陆地棉染色体片段代换系的构建及QTL定位初探
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摘要
海岛棉具有纤维品质好,纤维长、强、细,抗黄萎病等优点,如何有效地将存在于海岛棉中的优异基因转移到栽培的陆地棉中,是一项具有重要意义的工作。本研利用生产上大面积推广种植的陆地棉中221和品质优良的海岛棉栽培品系海1杂交,并与陆地棉高代回交,结合分子标记辅助选择建立一套具有陆地棉背景并渐渗有海岛棉基因的染色体片段代换系。并利用BC4F1、BC4F2家系和BC4F2单株的产量及纤维品质性状数据及SSR标记检测结果,采用复合区间作图法(CIM)对纤维品质性状和产量性状进行了QTL定位研究。通过对染色体渐渗片段的分析,为进一步筛选染色体单片段代换系、优质纤维品质基因的精细定位、基因克隆和基因聚合等研究准备材料。
1.性状表型分析
通过两个优良品种的杂交并回交,培育了一个BC4F1、BC4F2家系群体和鉴定了一个BC4F2单株群体,获得了大量在纤维品质和产量上超过轮回亲本的材料。与轮回亲本相比较,BC4F1、BC4F2家系群体和BC4F2单株群体的纤维品质性状各指标的平均值与轮回亲本中221接近,其中在纤维长度和纤维强度上略高于中221,但各性状超亲比率很高。在纤维长度上,3个群体的超亲比例均高于50%,其中,BC4F1家系的超亲比例达到了91.38%。在纤维强度上,超亲比例均超过60%,BC4F1家系最大,为72.41%。BC4F1、BC4F2家系和BC4F2单株的马克隆值最小值相对中221下调0.9～2.1,超亲比例(指马克隆值大于3.7小于中221马克隆值的比例)分别为57.76%、72.28%和67.69%。衣分在3个群体中的超亲比例都超过了50%,最高的达到93.1%。铃重和铃数的超亲比例除BC4F2家系铃重的超亲比例较低,只有2.5%外,其它集中在17.24～48.46%之间。
2.家系海岛棉染色体片段代换情况
筛选了覆盖棉花基因组2347.0 cM的276个SSR标记,检测家系的染色体片段渐渗情况。导入的海岛棉片段覆盖棉花基因组最长的达到969.34cM,覆盖检测长度2347.0cM的41.3%；最小的覆盖7.7%,覆盖长度为180.7cM;平均覆盖基因组502cM,占检测长度的21.4%。每个家系中代换片段最多的55个,最少的也有15个,平均32.9个,回交4代后家系中任然保留了大量的海岛棉染色体片段。各连锁群在每个BC4F1家系中平均包含0.1～8.3个海岛棉片段,平均覆盖每个连锁群0.2-54.2cM,占每条连锁群被检测长度0.5～207cM的10.3%～35.5%。
3.单株海岛棉代换情况的检测
208个BC4F2单株检测结果,回到轮回亲本背景的比例高达89%-100%,平均为96.7%,接近其理论值96.9%。有3株含纯合的海岛棉单片段,16株含杂合的单片段,共19株只含有一个海岛棉的片段；18株含2个片段；其它均含3个片段以上,但主要在3～8个片段,最多的含21个片段,单株导入的总片段平均覆盖棉花基因组的3.02%,最大的覆盖11.3%。每株中海岛棉杂合片段平均总长69.4cM,最长的294.4 cM,最短只有2.3cM,覆盖比率为0.1～11.0%,平均为2.5%。导入的纯合片段平均为13 cM,最短的1cM,最长的97 cM,覆盖比率为0.1～3.6%,平均为0.5%。各连锁群含海岛棉杂合片段平均2.7cM,占各连锁群检测区域的2.7%；含纯合片段平均0.5cM,占0.5%,合计3.2cM,为3.2%。
4.品质优良的单株海岛棉导入情况
对选出的5个家系后代中纤维长度和纤维强度优良的12个单株海岛棉渐渗片段为1-8个；控制纤维长度和纤维强度的QTLqFL-18-5、qFL-18-18、qFS-18-5和qFS-16-1(其增效基因全部来自于海岛棉)邻近标记NAU3447b、BNL1438和BNL3932存在于这些海岛棉代换片段上,说明这些家系和单株中可能导入了较多的海岛棉优良的品质基因,为进一步获得含有优良品质性状基因的染色体单片段片段代换系、基因克隆和基因聚合等奠定了基础。
5.QTL分析
共检测到51个QTL,单个QTL解释表型变异6.86%～39.60%。其中控制纤维品质相关性状的有28个,控制产量及抗黄萎病相关性状的有23个。有25个QTL的增效基因来自陆地棉；26个QTL增效基因来自海岛棉。所有QTL分布在15条染色体上,发现10组(25个)QTL在染色体上的特定位置成簇分布。3个QTL(qFS-24-4、qBW-5-9和qUN-3-9)在两个群体中被检测到,可进一步用于标记辅助选择。
The fiber quality of G. bardadense cultivars is generally superior with longer, stronger, and finer fiber than G. hirsutum. It is important to introgress fiber quality genes from G. bardadense to G. hirsutum. In the study, a cotton population with genetic background of G.hirsutum and some chromosome segments of G. barbadense were developed from advanced backcrosses between the recipient parent, CCRI221, a commercial cotton cultivar, and donor parent, hail,a barbadense cotton line by microsatellite marker-assisted selection (MAS). QTL analysis of yield-related trait and fiber quality using the phenotype and the genotype data in BC4F1, BC4F2 lines and BC4F2 individuals were proceeded by the Composite Interval Mapping method, which will provide a number of germplasm resources for further developing single segment substitution lines,locating accurately QTL,cloning genes and genes pyramiding.
1. Analysis of trait phenotype
We got a number of better fiber quality and higher yield CSSLs than their parents through intercross advanced backcrosses between two superior cultivars. Means of fiber quality in BC4F1, BC4F2 lines and BC4F2 individuals is near to that of CCRI 221 and means of fiber length and strength is a little better than that of CCRI 221. The rate of lines and individuals with longer fiber than CCRI 221 in the three populations is over 50%, specially, the rate is 91.38% in BC4F1. More than 60% of lines and individuals in each population are better than CCRI 221 for fiber strength, and specially, the rate is 72.41% in BC4F1. The minimum of fiber micronaire of BC4F1, BC4F2 lines and BC4F2 individuals is lower with decreasing 0.9-2.1 than that of CCRI 221,and the rate of lines and individuals with lower micronaire than CCRI221 is 57.76%, 72.28% and 67.69%. The rate of lines and individuals with higher lint percentage than CCRI 221 in the three populations is 50%-93.1%. The rate of better lines and individuals in boll weight and number of boll per plant than CCRI 221 in the three populations is 17.24～48.46%, but the rate of boll weight in BC4F2 lines is 2.5%.
2. Chromosome segment substitution of G. barbadense in lines
A total of 450 pairs of SSR primers were, selected with a marker erery 5 cM from 25 linkage groups developed by molecule breeding lab of cotton research institute of Chinese Academy of Agricultural Science to screen polymorphism between CCRI 221 and Hail, which resulted in 276 pairs of polymorphic primers, covered a total genetic distance of 2347.0 cM, approximately 52.7% of cotton genome. And maximum distance of the neighboring markers was 47.51 cM, the minimum distance was 1.2 cM, the average distance of neighboring markers was 7.4 cM.116 BC4F1 lines were screened using the 276 pairs of SSR primers. There are 15 to 55 donor segments of G barbadense in the lines with average of 32.92,which cover 180.7 cM to 969 cM of cotton genome. Average size of cotton genome covered by substitution seg-ments is 502 cM, covered 21.4% of cotton genome detected. Average donor segments are 0.1-8.3 in each linkage group in every BC4F1 line, which covered from 0.2 cM to 54.2 cM, with 10.3-35.5% of the average distance screened of each linkage group in every lines.
3. chromosome segment substitution of G. barbadense in the individuals
208 BC4F2 individuals, originated from 5 lines of BC4F1 (about 40 individuals in every line), were screened using the primers with polymorphism in the related line. The rate of chromosome of the recurrent parent, CCRI 221, was high, about 89%-100%, the average is 96.7%, near to theoretical value 96.9%. There are three individuals with a homozygous chromosome segment of G. barbadense,16 individuals with a heterozygous chromosome segment of G barbadense, and 18 individuals with only two chromosome segments of G. barbadense, and the others with more than 3 chromosome segments of G. barbadense, mainly about 3-8 chromosome segments, and maximum of 21 chromosome segment of G barbadense, which averaged coverage of 3.02% of cotton genome with the maximum of 11.3%. the average length of heterozygous chromosome segment of G. barbadense in every individual is 69.4 cM, ranged from 2.3 cM to 294.4 cM, which cover 0.1-11.0% of cotton genome, the average rate is 2.5%. The average length of homozygous chromosome segment of G. barbadense in every individual is 13 cM, ranged from 1 cM to 97 cM, which cover 0.1-3.6% of cotton genome with average of 0.5. The average length of heterozygous chromosome segment of G barbadense in every linkage group is 2.7 cM, covered 2.7% of every linkage group region screened. The average length of homozygous chromosome segment of G barbadense in every linkage group is 0.5 cM about 0.5% of cotton genome.The total of all chromosome segment substitution of G. barbadense is 3.2 cM, about 3.2% of cotton genome.
4. Analysis of QTL
51 QTL were mapped in BC4F1, BC4F2 lines and BC4F2 individuals by Composite Interval Mapping method with the software of Windows QTL Cartographer 2.5, which accounted for 6.86%-39.60% of phenotypic variance. There were 28 QTL for fiber quality traits and 23 QTL for yield-related and resistance to verticillium wilt. There were 26 QTL with positive additive effect, which indicated the additive genes from G. barbadense.10 groups of QTL were mapped assembly in special region of chromosome.3 QTL (qFS-24-4,qBW-5-9 and qUN-3-9) were found in two populations, which should be used for marker assisted selection.
5. Chromosome segment substitution of G. barbadense in 12 individuals with superior fiber quality
There were 1 to 8 chromosome substitution segments of G. barbadense in 12 BC4F2 individuals with longer and stronger fiber than CCRI 221. And there were 3 makers of NAU3447b,BNL1438 and BNL3932, which linked to the QTL qFL-18-5 of qFL-18-18, qFS-18-5 and qFS-16-1 for fiber length and strength in these chromosome segment, which indicated that we have introgressed fiber quality genes from G. bardadense to G. hirsutum. This will provide a number of germplasm resources for developing single segment substitution lines, accurately locating QTL, cloning and pyramiding genes.
1.性状表型分析
通过两个优良品种的杂交并回交,培育了一个BC4F1、BC4F2家系群体和鉴定了一个BC4F2单株群体,获得了大量在纤维品质和产量上超过轮回亲本的材料。与轮回亲本相比较,BC4F1、BC4F2家系群体和BC4F2单株群体的纤维品质性状各指标的平均值与轮回亲本中221接近,其中在纤维长度和纤维强度上略高于中221,但各性状超亲比率很高。在纤维长度上,3个群体的超亲比例均高于50%,其中,BC4F1家系的超亲比例达到了91.38%。在纤维强度上,超亲比例均超过60%,BC4F1家系最大,为72.41%。BC4F1、BC4F2家系和BC4F2单株的马克隆值最小值相对中221下调0.9～2.1,超亲比例(指马克隆值大于3.7小于中221马克隆值的比例)分别为57.76%、72.28%和67.69%。衣分在3个群体中的超亲比例都超过了50%,最高的达到93.1%。铃重和铃数的超亲比例除BC4F2家系铃重的超亲比例较低,只有2.5%外,其它集中在17.24～48.46%之间。
2.家系海岛棉染色体片段代换情况
筛选了覆盖棉花基因组2347.0 cM的276个SSR标记,检测家系的染色体片段渐渗情况。导入的海岛棉片段覆盖棉花基因组最长的达到969.34cM,覆盖检测长度2347.0cM的41.3%；最小的覆盖7.7%,覆盖长度为180.7cM;平均覆盖基因组502cM,占检测长度的21.4%。每个家系中代换片段最多的55个,最少的也有15个,平均32.9个,回交4代后家系中任然保留了大量的海岛棉染色体片段。各连锁群在每个BC4F1家系中平均包含0.1～8.3个海岛棉片段,平均覆盖每个连锁群0.2-54.2cM,占每条连锁群被检测长度0.5～207cM的10.3%～35.5%。
3.单株海岛棉代换情况的检测
208个BC4F2单株检测结果,回到轮回亲本背景的比例高达89%-100%,平均为96.7%,接近其理论值96.9%。有3株含纯合的海岛棉单片段,16株含杂合的单片段,共19株只含有一个海岛棉的片段；18株含2个片段；其它均含3个片段以上,但主要在3～8个片段,最多的含21个片段,单株导入的总片段平均覆盖棉花基因组的3.02%,最大的覆盖11.3%。每株中海岛棉杂合片段平均总长69.4cM,最长的294.4 cM,最短只有2.3cM,覆盖比率为0.1～11.0%,平均为2.5%。导入的纯合片段平均为13 cM,最短的1cM,最长的97 cM,覆盖比率为0.1～3.6%,平均为0.5%。各连锁群含海岛棉杂合片段平均2.7cM,占各连锁群检测区域的2.7%；含纯合片段平均0.5cM,占0.5%,合计3.2cM,为3.2%。
4.品质优良的单株海岛棉导入情况
对选出的5个家系后代中纤维长度和纤维强度优良的12个单株海岛棉渐渗片段为1-8个；控制纤维长度和纤维强度的QTLqFL-18-5、qFL-18-18、qFS-18-5和qFS-16-1(其增效基因全部来自于海岛棉)邻近标记NAU3447b、BNL1438和BNL3932存在于这些海岛棉代换片段上,说明这些家系和单株中可能导入了较多的海岛棉优良的品质基因,为进一步获得含有优良品质性状基因的染色体单片段片段代换系、基因克隆和基因聚合等奠定了基础。
5.QTL分析
共检测到51个QTL,单个QTL解释表型变异6.86%～39.60%。其中控制纤维品质相关性状的有28个,控制产量及抗黄萎病相关性状的有23个。有25个QTL的增效基因来自陆地棉；26个QTL增效基因来自海岛棉。所有QTL分布在15条染色体上,发现10组(25个)QTL在染色体上的特定位置成簇分布。3个QTL(qFS-24-4、qBW-5-9和qUN-3-9)在两个群体中被检测到,可进一步用于标记辅助选择。
The fiber quality of G. bardadense cultivars is generally superior with longer, stronger, and finer fiber than G. hirsutum. It is important to introgress fiber quality genes from G. bardadense to G. hirsutum. In the study, a cotton population with genetic background of G.hirsutum and some chromosome segments of G. barbadense were developed from advanced backcrosses between the recipient parent, CCRI221, a commercial cotton cultivar, and donor parent, hail,a barbadense cotton line by microsatellite marker-assisted selection (MAS). QTL analysis of yield-related trait and fiber quality using the phenotype and the genotype data in BC4F1, BC4F2 lines and BC4F2 individuals were proceeded by the Composite Interval Mapping method, which will provide a number of germplasm resources for further developing single segment substitution lines,locating accurately QTL,cloning genes and genes pyramiding.
1. Analysis of trait phenotype
We got a number of better fiber quality and higher yield CSSLs than their parents through intercross advanced backcrosses between two superior cultivars. Means of fiber quality in BC4F1, BC4F2 lines and BC4F2 individuals is near to that of CCRI 221 and means of fiber length and strength is a little better than that of CCRI 221. The rate of lines and individuals with longer fiber than CCRI 221 in the three populations is over 50%, specially, the rate is 91.38% in BC4F1. More than 60% of lines and individuals in each population are better than CCRI 221 for fiber strength, and specially, the rate is 72.41% in BC4F1. The minimum of fiber micronaire of BC4F1, BC4F2 lines and BC4F2 individuals is lower with decreasing 0.9-2.1 than that of CCRI 221,and the rate of lines and individuals with lower micronaire than CCRI221 is 57.76%, 72.28% and 67.69%. The rate of lines and individuals with higher lint percentage than CCRI 221 in the three populations is 50%-93.1%. The rate of better lines and individuals in boll weight and number of boll per plant than CCRI 221 in the three populations is 17.24～48.46%, but the rate of boll weight in BC4F2 lines is 2.5%.
2. Chromosome segment substitution of G. barbadense in lines
A total of 450 pairs of SSR primers were, selected with a marker erery 5 cM from 25 linkage groups developed by molecule breeding lab of cotton research institute of Chinese Academy of Agricultural Science to screen polymorphism between CCRI 221 and Hail, which resulted in 276 pairs of polymorphic primers, covered a total genetic distance of 2347.0 cM, approximately 52.7% of cotton genome. And maximum distance of the neighboring markers was 47.51 cM, the minimum distance was 1.2 cM, the average distance of neighboring markers was 7.4 cM.116 BC4F1 lines were screened using the 276 pairs of SSR primers. There are 15 to 55 donor segments of G barbadense in the lines with average of 32.92,which cover 180.7 cM to 969 cM of cotton genome. Average size of cotton genome covered by substitution seg-ments is 502 cM, covered 21.4% of cotton genome detected. Average donor segments are 0.1-8.3 in each linkage group in every BC4F1 line, which covered from 0.2 cM to 54.2 cM, with 10.3-35.5% of the average distance screened of each linkage group in every lines.
3. chromosome segment substitution of G. barbadense in the individuals
208 BC4F2 individuals, originated from 5 lines of BC4F1 (about 40 individuals in every line), were screened using the primers with polymorphism in the related line. The rate of chromosome of the recurrent parent, CCRI 221, was high, about 89%-100%, the average is 96.7%, near to theoretical value 96.9%. There are three individuals with a homozygous chromosome segment of G. barbadense,16 individuals with a heterozygous chromosome segment of G barbadense, and 18 individuals with only two chromosome segments of G. barbadense, and the others with more than 3 chromosome segments of G. barbadense, mainly about 3-8 chromosome segments, and maximum of 21 chromosome segment of G barbadense, which averaged coverage of 3.02% of cotton genome with the maximum of 11.3%. the average length of heterozygous chromosome segment of G. barbadense in every individual is 69.4 cM, ranged from 2.3 cM to 294.4 cM, which cover 0.1-11.0% of cotton genome, the average rate is 2.5%. The average length of homozygous chromosome segment of G. barbadense in every individual is 13 cM, ranged from 1 cM to 97 cM, which cover 0.1-3.6% of cotton genome with average of 0.5. The average length of heterozygous chromosome segment of G barbadense in every linkage group is 2.7 cM, covered 2.7% of every linkage group region screened. The average length of homozygous chromosome segment of G barbadense in every linkage group is 0.5 cM about 0.5% of cotton genome.The total of all chromosome segment substitution of G. barbadense is 3.2 cM, about 3.2% of cotton genome.
4. Analysis of QTL
51 QTL were mapped in BC4F1, BC4F2 lines and BC4F2 individuals by Composite Interval Mapping method with the software of Windows QTL Cartographer 2.5, which accounted for 6.86%-39.60% of phenotypic variance. There were 28 QTL for fiber quality traits and 23 QTL for yield-related and resistance to verticillium wilt. There were 26 QTL with positive additive effect, which indicated the additive genes from G. barbadense.10 groups of QTL were mapped assembly in special region of chromosome.3 QTL (qFS-24-4,qBW-5-9 and qUN-3-9) were found in two populations, which should be used for marker assisted selection.
5. Chromosome segment substitution of G. barbadense in 12 individuals with superior fiber quality
There were 1 to 8 chromosome substitution segments of G. barbadense in 12 BC4F2 individuals with longer and stronger fiber than CCRI 221. And there were 3 makers of NAU3447b,BNL1438 and BNL3932, which linked to the QTL qFL-18-5 of qFL-18-18, qFS-18-5 and qFS-16-1 for fiber length and strength in these chromosome segment, which indicated that we have introgressed fiber quality genes from G. bardadense to G. hirsutum. This will provide a number of germplasm resources for developing single segment substitution lines, accurately locating QTL, cloning and pyramiding genes.
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