率大多不超过10%,且很多QTL只在特定遗传群体或试验条件下才被检测得到。对种子含油量这一复杂性状的育种改良,需要有更多可操作的遗传位点、获得这些位点上相对高效的等位基因,而且还需要了解这些位点上特异基因型发挥功能的适宜条件,由此才有可能使育种实践的结果与预期相符。
本研究考察了一个甘蓝型油菜双单倍体(DH)遗传群体一-TN DH群体,以及由它衍生的重构F2(Reconstructed-F2)群体--TN RC-F2群体,在连续5年、设置于中国不同地域的、共计15个独立试验中的种子含油量变异。其中,该遗传群体的两亲本材料Tapidor和宁油7号均为中等含油量品系,在多个试验中的种子含油量均值分别为43.0±1.7%和41.7±2.1%,遗传群体内的变异为典型的连续分布,且呈现超亲分离,变异范围可达到从33%到51%,平均变幅为11%。
另外,通过对TNDH群体各个株系的主花序自下而上进行分节段收获角果(分为5段),我们发现不同株系在沿节段发育过程中种子含油量的变异形式有所不同,相邻节段间种子含油量或表现上升或表现下降。在一定的植株极性条件下,这种不同形式的变异有可能与不同节段上角果发育期间的环境条件有关。通过对相邻节段过渡上变异形式不同的株系分组,进行基因型与变异类型的独立性测验,得到基因组上103个标记位点与不同的变异类型相关。
将各个节段上的收获也作为独立试验,我们对上述共计20个独立试验分别进行了QTL定位和上位性互作对位点的检测,共得到195个、大部分存在重叠的原始QTL区间,以及92个涉及互作的标记位点。其中,105个原始QTL的显著性概率为p≤0.05,被定义为显著QTL,另90个为p≤0.5,被定义为微效QTL。涉及互作的标记位点在不同的试验间鲜有重叠,其中大部分包含在原始QTL区间以内,只有36个互作位点独立在原始QTL区间之外。而上述与节段上种子含油量变异类型相关的遗传位点中,也有一半包含在这些原始QTL区间内。
根据“元分析(meta-analysis)"的方法,我们将重叠的原始QTL进行了区间的整合与精炼。由此,195个原始QTL区间被归纳为59个独立的、不相重叠的“元QTL(meta-QTL)",和28个微效、独立的原始QTL；根据这些元QTL所包含的原始QTL的显著性概率,可将它们划分为50个显著元QTL(即包含的原始QTL至少有一个为显著QTL),和9个微效真实元QTL(即包含多个重叠的微效QTL)。由于每个独立QTL包含的若干原始QTL都对应若干特定的试验,而我们记录了各个独立试验中种子油份积累关键时期的光照、温度条件,因此,可以分析这些独立QTL的环境特异性。结果发现,有28个元QTL表现出一定的环境特异性,且其中一些只在特定试验中被检测得到的独立QTL的确是对应环境条件相对极端的试验。所以我们提出,不应该忽视那些环境特异性的QTL。
借助遗传图谱间的共同分子标记,我们实现了将之前报道过的其它几个具有代表性的甘蓝型油菜遗传群体中种子含油量QTL的定位结果部分映射到了TN遗传图谱上。通过比较分析发现,多环境试验有助于提高TN遗传群体中的QTL检出率,使其一个群体中所得的QTL可以覆盖其它三个群体所得QTL的80%；另一方面,其它群体定位的QTL也验证了TN群体中30%的QTL,其中包括25%的微效QTL,另外还有近20%独立于QTL区间之外的互作位点。
除了不同遗传群体间的QTL比较,我们还将由育种品系中关联分析所得的含油量变异关联位点对应到TN遗传图谱上,与定位的QTL进行了比较。关联位点验证了TN群体中约25%的QTL,其中一半为微效QTL,另外也还有近20%独立于QTL区间之外的互作位点被关联分析验证。同时,QTL效应大小也在比较分析中表现出与遗传背景或环境条件有关的明显的相对性特点。
本研究中所揭示的大量QTL将为育种改良提供丰富的目标位点,而所反映出来的QTL效应大小的相对性以及遗传背景或环境条件的特异性,将对特定的遗传改良设计中对于相关遗传位点上等位基因的选择与组合提供重要的参考。
Oilseed rape (Brassica napus L.) is one of the most improtant oil crops worldwide, the oil from its seeds could support both edible and industrial requirement. As the reservation of non-renewable crude oil resources gets more and more limeted, it is strategically crutial for national safty and development of improving the yield and quality of renewable oil resources. During the past decades, the seed yield per unit of oilseed rape has been doubled, the fatty acid composition could be optimized for requirement of different usage, but the improvement of total oil content in seeds was slow and minor. Since the oil content in seeds is typically a quantitative trait, it is feasible to exploit the related genetic loci, unreval the variation mechanisms, and facilitate breeding practice via the quantitative trait loci (QTL) mapping. There has been a lot of studies referring QTL mapping related to oil content in seeds of B. napus, but some problems from those studies hamperred their application in breeding practice, for example, the number of identified QTL from single independent experiment is quite limited and always below 20, most of the single QTL contributes less than 10% of the phenotypic variation, and many QTL could be only detected from particular population or environmental condition. So, to effectively improve the complex trait seed oil content, it requires more manipulable genetic loci and their corresponding superior alleles, and furthermore, it will make the results of breeding practice better accordant with expectation when understand the proper conditions that particular alleles need to well function.
In this study, seed oil content variation in a double haploid (DH) population, the TN DH population, and its derivate reconstruct-F2 (RC-F2) population were measured from totally 15 independent experiments including different years and different locations. The two parental lines of TN genetic population were both with moderate content of oil in seeds, that Tapidor had oil content 43.0±1.7% in seeds while Ningyou7 had 41.7±2.1%, on average. The oil content variation showed continuous distribution and transgressive segregation among the TN genetic population, the oil content varied from 33% to 51% extremely, and the average extent of oil content variation was 11%.
Moreover, a kind of main inflorescence section harvest experiment has been carried out based on the TN DH popoulation, the main inflorescence of each line was labelled as 5 sections from bottom to tip along flowering, and siliques from each section were seperately harvested. It is notable that the direction of oil content variation along adjacent sections was different among lines, either increasing or decreasing. When under a certain polarity of plant, the different types of oil content variation along adjacent sections may probably due to divergent responses of different genotypes to the specific environmental conditions, under which the siliques of each section developed. Grouping lines according to the opposite types of oil content variation between each adjacent sections, theχ2 test for independence was applied to evaluate the correlation of genotype and phenotypic variation type, and 103 marker loci were found related to the divergence of oil content variation types along adjacent sections.
Taking the seed oil content variation among TN DH population on each section as an independent experiment also, we did QTL mapping and epistatic interaction loci detection for each of the 20 independent experiments, respectively. A total of 195 original QTL, most intervals of which were overelapped with others, and 92 epistatic interaction involved loci, were identified from all the experiments. Among the original QTL,105 were defined as significant-large QTL (SL-QTL) which at the significant level p≤0.05, while the other 90 were defined as weak QTL which at the significant level p<0.5. Most of the epistatic loci were specifically identified from particular experiment; a large part of the epistatic loci were included in the original QTL intervals, while the other 36 epistatic loci were independent of QTL intervals. Half of the 103 marker loci, which were found related to the divergence of oil content variation types along adjacent sections, were also involved in the original QTL intervals.
Applying the "meta-analysis"method, we integrated the overlapped original QTL to get refined intervals. Then the 195 original QTL were concluded as 59 independent "meta-QTL" and 28 weak-singleton QTLs (WS-QTL). The meta-QTL were sorted according to the significant level of the original QTL they included,50 meta-QTL had at least one significant original QTL included and were defined as meta-SL-QTL, and 9 meta-QTL had multiple weak original QTL included and were defined as meta-MR(micro-real)-QTL. As each meta-QTL included one or several original QTL which were corresponding to particular experiment(s), and the environmental conditions, temperature and sunlight illumination time, were recorded during the pivotal phase of seeds filling in each experiment, then it is feasible to analyse the environmental specificity that the meta-QTL required to well function. We found 28 meta-QTLs were specifically detected from particular conditions of temprature, or sunlight illumination, or both aspects, and notably several independent QTL which were detected from only single experiment indeed with the particular experiment had extreme condition(s). So, we suggest that the environment specific QTL should not be neglected.
Through the common molecular markers on different genetic maps, we projected the reported QTL, which were related to seed oil content variation and identified from other genetic populations of B. napus, onto the TN genetic map that was used as reference. The multiple experiments showed significant improving on the detectable number of QTL:the QTL from only TN genetic population covered 80% of the QTL which were identified previously from other three populations. On the other hand, the QTL identified from other populations validated 30% of the QTL in TN population, including 25% of the weak QTL and 20% of the epistatic loci which were independent of QTL intervals.
In addition to the comparison among different genetic populations, we furthermore compared the QTL that were identified from genetic map based mapping with associated markers that were identified from naturally varied breeding lines. The corresponding loci on TN genetic map of the associated markers were identified first, and then we found 25% of the QTL in TN population were validated by the associated loci, including half of the weak QTL and also 20% of the epistatic loci which were independent of QTL intervals. The inconsistent effects of particular QTL between different genetic backgrounds or environmental conditions were apparent through the comparisons.
The unprecedented large amount of QTL related to seed oil content variation in B. napus that was revealled in this study, will provide rich targets for genetic improvement of such trait; While the uncovered relativity and specificity of QTL effects, are important references for choosing and combining of the most effective alleles in particular genetic design under a certain breeding condition.
本研究考察了一个甘蓝型油菜双单倍体(DH)遗传群体一-TN DH群体,以及由它衍生的重构F2(Reconstructed-F2)群体--TN RC-F2群体,在连续5年、设置于中国不同地域的、共计15个独立试验中的种子含油量变异。其中,该遗传群体的两亲本材料Tapidor和宁油7号均为中等含油量品系,在多个试验中的种子含油量均值分别为43.0±1.7%和41.7±2.1%,遗传群体内的变异为典型的连续分布,且呈现超亲分离,变异范围可达到从33%到51%,平均变幅为11%。
另外,通过对TNDH群体各个株系的主花序自下而上进行分节段收获角果(分为5段),我们发现不同株系在沿节段发育过程中种子含油量的变异形式有所不同,相邻节段间种子含油量或表现上升或表现下降。在一定的植株极性条件下,这种不同形式的变异有可能与不同节段上角果发育期间的环境条件有关。通过对相邻节段过渡上变异形式不同的株系分组,进行基因型与变异类型的独立性测验,得到基因组上103个标记位点与不同的变异类型相关。
将各个节段上的收获也作为独立试验,我们对上述共计20个独立试验分别进行了QTL定位和上位性互作对位点的检测,共得到195个、大部分存在重叠的原始QTL区间,以及92个涉及互作的标记位点。其中,105个原始QTL的显著性概率为p≤0.05,被定义为显著QTL,另90个为p≤0.5,被定义为微效QTL。涉及互作的标记位点在不同的试验间鲜有重叠,其中大部分包含在原始QTL区间以内,只有36个互作位点独立在原始QTL区间之外。而上述与节段上种子含油量变异类型相关的遗传位点中,也有一半包含在这些原始QTL区间内。
根据“元分析(meta-analysis)"的方法,我们将重叠的原始QTL进行了区间的整合与精炼。由此,195个原始QTL区间被归纳为59个独立的、不相重叠的“元QTL(meta-QTL)",和28个微效、独立的原始QTL；根据这些元QTL所包含的原始QTL的显著性概率,可将它们划分为50个显著元QTL(即包含的原始QTL至少有一个为显著QTL),和9个微效真实元QTL(即包含多个重叠的微效QTL)。由于每个独立QTL包含的若干原始QTL都对应若干特定的试验,而我们记录了各个独立试验中种子油份积累关键时期的光照、温度条件,因此,可以分析这些独立QTL的环境特异性。结果发现,有28个元QTL表现出一定的环境特异性,且其中一些只在特定试验中被检测得到的独立QTL的确是对应环境条件相对极端的试验。所以我们提出,不应该忽视那些环境特异性的QTL。
借助遗传图谱间的共同分子标记,我们实现了将之前报道过的其它几个具有代表性的甘蓝型油菜遗传群体中种子含油量QTL的定位结果部分映射到了TN遗传图谱上。通过比较分析发现,多环境试验有助于提高TN遗传群体中的QTL检出率,使其一个群体中所得的QTL可以覆盖其它三个群体所得QTL的80%；另一方面,其它群体定位的QTL也验证了TN群体中30%的QTL,其中包括25%的微效QTL,另外还有近20%独立于QTL区间之外的互作位点。
除了不同遗传群体间的QTL比较,我们还将由育种品系中关联分析所得的含油量变异关联位点对应到TN遗传图谱上,与定位的QTL进行了比较。关联位点验证了TN群体中约25%的QTL,其中一半为微效QTL,另外也还有近20%独立于QTL区间之外的互作位点被关联分析验证。同时,QTL效应大小也在比较分析中表现出与遗传背景或环境条件有关的明显的相对性特点。
本研究中所揭示的大量QTL将为育种改良提供丰富的目标位点,而所反映出来的QTL效应大小的相对性以及遗传背景或环境条件的特异性,将对特定的遗传改良设计中对于相关遗传位点上等位基因的选择与组合提供重要的参考。
Oilseed rape (Brassica napus L.) is one of the most improtant oil crops worldwide, the oil from its seeds could support both edible and industrial requirement. As the reservation of non-renewable crude oil resources gets more and more limeted, it is strategically crutial for national safty and development of improving the yield and quality of renewable oil resources. During the past decades, the seed yield per unit of oilseed rape has been doubled, the fatty acid composition could be optimized for requirement of different usage, but the improvement of total oil content in seeds was slow and minor. Since the oil content in seeds is typically a quantitative trait, it is feasible to exploit the related genetic loci, unreval the variation mechanisms, and facilitate breeding practice via the quantitative trait loci (QTL) mapping. There has been a lot of studies referring QTL mapping related to oil content in seeds of B. napus, but some problems from those studies hamperred their application in breeding practice, for example, the number of identified QTL from single independent experiment is quite limited and always below 20, most of the single QTL contributes less than 10% of the phenotypic variation, and many QTL could be only detected from particular population or environmental condition. So, to effectively improve the complex trait seed oil content, it requires more manipulable genetic loci and their corresponding superior alleles, and furthermore, it will make the results of breeding practice better accordant with expectation when understand the proper conditions that particular alleles need to well function.
In this study, seed oil content variation in a double haploid (DH) population, the TN DH population, and its derivate reconstruct-F2 (RC-F2) population were measured from totally 15 independent experiments including different years and different locations. The two parental lines of TN genetic population were both with moderate content of oil in seeds, that Tapidor had oil content 43.0±1.7% in seeds while Ningyou7 had 41.7±2.1%, on average. The oil content variation showed continuous distribution and transgressive segregation among the TN genetic population, the oil content varied from 33% to 51% extremely, and the average extent of oil content variation was 11%.
Moreover, a kind of main inflorescence section harvest experiment has been carried out based on the TN DH popoulation, the main inflorescence of each line was labelled as 5 sections from bottom to tip along flowering, and siliques from each section were seperately harvested. It is notable that the direction of oil content variation along adjacent sections was different among lines, either increasing or decreasing. When under a certain polarity of plant, the different types of oil content variation along adjacent sections may probably due to divergent responses of different genotypes to the specific environmental conditions, under which the siliques of each section developed. Grouping lines according to the opposite types of oil content variation between each adjacent sections, theχ2 test for independence was applied to evaluate the correlation of genotype and phenotypic variation type, and 103 marker loci were found related to the divergence of oil content variation types along adjacent sections.
Taking the seed oil content variation among TN DH population on each section as an independent experiment also, we did QTL mapping and epistatic interaction loci detection for each of the 20 independent experiments, respectively. A total of 195 original QTL, most intervals of which were overelapped with others, and 92 epistatic interaction involved loci, were identified from all the experiments. Among the original QTL,105 were defined as significant-large QTL (SL-QTL) which at the significant level p≤0.05, while the other 90 were defined as weak QTL which at the significant level p<0.5. Most of the epistatic loci were specifically identified from particular experiment; a large part of the epistatic loci were included in the original QTL intervals, while the other 36 epistatic loci were independent of QTL intervals. Half of the 103 marker loci, which were found related to the divergence of oil content variation types along adjacent sections, were also involved in the original QTL intervals.
Applying the "meta-analysis"method, we integrated the overlapped original QTL to get refined intervals. Then the 195 original QTL were concluded as 59 independent "meta-QTL" and 28 weak-singleton QTLs (WS-QTL). The meta-QTL were sorted according to the significant level of the original QTL they included,50 meta-QTL had at least one significant original QTL included and were defined as meta-SL-QTL, and 9 meta-QTL had multiple weak original QTL included and were defined as meta-MR(micro-real)-QTL. As each meta-QTL included one or several original QTL which were corresponding to particular experiment(s), and the environmental conditions, temperature and sunlight illumination time, were recorded during the pivotal phase of seeds filling in each experiment, then it is feasible to analyse the environmental specificity that the meta-QTL required to well function. We found 28 meta-QTLs were specifically detected from particular conditions of temprature, or sunlight illumination, or both aspects, and notably several independent QTL which were detected from only single experiment indeed with the particular experiment had extreme condition(s). So, we suggest that the environment specific QTL should not be neglected.
Through the common molecular markers on different genetic maps, we projected the reported QTL, which were related to seed oil content variation and identified from other genetic populations of B. napus, onto the TN genetic map that was used as reference. The multiple experiments showed significant improving on the detectable number of QTL:the QTL from only TN genetic population covered 80% of the QTL which were identified previously from other three populations. On the other hand, the QTL identified from other populations validated 30% of the QTL in TN population, including 25% of the weak QTL and 20% of the epistatic loci which were independent of QTL intervals.
In addition to the comparison among different genetic populations, we furthermore compared the QTL that were identified from genetic map based mapping with associated markers that were identified from naturally varied breeding lines. The corresponding loci on TN genetic map of the associated markers were identified first, and then we found 25% of the QTL in TN population were validated by the associated loci, including half of the weak QTL and also 20% of the epistatic loci which were independent of QTL intervals. The inconsistent effects of particular QTL between different genetic backgrounds or environmental conditions were apparent through the comparisons.
The unprecedented large amount of QTL related to seed oil content variation in B. napus that was revealled in this study, will provide rich targets for genetic improvement of such trait; While the uncovered relativity and specificity of QTL effects, are important references for choosing and combining of the most effective alleles in particular genetic design under a certain breeding condition.
引文
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18. Burns M J, Barnes S R, Bowman J G, et al. QTL analysis of an intervarietal set of substitution lines in Brassica napus:(i) seed oil content and fatty acid composition. Heredity,2003,90:39-48
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22. Ching A, Caldwell K S, Jung M, et al. SNP frequency, haplotype structure and linkage disequilibrium in elite maize inbred lines. BMC Genet,2002,3:19
23. Delourme R, Falentin C, Huteau V, et al. Genetic control of oil content in oilseed rape(Brassica napus L.). Theor Appl Genet,2006,113:1331-1345
24. Eastmond P J and Rawsthorne S. Coordinate changes in carbon partitioning and plastidial metabolism during the development of oilseed rape embryos. Plant Physiol,2000,122:767-774
25. Ecke W, Uzunova M and WeiBleder K. Mapping the genome of rapeseed(Brassica napus L.). II. Localization of genes controlling erucic acid synthesis and seed oil content. Theor Appl Genet, 1995,91:972-977
26. Falconer D S. The problem of environment and selection. The American Naturalist,1952, 86:293
27. Flint-Garcia S A, Thornsberry J M and Buckler E St. Structure of linkage disequilibrium in plants. Annu Rev Plant Biol,2003,54:357-374
28. Flint-Garcia S A, Thuillet A C, Yu J, et al. Maize association population:A high-resolution platform for quantitative trait locus dissection. Plant J,2005,44:1054-1064
29. Fourmann M, Barret P, Renard M, et al. The two genes homologous to Arabidopsis FAE1 co-segregate with the two loci governing erucic acid content in Brassica napus. Theor Appl Genet, 1998,96:852-858
30. Frandsen G I, Mundy J and Tzen J T. Oil bodies and their associated proteins, oleosin and caleosin. Physiol Plant,2001,112:301-307
31. Fu D, Uauy C, Distelfeld A, et al. A kinase-start gene confers temperature-dependent resistance to wheat stripe rust. Science,2009a,323:1357-1360
32. Fu J, Keurentjes J J, Bouwmeester H, et al. System-wide molecular evidence for phenotypic buffering in Arabidopsis. Nat Genet,2009b,41:166-167
33. Fu S X, Cheng H and Qi C. Microarray analysis of gene expression in seeds of Brassica napus planted in Nanjing (altitude:8.9 m), Xining (altitude:2261.2 m) and Lhasa (altitude:3658 m) with different oil content. Mol Biol Rep,2009c,36:2375-2386
34. Gegas V C, Nazari A, Griffiths S, et al. A genetic framework for grain size and shape variation in wheat. Plant Cell,2010,22:1046-1056
35. Goffinet B and Gerber S. Quantitative trait loci:A meta-analysis. Genetics,2000,155:463-473
36. Gupta V, Mukhopadhyay A, Arumugam N, et al. Molecular tagging of erucic acid trait in oilseed mustard(Brassica juncea) by QTL mapping and single nucleotide polymorphisms in FAE1 gene. Theor Appl Genet,2004,108:743-749
37. Harris H C, McWilliam J R and Mason W K. Influence of temperature on oil content and composition of sunflower seed. Aust J Agr Res,1978,29:1203-1212
38. Hasan M, Friedt W, Pons-Kuhnemann J, et al. Association of gene-linked SSR markers to seed glucosinolate content in oilseed rape(Brassica napus ssp. Napus). Theor Appl Genet,2008, 116:1035-1049
39. Hodgson A S. Repeseed adaptation in northern New South Wales. II. Predicting plant development of Brassica campestris L. and Brassica napus L. and its implications for planting time, designed to avoid water deficit and frost. Aust J Agr Res,1978,29:711-726
40. Hu X, Sullivan-Gilbert M, Gupta M, et al. Mapping of the loci controlling oleic and linolenic acid contents and development of FAD2 and FAD3 allele-specific markers in canola(Brassica napus L.). Theor Appl Genet,2006,113:497-507
41. Hu Y, Wu G, Cao Y, et al. Breeding response of transcript profiling in developing seeds of Brassica napus. BMC Mol Biol,2009,10:49
42. Huang X Q, Paulo M-J, Effgen S, et al. Constructing a complex NIL population derived from intercrossing 8 Arabidopsis accessions. Abstract P-385,18th International Conference on Arabidopsis Research,2007, Beijing, China
43. Jansen R C. Interval mapping of multiple quantitative trait loci. Genetics,1993,135:205-211
44. Jiang C and Zeng Z B. Multiple trait analysis of genetic mapping for quantitative trait loci. Genetics,1995,140:1111-1127
45. Kang F and Rawsthorne S. Starch and fatty acid synthesis in plastids from developing embryos of oilseed rape(Brassica napus L.). Plant J,1994,6:795-805
46. Kang F and Rawsthorne S. Metabolism of glucose-6-phosphate and utilization of multiple metabolites for fatty acid synthesis by plastids from developing oilseed rape embryos. Planta, 1996,199:321-327
47. Katavic V, Agrawal G K, Hajduch M, et al. Protein and lipid composition analysis of oil bodies from two Brassica napus cultivars. Proteomics,2006,6:4586-4598
48. Kearsey M J and Farquhar A G. QTL analysis in plants; where are we now? Heredity,1998, 80:137-142
49. Keurentjes J J, Fu J, de Vos C H, et al. The genetics of plant metabolism. Nat Genet,2006, 38:842-849
50. Kim S, Plagnol V, Hu T T, et al. Recombination and linkage disequilibrium in Arabidopsis thaliana. Nat Genet,2007,39:1151-1155
51. Knutzon D S, Thompson G A, Radke S E, et al. Modification of Brassica seed oil by antisense expression of a stearoyl-acyl carrier protein desaturase gene. Proc Natl Acad Sci. USA,1992, 89:2624-2628
52. Lander E S and Botstein D. Mapping mendelian factors underlying quantitative traits using RFLP linkage maps. Genetics,1989,121:185-199
53. Lassner M W, Levering C K, Davies H M, et al. Lysophosphatidic acid acyltransferase from meadowfoam mediates insertion of erucic acid at the sn-2 position of triacylglycerol in transgenic rapeseed oil. Plant Physiol,1995,109:1389-1394
54. Lebowitz R J, Soller M and Beckmann J S. Trait-based analyses for the detection of linkage between marker loci and quantitative trait loci in crosses between inbred lines. Theor Appl Genet, 1987,73:556-562
55. Li R J, Wang H Z, Mao H, et al. Identification of differentially expressed genes in seeds of two near-isogenic Brassica napus lines with different oil content. Planta,2006,224:952-962
56. Li X, Qian Q, Fu Z, et al. Control of tillering in rice. Nature,2003,422:618-621
57. Long Y, Shi J, Qiu D, et al. Flowering time quantitative trait loci analysis of oilseed Brassica in multiple environments and genomewide alignment with Arabidopsis. Genetics,2007, 177:2433-2444
58. Lowe A J, Moule C, Trick M, et al. Efficient large-scale development of microsatellites for marker and mapping applications in Brassica crop species. Theor Appl Genet,2004, 108:1103-1112
59. Michaels S D and Amasino R M. Loss of Flowering Locus C activity eliminates the late-flowering phenotype of FRIGIDA and autonomous pathway mutations but not responsiveness to vernalization. Plant Cell,2001,13:935-941
60. Monna, Lin, Kojima, et al. Genetic dissection of a genomic region for a quantitative trait locus, Hd3, into two loci, Hd3a and Hd3b, controlling heading date in rice. Theor Appl Genet,2002, 104:772-778
61. Mu J, Tan H, Zheng Q, et al. LEAFY COTYLEDON1 is a key regulator of fatty acid biosynthesis in Arabidopsis. Plant Physiol,2008,148:1042-1054
62. Niu Y, Wu G Z, Ye R, et al. Global analysis of gene expression profiles in Brassica napus developing seeds reveals a conserved lipid metabolism regulation with Arabidopsis thaliana. Mol Plant,2009,2:1107-1122
63. Nordborg M, Borevitz J O, Bergelson J, et al. The extent of linkage disequilibrium in Arabidopsis thaliana. Nat Genet,2002,30:190-193
64. Norton G J, Aitkenhead M J, Khowaja F S, et al. A bioinformatic and transcriptomic approach to identifying positional candidate genes without fine mapping:An example using rice root-growth QTL. Genomics,2008,92:344-352
65. O'Neill C, Morgan C, Kirby J, et al. Six new recombinant inbred populations for the study of quantitative traits in Arabidopsis thaliana. Theor Appl Genet,2008,116:623-634
66. Ohlrogge J and Browse J. Lipid biosynthesis. Plant Cell,1995,7:957-970
67. Ohlrogge J B. Design of new plant products:Engineering of fatty acid metabolism. Plant Physiol, 1994,104:821-826
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