烟草染色体片段代换系的构建与遗传评价

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童治军, 焦芳婵, 方敦煌, 陈学军, 吴兴富, 曾建敏, 谢贺, 张谊寒, 肖炳光. 烟草染色体片段代换系的构建与遗传评价. 作物学报, 2016, 42(11): 1609-1619
TONG Zhi-Jun, JIAO Fang-Chan, FANG Dun-Huang, CHEN Xue-Jun, WU Xing-Fu, ZENG Jian-Min, XIE He, ZHANG Yi-Han, XIAO Bing-Guang. Construction and Genetic Evaluation of Chromosome Segment Substitution Lines in Tobacco ( Nicotiana tabacum L.) . ACTA AGRONOMICA SINICA, 2016, 42(11): 1609-1619 复制到剪切板

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作物学报编辑部

烟草染色体片段代换系的构建与遗传评价

童治军, 焦芳婵, 方敦煌, 陈学军, 吴兴富, 曾建敏, 谢贺, 张谊寒, 肖炳光^*

云南省烟草农业科学研究院 / 烟草行业烟草生物技术育种重点实验室 / 国家烟草基因工程研究中心, 云南昆明 650021

^* 通讯作者(Corresponding author): 肖炳光, E-mail: xiaobg@263.net

第一作者联系方式: E-mail: tzj861@163.com

收稿日期:2016-03-09 接受日期:2016-07-11 网络出版日期:2016-07-28

基金:本项目由中国烟草总公司项目[2014TBB01,110201301006(JY-06)]和中国烟草总公司云南省公司项目(2014YN18, 2013YN01)资助

摘要

以综合性状优良的烤烟种质Y3为轮回亲本, 抗烟草黑胫病(0号和1号生理小种)和赤星病的雪茄烟种质Beinhart 1000-1及优质烤烟品种K326为供体亲本, 连续回交并结合分子标记辅助选择, 构建国内外第一套由256个具有烤烟Y3遗传背景并渐渗有Beinhart 1000-1和K326染色体片段株系代换系群体。该群体携带377个代换片段, 分布于烟草24条连锁群上。每个株系携带1~5个代换片段, 代换片段长度介于0.05~36.88 cM, 平均长度7.75 cM。代换片段重叠累加总长度为2922.57 cM, 是烟草基因组总长度的2.61倍。代换片段覆盖总长度为1114.32 cM, 烟草基因组覆盖率为99.45%。本研究构建的片段代换系可用于烟草基因定位、复杂性状的QTL分析和标记辅助选择育种。

关键词: 烟草( Nicotiana tabacumL.); 染色体片段代换系; SSR; 分子标记辅助选择

Construction and Genetic Evaluation of Chromosome Segment Substitution Lines in Tobacco ( Nicotiana tabacum L.)

TONG Zhi-Jun, JIAO Fang-Chan, FANG Dun-Huang, CHEN Xue-Jun, WU Xing-Fu, ZENG Jian-Min, XIE He, ZHANG Yi-Han, XIAO Bing-Guang^*

Yunnan Academy of Tobacco Agricultural Sciences / Key Laboratory of Tobacco Biotechnological Breeding / National Tobacco Genetic Engineering Research Center, Kunming 650021, China

Fund:This study was supported by China National Tobacco Company [2014TBB01, 110201301006(JY-06)] and Yunnan Tobacco Company (2014YN18, 2013YN01)

Abstract

A set of chromosome segment substitution lines (CSSLs) of tobacco ( Nicotiana tabacum L.) were first developed by molecular marker assisted selection (MAS) and successive backcrossing with Y3, the flue-cured tobacco germplasm with comprehensive traits as the recipient parent, and two common tobacco cultivars Beinhart1000-1 and K326 as the donor parents. The cigar tobacco cultivar Beinhart 1000-1 carries a variety of resistance traits including black shank (race 0 and race 1) and brown spot resistance, while the flue-cured tobacco K326 is a commercial cultivar with high quality. In 256 CSSLs, a total of 377 substituted segments derived from donor Beinhart 1000-1 and K326 with genetic background of Y3 distributed on 24 linkage groups. Each CSSL contained only 1-5 substituted segments and the length of substituted segments ranged from 0.05 to 36.88 cM with an average of 7.75 cM. The total length of the overlapped substituted segments was 2922.57 cM, which was 2.61 times of the whole tobacco genome length. And the CSSLs covered length was 1114.32 cM, with a covered ratio of 99.45% in the recurrent tobacco genome. The CSSLs constructed in this study are excellent genetic materials for gene mapping, QTL analysis of complicated traits, and developing varieties by marker assisted selection in Nicotiana tabacum L.

Keyword: Tobacco ( Nicotiana tabacum L.); Chromosome segment substitution lines (CSSLs); SSR; Molecular marker assisted selection (MAS)

Show Figures

烟草是我国重要的经济作物之一, 其品种的选育虽取得了一定成绩, 但与K326等国外引进的优良品种相比仍有一定差距, 无法完全满足国内烟叶生产对品种的需求^[1]。烟草的产量、品质和抗性等重要育种目标性状绝大部分为多基因控制的数量性状, 遗传方式复杂, 受多基因位点和环境的共同影响, 且数量性状位点(QTL)的鉴定难度较大^{[2, 3, 4, 5, 6]}。利用传统的分离群体, 因为数量性状本身的复杂性, 及分离群体遗传背景的复杂性, 很难取得QTL定位的满意结果。染色体片段代换系(chromosome segment substitution lines, CSSL)是在受体的遗传背景中代换入某个或某些供体亲本的染色体片段, 并利用回交和分子标记辅助选择(MAS)技术建立的覆盖作物整个基因组的一系列近等基因系^{[7, 8, 9]}。与分离群体相比, 其遗传背景简单、群体遗传稳定、QTL定位及遗传效应分析准确, 在番茄^{[10, 11, 12]}、油菜^[13]、莴苣^[14]、大麦^[15]、小麦^[16]、棉花^[17]、黄瓜^{[18, 19, 20]}、水稻^{[21, 22, 23]}等作物中得到了广泛应用, 同时, 也是QTL的精确定位、挖掘和利用新基因资源、实现分子标记技术和作物育种链接的理想材料^[24]。

迄今, 国内外烟草染色体片段代换系的相关研究尚未见报道。虽然烟草基因组测序为构建高密度烟草分子标记遗传图谱奠定了基础^{[25, 26, 27, 28]}, 但因其遗传基础狭窄, 栽培品种间多态性低^{[29, 30, 31, 32, 33]}, QTL定位等基础研究相对落后; 目前仅有的少数QTL定位^{[2, 3, 4, 5, 6, 34]}主要基于初级定位群体, 其结果无法直接应用于育种研究。鉴于此, 本研究利用优质烤烟品种K326和抗烟草黑胫病(0号和1号生理小种)、赤星病的雪茄烟种质Beinhart 1000-1为供体亲本, 优良烤烟品种Y3为受体亲本, 经多代回交、自交, 并结合SSR标记辅助选择构建一套烟草染色体片段代换系, 对K326和Beinhart 1000-1中优良品质和抗病的QTL (基因)定位和转育、烤烟品种改良及促进烟草分子育种具有重要意义。

1 材料与方法

1.1 供试材料

供试的三份烟草材料均由云南省烟草农业科学研究院提供(表1)。受体亲本Y3是从津巴布韦引进的烤烟种质资源, 该资源生育期适中、株型优良、节距较大、叶数多且宽、鲜烟叶柔软, 但叶片较薄、香气质中等、香气量欠足且易感烟草黑胫病和叶斑病害(如赤星病等)。供体亲本K326香气质好、香气量足、燃烧性强, 是目前世界上种植区域最广、种植面积较大的烤烟品种; 供体亲本Beinhart 1000-1是叶片肥厚且抗烟草黑胫病0、1号生理小种和烟草赤星病的雪茄烟种质。

表1 用于构建染色体片段代换系的亲本来源及类型 Table 1 Origin and type of parents used for development of CSSLs

1.2 染色体片段代换系的构建

如图1所示, 2013年在云南省烟草农业科学研究院研和试验基地(云南省玉溪市)温室, 以烤烟种质Y3为受体亲本, 与烤烟品种K326和雪茄烟种质Beinhart 1000-1分别杂交获得F₁, 再以Y3为轮回亲本回交, 获得2个分别含300个单株的回交一代群体。从2014年1月(BC₂F₁)开始对导入的供体片段筛选。即在前期已构建的两张烟草遗传图谱的24个连锁群上每10~20 cM选取一个标记, 共计100个SSR标记(表2), 对2个群体中的各单株分别进行代换片段的追踪和检测。仅含有1个代换片段的单株直接自交纯化建立单片段代换系, 而含有多个供体染色体片段( > 5个片段)的单株继续回交并结合MAS进行前景和背景筛选。2014年6月, 将从BC₂F₁挑选出的240个株系(每个群体120个株系)播种组成1200株BC₃F₁群体。2014年12月, 从1200个BC₃F₁株系中筛选出240个株系并种植成1200株BC₄F₁群体。2015年3月, 利用已构建的2张烟草图谱中共有的416个SSR标记对1200株BC₄F₁单株进行代换片段的追踪和检测。至2015年12月, 筛选获得256个含有1~5个供体片段的烟草染色体片段代换系并分别自交得到BC₄F₂群体。2016年2月, 再利用416个标记对2560个BC₄F₂单株(每个株系随机选10株)进行供体片段的追踪和检测, 直至筛选获得稳定、纯合的烟草染色体(单)片段代换系。

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	图1 烟草CSSLs群体构建程序Fig. 1 Procedure for developing CSSLs in tobacco

表2 受体与供体亲本间的烟草SSR标记多态性 Table 2 Polymorphism between the recipient and donors detected by tobacco SSR markers

1.3 基因组DNA的提取及代换系基因型的分析

取烟草幼嫩叶片, 提取基因组DNA参考Maguire等^[35]的方法略作修改。参照Tong等^{[2, 3, 28]}的SSR标记分析及非变性聚丙烯酰胺凝胶电泳检测方法。为了构建烟草染色体片段代换系, 前期利用公共数据库NCBI上公开的普通烟草基因组序列数据[Nicotiana tabacum(K326), http://www.ncbi.nlm. nih.gov/Traces/wgs/?val=AWOJ01]开发了13 645对烟草SSR标记^{[27, 28]}; 并以K326、Beinhart 1000-1为供体亲本与受体亲本Y3组配的BC₁F₁为作图群体, 构建了2张分别包含626个和562个标记位点的SSR标记遗传图谱(数据尚未公开), 2张图谱中共有SSR标记416个, 且较均匀分布于24个连锁群上(表2)。

1.4 代换系的评价

根据已构建的烟草遗传图谱上SSR标记间的遗传距离来估算染色体片段长度, 若标记的相邻基因型相同, 则忽略相邻标记区间的双交换事件, 两标记间的区段为相同的标记基因型; 若标记的相邻基因型不同, 则无多态性的标记与有多态性标记的中点为染色体代换片段的末端, 即认为这2个标记基因型分别组成这个区间的1/2, 代换片段的长度按已构建的遗传图谱中的遗传距离来计算^[36]。

利用MapChat 2.20软件构建全基因组的图示基因型, 直观显示代换片段的位置和大小, 并分别累计来自供体亲本Beinhart 1000-1和K326的染色体片段总长度, 计算其占受体基因组的比例。

2 结果与分析

2.1 SSR标记的选择

以已构建的2张遗传图谱为基础, 每隔10~20 cM左右选择1个SSR标记, 共选择了100个SSR标记, 在BC₂F₁和BC₃F₁世代中进行供体片段的追踪和检测。其中, 第5、第14、第17和第18连锁群上标记最少, 有3个标记。第2、第13、第19和第22连锁群上标记最多, 有11个标记。SSR标记间最小距离为0.76 cM, 最大距离为41.03 cM, 平均距离为11.89 cM。从BC₄F₁开始, 利用两张图谱中共有的416个标记进行代换片段的检测与筛选。其中, 第5连锁群上标记最少, 有7个标记; 第2连锁群上标记最多, 有35个标记。SSR标记间最小距离为0.05 cM, 最大距离为36.88 cM, 平均距离为7.75 cM。

2.2 代换片段的遗传评价

通过3个世代(BC₂F₁、BC₃F₁和BC₄F₁)的分子标记辅助选择, 筛选出由256个单株组成的一套分别含有烤烟K326和雪茄烟Beinhart 1000-1导入片段的烤烟染色体片段代换系(图2)。其中, 含有K326片段的代换系有131个株系, 含有Beinhart 1000-1片段的代换系有125个株系。在筛选出的256份BC₄F₁单株中, 单片段代换单株有31个, 2~4片段代换单株有183个, 5片段代换单株有42个。其中, 具有烤烟K326片段的代换系中, 单片段代换单株有17个, 2~4片段代换单株有95个, 5片段代换单株有19个; 具有雪茄烟Beinhart 1000-1片段的代换系中, 单片段代换单株有14个, 2~4片段代换单株有88个, 5片段代换单株有23个。

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	图2(1) 256份代换系的代换片段在烟草基因组上的位置黑色部分代表供体亲本K326的染色体片段; 绿色部分代表供体亲本Beinhart 1000-1的染色体片段。Fig. 2(1) Genomic positions of the substituted segment in 256 CSSLs Black bar represents the chromosomal segments of K326; Green bar represents the chromosomal segments of Beinhart 1000-1.

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	图2(2) 256份代换系的代换片段在烟草基因组上的位置黑色部分代表供体亲本K326的染色体片段; 绿色部分代表供体亲本Beinhart 1000-1的染色体片段。Fig. 2(2) Genomic positions of the substituted segment in 256 CSSLs Black bar represents the chromosomal segments of K326; Green bar represents the chromosomal segments of Beinhart 1000-1.

从表3和图2可看出, 烟草基因组导入的377个代换片段总长度为2922.57 cM, 长度范围为0.05~ 36.88 cM, 平均长度为7.75 cM, 最长的代换片段位于第13连锁群上, 最短的代换片段位于第20连锁群上的中下部。其中, 第4连锁群代换片段的总长最大, 为223.53 cM; 第16连锁群代换片段的总长最小, 为50.38 cM。具有烤烟K326片段的代换系中导入的215个代换片段总长度为1578.91 cM, 长度范围为0.05~25.69 cM, 平均长度为7.34 cM。具有雪茄烟Beinhart 1000-1片段的代换系中导入的162个代换片段总长度为1343.66 cM, 长度范围为0.06~36.88 cM, 平均长度为8.29 cM。

表3 烟草染色体片段代换系的遗传评价 Table 3 Genetic evaluation of chromosome segment substitution lines in tobacco (Nicotiana tabacum L.)

377个代换片段在烟草24条连锁群上的分布不等, 各连锁群平均导入15~16个片段。其中, 第2连锁群的代换片段数最多, 为35个, 其次是第1连锁群的29个片段; 第16号连锁群代换片段数目最少, 为5个。215个具有烤烟K326片段的代换系中, 第2连锁群的代换片段数最多, 为21个, 其次是第1连锁群的16个片段; 第16连锁群代换片段数目最少, 为2个。162个具有雪茄烟Beinhart 1000-1片段的代换系中, 同样是第2连锁群的代换片段数最多, 为14个, 其次是第1连锁群的13个片段; 第16连锁群代换片段数目最少, 仅有2个。

代换片段在烟草基因组24条连锁群上覆盖了1114.32 cM的距离, 覆盖率为99.45% (1114.32/ 1120.45)。各连锁群覆盖率虽有不同, 但均在87.5%以上。其中, 有21条连锁群上的代换片段重叠后覆盖率为100%, 而3个第8、第9和第19连锁群上分别存在12.49% (4.60 cM)、1.77% (1.00 cM)和1.94% (0.53 cM)的区域未被覆盖(表3)。具有烤烟K326片段的代换系中, 导入的供体片段覆盖烟草基因组长度为1048.28 cM, 覆盖率为93.56%。其中, 有11条连锁群上的代换片段重叠后覆盖率达100.00%, 剩余的13条连锁群中的覆盖率范围为64.65%~ 99.34%。具有雪茄烟Beinhart 1000-1片段的代换系中, 代换片段覆盖烟草基因组长度为989.20 cM, 覆盖率为88.29%。其中, 代换片段重叠后覆盖第1、第21和第24连锁群100%, 未能达到全覆盖的21条连锁群上代换片段覆盖率范围为59.52%~98.48% (表3)。

3 讨论

在染色体片段代换系构建过程中, 亲本选择十分重要, 一般要选择遗传背景差异较大的^[7]。已有的研究中的亲本的选择主要分为两类, 一类是选择供体亲本与受体亲本间亲缘关系较远、遗传背景差异较大且双亲(或其中之一)与生产应用品种间存在较大的性状差异, 此类染色体代换系群体较为适用于遗传学研究, 但不易用于育种实践^{[10, 11, 12, 14, 15, 18]}。另一类则是供体与受体亲本间亲缘关系较近且双亲均为当前大面积推广应用的主栽品种, 此类代换系群体虽可较容易选择出具有优良代换系株系、并直接供育种实践应用, 但双亲间因亲缘关系较近而不易筛选获得足够量的分子标记, 从而影响代换系群体构建的质量^{[8, 9, 21, 22, 23]}。本研究为获得既有较高质量又易于烤烟育种实践应用的烟草染色体片段代换系, 在亲本的选择上兼顾了以下3点。首先, 选择的亲本材料具较多优点, 没有突出或致命的缺点, 且在主要育种性状上优缺点能形成互补。受体亲本(轮回亲本) Y3是综合性状优良但香气量欠足、易感烟草黑胫病和叶斑病害的烤烟种质, 供体亲本则分别具有优良品质、高抗烟草黑胫病和赤星病。其次, 亲本(或亲本之一)是能适应当地生态条件且综合性状较好的推广品种。本研究的3个亲本中, 受体亲本Y3经引种试验, 证明其在我国的西南地区(尤其云南)具有较强的适应性且综合性状优良。供体亲本K326是目前世界上种植区域最广、面积较大且深受我国卷烟企业喜好的烤烟品种。供体亲本Beinhart 1000-1具有较高的烟草黑胫病(0号和1号生理小种)和赤星病抗性。最后, 亲本间的遗传差异较大。本研究所选用的3份亲本材料的生态类型差异较大, 亲缘关系相对较远。从地域上看, 亲本K326和Beinhart 1000-1来源于美国, 亲本Y3来源于津巴布韦。从烟草类型上看, 亲本Y3和K326属于烤烟, 而亲本Beinhart 1000-1属于雪茄烟(表1)。因此, 亲本间的多态率及共显性多态标记数目均较其他烤烟品种间高(表2)。

由于CSSLs与其轮回亲本(受体)间只存在代换片段的差异, 而遗传背景与受体亲本一致, 可将复杂性状分解为单个孟德尔因子, 对代换区段中的QTL分析时遗传背景干扰小, 因而受到研究者的重视^{[22, 24]}。Eshed和Zamir等^[10]最早建立了以栽培番茄为遗传背景、含有野生番茄单片段的代换系群体, 该群体含有50个单片段代换系, 可以覆盖野生番茄的全基因组。张桂权等^[37]利用优良水稻品种华粳籼

74为受体, 以来源于世界各地的26个品种(包括14个籼稻和12个粳稻)为供体, 通过回交和分子标记辅助选择方法建立了目前世界上规模最大、质量最好的水稻单片段代换系文库。该文库共有1529个染色体单片段代换系(CSSSL), 代换片段的平均长度为18.8 cM, 代换片段总长度达28 700 cM, 相当于约18.8倍的水稻基因组长度。通过对代换片段的基因鉴定和定位, 筛选获得了包括植株形态、稻米品质、产量、抗病性、生育期等重要性状共2000多个优良基因(QTL)。本研究已获得了以烤烟Y3为遗传背景的256个含有1~5个供体片段的烟草染色体片段代换系。其中, 含有K326片段的代换系有131个株系, 含有Beinhart 1000-1片段的代换系有125个株系。获得的代换系中代换片段重叠后总长度为2922.57 cM, 相当于烤烟全基因组总长的2.61倍(本研究中以Y3和K326为作图亲本构建的烤烟SSR标记遗传图谱全长为1120.45 cM), 代换片段覆盖烟草基因组的长度为1114.32 cM, 总覆盖率高达99.45%。因此, 与其他作物相关研究比较, 本研究构建的烟草CSSLs质量总体上还不高, CSSL所包含株系的数量、受体携带代换片段的数目、代换片段的大小(长度)及代换片段在基因组的覆盖率等方面均不能完全满足烟草育种的要求。究其原因, 可能是: (1)烟草育种过程中, 过度依赖极少数的主体亲本而使得许多非育种目标的多样化性状丢失, 导致烟草(尤其是烤烟)品种间的亲缘关系较近、遗传基础狭窄、遗传多样性降低。(2)烟草基础研究相对薄弱。虽然近些年随着烟草基因组计划的开展, 基础研究在分子水平有了质的提升, 但起步较晚, 与其他作物仍有相当大差距。基于此, 导致烟草CSSL在构建过程中既受材料(亲本)的限制又受手段(研究基础)的制约。目前, 本研究获得的代换系尽管不是覆盖烟草全基因组的单片段代换系, 但正在对BC₄F₁继续进行代换片段的追踪和检测, 直至获得一整套覆盖烟草全基因组的CSSSLs, 为复杂性状QTL分析及有效转育供体亲本K326和Beinhart 1000-1中优质和抗病性状的QTL/基因到其他烤烟品种提供理想材料。

4 结论

以综合性状优良的烤烟种质Y3为轮回亲本, 以抗黑胫病0号、1号生理小种和抗赤星病的Beinhart 1000-1及优质品种K326为供体亲本, 经杂交、回交、自交, 结合分子标记辅助选择建立了国内外首套烟草染色体片段代换系群体。该群体由256个不同代换系株系组成, 且每个代换系株系中仅含有1~5个代换片段。通过对Y3遗传背景中的Beinhart 1000-1和K326染色体导入片段的分析, 筛选染色体单片段代换系, 为复杂性状QTL分析、分子标记辅助选择育种等研究提供材料。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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1995

0.0

王元英, 周健. 中美主要烟草品种亲源分析与烟草育种. 中国烟草学报, 1995, 13(5): 11-22

Wang Y

, Zhou

. Genetic analysis of the main tobacco cultivars and the tobacco breeding in USA and China. Acta Tab Sin, 1995, 13(5): 11-22 (in Chinese with English abstract)

本文分析了中美３００多个烤烟，８０多个晒晾烟主要育成品种的亲源关系，从中可以看出，品质与抗病育种是烟草育种的主体，由Ｏｒｉｎｏｃｏ衍生的Ｈｉｃｋｓ、Ｖｉｒｇｉｎｉａｂｒｉｇｈｔｌｅａｆ和４００及后来育成的ＮＣ９５、Ｃｏｋｅｒ１３９和Ｃｏｋｅｒ３１９等是烤烟品质育种和抗病育种的主体亲本；由ＷｈｉｔｅＢｕｒｌｅｙ衍生的Ｋｅｎｔｕｃｋｙ１６、Ｂｕｒｌｅｙ２１和由ＭＤＲｏｂｉｎｓｏｎ育成的Ｍａｒｙｌａｎｄ６４、Ｍａｒｙｌａｎｄ６０９分别是白肋烟与马里兰烟品质、抗病育种的主体亲本；烟草抗黑胫病、青枯病、根结线虫病、根黑腐病、烟草普通花叶病和野火病的抗源分别来自Ｆｌｏｒｉｄａ３０１、ＴＩ４４８Ａ、ＴＩ７０６、Ｎ．ｄｅｂｎｅｘｙｉ、Ｎ．ｇｌｕｔｉｎｏｓａ和Ｎ．ｌｏｎｇｉｆｌｏｒａ等原始栽培品种或野生种。美国烟草育种经历了从品质到多抗，再到综合优良性状改良的阶梯式发展过程。而我国优质、多抗育种起步较晚，仍存在品质与抗性、品质与易烤性关系研究较少，外源优良基因利用较少，遗传基础狭窄等问题，本文就近期烟草育种的目标和途径进行了初步探讨。

... 烟草是我国重要的经济作物之一, 其品种的选育虽取得了一定成绩, 但与K326等国外引进的优良品种相比仍有一定差距, 无法完全满足国内烟叶生产对品种的需求^[1] ...

2012

0.0

... 烟草的产量、品质和抗性等重要育种目标性状绝大部分为多基因控制的数量性状, 遗传方式复杂, 受多基因位点和环境的共同影响, 且数量性状位点(QTL)的鉴定难度较大^[2,3,4,5,6] ...

... 目前仅有的少数QTL定位^{[2,3,4,5,6,34]}主要基于初级定位群体, 其结果无法直接应用于育种研究 ...

... 参照Tong等^[2,3,28]的SSR标记分析及非变性聚丙烯酰胺凝胶电泳检测方法 ...

2012

0.0

... 目前仅有的少数QTL定位^{[2,3,4,5,6,34]}主要基于初级定位群体, 其结果无法直接应用于育种研究 ...

... 参照Tong等^[2,3,28]的SSR标记分析及非变性聚丙烯酰胺凝胶电泳检测方法 ...

2003

0.0

... 目前仅有的少数QTL定位^{[2,3,4,5,6,34]}主要基于初级定位群体, 其结果无法直接应用于育种研究 ...

2006

0.0

... 目前仅有的少数QTL定位^{[2,3,4,5,6,34]}主要基于初级定位群体, 其结果无法直接应用于育种研究 ...

2010

0.0

... 目前仅有的少数QTL定位^{[2,3,4,5,6,34]}主要基于初级定位群体, 其结果无法直接应用于育种研究 ...

2007

0.0

廖长见, 王颖姮, 潘光堂. 作物染色体导入系的构建及其应用. 分子植物育种, 2007, 5(6): 139-144

Liao C

, Wang Y

, Pan G

. Construction and application of chromosome introgression lines in crops. Mol Plant Breed, 2007, 5(6): 139-144 (in Chinese with English abstract)

染色体导入系,也称染色体片段代换系、回交重组自交系或近等基因系,是借助分子辅助选择方法,通过回交和自交,从供体中导入一个或几个小的染色体片段进入轮回亲本.相对于传统的遗传群体,导入系群体大大降低了供体材料的遗传背景的干扰,提高了QTL分析的灵敏度和准确性,是QTL鉴定、精细定位、互作分析、图位克隆、性状改良及杂种优势利用和基因表达分析的理想的实验材料.本文就染色体导入系的研究情况进行了详细的综述.

... 染色体片段代换系(chromosome segment substitution lines, CSSL)是在受体的遗传背景中代换入某个或某些供体亲本的染色体片段, 并利用回交和分子标记辅助选择(MAS)技术建立的覆盖作物整个基因组的一系列近等基因系^[7,8,9] ...

... 3 讨论在染色体片段代换系构建过程中, 亲本选择十分重要, 一般要选择遗传背景差异较大的^[7] ...

1992

0.0

... 另一类则是供体与受体亲本间亲缘关系较近且双亲均为当前大面积推广应用的主栽品种, 此类代换系群体虽可较容易选择出具有优良代换系株系、并直接供育种实践应用, 但双亲间因亲缘关系较近而不易筛选获得足够量的分子标记, 从而影响代换系群体构建的质量^{[8,9,21,22,23]} ...

1992

0.0

1995

0.0

... 与分离群体相比, 其遗传背景简单、群体遗传稳定、QTL定位及遗传效应分析准确, 在番茄^[10,11,12]、油菜^[13]、莴苣^[14]、大麦^[15]、小麦^[16]、棉花^[17]、黄瓜^[18,19,20]、水稻^[21,22,23]等作物中得到了广泛应用, 同时, 也是QTL的精确定位、挖掘和利用新基因资源、实现分子标记技术和作物育种链接的理想材料^[24] ...

... 已有的研究中的亲本的选择主要分为两类, 一类是选择供体亲本与受体亲本间亲缘关系较远、遗传背景差异较大且双亲(或其中之一)与生产应用品种间存在较大的性状差异, 此类染色体代换系群体较为适用于遗传学研究, 但不易用于育种实践^{[10,11,12,14,15,18]} ...

... Eshed和Zamir等^[10]最早建立了以栽培番茄为遗传背景、含有野生番茄单片段的代换系群体, 该群体含有50个单片段代换系, 可以覆盖野生番茄的全基因组 ...

2000

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1998

0.0

1996

0.0

2004

0.0

2003

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2004

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2009

0.0

冯长辉, 张胜昔, 王志伟, 王夏青, 李夕梅. 棉花染色体单片段导入系的研究进展. 湖北农业科学, 2009, 48(12): 24-26

Feng C

, Zhang S

, Wang Z

, Wang X

, Li X

. Research progress on construction and application of SSILs in cotton. Hubei Agric Sci, 2009, 48(12): 24-26 (in Chinese with English abstract)

染色体单片段导入系具有片段单一、背景一致、遗传稳定等特点,能把复杂性状的基因分解为多个简单的孟德尔因子,将在作物数量性状和功能基因组的分子研究中发挥重要的作用.本文针对棉花中数量性状位点作图中所遇到的如遗传背景变异的干扰等难题,概述了培育染色体单片段导入系的原理和方法,以及在棉花中的应用情况,讨论了棉花染色体单片段导入系构建过程中的问题和在棉花遗传育种中的潜在价值. Abstract： QTL(Quantitative trait locus) can be divided into several simple Mendelian factors using SSILs (Single segment introgression lines) with the characterizations of single introgressed segment,purifying of genetic background,genetic stability and so on.SSILs will play an important role in molecular research of QTLs and function genome.In the light of problems as encountered in QTL mapping for cotton and other crops such as genetic background variation, we tried to make a detailed review that focused on the construction and application of SSILs in cotton in this paper. Furthermore,The potential value and the problem which existed in SSILs were discussed.

2012

0.0

孟佳丽, 娄群峰, 周晓慧, 史建磊, 陈劲枫. 黄瓜-酸黄瓜染色体片段导入系群体的构建及果实相关数量性状基因的定位. 中国农业科学, 2012, 45: 1558-1567

Meng J

, Lou Q

, Zhou X

, Shi J

, Chen J

. Construction of cucumber-sour cucumber chromosome introgression lines and location of fruit related QTLs. Sci Agric Sin, 2012, 45: 1558-1567 (in Chinese with English abstract)

【Objective】 A set of cucumber-sour cucumber chromosome introgression lines were constructed by using cultivated cucumber (Cucumis sativus L., 2n=14) cultivar ‘Beijingjietou’ as recurrent parent and the wild ‘sour cucumber’ (C. hystrix Chakr., 2n = 24) as donor parent by the way of marker-assisted selection, and QTL controlling cucumber fruit shape was detected initially. 【Method】 Fristly, through interspecific-backcrossing-selfing approach, a lot of cucumber-sour cucumber chromosome introgression lines were obtained. Then polymorphism of the parents was detected by 298 pairs of SSR markers which uniformly distributed in the cucumber genome. And then the cucumber-sour cucumber chromosome introgression lines which contained the wild sour cucumber chromosome segments from the strains were screened by using the detected differences between parents SSR markers. QTL analysis of three main traits affecting cucumber fruit shape(fruit length, fruit diameter, fruit length/diameter ratio) was conducted using t-test compared with the recurrent parent. 【Result】 A set of 50 cucumber-sour cucumber chromosome introgression lines was constructed. There are 149 chromosome introgression segments containing 61 different introgression segments were identified in the lines. The total length of the different segments was 259.95 cM, and the coverage was 45.37% in the cucumber genome. The length of the introgression segments ranged from 1.65 cM to 15.4 cM with an average of 5.41 cM. They distributed in 7 chromosomes of cucumber. A total of 13 QTL affecting cucumber fruit shape were detected in CSIL population initially. 【Conclusion】 In this research, a set of cucumber-sour cucumber introgression lines were created and QTL affecting cucumber fruit shape was detected initially. It provided new resources for utilization of genes from C. hystrix and also the material basis for fine mapping of quantitative traits in cucumber.

【目的】以栽培黄瓜（Cucumis sativus L. , 2n = 14)‘北京截头’为受体亲本，以野生酸黄瓜( C. hystrix Chakr, 2n = 24）为供体亲本，采用SSR标记辅助选择法构建黄瓜-酸黄瓜染色体片段导入系群体。初步定位控制黄瓜果实外形的数量性状基因。【方法】首先通过种间杂交-回交-自交获得大量的染色体片段导入系株系。然后选择均匀分布在黄瓜染色体组上的298对SSR标记对亲本进行多态性检测，使用检测出的亲本间差异引物对染色体片段导入系株系进行检测，筛选含有野生酸黄瓜染色体片段的植株。对该群体果实外形进行初步调查，利用t测验与轮回亲本比较，鉴定QTL。【结果】本研究构建了由50个株系组成的染色体片段导入系群体。在该群体中共检测到149个染色体导入片段，包含61个不同的导入片段，不同导入片段的总长度为259.95 cM，基因组覆盖率为45.37%。导入片段的长度在1.65—15.4 cM，平均长度为5.41 cM，分布于黄瓜的7条染色体上。利用该导入系群体初步定位了控制黄瓜果型的13个QTL。【结论】构建了一套以栽培黄瓜为背景，野生酸黄瓜为前景的染色体片段导入系群体，并利用该导入系初步定位了控制黄瓜果型的QTL，为开发利用野生酸黄瓜的优良基因提供了新的种质资源，也为今后定位黄瓜的数量性状遗传位点奠定了材料基础。

2012

0.0

林肖剑, 许学文, 钱红梅, 齐晓花, 徐强, 陈学好. 黄瓜抗白粉病染色体片段导入系的SSR鉴定. 园艺学报, 2012, 39: 485-492

Lin J

, Xu X

, Qian H

, Qi X

, Xu

, Chen X

. Analysis of cucumber chromosome segment introgression line with powdery mildew resistance based on SSR markers. Acta Hort Sin, 2012, 39: 485-492 (in Chinese with English abstract)

17 cucumber chromosome segment introgression lines（CSILs）were constructed from a cross between the powdery mildew susceptible cultivar（D8）and resistant cultivar（JIN5-508），and 63 plants with high powdery mildew resistance（disease index ≤ 10）were selected. In this study，324 pairs of SSR primers were applied to analysis the polymorphic level between JIN5-508 and D8，and 91 pairs of SSR primer which generated polymorphic bands were applied to analysis the distribution，segment size of the chromosome segments. The results indicated that 25 CSILs were single segment introgression line，which accounted for 39.68% of the total CSILs；Most of the CSILs were double segment introgression lines，take up to 50.79%；Four CSILs were imported 3 chromosome segments；No donor introgression segments were found in 2 lines. 4 donor segments occupied the highest frequency，which located on 23.4–41.3 cM of chromosome 1，51.1–54.0 cM of chromosome 3，9.3–11.3 cM of chromosome 4 and 1.4–3.2 cM of chromosome 6，respectively.

以黄瓜高抗白粉病品种JIN5-508为供体亲本，以高感白粉病品种D8为轮回亲本，通过多代回交后自交，并在每个世代结合白粉病抗性进行接种鉴定，获得了17个以D8为遗传背景但具有抗白粉病的染色体片段导入系。利用平均分布于黄瓜7条染色体上的324对SSR引物对17个导入系63个抗病单株（病情指数 ≤ 10）的染色体导入片段特征进行了鉴定。结果表明：74对SSR引物在亲本间表现多态性；63个抗病单株中单片段导入系共25株，占39.68%；导入双片段的共32株，占50.79%；有3个供体片段导入的为4株，占6.35%；无片段导入的2株，占3.17%。出现频率较高的导入片段共4个，分别位于1号染色体的23.4 ~ 41.3 cM、3号染色体的51.1 ~ 54.0 cM、4号染色体的9.3 ~ 11.3 cM和6号染色体的1.4 ~ 3.2 cM处，出现的频率分别为74.6%、44.4%、25.4%和7.9%。

2011

0.0

史建磊, 娄群峰, 钱春桃, 万红建, 周晓慧, 陈劲枫. 黄瓜染色体片段导入系的构建与遗传评价. 南京农业大学学报, 2011, 34(1): 20-24

Shi J

, Lou Q

, Qian C

, Wan H

, Zhou X

, Chen J

. Construction and genetic evaluation of chromosome segment introgression lines in cucumber. J Nanjing Agric Univ, 2011, 34(1): 20-24 (in Chinese with English abstract)

以综合性状优良的栽培黄瓜(Cucumis sativus L.,2n=14)北京截头为受休亲本,以携带有多种抗性性状的黄瓜珍稀野牛种酸黄瓜(Cucumis hystrix Chakr.,2n=24)为供体亲本,在本实验室前期获得种间杂交异源叫倍体的基础上,通过多代回交和自交,结合SSR标记辅助选择构建了以北京截头为遗传背景的黄瓜野生种染色体片段导入系,并对其导入片段的数日、分布、大小和覆盖率等进行了评价.结果表明:该导入系群体携带的38个供体片段不均匀地分布于黄瓜7条染色体上,其长度介于1.1～14.9 cM,平均长度为5.3 cM,总长度为201.9 cM,在黄瓜基因组上的覆盖率为23.5%.

2004

0.0

刘冠明, 李文涛, 曾瑞珍, 张泽民, 张桂权. 水稻单片段代换系片段的QTL鉴定. 遗传学报, 2004, 31: 1395-1400

Liu G

, Li W

, Zeng R

, Zhang Z

, Zhang G

. Identification of QTLs on substituted segments in single segment substitution lines of rice. Acta Genet Sin, 2004, 31: 1395-1400 (in Chinese with English abstract)

The quantitative trait loci (QTLs) for 21 traits of agronomy importance on ten substituted chromosomal segments were identified using single segment substitution lines (SSSLs) in rice, which were developed by the use of Taichung 65 as a recipient and Zhai-ye-qing and Dee-geo-woo-gen as donors respectively. Total length of the substituted segments in the SSSLs was 230.00 cM, which was 12.62% of rice genome. T-test was used to detect QTLs in the condition that the difference in phenotypes between a SSSl and the recipient parent (Taichung 65) was significant at the 0.1% level (P≤0.001). A total of 57 QTLs for 17 traits had been detected on the ten substituted segments, which were on chromosomes 1, 3, 5, 7, 8, 10 and 12 respectively. The additive effect percentages of the QTLs ranged from 1.10% to 89.73%. Among of 57 QTLs, fifteen QTLs had the additive effect percentage over 10%, thirty QTLs varied from 3% to 10% and other twelve QTLs were below 3%.

利用以台中65为受体，窄叶青和低脚乌尖为供体培育的10个水稻单片段代换系为材料，对代换片段上21个重要农艺性状的QTL进行了鉴定。10个代换片段的总长度为230.00 cM ，占水稻基因组总长度的12.62%。通过t 测验比较单片段代换系与受体亲本的表型差异，以P≤0.001为阈值，在10个代换片段上共鉴定出17个性状的57个QTL，平均每个性状鉴定出3.35个，这些QTL分布于第1、3、5、7、8、10和12染色体上。57个QTL的加性效应百分率在1.10%~89.73%之间，其中15个QTL的加性效应百分率大于10%，30个QTL在3%~10%之间，12个QTL小于3%。

2005

0.0

何风华, 席章营, 曾瑞珍, Akshay

, 张桂权. 利用高代回交和分子标记辅助选择建立水稻单片段代换系. 遗传学报, 2005, 32: 825-831

He F

, Xi Z

, Zeng R

, Akshay

, Zhang G

. Developing single segment substitution lines (SSSls) in rice (Oryza sativa L. ) using advanced backcrosses and MAS. Acta Genet Sin, 2004, 32: 825-831 (in Chinese with English abstract)

A novel population consisted of 86 single segment substitution lines (SSSLs) were developed from advanced backcrosses between the recipient, Huajingxian 74 and six donors by microsatellite marker-assisted selection (MAS). Fifty-two SSSLs were selected in BC3F2, and 34 others were selected in BC3F3. Every SSSL contains a single chromosome segment introgressed from one donor in the genetic background of Huajingxian 74. The substituted segments in SSSLs were distributed on 12 rice chromosomes. The estimated length of the substituted segments in SSSLs ranged from 1.5 cM to 56.3 cM with an average of 23.0 cM. Total of the substituted segments covered 57.1% of the rice genome.

以水稻品种华粳籼74为受体，以6个水稻品种为供体，通过高代回交和微卫星标记辅助选择相结合的方法，建立了水稻的一个单片段代换系群体。该群体由86个单片段代换系组成，其中52个在BC3F2中获得，34个在BC3F3中获得。每个单片段代换系只含有来自一个供体的一个染色体代换片段，而遗传背景与华粳籼74相同。这些单片段代换系的代换片段分布于水稻的12条染色体，代换片段的长度为1.5 ~ 56.3 cM，平均长度为23.0 cM。全部代换片段在水稻基因组上的覆盖率为57.1%。

... 由于CSSLs与其轮回亲本(受体)间只存在代换片段的差异, 而遗传背景与受体亲本一致, 可将复杂性状分解为单个孟德尔因子, 对代换区段中的QTL分析时遗传背景干扰小, 因而受到研究者的重视^[22,24] ...

2005

0.0

2008

0.0

王玉民, 席章营, 尚爱兰, 王帮太. 作物单片段代换系的构建及应用. 中国农学通报, 2008, 24(3): 67-71

Wang Y

, Xi Z

, Shang A

, Wang B

. Construction and application of single segment substitution lines in crops. Chin Agric Sci Bull, 2008, 24(3): 67-71 (in Chinese with English abstract)

准确鉴定和定位作物的数量性状基因位点(quantitative trait loci,QTLs)是分子标记辅助选择和基因克隆的基础,传统的分离群体遗传背景复杂,很难对单个数量性状基因位点进行准确鉴定和深入研究.单片段代换系具有片段单一、背景一致、遗传稳定等特点,能把复杂性状的基因分解为多个简单的孟德尔因子,在作物数量性状和基因功能的分子研究中将发挥重要的作用.文中概述了作物单片段代换系的构建方法和应用概况,讨论了单片段代换系库构建过程中存在的问题及潜在的利用价值.

2007

0.0

... 虽然烟草基因组测序为构建高密度烟草分子标记遗传图谱奠定了基础^{[25,26,27,28]}, 但因其遗传基础狭窄, 栽培品种间多态性低^{[29,30,31,32,33]}, QTL定位等基础研究相对落后 ...

2011

0.0

2014

0.0

... gov/Traces/wgs/?val=AWOJ01]开发了13 645对烟草SSR标记^[27,28] ...

2012

0.0

... 参照Tong等^[2,3,28]的SSR标记分析及非变性聚丙烯酰胺凝胶电泳检测方法 ...

... gov/Traces/wgs/?val=AWOJ01]开发了13 645对烟草SSR标记^[27,28] ...

2006

0.0

2000

0.0

2008

0.0

2009

0.0

2009

0.0

1995

0.0

... 目前仅有的少数QTL定位^{[2,3,4,5,6,34]}主要基于初级定位群体, 其结果无法直接应用于育种研究 ...

1994

0.0

... 3 基因组DNA的提取及代换系基因型的分析取烟草幼嫩叶片, 提取基因组DNA参考Maguire等^[35]的方法略作修改 ...

1989

0.0

... 若标记的相邻基因型不同, 则无多态性的标记与有多态性标记的中点为染色体代换片段的末端, 即认为这2个标记基因型分别组成这个区间的1/2, 代换片段的长度按已构建的遗传图谱中的遗传距离来计算^[36] ...

2005

0.0

张桂权, 曾瑞珍, 丁效华, 张泽民, 李文涛. 水稻单片段代换系文库的构建及其在分子育种上的应用. 见: 方宣钧, 黎志康主编. 2005植物分子育种国际学术研讨会论文集. 北京: 科学出版社, 2005. pp 106-113

Zhang G

, Zeng R

, Ding X

, Zhang Z

, Li W

. Construction of a library of single segment substitution lines for rice breeding by design. In: Fang X J, Li Z K, eds. Proceedings of the 2005 International Conference on Plant Molecular Breeding. Beijing: Science Press, 2005. pp 106-113

姝 We have constructed a library of single segment substitution lines (SSSLs) in rice.To construct thelibrary,Hua-jing-xian 74 (HJX74),an elite Indica variety from south China,was used as recipient and 24 acces-sions including 14 Indica and 10 Japonica collected worldwide as donors.Through marker-assisted selection with574 SSR markers,a total of 1 123 SSSLs were developed from the 24 populations.Each of the SSSLs containsonly one substituted chromosome segment from a donor in the HJX74 genetic background.The size of a substi-tuted segment in the SSSLs was from 0.2 to 109.7cM with an average of 19.3cM.The total length of substitutedsegments in the SSSL library was 21 674cM,which was the size of about 14 times office genome.The SSSL li-brary has been used for identification and mapping of the QTLs of agronomic importance.The SSSLs in the li-brary,which carried favorable alleles in the genes on the substituted segments,were selected as candidate linesfor breeding.An SSSL showing almost the same phenotype as HJX74 except having purple pericarp was selectedin the library.The SSSL carries a substituted segment on chromosome 4 with Pb gene.Recently,the SSSL wasnamed Hua-xiao-hei 1 and permitted to release to farmers.The SSSL library collected a large number of the natu-rally occurring allelic variation in rice genome.It will be a powerful tool for functional genomics and rice breed-ing by design.

... 张桂权等^[37]利用优良水稻品种华粳籼 ...