陆地棉TM-1背景的海岛棉染色体片段导入系的培育鉴定和纤维强度QTL精细定位
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摘要
棉花是世界性的重要经济作物。中国是世界上最大的棉花生产,消费和纺织国,随着我国人民生活水平的飞速提高和纺织品配额的取消,纺织品消费和出口必将快速提高进而对棉花的需求越来越大,品质要求也越来越高。
陆地棉(Gossypium hirsutum L.)和海岛棉(G. barbadense L.)是棉花的2个四倍体栽培种,陆地棉产量高,纤维品质中等,海岛棉产量低但纤维细强,可纺100-200支纱,是纺高支纱的原料。遗传基础狭窄已经成为目前限制陆地棉遗传改良的主要因素。在陆地棉和海岛棉间进行优良基因的高效转移渗透对二者的遗传改良具有极为重要的现实意义,而限制这一过程的主要因素是连锁累赘。染色体片段导入系了除目标基因所在座位的局部区域外,基因组其余部分都是相同的,在这样的材料间找到的多态性标记才可能与目标基因紧密连锁,是QTL精细定位和图位克隆的一类理想群体。本研究以海陆杂交群体为基础,通过回交和标记辅助选择构建了海岛棉的染色体片段导入系,并利用这过程产生的高代回交群体和片段导入系对纤维品质等性状进行了QTL标记定位研究。同时还研究了QTL的精细定位。研究的主要结果如下：
1.陆地棉遗传标准系TM-1背景海岛棉染色体片段导入系的培育
以纤维品质中等陆地棉遗传标准系TM-1为母本与优质抗黄萎病海岛棉“海7124”杂交,然后用TM-1回交,回交4-5次和自交一次,在BC5S1代借助330个SSR分子标记进行分子标记辅助选择技术(Maker-assisted Selection, MAS).BC5S1代杂合的单株自交1-2代得BC5S2-3继续进行MAS选择,BC4S1代的单株自交直到BC4S2-3继续进行MAS,从而培育了国内首套陆地棉标准系TM-1背景的海岛棉染色体片段导入系。
这套导入系共有169个家系,单片段导入系有51个,占30.18%,有的家系最多导入了5个片段,同时一部分家系的替换片段是杂合的。导入片段最短的为3.54cM,最长的是46.03cM,平均长度为17.70cM,总的长度是3526.34cM,覆盖率是80.9%。
A5 (Chr.5)和A8 (Chr.8)染色体被供体片段完全覆盖,D1 (Chr.15)没有被供体片段覆盖的部分最大,达到了50.8cM, D7(Chr.16)没有被供体片段覆盖的部分最小,只有2.2cM。供体片段没有覆盖的部分主要集中在染色体的两端和中间部分。
2.高代回交群体和染色体片段导入系进行QTL的定位
2005年夏在南京江浦棉花试验站种植BC5S1和BC4S1,每隔20行种植对照TM-1；2006年夏在南京江浦棉花试验站种植BC5S2和BC4S2家系,每隔10行种植对照TM-1；2008年夏在南京江浦棉花试验站种植全部染色体片段导入系,完全随机区组试验,两次重复,每隔10行种植对照TM-1。调查BC5S1的叶型、花冠颜色、花瓣基部红心、花药颜色；调查染色体片段导入系各行的铃型、茎秆颜色和茸毛。对三个世代的产量组分(铃重和衣分)及纤维品质性状(纤维长度,纤维强度,马克隆值,整齐度,短纤维率,伸长率及成熟度)进行调查。
通过调查发现了染色体片段导入系中出现的几种表型变异,包括类im(immaturity,不成熟纤维)、叶脉融合、鸡脚叶、青茎、茎秆光滑和铃形的变异等。
用MapManager QTXb20在p<0.001的设定下,进行单标记分析。在BC5群体中检测到23个显著性标记,其中纤维品质的17个,产量组分(铃重和衣分)6个。在BC5S1群体中检测到27显著性标记,其中纤维品质的22个,产量组分(铃重和衣分)5个。在CSIL群体中检测到14个显著性标记,其中纤维品质的12个,产量组分(铃重和衣分)2个。
根据某性状的单标记分析结果进行1000次重排计算(Permutation test)估计各个性状的LR阈值,针对特定的染色体进行区间作图。通过1000次重排计算,得到了低显著水平,显著水平及高显著水平等3个水平的LR阈值。在低显著水平下检测到了所有的显著性标记。同时在三个群体中用区间作图检测到影响多个性状的标记区间,在BC5群体中检测到3个区间影响多个性状；在BC5S1群体中检测到5个区间影响多个性状；在CSIL群体中检测到3个区间影响多个性状。
3.利用染色体片段导入系精细定位纤维品质的QTL
在培育染色体片段导入系的过程中,发现一个纯合的单片段导入系在纤维强度上与轮回亲本存在明显差异,从而认为该片段上存在控制纤维强度的QTL。然后该家系与陆地棉遗传标准系TM-1杂交得到F1,种植600个F2和F2：3家系进行连锁作图,进行导入片段的分析和纤维强度QTL的精细定位。得到的结果如下：
经过F2分离群体重新作图得到了导入片段的长度是70.870cM,是通过已发表图谱计算得到的片段长度(31.10cM)的2.84倍。在F2分离群体中共检测到2个QTL。一个纤维强度的QTL,解释的表型变异是6.15%。1个马克隆值的QTL解释的表型变异是9.27%。F2：3群体中检测到5个QTL,其中纤维强度3个QTL,整齐度2个QTL。纤维强度的3个QTL解释表型变异分别是7.49%,4.96%,5.97%。整齐度2个QTL解释表型变异分别是5.89%,9.61%。
Cotton (Gossypium spp.) is one of the most important cash crops worldwide. Improvements in the standard of living in the country and the elimination of textile quotas are expected to rapidly increase the consumption and export of textile products.
Upland and Sea Island cotton are two cultivated tetraploid cotton varieties. Upland cotton (Gossypium hirsutum L.) is characterized by high yield and moderate fiber quality performance. Sea Island cotton (G barbadense L.) is characterized by low yield, increased fiber fineness and strength. Sea Island cotton is used as the raw material for fine count yarn. Intraspecific narrow genetic base has become a main problem for genetic improvement of Gossypium hirsutum L.Transmission of valuable genes extensively and efficiently between G.hirsutum Land G.barbadense L.is essential for their breeding improvements, which is limited mainly by linkage drag. Chromosome segment introgression lines (CSIL) consist of a battery of near-isogenic lines covering whole genome of crop. Except for one homozygous chromosome segment transferred from donor parent each line, the remaining parts of genome are the same as the recipient parent. It is an ideal material for genome research and especially for QTL mapping. Based on a advanced backcross population between G.hirsutum L.xG.barbadense L, Sea Island cotton chromosome segment introgression lines of cotton in background of genetic standard line of Upland cotton TM-1 were developed. QTLs were mapped by advanced backcross population and CSIL population. Fine mapping QTL for fiber strength by F2 and F2:3 populations. The main results were summarized as follows:
1. Development of Gossypium barbadense chromosome segment substitution lines in the genetic standard line TM-1 of Gossypium hirsutum
In the present research, we firstly developed one set of CSILs lines. The development of the CSILs lines:the F1 was generated by a cross using TM-1 as the recipient parent and Hai7124 as the donor parent. The TM-1 x Hai7124 F1 progeny were backcrossed with TM-1 untill BC5 and self-pollinated to produce BC5S1. MAS was conducted to identify individual BC5S1 genotypesan having 1-2 donor chromosome segments in the TM-1 background. The BC5S1 heterozygous individuals were self-pollinated to generate homozygous BC5S2-3. BC4S3 individuals were also identified by MAS.
The CSILs consist of 169 different lines. Single segment introgression lines have 51 lines,30.18% percentage of total CSIL lines. There are more than 5 segments or heterozygous segments in few lines. The length of the substituted segments covered 3526.34cM in total with an average distance of segment by 17.70cM. The substituted segments of each line are different in length, ranging from the shortest segment 3.5 4cM to the longest 46.03cM.
There are no absent segments in Chr.A5 (Chr.5) and Chr.A8 (Chr.8).50.8cM, most length segment, are absent in the Chr.Dl (Chr.15) and 2.2cM, shortest length segment are absent in the Chr.D7 (Chr.16). The absent segments were in the terminal and middle parts of each chromosome.
2. QTLs mapping by advanced backcross population and CSIL population
BC5S1 and BC4S1 seeds were planted at intervals in twenty rows, including one row of TM-1 as a control in 2005. BC5S2and BC4S2seeds were planted at intervals in ten rows, including one row of TM-1 as a control in 2006. CSIL seeds were planted using a randomized block design having 2 replications at intervals in ten rows, including one row of TM-1 as a control in 2008. The phenotype variance of BC5S1 and CSIL individuals were evaluated including leaf shape, corollaceous colour, bottom red corolla, smooth stem and boll shape. Two yield components traits including boll weigth, lint percentage and seven fiber quality traits including fiber length, fiber strength, fiber maturity, micronaire, short fiber index, fiber uniformity ratio, fiber elongation were measured from BC5S1, BC5S2and CSIL individuals.
In the present research, similar Immaturity fiber, nervate amalgamation, chicken-paw leaves, green stem, smooth stem and boll shape lines were indentified from the CSIL population.
Marker regression was done by MapManagerQTXb20 at p<0.001. Significant makers were detected in the BC5 population including 17 significant makers for fiber quality and 6 significant makers for components of yield (boll weight and lint percentage).27 significant makers were detected in the BC5S1 population including 22 significant makers for fiber quality and 5 significant makers for components of yield (boll weight and lint percentage).14 significant makers were detected in the CSIL population including 12 significant makers for fiber quality and 2 significant makers for components of yield (boll weight and lint percentage).
Interval mapping was done by MapManagerQTXb20. All the significant markers could be detected with interval mapping.3 QTLs identified in the BC5 population could affect more than one trait.5 QTLs identified in the BC5S1 population could affect more than one trait.3 QTLs identified in the CSIL population could affect more than one trait.
3. Fine mapping QTLs for fiber strength using CSIL
One CSIL line was found during the development of the CSIL, which carried single segment for fiber strength. The CSIL line was crossed with TM-1 in summer at Jiangpu Cotton Breeding Station, NAU. Seven fiber quality traits including fiber length, fiber strength, fiber maturity, micronaire, short fiber index, fiber uniformity ratio, fiber elongation were measured from 600 F2 and F2:3 individuals.
2 QTLs for fiber quality were identified in F2 population including 1QTL for fiber strength,1 QTL for micronaire. They separately explained 6.51%、9.27% of the phenotypic variance.5 QTLs for fiber quality were identified in F2:3 population including3 QTLs for fiber strength,2 QTLs for fiber uniformity ratio. They separately explained 7.49%,4.96%,5.97%,5.89% and 9.61% of the phenotypic variance.
陆地棉(Gossypium hirsutum L.)和海岛棉(G. barbadense L.)是棉花的2个四倍体栽培种,陆地棉产量高,纤维品质中等,海岛棉产量低但纤维细强,可纺100-200支纱,是纺高支纱的原料。遗传基础狭窄已经成为目前限制陆地棉遗传改良的主要因素。在陆地棉和海岛棉间进行优良基因的高效转移渗透对二者的遗传改良具有极为重要的现实意义,而限制这一过程的主要因素是连锁累赘。染色体片段导入系了除目标基因所在座位的局部区域外,基因组其余部分都是相同的,在这样的材料间找到的多态性标记才可能与目标基因紧密连锁,是QTL精细定位和图位克隆的一类理想群体。本研究以海陆杂交群体为基础,通过回交和标记辅助选择构建了海岛棉的染色体片段导入系,并利用这过程产生的高代回交群体和片段导入系对纤维品质等性状进行了QTL标记定位研究。同时还研究了QTL的精细定位。研究的主要结果如下：
1.陆地棉遗传标准系TM-1背景海岛棉染色体片段导入系的培育
以纤维品质中等陆地棉遗传标准系TM-1为母本与优质抗黄萎病海岛棉“海7124”杂交,然后用TM-1回交,回交4-5次和自交一次,在BC5S1代借助330个SSR分子标记进行分子标记辅助选择技术(Maker-assisted Selection, MAS).BC5S1代杂合的单株自交1-2代得BC5S2-3继续进行MAS选择,BC4S1代的单株自交直到BC4S2-3继续进行MAS,从而培育了国内首套陆地棉标准系TM-1背景的海岛棉染色体片段导入系。
这套导入系共有169个家系,单片段导入系有51个,占30.18%,有的家系最多导入了5个片段,同时一部分家系的替换片段是杂合的。导入片段最短的为3.54cM,最长的是46.03cM,平均长度为17.70cM,总的长度是3526.34cM,覆盖率是80.9%。
A5 (Chr.5)和A8 (Chr.8)染色体被供体片段完全覆盖,D1 (Chr.15)没有被供体片段覆盖的部分最大,达到了50.8cM, D7(Chr.16)没有被供体片段覆盖的部分最小,只有2.2cM。供体片段没有覆盖的部分主要集中在染色体的两端和中间部分。
2.高代回交群体和染色体片段导入系进行QTL的定位
2005年夏在南京江浦棉花试验站种植BC5S1和BC4S1,每隔20行种植对照TM-1；2006年夏在南京江浦棉花试验站种植BC5S2和BC4S2家系,每隔10行种植对照TM-1；2008年夏在南京江浦棉花试验站种植全部染色体片段导入系,完全随机区组试验,两次重复,每隔10行种植对照TM-1。调查BC5S1的叶型、花冠颜色、花瓣基部红心、花药颜色；调查染色体片段导入系各行的铃型、茎秆颜色和茸毛。对三个世代的产量组分(铃重和衣分)及纤维品质性状(纤维长度,纤维强度,马克隆值,整齐度,短纤维率,伸长率及成熟度)进行调查。
通过调查发现了染色体片段导入系中出现的几种表型变异,包括类im(immaturity,不成熟纤维)、叶脉融合、鸡脚叶、青茎、茎秆光滑和铃形的变异等。
用MapManager QTXb20在p<0.001的设定下,进行单标记分析。在BC5群体中检测到23个显著性标记,其中纤维品质的17个,产量组分(铃重和衣分)6个。在BC5S1群体中检测到27显著性标记,其中纤维品质的22个,产量组分(铃重和衣分)5个。在CSIL群体中检测到14个显著性标记,其中纤维品质的12个,产量组分(铃重和衣分)2个。
根据某性状的单标记分析结果进行1000次重排计算(Permutation test)估计各个性状的LR阈值,针对特定的染色体进行区间作图。通过1000次重排计算,得到了低显著水平,显著水平及高显著水平等3个水平的LR阈值。在低显著水平下检测到了所有的显著性标记。同时在三个群体中用区间作图检测到影响多个性状的标记区间,在BC5群体中检测到3个区间影响多个性状；在BC5S1群体中检测到5个区间影响多个性状；在CSIL群体中检测到3个区间影响多个性状。
3.利用染色体片段导入系精细定位纤维品质的QTL
在培育染色体片段导入系的过程中,发现一个纯合的单片段导入系在纤维强度上与轮回亲本存在明显差异,从而认为该片段上存在控制纤维强度的QTL。然后该家系与陆地棉遗传标准系TM-1杂交得到F1,种植600个F2和F2：3家系进行连锁作图,进行导入片段的分析和纤维强度QTL的精细定位。得到的结果如下：
经过F2分离群体重新作图得到了导入片段的长度是70.870cM,是通过已发表图谱计算得到的片段长度(31.10cM)的2.84倍。在F2分离群体中共检测到2个QTL。一个纤维强度的QTL,解释的表型变异是6.15%。1个马克隆值的QTL解释的表型变异是9.27%。F2：3群体中检测到5个QTL,其中纤维强度3个QTL,整齐度2个QTL。纤维强度的3个QTL解释表型变异分别是7.49%,4.96%,5.97%。整齐度2个QTL解释表型变异分别是5.89%,9.61%。
Cotton (Gossypium spp.) is one of the most important cash crops worldwide. Improvements in the standard of living in the country and the elimination of textile quotas are expected to rapidly increase the consumption and export of textile products.
Upland and Sea Island cotton are two cultivated tetraploid cotton varieties. Upland cotton (Gossypium hirsutum L.) is characterized by high yield and moderate fiber quality performance. Sea Island cotton (G barbadense L.) is characterized by low yield, increased fiber fineness and strength. Sea Island cotton is used as the raw material for fine count yarn. Intraspecific narrow genetic base has become a main problem for genetic improvement of Gossypium hirsutum L.Transmission of valuable genes extensively and efficiently between G.hirsutum Land G.barbadense L.is essential for their breeding improvements, which is limited mainly by linkage drag. Chromosome segment introgression lines (CSIL) consist of a battery of near-isogenic lines covering whole genome of crop. Except for one homozygous chromosome segment transferred from donor parent each line, the remaining parts of genome are the same as the recipient parent. It is an ideal material for genome research and especially for QTL mapping. Based on a advanced backcross population between G.hirsutum L.xG.barbadense L, Sea Island cotton chromosome segment introgression lines of cotton in background of genetic standard line of Upland cotton TM-1 were developed. QTLs were mapped by advanced backcross population and CSIL population. Fine mapping QTL for fiber strength by F2 and F2:3 populations. The main results were summarized as follows:
1. Development of Gossypium barbadense chromosome segment substitution lines in the genetic standard line TM-1 of Gossypium hirsutum
In the present research, we firstly developed one set of CSILs lines. The development of the CSILs lines:the F1 was generated by a cross using TM-1 as the recipient parent and Hai7124 as the donor parent. The TM-1 x Hai7124 F1 progeny were backcrossed with TM-1 untill BC5 and self-pollinated to produce BC5S1. MAS was conducted to identify individual BC5S1 genotypesan having 1-2 donor chromosome segments in the TM-1 background. The BC5S1 heterozygous individuals were self-pollinated to generate homozygous BC5S2-3. BC4S3 individuals were also identified by MAS.
The CSILs consist of 169 different lines. Single segment introgression lines have 51 lines,30.18% percentage of total CSIL lines. There are more than 5 segments or heterozygous segments in few lines. The length of the substituted segments covered 3526.34cM in total with an average distance of segment by 17.70cM. The substituted segments of each line are different in length, ranging from the shortest segment 3.5 4cM to the longest 46.03cM.
There are no absent segments in Chr.A5 (Chr.5) and Chr.A8 (Chr.8).50.8cM, most length segment, are absent in the Chr.Dl (Chr.15) and 2.2cM, shortest length segment are absent in the Chr.D7 (Chr.16). The absent segments were in the terminal and middle parts of each chromosome.
2. QTLs mapping by advanced backcross population and CSIL population
BC5S1 and BC4S1 seeds were planted at intervals in twenty rows, including one row of TM-1 as a control in 2005. BC5S2and BC4S2seeds were planted at intervals in ten rows, including one row of TM-1 as a control in 2006. CSIL seeds were planted using a randomized block design having 2 replications at intervals in ten rows, including one row of TM-1 as a control in 2008. The phenotype variance of BC5S1 and CSIL individuals were evaluated including leaf shape, corollaceous colour, bottom red corolla, smooth stem and boll shape. Two yield components traits including boll weigth, lint percentage and seven fiber quality traits including fiber length, fiber strength, fiber maturity, micronaire, short fiber index, fiber uniformity ratio, fiber elongation were measured from BC5S1, BC5S2and CSIL individuals.
In the present research, similar Immaturity fiber, nervate amalgamation, chicken-paw leaves, green stem, smooth stem and boll shape lines were indentified from the CSIL population.
Marker regression was done by MapManagerQTXb20 at p<0.001. Significant makers were detected in the BC5 population including 17 significant makers for fiber quality and 6 significant makers for components of yield (boll weight and lint percentage).27 significant makers were detected in the BC5S1 population including 22 significant makers for fiber quality and 5 significant makers for components of yield (boll weight and lint percentage).14 significant makers were detected in the CSIL population including 12 significant makers for fiber quality and 2 significant makers for components of yield (boll weight and lint percentage).
Interval mapping was done by MapManagerQTXb20. All the significant markers could be detected with interval mapping.3 QTLs identified in the BC5 population could affect more than one trait.5 QTLs identified in the BC5S1 population could affect more than one trait.3 QTLs identified in the CSIL population could affect more than one trait.
3. Fine mapping QTLs for fiber strength using CSIL
One CSIL line was found during the development of the CSIL, which carried single segment for fiber strength. The CSIL line was crossed with TM-1 in summer at Jiangpu Cotton Breeding Station, NAU. Seven fiber quality traits including fiber length, fiber strength, fiber maturity, micronaire, short fiber index, fiber uniformity ratio, fiber elongation were measured from 600 F2 and F2:3 individuals.
2 QTLs for fiber quality were identified in F2 population including 1QTL for fiber strength,1 QTL for micronaire. They separately explained 6.51%、9.27% of the phenotypic variance.5 QTLs for fiber quality were identified in F2:3 population including3 QTLs for fiber strength,2 QTLs for fiber uniformity ratio. They separately explained 7.49%,4.96%,5.97%,5.89% and 9.61% of the phenotypic variance.
引文
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