向日葵高密度遗传连锁图谱构建及两种水分条件下芽期性状的QTL分析

引用本文

吕品, 于海峰, 于志贤, 张永虎, 张艳芳, 王婷婷, 侯建华. 向日葵高密度遗传连锁图谱构建及两种水分条件下芽期性状的QTL分析.作物学报, 2016, 43(1): 19-30
LYU Pin, YU Hai-Feng, YU Zhi-Xian, ZHANG Yong-Hu, ZHANG Yan-Fang, WANG Ting-Ting, HOU Jian-Hua. Construction of High-density Genetic Map and QTL Mapping for Seed Germination Traits in Sunflower under Two Water Conditions. Acta Agronomica Sinica, 2016, 43(1): 19-30 复制到剪切板

Doi:10.3724/SP.J.1006.2017.00019
Permissions

作物学报编辑部

向日葵高密度遗传连锁图谱构建及两种水分条件下芽期性状的QTL分析

吕品¹, 于海峰², 于志贤¹, 张永虎², 张艳芳², 王婷婷¹, 侯建华^1,^*

¹内蒙古农业大学, 内蒙古呼和浩特 010019

²内蒙古农牧业科学院, 内蒙古呼和浩特 010031

^* 通讯作者(Corresponding author): 侯建华, E-mail:houjh68@163.com

第一作者联系方式: E-mail:18847123096@163.com

收稿日期:2016-09-18

基金:本研究由国家自然科学基金项目(31160288, 31540043)资助

摘要

干旱胁迫对向日葵发芽出苗有重要影响。以K55×K58组合衍生的187个F₆重组自交系为材料, 利用SSR、SRAP、AFLP标记构建向日葵高密度遗传连锁图谱, 设置正常水分(CK)和模拟干旱(18%聚乙二醇PEG-6000)两种水分条件, 调查9个芽期数量性状, PCR扩增株系, 构建一张包含17个连锁群、1105个标记(368个SSR、368个SRAP和369个AFLP)的高密度遗传连锁图谱。该图谱覆盖基因组长度3846.0 cM, 平均图距3.48 cM, 连锁群长度147.6~295.5 cM, 每个连锁群标记数10~165个。两种条件下检测到33个QTL, 其中干旱条件下检测到发芽指数、发芽率、胚芽长、胚根长、胚芽鲜重和胚根鲜重6个性状的14个QTL, 可解释6.1%~14.0%的表型变异; 正常水分(CK)条件下检测到发芽势、胚根长、胚芽鲜重、胚根鲜重、胚根干重和胚芽干重6个性状的19个QTL, 可解释6.1%~25.8%的表型变异。两种水分条件下检测到 Qefw5-1、Qefw5-2、Qefw5-4、Qrfw5、Qrfw10和 Qrl9共6个QTL的遗传贡献率超过10%, 此外, 还检测到9个影响干旱胁迫与正常水分条件下性状差值的QTL, 可能对抗旱性有直接贡献。这些QTL可为向日葵芽期抗旱分子设计育种研究提供重要参考。

关键词: 向日葵; 遗传连锁图谱; 耐旱性; 芽期性状; QTL

Construction of High-density Genetic Map and QTL Mapping for Seed Germination Traits in Sunflower under Two Water Conditions

LYU Pin¹, YU Hai-Feng², YU Zhi-Xian¹, ZHANG Yong-Hu², ZHANG Yan-Fang², WANG Ting-Ting¹, HOU Jian-Hua^1,^*

¹Inner Mongolia Agricultural University, Huhhot 010010, China

² Inner Mongolia Academy of Agricultural and Animal Husbandry Sciences, Huhhot 010031, China

Fund:The study was supported by the National Natural Science Foundation of China (31160288, 31540043)

Abstract

Seed germination and emergence of sunflower are seriously affected by water stress. In this study, SSR, SRAP, and AFLP markers were applied to construct a genetic linkage map by using the F6 population derived from a cross of K55 (drought sensitive) × K58 (drought resistant). For mapping quantitative trait loci (QTLs) for nine traits of seed germination in sunflower, the parents and 187 F6 family lines were used to investigate seed germination traits under normal condition (CK) and 18% polyethylene glycol (PEG-6000) (drought stress). A genetic map consisting of 17 linkage groups was constructed with 1105 loci (368 SSR, 368 SRAP, 369 AFLP) which covers 3846.0 cM and the length of each linkage group varies from 147.6 to 295.5 cM, the number of markers in each linkage group varies from 10 to 165 with an average distance of 3.48 cM. As a result, a total of 33 QTLs were detected. We identified fourteen additive QTLs for germination index (GI), germination rate (GR), embryo length (EL), radicle length (RL), embryo fresh weight (EFW), radicle fresh weight (RFW) under18% PEG condition with explained phenotypic variance ranging from 6.1% to 14.0%. Nineteen additive QTLs were identified for germination energy(GE), radicle length (RL), embryo fresh weight (EFW), radicle fresh weight (RFW), embryo dry weight (EDW), radicle dry weight (RDW) under normal condition with explained phenotypic variance ranging from 6.1% to 25.8%. Each of Qefw5-2, Qefw5-1, Qefw5-4, Qrfw10, Qrfw5, and Qrl9 could explain phenotypic variance over 10%. Nine QTLs affecting trait differences between stress treatment and control were identified, which are considered to directly contribute to drought tolerance. These QTLs identified could provide important reference to molecular breeding for drought-resistance during seed germination in sunflower.

Keyword: Sunflower; Genetic linkage map; Drought tolerance; Seed germination traits; QTL

Show Figures

向日葵是世界第四大油料作物, 也是我国主要油料作物。由于其具有耐旱、耐瘠薄的特点, 广泛种植于干旱、半干旱地区, 但这些地区降水量不足且时空分布不均, 干旱问题频发, 已成为向日葵减产的最主要非生物胁迫因素。发芽出苗期作为向日葵生长发育的第一阶段, 芽期抗旱性直接关系到出苗情况, 进而影响后续的生长发育及产量^[1], 发芽率、发芽势、发芽指数、胚芽、胚根的长度、鲜重、干重均为衡量种子萌发状况的主要指标, 与芽期抗旱性密切相关^{[2, 3, 4]}。作物抗旱性属多基因控制的复杂性状, 分子标记技术及QTL定位的发展为复杂数量性状的研究及遗传改良提供了有力的工具^[5]。

高密度分子遗传连锁图谱的构建是进行QTL定位的基础, 自第一张向日葵图谱使用RAPD标记构建完成以来^[6], RFLP、AFLP、SSR等标记被相继成功应用于向日葵遗传连锁图谱构建, 其中较具参考价值的是Tang等^[7]利用F₇重组自交系群体构建的含有459个SSR标记、平均距离3.1 cM、总长度1368.3 cM的遗传连锁图谱。以SNP标记为代表的第3代分子标记用于向日葵图谱构建方面, John等^[8]利用4个群体构建了一张含有10 080个SNP标记的向日葵遗传连锁图谱, Zahirul等^[9]利用3个F₂群体构建一张向日葵遗传连锁图谱, 该图谱包含SNP标记5019个、SSR标记118个。国内向日葵图谱构建研究进展缓慢, 黄先群等^[10]于2012年利用SSR标记加密了遗传图谱, 本实验室利用SSR和AFLP标记构建了一张包含495个标记、平均距离5.6 cM、总长度为2759.4 cM的遗传连锁图谱^[11]。向日葵相关性状QTL的定位研究来自国外的较多, 不同水分条件下的株高、茎粗、开花期、水分利用率等性状的QTL均被定位到图谱上^{[12, 13, 14]}, 但有关不同水分条件下芽期性状QTL定位研究国内外鲜有报道。本研究拟利用SRAP和SSR标记整合本实验室已有的SSR和AFLP标记^[11], 构建一张高密度遗传连锁图谱, 结合2种水分条件下芽期9个性状的表型值, 进行QTL定位分析, 为解析向日葵水、旱条件下芽期相关性状QTL及遗传效应和分子标记辅助育种奠定坚实的研究基础。

1 材料与方法

1.1 试验材料

以抗旱性差异显著的油用向日葵自选自交系K55 (弱抗旱性)、K58 (强抗旱性)为亲本, 杂交获得F₁代, 采用单粒传的方法经过连续自交获得F₆重组自交系(RILs)群体, 共187个株系。

1.2 表型性状测定

按照陈雪等^[15]的方法, 设置正常水分(CK)和模拟干旱胁迫(18%聚乙二醇PEG-6000) 两种水分条件, 调查9个芽期性状。选取亲本及187个株系各50粒饱满种子, 经消毒处理(75%酒精消毒30 s, 0.1%的HgCl₂消毒8 min, 蒸馏水漂洗2次), 置铺有2层滤纸的培养皿(150 mm)中, 分别加5 mL的18% PEG-6000溶液和蒸馏水, 每个处理重复3次, 于25℃恒温光照培养箱中(光照8 h/黑暗16 h)培养10 d, 期间每天更换滤纸以保持恒定PEG浓度。种子置床后第2天开始, 每天统计种子发芽数, 第4天计算发芽势(%)(germination energy, GE), 发芽结束后计算发芽率(%)(germination rate, GR)、发芽指数(germination index, GI), 从每个重复培养皿中随机选取20株测量胚芽长度(cm)(embryo length, EL)、胚根长度(cm)(radicle length, RL), 称量胚芽鲜重(g)(embryo fresh weight, EFW)、胚根鲜重(g)(radicle fresh weight, RFW), 将称过鲜重的胚芽、胚根105℃烘干30 min, 85℃烘干8 h后称量胚芽干重(g) (embryo dry weight, EDW)、胚根干重(g)(radicle dry weight, RDW)。

发芽率(%)=发芽结束时正常幼苗数/供试种子总数× 100%

发芽势(%)=第4天时发芽的种子粒数/供试种子数× 100%

发芽指数=∑ Gt/Dt, Gt为第t天发芽种子数, Dt为相对应的发芽试验天数。

1.3 DNA提取及PCR扩增

选取向日葵苗期鲜嫩真叶, 用TIANGEN试剂盒提取双亲及187个株系的基因组DNA。SSR引物ORS系列来自NCBI (http://www. ncbi.nlm.nih.gov/), Ha系列来自Paniego等^[16]的文献; SRAP引物来自于Li和Quiros等^[17]的文献, 引物由上海生工生物工程有限公司合成。参照房冬梅等^[18]和于志贤等^[19]改进的PCR体系及程序。

1.4 分子标记统计分析

在统计位点时, 对SSR和SRAP标记均采用A, B矩阵, 即和亲本K55带型一样的记作A, 和亲本K58带型一样的记作B, 缺失或模糊不清的带用“ -” 表示。采用引物组合名称命名标记, 同一引物扩增的不同片段加1、2、3代表片段由大到小, 对分离带型以孟德尔分离比1∶ 1进行卡方测验。

1.5 遗传图谱构建及QTL定位

应用JoinMap 4.0软件构建图谱, 采用Kosambi函数计算图谱距离, LOD值定为4.0。分析芽期9个性状3次重复的平均值, 利用SPSS17.0软件进行性状表型分析及正态分布检测, 以两种条件下的性状表型值及其干旱与对照的差值为输入数据, 利用Map QTL4.0软件进行QTL定位及效应分析, 将LOD≥ 2.5作为QTL存在的判定标准。

2 结果与分析

2.1 多态性标记的筛选

利用560对SSR引物和388对SRAP引物在双亲中筛选, 得到多态性引物266对, 其中SSR引物164对, SRAP引物102对, 引物多态性比率分别为29.29%和26.29%。利用多态性引物在重组自交系群体187个株系中, 共检测到SSR标记位点1025个, 其中多态性位点405个, 多态率为39.51%, 平均每对引物组合扩增出多态性位点2.53个; SRAP标记位点997个, 其中多态性位点430个, 多态率为43.13%, 平均每对引物组合扩增出4.22个。

2.2 遗传图谱的构建

根据在重组自交系群体F₆检测到的835个(405个SSR和430个SRAP)多态性标记, 结合实验室前期在F_5:6重组自交系群体检测得到的577个多态性标记(184个SSR标记及393个AFLP标记)共计1412个标记数据, 利用 JoinMap 4.0软件进行遗传连锁分析, 获得一张包含17个连锁群的高密度多标记油葵分子遗传图谱(图1), 该图谱总长度为3846.0 cM, 连锁群长度在147.6~295.5 cM之间, 平均每个连锁群长度为226.2 cM; 图谱含1105个标记(368个SSR、368个SRAP和369个AFLP), 每个连锁群标记数有10~165个不等、平均为65个, 相邻两两标记间的平均距离为3.5 cM。图谱中标记分布不均匀(表1), 标记数最多的连锁群为第8连锁群, 有165个标记, 最少的连锁群是第15和17连锁群, 有10个标记; 相邻标记间的距离也不同, 图谱中相邻两标记之间距离≤ 10 cM的有1016个, 占93.38%, 相邻标记间距离最小为0 cM; 相邻两标记之间距离≥ 20 cM的有30个, 距离最大为68.4 cM, 位于第15连锁群上的ORS-734-2和ORS-1143-2之间。连锁群中共有333个偏分离标记, 占图谱总标记数的31.68%, 这些偏分离标记在图谱上共形成共有27个偏分离热点区域(SDR), 分布于1、2、4、5、6、7、8、9、11、12、15、16连锁群上。

新构建的图谱与AFLP和SSR标记构建的原图谱[11]比较, 长度增加了1086.6 cM, 新增位点610个, 平均距离缩小了2.1 cM, 偏分离比率降低了2.66%。

表1 图谱中连锁群标记及其分布 Table 1 Markers of linkage groups and their distribution

2.3 两种水分条件下芽期相关性状表型变异分析

由表2可见, 模拟干旱条件下, 亲本及后代重组自交系群体的发芽率、发芽势等9个芽期相关性状均表现显著降低, 其中亲本K58各性状平均降低48.01%, K55降低58.81%, 说明干旱胁迫明显抑制向日葵种子的萌发, 且亲本K58对干旱胁迫的敏感性显著低于K55。干旱胁迫下, 除胚根干重外, 其他8个性状的变异系数都有不同程度的增加, 其中发芽率增加幅度最大, 为65%, 说明各株系的不同性状间对干旱胁迫的响应差异较大。除胚根长度差值偏度略大于1外, 其余各性状的偏度和峰度的绝对值均小于1, 分离的连续性符合正态分布, 具有数量性状特点, 适合作进一步的QTL定位分析。

表2 亲本及RIL群体水、旱条件下芽期相关性状的表现 Table 2 Phenotypic performance of seed germination traits in RILs under water and drought conditions

2.4 QTL定位分析

2.4.1 正常水分(CK)条件下芽期相关性状QTL定位分别检测到发芽势、胚根长、胚芽鲜重、胚根鲜重、胚根干重和胚芽干重6个性状的19个QTL, 分别解释8.3%、20.7%、64.9%、26.2%、7.6%和22.1%的表型变异。其中, 1个发芽势QTL, 位于第5连锁群, 可解释表型变异的8.3%, 加性效应0.07, 增效基因来自K55。3个胚根长QTL, 分别位于第2、第6、第11连锁群上, 可解释表型变异的6.2%、6.6%和7.9%, 加性效应0.46、0.48和0.52, 增效基因均来自K55。6个胚芽鲜重QTL, 分别位于第2、第3、第5、第9连锁群, 可解释表型变异的6.2%、9.7%、25.8%、6.5%、10.1%和6.6%, 加性效应0.46、0.59、0.99、0.52、0.62和-0.59, 除Qefw6增效基因来自K58外, 其余QTL的增效基因均来自K55。5个胚根鲜重QTL, 分别位于第2、第3、第5、第6连锁群, 可解释表型变异的6.2%、9.0%、11.0%、6.8%和7.2%, 加性效应0.46、0.13、0.14、-0.11和0.11, 除Qrfw6增效基因来自K58外, 其余QTL的增效基因均来自K55。3个胚芽干重QTL, 分别位于第10、第11连锁群, 可解释表型变异的6.1%、9.8%和6.2%, 加性效应0.06、0.07和-0.06, Qedw10-1和Qedw10-2增效基因来自K55, Qedw11增效基因来自K58。1个胚根干重QTL, 位于第8连锁群, 可解释表型变异的7.6%, 加性效应0.02, 增效基因来自K55。

2.4.2 模拟干旱条件下芽期相关性状QTL定位

检测到发芽指数、发芽率、胚芽长、胚根长、胚芽鲜重和胚根鲜重6个性状的14个QTL, 分别解释8.6%、6.1%、49.9%、18.9%、10.8%和17.9%的表型变异。其中, 1个发芽指数QTL, 位于第8连锁群, 可解释表型变异的8.6%, 加性效应3.99, 增效基因来自K55; 1个发芽率QTL, 位于第9连锁群, 可解释表型变异的6.1%, 加性效应0.05, 增效基因来自K55; 5个胚芽长QTL, 分别位于第1、第2、第4、第8、第12连锁群, 可解释表型变异的7.5%、7.5%、9.8%、7.1%和8.0%, 加性效应-0.13、-0.13、-0.15、0.13和-0.14, 除Qel8增效基因来自K55外, 其余QTL的增效基因均来自K58。3个胚根长QTL, 分别位于第5、第7、第9连锁群, 可解释表型变异的6.2%、6.7%和14.0%, 加性效应0.18、-0.18和-0.26, Qrl5的增效基因来自K55, Qrl7、Qrl9增效基因来自K58。3个胚芽鲜重QTL, 分别位于第1、第2、第5连锁群, 可解释表型变异的7.4%、7.4%和13.1%, 加性效应-0.04、-0.04和-0.05, 增效基因来自K58。1个胚根鲜重QTL, 位于第10连锁群, 可解释表型变异的10.8%, 加性效应-0.01, 增效基因来自K58。

2.4.3 两种水分条件下QTL比较分析检测到的QTL中, 3个性状的5个QTL只在正常水分(CK)条件下被检测到, 包括胚根鲜重QTL Qrdw8、Qedw10-1、Qedw10-2、Qedw11和胚芽鲜重QTL Qge5, 占总QTL的15%; 3个性状的7个QTL只在18%PEG模拟干旱条件下被检测到, 包括胚芽长QTLQel1、Qel2、Qel4、Qel8、Qel12, 发芽指数QTLQgi8和发芽率QTLQgr9, 占总QTL的21%; 3个性状的21个QTL在两种条件下均被检测到, 包括胚芽鲜重QTL Qefw1、Qefw2-1、Qefw2-2、Qefw3、Qefw5-1~Qefw5-4、Qefw6, 胚根长QTL Qrl2、Qrl5、Qrl6、Qrl7、Qrl9、Qrl11, 胚根鲜重QTL Qrfw2、Qrfw3、Qrfw5、Qrfw6、Qrfw10、Qrfw12, 占总QTL的64%, 其中6个遗传贡献率超过10%的主效QTL分别是Qefw5-1 (25.8%)、Qefw 5-2 (13.1%)、Qefw5-4 (10.1%)、Qrfw5 (11.0%)、Qrfw10 (10.8%)、Qrl9 (14.0%), 占总QTL的18.2%。两种水分条件下未检测到位点一致QTL, 仅胚芽鲜重QTL (Qefw5-2、Qefw5-3)在水、旱两种条件下定位在第5连锁群Me4/Em8-3的相近位置。正常水分(CK)条件下检测到的19个QTL中, 16个QTL加性效应为正值, 表明多数的增效基因来自于K55, 干旱胁迫条件下检测到的14个QTL中, 10个QTL加性效应负值, 表明多数的增效基因来自于K58。检测到的33个QTL在第1、第2、第5和第8连锁群上有成簇分布现象。

2.4.4 差值QTL 共检测到9个干旱胁迫与正常水分条件差值的QTL, Qrl6对胚根长在干旱与对照条件下的差值有贡献, Qefw5-1、Qefw6对胚芽鲜重在两种条件下的差值有贡献, Qrfw2、Qrfw3、Qrfw5、Qrfw6、Qrfw12对胚根鲜重在两种条件下的差值有贡献, Qedw11对胚芽干重在两件条件下的差值有贡献, 其中K58等位基因在Qefw6、Qrfw6、Qedw11位点降低两种水分条件的差值, K55等位基因在Qrl6、Qefw5-1、Qrfw2、Qrfw3、Qrfw5、Qrfw12位点降低两种水分条件的差值即增加性状在不同环境下的稳定性。

表4 两种条件下芽期相关性状QTL定位结果 Table 4 QTL analysis of seed germination traits under two conditions

	Figure Option View Download New Window
	图1 向日葵遗传图谱及水、旱条件下芽期相关性状QTL分布 Fig. 1 Sunflower genetic linkage map and distribution of QTLs under water and drought conditions

3 讨论

3.1 多标记遗传图谱分析

本研究在前期用SSR和AFLP标记构建向日葵遗传图谱的基础上, 增加SRAP标记, 探讨多种标记结合构建遗传图谱的效果。SRAP标记主要对开放阅读框(ORF)扩增, 而对基因组相对较少的着丝粒附近以及端粒的扩增薄弱, SSR标记则可以对这些区域扩增, 并且通常用作锚定标记。AFLP标记利用酶切组合对DNA酶切产生多态性, 多态性高, 在图谱构建时通常与其他标记相结合, 对图谱加密^[20]。将3种分子标记相结合, 易获得均匀覆盖整个基因组的饱和连锁图^[21]。本试验利用SRAP、SSR及AFLP标记相结合构建的图谱覆盖度明显提高, 长度由原来的2759.4 cM^[11]增加到3846.0 cM, 增加了1086.6 cM, 位点由495个标记增加到1105个标记, 平均距离由5.57 cM缩短到3.48 cM, 偏分离比例从34.34%降低到31.68%。原图谱中有较多处分子标记聚集区域, 新构建图谱上分子标记分布较为均匀, 饱和度更高, 能够满足QTL定位要求。已有较多选用2~3个标记组合构建遗传连锁图谱的研究报道^{[21, 22, 23]}, 与本研究结果相同, 多数研究指出, 多个标记结合使用可以充分利用标记间互补优势, 更有利于构建标记均匀分布、高密度、高饱和性的分子遗传图谱。

3.2 QTL的稳定性

不同性状在不同环境下QTL的表达水平有所不同。本研究检测到的两种条件下9个性状的33个QTL中, 21%和15%分别在18%PEG干旱胁迫下和正常水分条件下特异表达, 64%在两种水分条件下均表达, 但没有检测到稳定表达的一致QTL, 说明本试验中不同水分处理下向日葵芽期相关性状QTL稳定性较差。多数研究认为, 只有在两种环境下表现相似的QTL可能对抗旱性有直接贡献, 而受干旱诱导的QTL 可能与耐旱性反应有关, 但未必对耐旱性有实质性贡献^{[24, 25, 26]}, 本研究中, 胚芽鲜重、胚根鲜重及胚根长3个性状的QTL在两种水分条件下均被检测到, 其中位于第5连锁群控制胚芽鲜重QTL Qefw5-1、Qefw 5-2、Qefw5-4, 位于第5和第10连锁群控制胚根鲜重QTL Qrfw5、Qrfw10以及第9连锁群控制胚根长QTL Qrl9遗传贡献率都大于10%, 这些受环境影响较小且效应值较大的主效QTL有望应用于芽期抗旱分子标记辅助选择。此外, 第5连锁群18%PEG胁迫下定位到的Qefw5-2与正常水分下定位到的Qefw5-3位点在位置上非常接近, 有待于探讨是否为同一位点。另外检测到的9个QTL包括5个RFW(Qrfw2、Qrfw3、Qrfw5、Qrfw6、Qrfw12)、2个EFW (Qefw5-1、Qefw6)、1个RL (Qrl6)、1个EDW (Qedw11), 对各芽期性状在2种条件下的差值有贡献, 研究表明, 这些QTL能够解释干旱胁迫所造成表型变化的净值, 对抗旱有直接贡献^[24]。

3.3 QTL的“ 成簇” 分布

QTL的“ 成簇” 分布现象在动植物中普遍存在, 本研究检测到6处QTL“ 成簇” 分布区域。第1连锁群控制胚芽长的Qel1、控制胚芽鲜重的Qefw1均位于P35-M78-8~P39-M83-4标记区间且与P42-M83-4紧密连锁, 说明该区间可能存在控制胚芽生长的QTL; 第2连锁群控制胚根长的Qrl2、控制胚芽鲜重的Qefw2-1、控制胚根鲜重的Qrfw2均位于ORS547-1-P28-M78-11标记区间且与Me7/Em3-2紧密连锁, 说明该区间可能存在同时控制胚芽、胚根生长的QTL, 控制胚芽长的Qel2、控制胚芽鲜重的Qefw2-2均位于Ha1360-3-P38-M79-2标记区间且与P39-M85-4紧密连锁, 说明该区间可能存在控制胚芽生长的QTL; 第5连锁群控制发芽势的Qge5、控制胚根长的Qrl5均位于Me14/Em7-1-P33-M76-2标记区间且与Me3/Em7-3紧密连锁, 说明该区间可能存在控制发芽势及胚根生长的QTL, 控制胚芽鲜重的Qefw5-1、控制胚根鲜重的Qrfw5均位于Me12/Em1-3-Me1/Em12-4标记区间且均为贡献率大于10%的主效QTL, 表明该区间可能存在效应值较大的多效性主效QTL。第8连锁群控制胚芽长的Qel8、控制发芽指数的Qgi8均位于P38-M79-7- Me8/Em7-5标记区间, 说明该区间可能存在控制发芽指数及胚根生长的QTL。QTL“ 成簇” 分布的现象从遗传学角度通常表现为一因多效、基因连锁或重叠, 可能由于一些关键因子的存在而同时控制多种代谢途径, 从而表现出多效性, 使相关性状在自然选择过程中形成相互适应的有利性状^[27]。下一步对这些成簇分布的QTL精细定位, 在分子标记辅助选择时可以同时对多个性状进行遗传改良, 提高抗旱育种的选择效率, 但也应注意可能存在遗传累赘, 尽量打破不利连锁。

4 结论

利用K55× K58的RIL群体构建了覆盖向日葵基因组3846.0 cM的分子遗传连锁图谱, 图谱共有17个连锁群, 1105个标记位点, 平均图距为3.5 cM。每个连锁群上标记数在10~165个之间, 长度在147.6~295.5 cM之间。在两种水分条件下对发芽率、发芽势、发芽指数、胚芽长、胚根长、胚芽鲜/干重和胚根鲜/干重9个芽期相关性状进行了QTL定位及分析, 共检测到33个QTL, 包括18%PEG模拟干旱胁迫条件下6个性状的14个QTL, 正常水分(CK)条件下6个性状的19个QTL。检测到Qefw 5-1、Qefw 5-2、Qefw 5-4、Qrfw 5、Qrfw 10和Qrl9共6个主效QTL, 同时检测到9个差值QTL, 分别为Qrl 6、Qefw 5-1、Qefw 6、Qrfw 2、Qrfw 3、Qrfw 5、Qrfw 6、Qrfw 12和Qedw 11, 对抗旱性有直接贡献。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

参考文献

View Option

[1]	刘天鹏, 董孔军, 何继红, 任瑞玉, 张磊, 杨天育. 糜子育成品种芽期抗旱性鉴定与评价研究. 植物遗传资源学报, 2014, 15: 746-752 Liu T P, Dong K J, He J H, Ren R Y, Zhang L, Yang T Y. Identification and evaluation on the drought resistance of broomcorn millet bred cultivars at germinating stage. J Plant Genet Resour, 2014, 15: 746-752 (in Chinese with English abstract) [本文引用:1]
[2]	Al-Chaarani G R, Gentzbittel L, Wedzony M, Sarrafi A. Identification of QTLs for germination and seedling development in sunflower (Helianthus annuus L. ). Plant Sci, 2005, 6: 1-7 [本文引用:1]
[3]	Davar R, Majd A, Darvishzadeh R, Sarrafi A. Mapping quantitative trait loci for seedling vigour and development in sunflower (Helianthus annuus L. ) using recombinant inbred line population. Plant Omics J, 2001, 4: 418-427 [本文引用:1]
[4]	袁倩倩, 李卓坤, 田纪春, 韩淑晓. 不同水分胁迫下小麦胚芽鞘和胚根长度的QTL分析. 作物学报, 2011, 37: 294-301 Yuan Q Q, Li Z K, Tian J C, Han S X. QTL Mapping for coleoptile length and radicle length in wheat under different simulated moisture stresses. Acta Agron Sin, 2011, 37: 294-301 (in Chinese with English abstract) [本文引用:1]
[5]	Bernardo R. Molecular markers and selection for complex traits in plants: learning from the last 20 years. Crop Sci, 2008, 48: 1649-1664 [本文引用:1]
[6]	Rieseberg L H, Choi H, Chan R. Genomic map of a diploid hybrid species. Heredity, 1993, 70: 285-293 [本文引用:1]
[7]	Tang S, Yu J K, Slabaugh M B, Shintani D K, Knapp S J. Simple sequence repeat map of the sunflower genome. Theor Appl Genet, 2002, 105: 1124-1136 [本文引用:1]
[8]	Bowers J E, Bachlava E, Brunick R L, Rieseberg L H, Knapp S J, Burke J M. Development of a 10000 locus genetic map of the sunflower genome based on multiple crosses. Genes Genom Genet, 2012, 2: 721-729 [本文引用:1]
[9]	Talukder Z I, Gong L, Hulke B S, Pegadaraju V, Song Q J, Schultz Q, Qi L. A high-density SNP map of sunflower derived from RAD-sequencing facilitating fine-mapping of the rust resistance gene R12. PLoS One, 2014, 7: 1-14 [本文引用:1]
[10]	黄先群, Genzbitelle L, Fabre F, Saraffi A. SSR分子标记丰富向日葵(Helianthus annuus L. )遗传图谱的研究. 西南农业学报, 2012, 25: 1031-1038 Huang X Q, Genzbitelle L, Fabre F, Saraffi A. Indication of genetic linkage map for sunflower by SSR markers. Southwest China J Agric Sci, 2012, 25: 1031-1038 (in Chinese with English abstract) [本文引用:1]
[11]	张永虎, 于海峰, 侯建华, 李素萍, 吕品, 于志贤. 利用向日葵重组自交系构建遗传图谱. 遗传, 2014, 36: 1036-1042 Zhang Y H, Yu H F, Hou J H, Li S P, Lyu P, Yu Z X. Construction of a genetic map of sunflower using a population of recombinant inbred lines (RILs), Hereditas, 2014, 36: 1036-1042 (in Chinese with English abstract) [本文引用:3]
[12]	Abdi N, Darvishzadeh R, Jafari1 M, Pirzad A, Haddadi P. Genetic analysis and QTL mapping of agro-morphological traits in sunflower (Helianthus annuus L. ) under two contrasting water treatment conditions. Plant Omics J, 2012, 5: 149-158 [本文引用:1]
[13]	Haddadi P, Yazdi-samadi B, Naghavi M R, Kalantari A, Maury P, Sarrafi A. QTL analysis of agronomic traits in recombinant inbred lines of sunflower under partial irrigation. Plant Biotechnol Rep, 2011, 5: 135-146 [本文引用:1]
[14]	Adiredjo A L, Navaud O, Munos S, Langlade N B, Lamaze T, Grieu P. *Genetic control of water use efficiency and leaf carbon isotope discrimination in sunflower ( Helianthus annuuL. ) subjected to two drought scenarios. PLoS One* , 2014, 101371/journal. pone. 0101218 [本文引用:1]
[15]	陈雪, 于海峰, 侯建华, 安玉麟. 向日葵芽期、苗期抗旱性鉴定方法研究. 中国油料作物学报, 2009, 31: 344-348 Chen X, Yu H F, Hou J H, An Y L. Drought resistance of sunflower in bud and seedling emergences stage. Chin J Oil Crop Sci, 2009, 31: 344-348 (in Chinese with English abstract) [本文引用:1]
[16]	Paniego N, Echaide M, Muñoz M, Fernand ez L, Torales S, Faccio P, Fuxan I, Carrera M, Zand omeni R, Suarez E Y, Hopp H E. Microsatellite isolation and characterization in sunflower (Helianthus annuus L. ). Genome, 2002, 45: 34-43 [本文引用:1]
[17]	Li G, Quiros C F. Sequence-related amplified polymorphism (SRAP), a new marker system based on a simple PCR reaction: its application to mapping and gene tagging in Brassica. Theor Appl Genet, 2001, 103: 455-461 [本文引用:1]
[18]	房冬梅, 吕品, 侯建华. 油葵SSR-PCR反应体系的优化及引物筛选. 中国农学通报, 2015, 31: 205-209 Fang D M, Lyu P, Hou J H. Optimization of SSR-PCR reaction system and primer screening in oil sunflower. Chin Agric Sci Bull, 2001, 103: 455-461 (in Chinese with English abstract) [本文引用:1]
[19]	于志贤, 侯建华. 油葵SRAP-PCR反应体系的建立与优化. 中国农学通报, 2013, 29: 296-301 Yu Z X, Hou J H. Establishment and optimization of SRAP-PCR reaction system in sunflower. Chin Agric Sci Bull, 2013, 29: 296-301 (in Chinese with English abstract) [本文引用:1]
[20]	昝逢刚, 吴才文, 陈学宽, 赵培方, 赵俊, 刘家勇. 118份甘蔗种质资源遗传多样性的AFLP分析. 作物学报, 2014, 40: 1877-1883 Zan F G, Wu C W, Chen X K, Zhao P F, Zhao J, Liu J Y. Genetic diversity of 118 sugarcane germplasm using AFLP markers. Acta Agron Sin, 2014, 40: 1877-1883 (in Chinese with English abstract) [本文引用:1]
[21]	李媛媛, 沈金雄, 王同华, 傅廷栋, 马朝芝. 利用SRAP、SSR和AFLP标记构建甘蓝型油菜遗传连锁图谱. 中国农业科学, 2007, 40: 1118-1126 Li Y Y, Shen J X, Wang T H, Fu T D, Ma C Z. Construction of a linkage map using SRAP, SSR and AFLP markers in Brassica napus L. Sci Agric Sin, 2007, 40: 1118-1126 (in Chinese with English abstract) [本文引用:2]
[22]	吴传书, 王丽侠, 王素华, 陈红霖, 吴健新, 程须珍, 杨晓明. 绿豆高密度分子遗传图谱的构建. 中国农业科学, 2014, 47: 2088-2098 Wu C S, Wang L X, Wang S H, Chen H L, Wu J X, Cheng X Z, Yang X M. Construction of a genetic linkage map in mungbean. Sci Agric Sin, 2014, 47: 2088-2098 (in Chinese with English abstract) [本文引用:1]
[23]	王茂芊, 李博, 王华忠. 甜菜遗传连锁图谱初步构建. 作物学报, 2014, 40: 222-230 Wang M Q, Li B, Wang H Z. Construction of molecular genetic linkage map of sugarbeet. Acta Agron Sin, 2014, 40: 222-230 (in Chinese with English abstract) [本文引用:1]
[24]	赵秀琴, 朱苓华, 徐建龙, 黎志康. 灌溉与自然降雨条件下水稻高代回交导入系产量QTL的定位. 作物学报, 2007, 33: 1536-1542 Zhao X Q, Zhu L H, Xu J L, Li Z K. QTL Mapping of yield under irrigation and rainfed field conditions for advanced backcrossing introgression lines in rice. Acta Agron Sin, 2007, 33: 1536-1542 (in Chinese with English abstract) [本文引用:2]
[25]	彭勃, 王阳, 李永祥, 刘成, 刘志斋, 王迪, 谭巍巍, 张岩, 孙宝成, 石云素, 宋燕春, 王天宇, 黎裕. 不同水分环境下玉米产量构成因子及籽粒相关性状的QTL分析. 作物学报, 2010, 36: 1832-1842 Peng B, Wang Y, Li Y X, Liu C, Liu Z Z, Wang D, Tan W W, Zhang Y, Sun B C, Shi Y S, Song Y C, Wang T Y, Li Y. QTL Analysis for yield components and kernel-related traits in maize under different water regimes. Acta Agron Sin, 2010, 36: 1832-1842 (in Chinese with English abstract) [本文引用:1]
[26]	谭巍巍, 李永祥, 王阳, 刘成, 刘志斋, 彭勃, 王迪, 张岩, 孙宝成, 石云素, 宋燕春, 杨德光, 王天宇, 黎裕. 在干旱和正常水分条件下玉米穗部性状QTL分析. 作物学报, 2011, 37: 235-248 Tan W W, Li Y X, Wang Y, Liu C, Liu Z Z, Peng B, Wang D, Zhang Y, Sun B C, Shi Y S, Song Y C, Yang D G, Wang T Y, Li Y. QTL mapping of ear traits of maize under different water regimes. Acta Agron Sin, 2008, 34: 1199-1205 (in Chinese with English abstract) [本文引用:1]
[27]	吴亚辉, 陶星星, 肖武名, 郭涛, 刘永柱, 王慧, 陈志强. 水稻穗部性状的QTL分析. 作物学报, 2014, 40: 214-221 Wu Y H, Tao X X, Xiao W M, Guo T, Liu Y Z, Wang H, Chen Z Q. Dissection of QTLs for panicle traits in rice (Oryza sativa). Acta Agron Sin, 2014, 40: 214-221 (in Chinese with English abstract) [本文引用:1]

2014

0.0

刘天鹏, 董孔军, 何继红, 任瑞玉, 张磊, 杨天育. 糜子育成品种芽期抗旱性鉴定与评价研究. 植物遗传资源学报, 2014, 15: 746-752

Liu T

, Dong K

, He J

, Ren R

, Zhang

, Yang T

Identification and evaluation on the drought resistance of broomcorn millet bred cultivars at germinating stage.

J Plant Genet Resour, 2014, 15: 746-752 (in Chinese with English abstract)

Abstract：Drought resistance at germinating stage of 56 Broomcorn Millet selective bred varieties from different province were identified using PEG-6000 treated, and drought resistance also were evaluated by direct and comprehensive assessment methods. The result showed that 56 varieties existed great differences in investigated traits and drought resistance between 20%(w/v) PEG-6000 treatment and CK. By two kinds of methods , three varieties with strong drought resistance have screened，namely Jinshu No.2, Longmi No.5, Longmi No.8. The result also showed that relative germination rate，relative shoot dry weight, relative root dry weight and storage matter relative transfer rate can be used as the main drought resistance identification indexes of broomcorn millet at germinating stage；Relative germination rate and weighted subordinate function value method are an ideal method of drought resistance evaluation at broomcorn millet germinating stage.

摘要：采用PEG-6000胁迫处理对国内不同地区育成的56个糜子品种的芽期抗旱性进行了鉴定，并用直接评价和综合评价法对其抗旱性进行了评价。结果表明：经20%（w/v）PEG-6000处理的56个品种与对照相比性状之间差异显著，品种抗旱性存在较大差异。用直接评价法和综合评价法相结合鉴定出强抗旱性品种3份，分别是晋黍2号、陇糜5号和陇糜8号。相对发芽率、相对芽干重、相对根干重、贮藏物质相对转运率4项指标可作为糜子芽期抗旱性鉴定评价指标，相对发芽率法和加权隶属函数值法是比较理想的糜子芽期抗旱性评价方法。

... 发芽出苗期作为向日葵生长发育的第一阶段, 芽期抗旱性直接关系到出苗情况, 进而影响后续的生长发育及产量^[1], 发芽率、发芽势、发芽指数、胚芽、胚根的长度、鲜重、干重均为衡量种子萌发状况的主要指标, 与芽期抗旱性密切相关^[2,3,4] ...

2005

0.0

2001

0.0

2011

0.0

袁倩倩, 李卓坤, 田纪春, 韩淑晓. 不同水分胁迫下小麦胚芽鞘和胚根长度的QTL分析. 作物学报, 2011, 37: 294-301

Yuan Q

, Li Z

, Tian J

, Han S

QTL Mapping for coleoptile length and radicle length in wheat under different simulated moisture stresses.

Acta Agron Sin, 2011, 37: 294-301 (in Chinese with English abstract)

Coleoptile length and Radicle length are the important indexes to evaluate resistance of wheat ( Triticum aestivum L.) seedlings to stresses. For mapping quantitative trait loci (QTLs) for lengths of coleoptile and Radicle in wheat, a set of immortalized F 2 (IF 2 ) population (168 lines) from Huapei 3 ´ Yumai 57 double haploid (DH) lines was treated with distilled water (normal condition) and 10%, 20%, and 30% of polyethylene glycol (PEG-6000). The coleoptile length (CL) and radicle length (RL) of the parents and the 168 IF 2 lines were measured after 7 d of treatment. QTLs for CL and RL were detected using 323 SSR markers, which were distributed in the whole genome of wheat. Based on inclusive composite interval mapping (ICIM) method, we identified 11 additive QTLs for CL and 12 additive QTLs for RL under normal and the three stress conditions. Each locus explained 4.93%–35.37% of phenotypic variance. In the interval between Xcfd39.2and Xcfd22.2on chromosome 4B, QTL QCl4B had the phenotypic contribution of 35.37%. Another QTL QCl3D-a located between Xcfd223 and Xbarc323 on chromosome 3D was detected in both normal and 20% PEG-6000 treatments, and explained phenotypic variances of 7.83% and 11.74%, respectively. QTL QCl3D-b was located on the same chromosome and close to QCl3D-a . In the linkage groups 1A and 5A1, three and two QTLs associated with RL were detected respectively. On chromosome 6D, two QTLs for CL and RL were found in the interval between Xswes679.1 and Xcfa2129 and the interval between Xwmc412.1and Xcfd49, respectively. The major QTLs identified can be applicable in marker-assisted selection in wheat breeding for coleoptile and root.

小麦胚芽鞘和胚根在不同渗透溶液下的长度变化是鉴评小麦幼苗抗逆性的重要指标。以小麦花培3号×豫麦57的DH株系衍生的含168个组合的永久F 2 (immortalized F 2 , IF 2 )群体为材料，在蒸馏水(正常条件)以及10%、20%和30%聚乙二醇(PEG-6000)模拟水分胁迫处理下，进行胚芽鞘长和胚根长度的数量性状基因(QTL)定位分析。利用完备区间作图法，共检测到影响胚芽鞘和胚根长度的23个QTL，单个QTL对表型的贡献率为4.93%~35.37%。位于4B染色体区间Xcfd39.2–Xcfd22.2上影响胚芽鞘长度的位点 QCl4B ，具有最大的遗传效应，贡献率为35.37%；在3D染色体Xcfd223–Xbarc323区段，正常条件和20% PEG-6000处理下同时检测到影响胚芽鞘长度的QTL ， QCl3D-a ，其贡献率分别为7.83%和11.74%。另外，在10% PEG-6000处理下，3D染色体上的相近区域还定位出了影响胚芽鞘长度的 QCl3D-b 位点；在染色体1A和染色体5A1上各检测出与胚根长度有关的2个和3个不同的QTL；在6D染色体Xswes679.1–Xcfa2129和Xwmc412.1–Xcfd49区间分别检测到2个影响胚芽鞘长度和胚根长度的QTL。这些主效QTL可用于胚芽鞘和根系的分子标记辅助选择。

2008

0.0

... 作物抗旱性属多基因控制的复杂性状, 分子标记技术及QTL定位的发展为复杂数量性状的研究及遗传改良提供了有力的工具^[5] ...

1993

0.0

... 高密度分子遗传连锁图谱的构建是进行QTL定位的基础, 自第一张向日葵图谱使用RAPD标记构建完成以来^[6], RFLP、AFLP、SSR等标记被相继成功应用于向日葵遗传连锁图谱构建, 其中较具参考价值的是Tang等^[7]利用F₇重组自交系群体构建的含有459个SSR标记、平均距离3 ...

2002

0.0

2012

0.0

... 以SNP标记为代表的第3代分子标记用于向日葵图谱构建方面, John等^[8]利用4个群体构建了一张含有10 080个SNP标记的向日葵遗传连锁图谱, Zahirul等^[9]利用3个F₂群体构建一张向日葵遗传连锁图谱, 该图谱包含SNP标记5019个、SSR标记118个 ...

2014

0.0

2012

0.0

黄先群, Genzbitelle

, Fabre

, Saraffi

SSR分子标记丰富向日葵(Helianthus annuus L. )遗传图谱的研究. 西南农业学报, 2012, 25: 1031-1038

Huang X

, Genzbitelle

, Fabre

, Saraffi

Indication of genetic linkage map for sunflower by SSR markers.

Southwest China J Agric Sci, 2012, 25: 1031-1038 (in Chinese with English abstract)

为了提高向日葵遗传图谱的密度和实用性,以125个来源于 PAC-2和RHA-266杂交的F(8)代重组自交系(RIIs)群体为材料,利用筒单序列重复(Simple sequence repeat,SSR)标记,采用MAPMARKER软件对向日英遗传图谱进行标注,并从300对SSR引物中筛选出51对多态性引物对群体进行标记.结果表明:①51对多态性引物中有19对引物无多态性或条带不清晰,32对引物表现多态性;②共检测到35个多态性位点,分布在图谱的15条连锁群上.③标记后的图谱总长度为2914.5 Cm,比原来的图谱增长7.5 Cm.④标记间平均距离由9.0 Cm缩短为8.1 Cm.

... 国内向日葵图谱构建研究进展缓慢, 黄先群等^[10]于2012年利用SSR标记加密了遗传图谱, 本实验室利用SSR和AFLP标记构建了一张包含495个标记、平均距离5 ...

2014

0.0

张永虎, 于海峰, 侯建华, 李素萍, 吕品, 于志贤. 利用向日葵重组自交系构建遗传图谱. 遗传, 2014, 36: 1036-1042

Zhang Y

, Yu H

, Hou J

, Li S

, Lyu

, Yu Z

Construction of a genetic map of sunflower using a population of recombinant inbred lines (RILs),

Hereditas, 2014, 36: 1036-1042 (in Chinese with English abstract)

A genetic linkage map of sunflower was constructed by combined applying the SSR and AFLP markers using 187 F 5:6 individuals of recombinant inbred lines (RILs) which derived from the cross between Helianthus annuus K55 and Helianthus annuus K58 through single-seed descent (SSD). Using 78 pairs of SSR primers and 48 pairs of AFLP primer, 341 and 1119 bands were amplified, respectively. Among these 1460 bands, 557 bands (39.52%) were polymorphic, including 184 bands by SSR markers and 393 bands by AFLP markers. In the group of these polymorphic bands, 84 bands from SSR markers and 108 bands from AFLP markers showed the genetic distortion ( P = 0.05). A total of 192 segregation distortion markers were obtained in this study. By using the JoinMap 4.0 software to do the linkage analysis, a genetic linkage map was established with length of 2759.4 cM, consisted of 17 linkage groups, and comprised of 495 polymorphic molecular markers including 170 segregation distortion markers. The mean marker interval distance is 5.57 cM between markers. In addition, the number of markers in the linkage groups varied from 5 to 72, and the length of linkage groups were from 68.88 cM to 250.17 cM. The genetic map developed in the present study could be used for QTL mapping and gene cloning of sunflower important genes.

以向日葵自选系K55为母本、K58为父本杂交组合,通过单粒传得到的187个F 5：6 代重组自交系群体为作图材料,联合应用SSR和AFLP标记构建遗传连锁图谱。经过78对SSR引物和48对AFLP引物组合选择性扩增,分别得到341和1119条带,共1460条,分别获得多态性条带184条和393条,共577条多态性条带,占所有条带的39.52%。SSR和AFLP标记各有84个和108个多态性标记偏离孟德尔分离比例( P =0.05),共192个偏分离标记。采用JoinMap4.0软件进行连锁分析,构建了1张总长度为2759.4 cM、包含17个连锁群、连锁495个多态性标记的遗传图谱,其中偏分离标记170个,标记间的平均图距为5.57 cM。每个连锁群上分布有5~72个标记,长68.88~250.17 cM。本图谱为向日葵永久性图谱,为向日葵重要性状QTL定位和基因克隆奠定基础。

... 4 cM的遗传连锁图谱^[11] ...

... 本研究拟利用SRAP和SSR标记整合本实验室已有的SSR和AFLP标记^[11], 构建一张高密度遗传连锁图谱, 结合2种水分条件下芽期9个性状的表型值, 进行QTL定位分析, 为解析向日葵水、旱条件下芽期相关性状QTL及遗传效应和分子标记辅助育种奠定坚实的研究基础 ...

... 4 cM^[11]增加到3846 ...

2012

0.0

... 向日葵相关性状QTL的定位研究来自国外的较多, 不同水分条件下的株高、茎粗、开花期、水分利用率等性状的QTL均被定位到图谱上^[12,13,14], 但有关不同水分条件下芽期性状QTL定位研究国内外鲜有报道 ...

2011

0.0

2014

0.0

2009

0.0

陈雪, 于海峰, 侯建华, 安玉麟. 向日葵芽期、苗期抗旱性鉴定方法研究. 中国油料作物学报, 2009, 31: 344-348

Chen

, Yu H

, Hou J

, An Y

Drought resistance of sunflower in bud and seedling emergences stage.

Chin J Oil Crop Sci, 2009, 31: 344-348 (in Chinese with English abstract)

通过室内发芽试验和室外盆栽试验对11个抗旱性不同的油用向日葵育种材料进行研究,测定与抗旱性有关的相对发芽指数、萌发抗旱指数和株高等14项鉴定指标。以种子萌发指数与幼苗成活率的乘积作为灰色关联度分析的参考数列X0,运用灰色关联度理论,分析各指标与苗期抗旱性的关系,从中筛选出关联度较大的相对发芽率、相对发芽指数、芽相对生长量、根相对生长量、苗期株高、鲜重、相对电导率、相对含水量8个指标,然后通过隶属函数法对抗旱性进行分析,定量地评价11个油葵育种材料芽苗期的抗旱性。本研究为油葵芽苗期抗旱性鉴定提供了一套科学的评价方法,对油葵抗旱育种有一定的实践意义和参考价值。

... 2 表型性状测定按照陈雪等^[15]的方法, 设置正常水分(CK)和模拟干旱胁迫(18%聚乙二醇PEG-6000) 两种水分条件, 调查9个芽期性状 ...

2002

0.0

... gov/), Ha系列来自Paniego等^[16]的文献 ...

2001

0.0

... SRAP引物来自于Li和Quiros等^[17]的文献, 引物由上海生工生物工程有限公司合成 ...

2015

0.0

房冬梅, 吕品, 侯建华. 油葵SSR-PCR反应体系的优化及引物筛选. 中国农学通报, 2015, 31: 205-209

Fang D

, Lyu

, Hou J

Optimization of SSR-PCR reaction system and primer screening in oil sunflower.

Chin Agric Sci Bull, 2001, 103: 455-461 (in Chinese with English abstract)

摘　要：为了获得适合油葵的SSR-PCR反应体系,通过采用控制变量法对影响SSR反应体系的主要因素进行优化,研究Mg^2＋浓度、d NTPs浓度、Taq酶浓度、引物浓度、退火温度等因素。结果表明各因素的终浓度分别在Mg2＋浓度2.5 mmol/L、d NTPs浓度0.2 mmol/L、Taq酶量0.5 U、引物浓度0.4μmol/L及退火温度为53.9℃时最为适合。通过试验优化后的体系筛选引物,最终从50对引物筛得19对多态性好的引物可进行下一步的研究。

... 参照房冬梅等^[18]和于志贤等^[19]改进的PCR体系及程序 ...

2013

0.0

于志贤, 侯建华. 油葵SRAP-PCR反应体系的建立与优化. 中国农学通报, 2013, 29: 296-301

Yu Z

, Hou J

Establishment and optimization of SRAP-PCR reaction system in sunflower.

Chin Agric Sci Bull, 2013, 29: 296-301 (in Chinese with English abstract)

In this study, one-factor experimental design had been used to establish the SRAP-PCR reaction system of sunflower, which including several different gradients of Mg2+, dNTPs, primer concentration, Taq DNA Polymerase, template DNA and filter out the suitable range, on this basis, then optimize the five factors which influence the SRAP-PCR through the orthogonal experimental design of L16(45). Eventually brought about an optimized system for sunflower, which was as follows: 20 μL capacity of reaction system contains 10×Buffer 2 μL, Mg2+ 2.75 mmol/L, dNTPs 0.18 mmol/L, Taq DNA Polymerase 1.25 U, forward primer and reverse primer 0.3 μmol/L, template DNA 60 ng, the best annealing temperature 52.2℃. Finally, 22 varieties of sunflower were used to test the optimized PCR reaction system stability. The results showed that, the bands of the amplification clear and high polymorphism, so the system was reliable which could be applied in the germplasm identification and the genetic map construction of sunflower effectively.

为建立油葵SRAP-PCR的反应体系，采用单因素试验法，对Mg2+、dNTPs、引物浓度、Taq DNA聚合酶、模板DNA分别设置5~7个水平梯度，筛选出适宜的用量范围，以此为基础，再通过L16(45)正交试验设计，对影响SRAP-PCR的5个因素进行优化，建立了油葵SRAP-PCR的最佳反应体系：20 μL体系中含10×Buffer 2 μL，Mg2+ 2.75 mmol/L，dNTPs 0.18 mmol/L，Taq DNA聚合酶1.25 U，正反引物各0.3 μmol/L，模板DNA 60 ng，最佳退火温度为52.2℃。用22份油葵材料对该体系进行验证，结果显示扩增条带清晰、多态性高，说明该体系稳定可靠，可有效的用于油葵种质资源的鉴定、遗传图谱构建等研究。

... 参照房冬梅等^[18]和于志贤等^[19]改进的PCR体系及程序 ...

2014

0.0

昝逢刚, 吴才文, 陈学宽, 赵培方, 赵俊, 刘家勇. 118份甘蔗种质资源遗传多样性的AFLP分析. 作物学报, 2014, 40: 1877-1883

Zan F

, Wu C

, Chen X

, Zhao P

, Zhao

, Liu J

Genetic diversity of 118 sugarcane germplasm using AFLP markers.

Acta Agron Sin, 2014, 40: 1877-1883 (in Chinese with English abstract)

AFLP markers were used to assess the genetic diversity of 118 sugarcane germplasm from China, Australia, USA, Philippines, Brazil, and France. A total of 1310 bands were amplified with ten pairs of primers, 1195 of which were polymorphic bands, the percentage of polymorphic loci was 91.22%. The genetic similarity coefficient among 118 sugarcane germplasm ranged from 0.583 to 0.929 with an average of 0.750, the polymorphism information content was 0.2332, and the effective number of alleles per loci was 1.3789. The results showed 118 germplasm were divided into two groups at 0.69 of the similarity coefficient. The group I consisted of YT00-236, YZ04-622 and SP80-1816, which shared lower genetic similarity with the rest of 115 germplasm in another group . The 115 germplasm were divided into five sub-groups at 0.74 of the similarity coefficient. The sugarcane germplasm shared high genetic similarity among countries, and there existed much higher genetic diversity within germplasm introduced from USA.

采用 AFLP 标记技术，对来自中国、澳大利亚、美国、菲律宾、巴西和法国的 118 份甘蔗种质进行遗传多样性分析。采用 10 对引物组合共扩增出 1310 条谱带，其中多态性条带为 1195 条，多态性条带比率为 91.2% 。 118 份种质间的相似性系数在 0.583~0.929 之间，平均为 0.750 ，多态信息量为 0.2332 ，每个位点的有效等位基因数为 1.3789 。结果显示在相似性系数 0.69 处切割时，可划分为 2 个类群，第 I 类群包括粤糖 00-236 、云蔗 04-622 和 SP80-1816 ；第 II 类群有 115 份种质，在相似性系数 0.74 处切割时，可将第 II 类群划分为 5 个亚群。各国甘蔗种质亲缘关系较近，其中美国种质遗传多样性相对较丰富。

... AFLP标记利用酶切组合对DNA酶切产生多态性, 多态性高, 在图谱构建时通常与其他标记相结合, 对图谱加密^[20] ...

2007

0.0

李媛媛, 沈金雄, 王同华, 傅廷栋, 马朝芝. 利用SRAP、SSR和AFLP标记构建甘蓝型油菜遗传连锁图谱. 中国农业科学, 2007, 40: 1118-1126

Li Y

, Shen J

, Wang T

, Fu T

, Ma C

Construction of a linkage map using SRAP, SSR and AFLP markers in Brassica napus L.

Sci Agric Sin, 2007, 40: 1118-1126 (in Chinese with English abstract)

【目的】建立甘蓝型油菜的高质量遗传图谱,为在分子水平上研究复杂性状打下基础并提供保证。【方法】以甘蓝型油菜自交不亲和系“SI-1300”及其恢复系“Eagle”组配得到的184个F2单株为群体,利用SRAP、SSR和AFLP标记技术构建甘蓝型油菜遗传图谱。【结果】该图谱共包含21个连锁群,涉及137个SRAP标记、143个SSR标记和118个AFLP标记。图谱总长1949.8cM,标记间平均图距为4.9cM。以SSR标记为锚定标记,该图谱与国际标准图谱进行了初步对应。研究共发现了8个偏分离区域。【结论】根据对标记分布的均匀程度,图谱覆盖程度以及标记密度等方面的分析,本研究构建的甘蓝型油菜分子遗传图谱质量较高,并且SRAP标记可能是比AFLP标记更适合于图谱构建的标记体系。

... 将3种分子标记相结合, 易获得均匀覆盖整个基因组的饱和连锁图^[21] ...

... 已有较多选用2~3个标记组合构建遗传连锁图谱的研究报道^[21,22,23], 与本研究结果相同, 多数研究指出, 多个标记结合使用可以充分利用标记间互补优势, 更有利于构建标记均匀分布、高密度、高饱和性的分子遗传图谱 ...

2014

0.0

吴传书, 王丽侠, 王素华, 陈红霖, 吴健新, 程须珍, 杨晓明. 绿豆高密度分子遗传图谱的构建. 中国农业科学, 2014, 47: 2088-2098

Wu C

, Wang L

, Wang S

, Chen H

, Wu J

, Cheng X

, Yang X

Construction of a genetic linkage map in mungbean.

Sci Agric Sin, 2014, 47: 2088-2098 (in Chinese with English abstract)

【Objective】On the basis of previous studies, a genetic map of mungbean was constructed by using genome SSR, EST-SSR, STS primers of mungbean and common bean to build a platform for positioning important traits related genes, cloning and molecular marker-assisted breeding of new varieties of mungbean.【Method】A total of 6 686 SSR, EST-SSR, STS primers of mungbean and common bean, including 6 100 genome SSR, 149 EST-SSR, 13 STS primer pairs of mungbean, and 424 genome SSR primers of common bean, were used for PCR amplification to screen polymorphic markers between Australia-imported Berken (highly susceptible cultivar)×ACC41 (highly resistant wild species) and a RIL derived from these two genotypes were tested with the polymorphic markers. Combined with the molecular marker data of previous studies, Mapmarker/Exp 3.0 software was used for map construction and set LOD≥3.0, the maximum figure at 50.00 cM. Finally, Joinmap 4.0 software was used for map integration.【Result】In this study, from the two parents, 6 686 SSR primers were screened, a total of 3 691 pairs of primers were amplified stable products, 588 pairs of polymorphic primers were obtained. Among them, mungbean SSR primers 6 100 pairs, effective amplification 3 459 pairs, the effective rate of 56.7%, obtained 559 pairs of polymorphic primers; Mungbean EST-SSR primers 149 pairs, effective amplification 126 pairs, the effective rate of 84.6%, obtained 21 pairs of polymorphic primers; Common bean SSR primers 424 pairs, effective amplification 97 pairs, the effective rate of 22.9%, obtained 6 pairs of polymorphic primers; Mungbean STS primers 13 pairs, effective amplification 9 pairs, the effective rate of 69.2%, obtained 2 pairs of polymorphic primers; These results indicated that different sources and types of SSR primers to amplify the effective rate of the parent were significantly different. Mungbean EST-SSR primers (84.6%) were the highest, mungbean STS primers (69.2%) and mungbean SSR primers (55.7%) followed, common bean SSR primers (22.9%) were the lowest. An integrated genetic linkage map of mungbean containing 585 markers was constructed (including 499 SSR markers, 74 RAPD markers, 9 STS markers and 3 RAPD markers). The total length of the map was 732.9 cM and covered 11 linkage groups. The average distance between markers was 1.25 cM. The average distance of each linkage group spanned 66.63 cM. The average number of markers was 53.18 for each of 11 chromosomes. The length of each linkage group ranged from 35 to 92 markers was from 45.2 cM to 112.8 cM. LG1 linkage group contained the largest number of 92 markers, the length was 112.8 cM. LG11 linkage group contained the minimum number of 35 markers, the length was 48.7 cM. Of the 585 markers loci mapping conducted χ2 tests under P＜0.05 and P＜0.01 conditions, respectively, 79 and 151 markers showed a segregation distortion, the total number of sites marked 39.3%.【Conclusion】A linkage map of mungbean with a maximum and the highest density genetic markers was constructed compared with that published at home and abroad at present.

【目的】在前期研究的基础上，进一步利用绿豆基因组SSR、EST-SSR、STS和普通菜豆基因组SSR等标记构建绿豆遗传连锁图谱，为绿豆重要性状相关基因的定位、克隆及分子标记辅助选育新品种等研究搭建技术平台。【方法】利用澳大利亚引进的Berken（高感豆象绿豆栽培种）× ACC41（高抗豆象绿豆野生种）及其重组自交系（recombinant inbreed line，RIL）群体，对6 686对引物进行PCR扩增及多态性筛选，包括6 100对绿豆基因组SSR、149对EST-SSR、13对STS和424对普通菜豆基因组SSR引物，将亲本间多态性引物，进一步分析重组自交系群体。结合前期研究的分子标记数据，利用Mapmarker/Exp 3.0软件构建遗传图谱，并设置LOD≥3.0，最大图距50.00 cM。用Joinmap 4.0软件进行图谱整合。【结果】用2个亲本共筛选了6 686对SSR引物，共有3 691对引物有稳定的扩增产物，得到有多态的引物有588对。其中，通过磁珠富集法开发的绿豆SSR引物6 100对，有效扩增3 459对，有效扩增率56.7%，得到多态性引物559对；通过转录组测序开发的绿豆MGCP引物149对，有效扩增126对，有效扩增率84.6%，得到多态性引物21对；通过磁珠富集法开发的菜豆SSR引物424对，有效扩增97对，有效扩增率22.9%，得到多态性引物6对；绿豆STS引物13对，有效扩增9对，有效扩增率69.2%，得到多态性引物2对。表明不同来源和种类的SSR引物在RIL群体亲本中的有效扩增率有明显差别，绿豆EST-SSR引物（84.6%）最高，绿豆STS引物（69.2%）和SSR引物（55.7%）次之，菜豆SSR引物（22.9%）最低。获得一张含有585个标记（499个SSR标记、74个RFLP标记、9个STS标记和3个RAPD标记）的绿豆遗传图谱，图谱总长732.9 cM，包括11个连锁群，每个标记间的平均距离为1.25 cM，平均长度为66.63 cM。每个连锁群长度为45.2—112.8 cM，每条染色体上面的标记数为35—92个，平均53.18个。标记位点数最多的连锁群LG1含92个标记，长度为112.8 cM；标记位点数最少的连锁群LG11仅含有35个标记，长度为48.7 cM。对图谱的585个标记位点进行χ2测验，在P＜0.05和P＜0.01条件下，分别有79个和151个标记表现为偏分离，占总标记位点数的39.3%。【结论】构建了一张目前国内外发表的标记数最多、密度最高的绿豆遗传连锁图谱。

2014

0.0

王茂芊, 李博, 王华忠. 甜菜遗传连锁图谱初步构建. 作物学报, 2014, 40: 222-230

Wang M

, Li

, Wang H

Construction of molecular genetic linkage map of sugarbeet.

Acta Agron Sin, 2014, 40: 222-230 (in Chinese with English abstract)

Molecular genetic map is a fundamental tool for genomic research. In this study, a molecular genetic map was constructed using an F 2 population consisting of 200 individuals from a cross between sugarbeet JV34-2 (the high beet yield and low sugar content line) and sugarbeet 2B023 (the low beet yield and high sugar content line) and a total of 561 SRAP primers and 114 SSR primers. Approximately 56 out of 561 and 20 out of 114 primers, respectively, showed polymorphisms. Each of these polymorphic primers produced at least one scorable polymorphic DNA band which was visible enough for detection and scoring. The map consisted of nine linkage groups which included 141 (123 SRAP and 18 SSR) markers, and covered 1399.88 cM with an average distance of 9.92 cM. Three markers were still unlinked. All nine linkage groups consisting of 3–26 markers were 15.69–237.21 cM in length. Twenty point five six percent partially segregated markers distributed in the map, were mainly located on Ch3 linkage groups, and the others were mapped on Ch1, Ch2, Ch8, and Ch9. This is the first gene map of sugarbeet, constructed by SRAP and SSR markers in China. It will provide the basis for gene mapping of important characters and cloning of excellent gene in sugarbeet.

以甜菜高产低糖型 JV34-2 和低产高糖型 2B023 两材料杂交 , 构建了 200 个单株的 F 2 作图群体 , 利用所筛选出的 56 对 SRAP 引物组合和 20 对 SSR 引物 , 对 F 2 作图群体进行 PCR 扩增和遗传连锁分析 , 初步构建了一张包含 9 个连锁群、 141 个 (123 个 SRAP 和 18 个 SSR) 标记位点的甜菜遗传连锁图谱。该图谱覆盖长度为 1399.88 cM , 平均图距 9.92 cM 。未进入连锁群的有 4 个标记。 9 个连锁群包含 3~26 个标记不等 , 连锁群遗传距离 15.69~237.21 cM 。连锁群上有 20.56% 的标记出现偏分离 , 主要集中在 Ch3 连锁群上 , 其余分散在 Ch1 、 Ch2 、 Ch8 和 Ch9 中。该图谱是我国甜菜领域利用 SRAP 和 SSR 相结合方法 , 构建的第一个较精密的分子遗传图谱 , 为重要性状的基因定位和优良基因的克隆奠定了基础。

2007

0.0

赵秀琴, 朱苓华, 徐建龙, 黎志康. 灌溉与自然降雨条件下水稻高代回交导入系产量QTL的定位. 作物学报, 2007, 33: 1536-1542

Zhao X

, Zhu L

, Xu J

, Li Z

QTL Mapping of yield under irrigation and rainfed field conditions for advanced backcrossing introgression lines in rice.

Acta Agron Sin, 2007, 33: 1536-1542 (in Chinese with English abstract)

In rice breeding community, drought tolerance (DT) is becoming one of the most important target traits for variety improvement under ever-increasing severe drought situation all over the world. Yield is the most important target focused by breeders, so identification of favorable alleles related to yield and yield components for introgression into rice varieties that suit the specific environments is an efficient practice in plant MAS breeding. In the present study, the QTL conditioning the grain yield per plant (GY) and its component traits including 1000-grain weight (GW) and filled grain number per panicle (GN) in irrigation(control) and rainfed (drought stress) fields were investigated using 254 advanced backcrossing introgression lines (IL) of Lemont with Teqing genetic background and the two parents. Total of 32 main-effect QTLs for GY, GW, and GN were identified in the two conditions which can be grouped into three types based on their behaviors. TypeⅠincluded 10 QTLs which were detected both in the two conditions, type Ⅱ consisted of 14 QTLs which were mapped only in control condition, and type Ⅲ comprised 8 QTLs which were induced by drought and detected only under stress. In addition, nine QTLs affecting trait differences between stress and control were identified ( QGn5 , QGn6 , QGn11a , QGw2 , QGw8 , QGw11a , QGw11b , QGy1 , and QGy11 ). There were three QTLs ( QGn11b , QGn12 , and QGn11b ) which expressed both in the two conditions with same direction and magnitude of gene effect. Therefore the three GN-QTLs and nine QTLs affecting trait differences were considered to directly contribute to drought tolerance (DT). Teqing alleles at the 12 DT-QTLs except QGn5 and QGy1 were associated with DT. As compared with the results from different mapping populations derived from the same parent, Lemont and Teqing, in different environments, seven QTLs including QGn3b , QGw1 , QGw5 , QGy1 , QGy5 , QGy8 , and QGy10 which stably expressed across different genetic backgrounds and environments were identified. The strategy of applying QTL mapping results in rice improvement for DT by marker-assisted selection was discussed.

利用254个Lemont导入到特青背景的高代回交导入系定位了灌溉（对照）与自然降雨（干旱胁迫）环境下影响单株籽粒产量及其穗部相关性状的QTL。在两种环境下共检测到32个影响单株粒籽产量、千粒重和每穗实粒数的主效QTL，根据不同环境下表达的情况将其分成3类，第1类10个QTL，在两种环境下均被检测到；第2类14个QTL，只在对照条件下检测到；第3类8个QTL，受干旱胁迫诱导，只在胁迫条件下被检测到。此外还检测到9个影响胁迫与对照条件下性状差值的QTL。认为在两种条件下均检测到的相对稳定的3个QTL（ QGn11b 、 QGn12 和 QGn11b ）及影响两种条件下性状差值（即性状稳定性）的9个QTL可能对耐旱性有直接贡献。在所有12个耐旱QTL中，除在 QGn5 和 QGy1 的Lemont等位基因减小性状差值（即增强耐旱性）外，其余位点上增强耐旱性的等位基因均来自特青。另外通过与源自相同亲本的不同定位群体在不同环境下定位结果的比较，鉴别出一些受遗传背景和环境影响较小的QTL如 QGn3b 、 QGw1 、 QGw5 、 QGy1 、 QGy5 、 QGy8 和 QGy10 。对应用QTL定位结果进行标记辅助选择培育耐旱品种进行了探讨。

... 多数研究认为, 只有在两种环境下表现相似的QTL可能对抗旱性有直接贡献, 而受干旱诱导的QTL 可能与耐旱性反应有关, 但未必对耐旱性有实质性贡献^[24,25,26], 本研究中, 胚芽鲜重、胚根鲜重及胚根长3个性状的QTL在两种水分条件下均被检测到, 其中位于第5连锁群控制胚芽鲜重QTL Qefw5-1、Qefw 5-2、Qefw5-4, 位于第5和第10连锁群控制胚根鲜重QTL Qrfw5、Qrfw10以及第9连锁群控制胚根长QTL Qrl9遗传贡献率都大于10%, 这些受环境影响较小且效应值较大的主效QTL有望应用于芽期抗旱分子标记辅助选择 ...

... 另外检测到的9个QTL包括5个RFW(Qrfw2、Qrfw3、Qrfw5、Qrfw6、Qrfw12)、2个EFW (Qefw5-1、Qefw6)、1个RL (Qrl6)、1个EDW (Qedw11), 对各芽期性状在2种条件下的差值有贡献, 研究表明, 这些QTL能够解释干旱胁迫所造成表型变化的净值, 对抗旱有直接贡献^[24] ...

2010

0.0

彭勃, 王阳, 李永祥, 刘成, 刘志斋, 王迪, 谭巍巍, 张岩, 孙宝成, 石云素, 宋燕春, 王天宇, 黎裕. 不同水分环境下玉米产量构成因子及籽粒相关性状的QTL分析. 作物学报, 2010, 36: 1832-1842

Peng

, Wang

, Li Y

, Liu

, Liu Z

, Wang

, Tan W

, Zhang

, Sun B

, Shi Y

, Song Y

, Wang T

, Li

QTL Analysis for yield components and kernel-related traits in maize under different water regimes.

Acta Agron Sin, 2010, 36: 1832-1842 (in Chinese with English abstract)

Maize yield and yield-related traits are seriously affected by water stress. Therefore, detecting quantitative trait locus (QTL) for yield components and kernel-related traits, analyzing the stability of QTLs and exploiting constitutive QTLs under different water regimes are of great importance in marker-assisted breeding for drought tolerance in maize. In this study two F 2:3 populations derived from Qi319×Huangzaosi (Q/H) and Ye478×Huangzaosi (Y/H) were used to investigate the genetic basis of yield components and kernel-related traits under different water regimes in Xinjiang (including well-water and water-stress environments) by stepwise joint QTL mapping method. The results showed that above 70% of the QTLs for yield components and kernel-related traits expressed stably under the same water regime across the two years. The QTLs detected in water-stress environments were less stable than those in well-water environments across the two years in Xinjiang. The joint analysis combining data of all environments indicated that the stability of the QTLs for all traits decreased, but above 60% of them still expressed stably. A total of 11 constitutive QTLs (with contribution rate more than10% in at least one environment, detected in more than two environments based on single environment analysis) distributed on bin1.10, 2.00, 4.09, 7.02, 9.02, 10.04 and 10.07 were detected in the two populations, and all of them except bin10.04 were stable across all environments. Consequently, most of the QTLs for yield components and kernel-related traits stably expressed under the same water regime across different years, and even under different water regimes in Xinjiang. These constitutive QTLs may provide references for molecular breeding and further basic studies.

干旱胁迫对玉米产量及其相关性状有重要影响。本研究以我国玉米育种骨干亲本齐319和掖478分别和黄早四组配构建的两个F 2:3 群体为材料，应用逐步联合分析的QTL定位方法，剖析新疆不同水分环境下(包含水区和旱区)玉米产量构成因子及籽粒相关性状的遗传基础。结果表明，在相同水分处理不同年份间产量构成因子和籽粒相关性状超过70%的QTL可稳定表达，旱区QTL的稳定性明显低于水区，当全部环境联合分析时，各性状QTL稳定性呈现一定程度的降低，但超过60%的QTL仍然稳定表达。两群体中共检测到11个环境钝感的主效QTL(在2个以上环境中检测到，且至少在一个环境下的贡献率大于10%)，分布在bin1.10、2.00、4.09、7.02、9.02、10.04和10.07共7个基因组区段上，除bin10.04外所有环境钝感的主效QTL在全部环境下稳定表达。因此，玉米产量构成因子和籽粒相关性状的QTL在新疆相同水分处理不同年份间，甚至不同水分条件下大部分均可稳定表达，这些主效QTL位点可为抗旱分子育种和进一步精细定位提供参考。

2011

0.0

谭巍巍, 李永祥, 王阳, 刘成, 刘志斋, 彭勃, 王迪, 张岩, 孙宝成, 石云素, 宋燕春, 杨德光, 王天宇, 黎裕. 在干旱和正常水分条件下玉米穗部性状QTL分析. 作物学报, 2011, 37: 235-248

Tan W

, Li Y

, Wang

, Liu

, Liu Z

, Peng

, Wang

, Zhang

, Sun B

, Shi Y

, Song Y

, Yang D

, Wang T

, Li

QTL mapping of ear traits of maize under different water regimes.

Acta Agron Sin, 2008, 34: 1199-1205 (in Chinese with English abstract)

Ear traits are closely associated with maize yield. Therefore, genetic dissection of ear traits can provide clues to maize high-yield breeding and is especially important in breeding for drought tolerance. In this study, seven ear related traits including ear length (EL), ear diameter (ED), row number (KRN), kernel number per row (KRE), grain yield per ear (GW), cob diameter (AD) and weight per ear (EW), were investigated using two sets of F 2:3 populations, which were derived from crosses of Ye478×Huangzaosi (Y/H) and Qi319×Huangzaosi (Q/H), respectively. The two populations were evaluated under different water conditions in Xinjiang in 2007 and 2008. The results showed that those ear traits under drought were phenotypically lower than those under normal water regime and ear length, ear diameter, weight per ear had positive correlation with GW. A total of 75 QTL were identified under different water regimes using mixed linear model with two methods of single-experiment analysis and water regime joint analysis, including 20 QTL detected under normal water regime and 55 QTL detected under drought stress. The QTL detected in Y/H were located on chromosome 1, 2, 5, 6, 7, and 10, and the QTL detected in Q/H were distributed on chromosome 2, 3, 4, 5, 6, 7, 9, and 10. However, only four and nineteen QTL was identified in the two populations under drought stress respectively, significantly lower than that detected under normal water regime. Meanwhile, in the joint analysis, only three QTL had significant interaction with environments and six QTL had epistatic interaction, showing that the genetic feature of ear traits was complex. In Y/H, two congruent QTL, qKRE1-5-1 and qKRE1-7-1 , were detected under different water levels, with the phenotypic contribution from 6.15% to 19.48%, while in Q/H, three congruent QTL, qKRE2-5-1 , qGW2-10-1 , and qKRE2-3-1 , were detected under different water levels, with the phenotypic contribution from 7.14% to 16.65%. These results implied that these QTL were little influenced by environment and could stably expressed, which can be used in marker-assisted selection.

穗部性状与产量密切相关，因此对其进行遗传剖析可为玉米高产育种提供理论基础，尤其是对干旱胁迫下的稳产有重要意义。本研究以玉米骨干亲本黄早四分别与自交系掖478和齐319进行杂交，构建了两套F 2:3 群体(分别记为Y/H和Q/H)。在正常水分灌溉和干旱胁迫下对穗长、穗粗、轴粗、穗行数、行粒数、穗粒重和穗重等7个穗部性状进行了表型鉴定，采用基于混合线性模型的单环境分析和相同处理水平的联合分析方法进行了QTL分析。结果表明，在干旱胁迫下，2个群体的亲本及F 2:3 家系的各性状值均低于正常水分条件，且穗粒重与穗长、穗重、穗粗呈正相关。在干旱胁迫下和正常水分条件下，通过两种检测方法共定位到75个玉米穗部性状QTL，其中Y/H群体共定位了20个QTL，分布在第1、第2、6、第5、第7、第10染色体上；Q/H群体共定位了55个QTL，分布在第2、第3、第4、第5、第6、第7、第9、第10染色体上；但是在干旱条件下两群体分别只检测到4个和19个QTL，明显低于正常水分条件下检测到的QTL数目。通过联合分析只检测到3个QTL与环境发生显著互作和6对QTL存在上位性互作效应，说明玉米穗部性状的遗传基础较为复杂。同时还发现，Y/H群体在正常灌溉与干旱条件下检测到2个一致性的QTL，分别是 qKRE1- 5-1 和 qKRE1- 7-1 ，对表型变异解释的变化范围是6.15%~19.48%；Q/H群体检测到3个一致性QTL，分别是 qKRE2-5-1 、 qGW2-10-1 和 qKRE2-3-1 ，对表型变异解释的变化范围是7.14%~16.65%，说明这些QTL受环境影响较小，能够稳定遗传，可以作为分子标记辅助选择的候选区间应用于玉米穗部性状抗旱性改良。

2014

0.0

吴亚辉, 陶星星, 肖武名, 郭涛, 刘永柱, 王慧, 陈志强. 水稻穗部性状的QTL分析. 作物学报, 2014, 40: 214-221

Wu Y

, Tao X

, Xiao W

, Guo

, Liu Y

, Wang

, Chen Z

Dissection of QTLs for panicle traits in rice (Oryza sativa).

Acta Agron Sin, 2014, 40: 214-221 (in Chinese with English abstract)

Rice yield potential is closely related to panicle traits. QTLs for five panicle traits including panicle length, number of primary branches, number of secondary branches, spikelets per panicle and seed setting density were identified by using 172 plants and 138 plants from a F 2 population derived from a cross between Nipponbare and H71D in 2011 and 2012 respectively. A total of 38 QTLs were detected in the two trials, including 21 QTLs in 2011 and 17 QTLs in 2012, of them four QTLs (only 10.5%) were repeatedly detected in both years. Some QTLs controlling different traits shared the same maker interval on the chromosome with each other, which was consistent with their significant phenotypic correlations. QTLs with large effects are easily to be detected in kinds of populations and different environments. These QTLs provide useful information for meta-analysis and fine mapping, as well as MAS for high-yield rice breeding.

水稻穗部性状与产量关系密切。以粳稻品种日本晴和一个大穗籼稻材料 H71D 为亲本构建 F 2 群体，于 2011 和 2012 年分别以 172 个单株和 138 个单株，对穗长、一次枝梗数、二次枝梗数、总粒数、穗着粒密度等 5 个穗部性状进行 QTL 定位。以 LOD≥3 为阈值，两年共检测到 38 个 QTL ，其中 2011 年 21 个， 2012 年 17 个， 2 年重复检测到的 QTL 为 4 个，占总数的 10.5% 。 5 个性状之间大都具有显著的表型相关性，相关性较强的性状之间具有较多相同或紧密连锁的 QTL ，效应值较大的 QTL 易于在不同群体和不同环境中被重复检测到。检测到的 QTL 为进一步进行元分析和精细定位打下了基础，也为通过分子标记辅助选择提高产量提供了有用信息。

... 分布的现象从遗传学角度通常表现为一因多效、基因连锁或重叠, 可能由于一些关键因子的存在而同时控制多种代谢途径, 从而表现出多效性, 使相关性状在自然选择过程中形成相互适应的有利性状^[27] ...