短季棉遗传多样性分析及其重要农艺和经济性状的QTL定位
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摘要
棉花是重要的经济作物，为纺织工业提供了大部分的天然纤维，是重要的纺织工业原料，在世界及我国国民经济中占有重要的战略地位。短季棉作为一种特殊的生态类型，在我国未来棉业可持续发展中有着不可替代的作用。利用分子标记可以对短季棉种质资源进行快速、准确的评价。本研究利用分子标记—RAPD评价了短季棉的遗传多样性，使育种工作者能更全面地了解早熟短季棉的亲缘关系和遗传基础，并且为指导早熟短季棉育种提供参考数据。
本研究利用RAPD标记对我国29份不同年代不同生态区的早熟短季棉品种进行了遗传多样性的比较分析。从600条RAPD引物中筛选出了122条带型清晰且重复性好的多态性引物，在29个短季早熟棉品种中共产生181个多态性位点。采用Jaccard's相似系数，使用NTSYS-pc 2.10数据分析软件，非加权组平均法(UPGMA)对数据进行聚类，对不同年代品种间平均相似系数进行了分析以及和系谱进行了比较分析，结果表明我国生产上推广的短季棉品种的遗传基础很窄，短季棉育种要取得新突破，必须收集与创新优良种质资源。
随着纺织技术的不断更新(环锭纺纱发展到气流转子纺纱到正在发展的喷气纺纱)，以及人民生活水平的提高，现有的棉花品种已经不能满足要求。因此，培育高产、优质的棉花品种成为当务之急。然而棉花的产量和纤维性状是复杂的数量性状，而且是负相关的；另外，陆地棉遗传基础狭窄，迫切需要引进海岛棉以及其它野生棉的优质基因。但是陆海种间杂交又存在连锁累赘等问题，传统育种很难打破性状间的连锁和同时兼顾产量和品质的改良。利用分子标记可以构建高密度遗传连锁图谱，可以对重要性状进行QTL定位，并利用与目标性状紧密连锁的标记进行辅助选择育种。
本研究利用多种类型(TRAP、SRAP、AFLP和SSR)的分子标记，以陆地棉×海岛棉F_2为作图群体，构建高密度的栽培棉的连锁图谱，并对纤维、产量、早熟相关性状进行了QTL定位分析。研究的主要结果如下：
1．以陆地棉“中棉所36”为母本，海岛棉品种“海7124”为父本配置杂交组合F_1，F_1自交后代的222个F_2单株(“永久”F_2群体)的宿根活体长期保存于海南三亚中国农业科学院棉花研究所野生棉种植园，选取186个F_2单株作为作图群体。
2．利用不同类型的分子标记对两个作图亲本进行多态性筛选，进而利用多态性引物分析186个F_2单株的基因型，共得到1250个分子标记。包括198个TRAPs，758个SSRs，143个AFLPs，151个SRAPs。同时考察了两个形态标记：黄色花药和基部红斑。将1252个标记位点进行X~2适合性检验，在5％的水平上共有249个(19.9％)标记位点不符合孟德尔分离比例，表现显著偏分离。SSR、TRAP、SRAP和AFLP的偏分离标记分别有124、38、24和27个，偏分离比例分别为19.2％、18.9％、15.9％和16.3％。
3．利用MapMaker／EXP.3.0对所得到的1252个标记进行连锁分析，最终有1097个标记进入35个连锁群，包括SSR 697个，TRAP 171个，SRAP 129个，AFLP 98个和两个形态标记。图谱总长4536.8 cM.，标记间平均距离4.1cM。根据锚定或公共的SSR标记确定连锁群，最后确定了和26条染色体对应的大连锁群和4个未知的小连锁群。每个连锁群的长度在3.5-235.1 cM，标记位点个数为2到63个不等，标记间平均距离为1.8-11.9 cM。其中最长的连锁群染色体9，长度为235.1 cM，含有56个标记。NL1-NL8这8个小的连锁群共有17个标记，长118.6 cM。染色体A亚组有590个标记，遗传距离2215.2 cM，平均距离3.8 cM；D染色体亚组有490个标记，共2203 cM，标记间平均距离4.1 cM。其中45个SSR重复位点建立了异源四倍体棉花13对同源染色体中的11对同源关系。构建到连锁群上的偏分离标记有198个，占图谱总标记的18.1％，其中SSR标记116个、TRAP标记42个、SRAP标记18个、AFLP标记22个，两个形态标记也是偏分离。大部分偏分离标记位点在连锁群上表现为明显的不同程度聚集现象，主要聚集在连锁群的中上部或中下部或末端。有几个可能是偏分离热点区。
4．本研究首次利用新型标记TRAP构建棉花的遗传连锁图谱。TRAP(Target Reglon Amplification polymorphism)是一种新型的基于PCR的标记。本研究根据与棉花纤维发育相关的cDNAs或ESTs设计了57条TRAP锚定引物，和SRAP的随机引物随机组合，其中93对引物组合共得到198个多态性位点，每个引物组合可产生1—7个不等的多态性位点，平均每个引物组合产生2.1个多态性位点。共显性标记有19个(9.6％)，偏分离标记有45个(22.7％)。最终，有171个TRAP(86.4％)标记进入所构建的连锁图谱上，除了三个小的连锁群上没有TRAP标记外，其它连锁群上都有TRAP标记，主要分布在染色体c2、c4、c5、c6、c9、c10、c14、c20、A02、A03等染色体上。
5．考查F_2单株纤维产量及品质性状以及F_(2：3)家系的产量(因为未收到足够的纤维，F_(2：3)没能考察纤维品质性状)和早熟性状(包括果枝始节和各个生育期)。利用软件QTL Cartographer V2.0，用复合区间作图法进行QTL检测，当LOD=3.0时，共检测出79个QTLs：纤维品质性状25个QTLs，能解释8.69％(qFU-c6-1)到23.19％(qFL-c1-2)表型变异；产量性状共检测出36个；F_2检测出16个QTLs，F_3共检测出20个QTLs，每个性状有1到6个QTLs不等。早熟农艺性状共检测到18个QTLs。对于同一性状，等位基因的增效作用既有来自母本方的，也有来自父本方的，即高值或低值亲本都可能存在对性状增效的等位基因。
6．与纤维品质QTLs相连锁的TRAP标记。8个TRAP标记分别和纤维长度、整齐度和马克隆值相关的QTLs连锁。有的TRAP标记和QTL紧密连锁，例如，染色体c1上的关于纤维长度的一个QTL(qFL-c1-2)和TRAP标记T16E2c紧密连锁(0.01cM内)，TRAP标记T16E2c的T16是根据参与棉花纤维发育相关的一个β-微管蛋白基因GhTub1设计的锚定引物而来，而GhTub1基因可能在纤维细胞延伸阶段的极性延伸中具有功能；而有的位于QTL区间内。例如与纤维整齐度相关的QTL(qFU-c17-1)，位于QTL的标记T9E10b-JESP195区间。T9是根据棉花纤维优势表达基因。GhCAP设计的，GhCAP基因是编码腺苷环化酶相关蛋白CAP，可能是细胞骨架改建中的信号。说明了利用TRAP标记不仅能增加图谱的饱和度，而且能增加目标QTL区域的标记饱和度，并且和性状相关联。
Cotton is one of the most important economic crops in the world; it provides the mostimportant material, natural fiber, for textile industry. So, cotton has an important strategicstatus in national economy. Short-season upland cotton plays an important role in thecultivation reform in china. We used molecular marker to detect the genetic diversityamong the short-season upland cotton varieties.
In this research, RAPDs analysis was used to assess the genetic diversity of 29 eliteshort-seasoned cotton cultivars. From 600 10-mer arbitrary primers, we selected 122primers that could produce legible and repetitive bands. 29 cotton varieties were screenedwith these 122 primers to generate 181 polymorphism loci. Cluster analysis and UPGMA(with NTSYS-PC2.10) showed that these 29 varieties could be classified into two groups.The correlation coefficients between species mostly range from 0.60 to 0.79; only 0.99percent was between 0.30 and 0.39. And this is agreed with the pedigree. In a word, thegenetic basis of popular short season upland cotton in China is narrow, and the Kings isthe main sources of earliness gene.
For cotton breeders, the major task is to improve the yield and the fiber quality, andduring the past decades of years, under the continuous efforts of the breeders, the yieldand fiber quality has been greatly improved. But, with the development of textile industryand increased consumption demands on cotton fiber commodity, most of the cottoncultivars are out of date, especially for cotton quantity. Consequently, it is eager to breedthe cultivars with high yield and good quality. However, cotton yield traits and qualitytraits are complicated quantitative traits; what is more, yield traits are negativelyassociated with quality trait, so it is difficult to improve yield and quantity simultaneity bytraditional breeding ways. The appearance of molecular markers and the development ofthe molecular quantitative genetics, make it possible to build high density linkage mapand QTL mapping, and then by using the linked markers, marker-assisted selection can beperformed during breeding process.
Based on a F_2 population from cross between Ghirsutum L.×G.barbadense L., acombined TRAP-SSR-AFLP-SRAP genetic map of tetraploid cotton was constructed andQTL mapping was performed. The main results were summarized as follows:
1. An interspecific F_2 population of 222 individuals between Ghirsutum L. cv CRI36×G.barbadense L. cv Hai7124 was planted in Hainan, from which 186 individuals werepicked out and used as mapping population.
2. Four kinds of molecular markers were used to identify polymorphism between twoparents. Finally, a total of 1250 polymorphic loci were obtained including 198TRAPs, 758 SSRs, 143 AFLPs and 151 SRAPs. Also, two morphologic markers,yellow pollen and red spot, were investigated. The 1252 polymorphic loci weresubsequently analyzed by X~2 test, at the 5% significant level, 249 loci deviated fromthe expected ratio (1:2:1 or 3:1), which included 124 SSRs (19.2%), 38 TRAPs(18.9%), 24 SRAPs (15.9) and 27 AFLPs (16.3%).
3. 1252(A thousand two hundred and fifty-two) marker loci were used to constructgenetic linkage groups using MapMaker/EXP. 3.0. The map consisted of 1097markers mapped into 35 linkage groups, including 697 SSRs, 171 TRAPs, 129SRAPs, 98 AFLPs, and two morphological markers, and spanned 4,536.7 cM with anaverage genetic distance of 4.1 cM per marker. According to the assigned SSRs orcommon loci, linkage groups were assigned to the 26 chromosomes of the tertraploidgenome and eight novel small linkage groups. The distance of single linkage groupwas from 106.5 cM to 235.1 cM, the loci number in every group was from 2 to 63,and the average distance between each locus was from 1.8 cM to 11.9 cM. The 13A-genome groups span 2,215.2 cM containing 590 makers with an average distanceof 3.8 cM, the 13 D-genome groups span 2,203 cM with 490 markers with an averagedistance of 4.1 cM. In this research, 11 of 13 pairs of homoeologous chromosomeswere bridged with a total of 45 SSR duplicated loci, except for c2-c14 and c3-c17.200 distorted segregation loci (116 SSRs, 42 TRAPs, 18 SRAPs, 22 AFLPs and 2morphological markers) were mapped in the 26 groups of allotetroploid cotton. At thepresent map, skewed markers were clustered in different degrees, which were usuallyobserved at the end or in the middle of the group. And maybe, some of them wereSegregation Distortion Regions(SDR).
4. This is the first time that the novel molecular marker, TRAP (Target RegionAmplification polymorphism), was used to construct genetic map in cotton. A total of57 fixed primers were designed on EST sequences using Primer Premier 5.0.Combined with the random primers of SRAP, 198 polymorphic loci were observedby using 93 primer pairs. Every primer combination could generate 1-7 polymorphicloci, with an average 2.1 polymorphic loci per primer pair. 19 of the 198 loci wereco-dominant loci, and 45 were distorted segregation loci. Totally, 171 loci weremapped on 26 linkage groups. Except for three short linkage groups (NL6, NL7, NL8), it could be found in the remained groups, and most distributed on the c2, c4, c5,c6, c9, c10, c14, c20, A02, A03.
5. The fiber yield, fiber quality, and some traits related to short-season of F_2 plants andcorresponding F_(2:3) families were investigated; but the fiber quality trait of F_(2:3) was notobtained because not enough fiber was harvest. QTLs were scanned by the method ofcomposite interval mapping (CIM) in program QTL Cartographer V2.0. When the logodds-ration threshold of QTL detection was 3.0, 79 QTLs could be detected,including 25 QTLs related to fiber quality traits explained from 8.69% (qFU-c6-1)to 23.19% (qFL-c1-2) of phenotypic variation (PV); 36 QTLs related to fiber yield(16 QTLs in F_2 plants, 20 QTLs in F_3 plants); and 18 QTLs related to short-season.
6. 8 TRAP markers linked to the QTLs of fiber length, fiber uniformity ratio andmacronaire value, were detected. For example, T16E2c was linked to a QTL(qFL-c1-2) in chromosome 1 (within 0.01cM), and a QTL (qFU-c17-1) related tofiber uniformity ratio was positioned at the interval of T9E10b-JESP195. And thesetwo primers were designed from the genes related to fiber extension and fiberformation respectively. So, TRAP marker is not only benefiting for increasing thegenetic map density, but also it is easy to link with the related QTLs.
本研究利用RAPD标记对我国29份不同年代不同生态区的早熟短季棉品种进行了遗传多样性的比较分析。从600条RAPD引物中筛选出了122条带型清晰且重复性好的多态性引物，在29个短季早熟棉品种中共产生181个多态性位点。采用Jaccard's相似系数，使用NTSYS-pc 2.10数据分析软件，非加权组平均法(UPGMA)对数据进行聚类，对不同年代品种间平均相似系数进行了分析以及和系谱进行了比较分析，结果表明我国生产上推广的短季棉品种的遗传基础很窄，短季棉育种要取得新突破，必须收集与创新优良种质资源。
随着纺织技术的不断更新(环锭纺纱发展到气流转子纺纱到正在发展的喷气纺纱)，以及人民生活水平的提高，现有的棉花品种已经不能满足要求。因此，培育高产、优质的棉花品种成为当务之急。然而棉花的产量和纤维性状是复杂的数量性状，而且是负相关的；另外，陆地棉遗传基础狭窄，迫切需要引进海岛棉以及其它野生棉的优质基因。但是陆海种间杂交又存在连锁累赘等问题，传统育种很难打破性状间的连锁和同时兼顾产量和品质的改良。利用分子标记可以构建高密度遗传连锁图谱，可以对重要性状进行QTL定位，并利用与目标性状紧密连锁的标记进行辅助选择育种。
本研究利用多种类型(TRAP、SRAP、AFLP和SSR)的分子标记，以陆地棉×海岛棉F_2为作图群体，构建高密度的栽培棉的连锁图谱，并对纤维、产量、早熟相关性状进行了QTL定位分析。研究的主要结果如下：
1．以陆地棉“中棉所36”为母本，海岛棉品种“海7124”为父本配置杂交组合F_1，F_1自交后代的222个F_2单株(“永久”F_2群体)的宿根活体长期保存于海南三亚中国农业科学院棉花研究所野生棉种植园，选取186个F_2单株作为作图群体。
2．利用不同类型的分子标记对两个作图亲本进行多态性筛选，进而利用多态性引物分析186个F_2单株的基因型，共得到1250个分子标记。包括198个TRAPs，758个SSRs，143个AFLPs，151个SRAPs。同时考察了两个形态标记：黄色花药和基部红斑。将1252个标记位点进行X~2适合性检验，在5％的水平上共有249个(19.9％)标记位点不符合孟德尔分离比例，表现显著偏分离。SSR、TRAP、SRAP和AFLP的偏分离标记分别有124、38、24和27个，偏分离比例分别为19.2％、18.9％、15.9％和16.3％。
3．利用MapMaker／EXP.3.0对所得到的1252个标记进行连锁分析，最终有1097个标记进入35个连锁群，包括SSR 697个，TRAP 171个，SRAP 129个，AFLP 98个和两个形态标记。图谱总长4536.8 cM.，标记间平均距离4.1cM。根据锚定或公共的SSR标记确定连锁群，最后确定了和26条染色体对应的大连锁群和4个未知的小连锁群。每个连锁群的长度在3.5-235.1 cM，标记位点个数为2到63个不等，标记间平均距离为1.8-11.9 cM。其中最长的连锁群染色体9，长度为235.1 cM，含有56个标记。NL1-NL8这8个小的连锁群共有17个标记，长118.6 cM。染色体A亚组有590个标记，遗传距离2215.2 cM，平均距离3.8 cM；D染色体亚组有490个标记，共2203 cM，标记间平均距离4.1 cM。其中45个SSR重复位点建立了异源四倍体棉花13对同源染色体中的11对同源关系。构建到连锁群上的偏分离标记有198个，占图谱总标记的18.1％，其中SSR标记116个、TRAP标记42个、SRAP标记18个、AFLP标记22个，两个形态标记也是偏分离。大部分偏分离标记位点在连锁群上表现为明显的不同程度聚集现象，主要聚集在连锁群的中上部或中下部或末端。有几个可能是偏分离热点区。
4．本研究首次利用新型标记TRAP构建棉花的遗传连锁图谱。TRAP(Target Reglon Amplification polymorphism)是一种新型的基于PCR的标记。本研究根据与棉花纤维发育相关的cDNAs或ESTs设计了57条TRAP锚定引物，和SRAP的随机引物随机组合，其中93对引物组合共得到198个多态性位点，每个引物组合可产生1—7个不等的多态性位点，平均每个引物组合产生2.1个多态性位点。共显性标记有19个(9.6％)，偏分离标记有45个(22.7％)。最终，有171个TRAP(86.4％)标记进入所构建的连锁图谱上，除了三个小的连锁群上没有TRAP标记外，其它连锁群上都有TRAP标记，主要分布在染色体c2、c4、c5、c6、c9、c10、c14、c20、A02、A03等染色体上。
5．考查F_2单株纤维产量及品质性状以及F_(2：3)家系的产量(因为未收到足够的纤维，F_(2：3)没能考察纤维品质性状)和早熟性状(包括果枝始节和各个生育期)。利用软件QTL Cartographer V2.0，用复合区间作图法进行QTL检测，当LOD=3.0时，共检测出79个QTLs：纤维品质性状25个QTLs，能解释8.69％(qFU-c6-1)到23.19％(qFL-c1-2)表型变异；产量性状共检测出36个；F_2检测出16个QTLs，F_3共检测出20个QTLs，每个性状有1到6个QTLs不等。早熟农艺性状共检测到18个QTLs。对于同一性状，等位基因的增效作用既有来自母本方的，也有来自父本方的，即高值或低值亲本都可能存在对性状增效的等位基因。
6．与纤维品质QTLs相连锁的TRAP标记。8个TRAP标记分别和纤维长度、整齐度和马克隆值相关的QTLs连锁。有的TRAP标记和QTL紧密连锁，例如，染色体c1上的关于纤维长度的一个QTL(qFL-c1-2)和TRAP标记T16E2c紧密连锁(0.01cM内)，TRAP标记T16E2c的T16是根据参与棉花纤维发育相关的一个β-微管蛋白基因GhTub1设计的锚定引物而来，而GhTub1基因可能在纤维细胞延伸阶段的极性延伸中具有功能；而有的位于QTL区间内。例如与纤维整齐度相关的QTL(qFU-c17-1)，位于QTL的标记T9E10b-JESP195区间。T9是根据棉花纤维优势表达基因。GhCAP设计的，GhCAP基因是编码腺苷环化酶相关蛋白CAP，可能是细胞骨架改建中的信号。说明了利用TRAP标记不仅能增加图谱的饱和度，而且能增加目标QTL区域的标记饱和度，并且和性状相关联。
Cotton is one of the most important economic crops in the world; it provides the mostimportant material, natural fiber, for textile industry. So, cotton has an important strategicstatus in national economy. Short-season upland cotton plays an important role in thecultivation reform in china. We used molecular marker to detect the genetic diversityamong the short-season upland cotton varieties.
In this research, RAPDs analysis was used to assess the genetic diversity of 29 eliteshort-seasoned cotton cultivars. From 600 10-mer arbitrary primers, we selected 122primers that could produce legible and repetitive bands. 29 cotton varieties were screenedwith these 122 primers to generate 181 polymorphism loci. Cluster analysis and UPGMA(with NTSYS-PC2.10) showed that these 29 varieties could be classified into two groups.The correlation coefficients between species mostly range from 0.60 to 0.79; only 0.99percent was between 0.30 and 0.39. And this is agreed with the pedigree. In a word, thegenetic basis of popular short season upland cotton in China is narrow, and the Kings isthe main sources of earliness gene.
For cotton breeders, the major task is to improve the yield and the fiber quality, andduring the past decades of years, under the continuous efforts of the breeders, the yieldand fiber quality has been greatly improved. But, with the development of textile industryand increased consumption demands on cotton fiber commodity, most of the cottoncultivars are out of date, especially for cotton quantity. Consequently, it is eager to breedthe cultivars with high yield and good quality. However, cotton yield traits and qualitytraits are complicated quantitative traits; what is more, yield traits are negativelyassociated with quality trait, so it is difficult to improve yield and quantity simultaneity bytraditional breeding ways. The appearance of molecular markers and the development ofthe molecular quantitative genetics, make it possible to build high density linkage mapand QTL mapping, and then by using the linked markers, marker-assisted selection can beperformed during breeding process.
Based on a F_2 population from cross between Ghirsutum L.×G.barbadense L., acombined TRAP-SSR-AFLP-SRAP genetic map of tetraploid cotton was constructed andQTL mapping was performed. The main results were summarized as follows:
1. An interspecific F_2 population of 222 individuals between Ghirsutum L. cv CRI36×G.barbadense L. cv Hai7124 was planted in Hainan, from which 186 individuals werepicked out and used as mapping population.
2. Four kinds of molecular markers were used to identify polymorphism between twoparents. Finally, a total of 1250 polymorphic loci were obtained including 198TRAPs, 758 SSRs, 143 AFLPs and 151 SRAPs. Also, two morphologic markers,yellow pollen and red spot, were investigated. The 1252 polymorphic loci weresubsequently analyzed by X~2 test, at the 5% significant level, 249 loci deviated fromthe expected ratio (1:2:1 or 3:1), which included 124 SSRs (19.2%), 38 TRAPs(18.9%), 24 SRAPs (15.9) and 27 AFLPs (16.3%).
3. 1252(A thousand two hundred and fifty-two) marker loci were used to constructgenetic linkage groups using MapMaker/EXP. 3.0. The map consisted of 1097markers mapped into 35 linkage groups, including 697 SSRs, 171 TRAPs, 129SRAPs, 98 AFLPs, and two morphological markers, and spanned 4,536.7 cM with anaverage genetic distance of 4.1 cM per marker. According to the assigned SSRs orcommon loci, linkage groups were assigned to the 26 chromosomes of the tertraploidgenome and eight novel small linkage groups. The distance of single linkage groupwas from 106.5 cM to 235.1 cM, the loci number in every group was from 2 to 63,and the average distance between each locus was from 1.8 cM to 11.9 cM. The 13A-genome groups span 2,215.2 cM containing 590 makers with an average distanceof 3.8 cM, the 13 D-genome groups span 2,203 cM with 490 markers with an averagedistance of 4.1 cM. In this research, 11 of 13 pairs of homoeologous chromosomeswere bridged with a total of 45 SSR duplicated loci, except for c2-c14 and c3-c17.200 distorted segregation loci (116 SSRs, 42 TRAPs, 18 SRAPs, 22 AFLPs and 2morphological markers) were mapped in the 26 groups of allotetroploid cotton. At thepresent map, skewed markers were clustered in different degrees, which were usuallyobserved at the end or in the middle of the group. And maybe, some of them wereSegregation Distortion Regions(SDR).
4. This is the first time that the novel molecular marker, TRAP (Target RegionAmplification polymorphism), was used to construct genetic map in cotton. A total of57 fixed primers were designed on EST sequences using Primer Premier 5.0.Combined with the random primers of SRAP, 198 polymorphic loci were observedby using 93 primer pairs. Every primer combination could generate 1-7 polymorphicloci, with an average 2.1 polymorphic loci per primer pair. 19 of the 198 loci wereco-dominant loci, and 45 were distorted segregation loci. Totally, 171 loci weremapped on 26 linkage groups. Except for three short linkage groups (NL6, NL7, NL8), it could be found in the remained groups, and most distributed on the c2, c4, c5,c6, c9, c10, c14, c20, A02, A03.
5. The fiber yield, fiber quality, and some traits related to short-season of F_2 plants andcorresponding F_(2:3) families were investigated; but the fiber quality trait of F_(2:3) was notobtained because not enough fiber was harvest. QTLs were scanned by the method ofcomposite interval mapping (CIM) in program QTL Cartographer V2.0. When the logodds-ration threshold of QTL detection was 3.0, 79 QTLs could be detected,including 25 QTLs related to fiber quality traits explained from 8.69% (qFU-c6-1)to 23.19% (qFL-c1-2) of phenotypic variation (PV); 36 QTLs related to fiber yield(16 QTLs in F_2 plants, 20 QTLs in F_3 plants); and 18 QTLs related to short-season.
6. 8 TRAP markers linked to the QTLs of fiber length, fiber uniformity ratio andmacronaire value, were detected. For example, T16E2c was linked to a QTL(qFL-c1-2) in chromosome 1 (within 0.01cM), and a QTL (qFU-c17-1) related tofiber uniformity ratio was positioned at the interval of T9E10b-JESP195. And thesetwo primers were designed from the genes related to fiber extension and fiberformation respectively. So, TRAP marker is not only benefiting for increasing thegenetic map density, but also it is easy to link with the related QTLs.
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