利用2个F2群体整合中国豌豆高密度SSR遗传连锁图谱

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刘荣^,¹^,^**, 王芳^,¹^,^**, 方俐^,¹^,^**, 杨涛¹, 张红岩¹, 黄宇宁¹, 王栋¹^,³, 季一山¹, 徐东旭², 李冠¹, 郭瑞军¹, 宗绪晓^,¹^,^*

¹ 中国农业科学院作物科学研究所作物种质资源中心, 北京 100081

² 张家口市农业科学院食用豆类研究所, 河北张家口 075000

³ 山东省作物种质资源中心, 山东济南 250100

An integrated high-density SSR genetic linkage map from two F₂ population in Chinese pea

LIU Rong^,¹^,^**, WANG Fang^,¹^,^**, FANG Li^,¹^,^**, YANG Tao¹, ZHANG Hong-Yan¹, HUANG Yu-Ning¹, WANG Dong¹^,³, JI Yi-Shan¹, XU Dong-Xu², LI Guan¹, GUO Rui-Jun¹, ZONG Xu-Xiao^,¹^,^*

¹ Center for Crop Germplasm Resources, Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China

² Institute of Legumes Crop, Zhangjiakou Academy of Agricultural Sciences, Zhangjiakou 075000, Hebei, China

³ Shandong Center of Crop Germplasm Resources, Jinan 250100, Shandong, China

通讯作者: * 宗绪晓, E-mail: zongxuxiao@caas.cn

** 同等贡献(Contributed equally to this work)
收稿日期:2020-02-8接受日期:2020-04-15网络出版日期:2020-05-09

基金资助:

国家自然科学基金项目.31801428
国家现代农业产业技术体系建设专项.CARS-08
农作物种质资源保护与利用专项.2019NWB036-07
中国农业科学院科技创新工程, 国家农作物种质资源共享服务平台.NICGR2019
国家重点研发计划项目.2017YFE0105100

Received:2020-02-8Accepted:2020-04-15Online:2020-05-09

Fund supported:

National Natural Science Foundation of China.31801428
China Agriculture Research System.CARS-08
Crop Germplasm Resources Protection.2019NWB036-07
Agricultural Science and Technology Innovation Program (ASTIP) in CAAS.NICGR2019
National Infrastructure for Crop Germplasm Resources Project from the Ministry of Science and Technology of China.2017YFE0105100

作者简介 About authors
刘荣, E-mail: liurong@caas.cn;

王芳, E-mail: fwang11@huskers.unl.edu;

方俐, E-mail: hathor_fang@163.com

摘要
豌豆(Pisum sativum L.)是一种重要的食用豆类作物, 在全世界范围内广泛种植, 既可作为人类食物, 也可作为牲畜饲料。用SSR标记构建的遗传连锁图谱在豌豆和其他作物的标记辅助育种中发挥着重要的作用。尽管对豌豆遗传连锁作图的研究已有悠久历史, 但公众可获得且可转移的SSR标记以及基于遗传独特的中国豌豆种质的高密度遗传连锁图谱仍然有限。为了获得更多可转移的SSR标记和中国豌豆的高密度遗传连锁图谱, 本研究首先从自主开发和文献获取的12,491个全基因组SSR标记中筛选了617个多态性SSR标记, 并用于G0003973×G0005527 F₂群体遗传连锁图谱的加密。加密后的图谱全长扩展到5330.6 cM, 包含603个SSR标记, 标记平均间距离8.8 cM, 相比之前的图谱有明显改善。基于上述结果, 我们又筛选了119个具有多态性的SSR标记, 用于构建大样本W6-22600×W6-15174 F₂群体的遗传连锁图谱, 新图谱累积长度为1127.1 cM, 包含118个SSR标记, 装配在7条连锁群上。最后, 将来自以上2个遗传图谱的数据进行整合, 得到了一张覆盖范围6592.6 cM的整合图谱, 包含668个SSR标记, 由509个基因组SSR、134个EST-SSR和25个锚定标记组成, 分布在7条连锁群上。这些SSR标记和遗传连锁图谱将为豌豆的遗传研究和标记辅助育种提供有力工具。
关键词： 豌豆;SSR;遗传连锁图谱;整合图谱;标记辅助育种

Abstract

Pea (Pisum sativum L.) is an important food legume crop grown widely throughout the world for humans or livestock consumption. Genetic linkage map constructed with SSR markers have played a vital role in marker-assisted breeding of many crops including pea. Public available and transferable SSR markers and genetic linkage map with sufficient SSR markers based on genetically distinct Chinese pea germplasm are limited despite a long study history on genetic linkage mapping in pea. In this study, in order to obtain more transferable SSR markers and high resolution genetic linkage maps for Chinese pea, 617 polymorphic SSR markers were firstly screened from 12,491 genome-wide SSR markers and some related literatures by our laboratory, and these SSR markers were used to construct an enhanced genetic linkage map for the G0003973 × G0005527 F₂ population. The enhanced genetic linkage map covered 5330.6 cM in total length containing 603 SSR markers with an average inter-marker distance of 8.8 cM, which was significantly improved both in marker number and in density compared with the previous map. 119 polymorphic SSR markers were screened based on the above results to develop a new map for a large W6-22600 × W6-15174 F₂ population including 118 SSR markers with a cumulative length of 1127.1 cM assembled into seven genetic linkage groups. Furthermore, data from the above two genetic maps were combined to build an integrated map of 6592.6 cM, comprising 668 SSR markers, 509 genomic SSRs, 134 EST-SSRs and 25 anchor markers distributed in seven genetic linkage groups. These SSR markers and genetic linkage maps will provide a valuable tool for the genetic study and marker-assisted breeding in pea.

Keywords：pea (Pisum sativum L.);SSR;genetic linkage map;integrated map;marker-assisted breeding

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刘荣, 王芳, 方俐, 杨涛, 张红岩, 黄宇宁, 王栋, 季一山, 徐东旭, 李冠, 郭瑞军, 宗绪晓. 利用2个F₂群体整合中国豌豆高密度SSR遗传连锁图谱[J]. 作物学报, 2020, 46(10): 1496-1506. doi:10.3724/SP.J.1006.2020.04028
LIU Rong, WANG Fang, FANG Li, YANG Tao, ZHANG Hong-Yan, HUANG Yu-Ning, WANG Dong, JI Yi-Shan, XU Dong-Xu, LI Guan, GUO Rui-Jun, ZONG Xu-Xiao. An integrated high-density SSR genetic linkage map from two F₂ population in Chinese pea[J]. Acta Agronomica Sinica, 2020, 46(10): 1496-1506. doi:10.3724/SP.J.1006.2020.04028

豌豆(Pisum sativum L.)属于豆科(Leguminosae/ Fabaceae), 野豌豆族(Vicieae), 豌豆属(Pisum), 染色体数为2n = 2x = 14, 基因组大小约为4.45 Gb^[1,2]。豌豆富含蛋白质和多种营养元素, 是经济上最重要的食用豆类作物之一, 在世界范围内广泛种植, 既可以作为谷物和蔬菜供人类食用, 又可作为牲畜的饲料^[3,4]。根据FAO的统计, 2018年, 豌豆(包括青豌豆和干豌豆)在全球食用豆类作物中的总产量仅次于普通菜豆; 同时, 中国青豌豆的总产量居世界首位, 而干豌豆的总产量仅次于加拿大和俄罗斯^[5]。此外, 豌豆因其固氮能力而被认为是一种环境友好型作物, 在可持续农业系统中起着至关重要的作用^[6]。

高密度遗传连锁图谱是功能基因定位、比较基因组学以及分子辅助育种等研究的重要工具^[7]。以前, 人们一直致力于利用包括RFLP、RAPD、SSR和SNP在内的多种分子标记基于不同类型的群体来构建豌豆的遗传连锁图谱^{[8,9,10,11,12,13,14,15]}。最近, 有****针对豌豆开发了基于高密度SNP的遗传连锁图谱, 并为鉴定重要农艺性状的遗传基础提供了强大的工具^{[16,17,18,19]}。此外, 新近公布的豌豆参考基因组也为理解豌豆关键农艺性状的分子基础并促进其育种改良奠定了重要基础^[20]。

SSR标记因其具有信息量丰富、共显性遗传、多等位基因、基因组覆盖广等特性, 同时在相近物种之间具有可重复性和可移植性^[21,22], 在遗传多样性评估和物种亲缘关系鉴定^[23,24]、遗传连锁图谱构建^[25,26]、标记辅助选择^[27,28]、DNA指纹图谱鉴定^[29,30]等方面具有显著优势。相比基因组SSR, 位于基因区的EST-SSR因其具有更高的可转移性、较低的开发成本以及与基因的密切关系, 而越来越受到人们的重视^[11,21]。然而, 尽管针对豌豆的遗传连锁作图研究已有很长的历史, 并且在豌豆中已经构建了几十种具有不同标记的遗传连锁图谱^[31], 但公众可获得的可用于豌豆遗传研究的SSR上图标记较少, 同时基于遗传独特的中国豌豆种质^[1,32-33]的遗传连锁图谱仍然有限。值得注意的是, 过去基于中国豌豆种质构建的遗传连锁图谱包括157个SSR标记, 分布在11个连锁群中, 全长1518 cM, 标记数量较少, 需要进一步加密并完善至7个连锁群^[15]。

与单个遗传连锁图谱相比, 整合遗传连锁图谱由于整合了多个群体的信息而具有多种优势^[34,35], 例如具有更高的标记密度, 更完整的基因组覆盖范围, 可对不同群体进行标记共线性比较等, 在许多作物包括豌豆中均有应用^{[16,20,36-37]}。因此, 本研究的目的如下: 1)筛选豌豆中可移植转换的SSR标记, 用于豌豆的遗传研究和分子作图。2)对我们以往基于G0003973×G0005527 F₂群体, 构建的遗传连锁图谱进行加密。3)基于W6-22600×W6-15174 F₂群体, 构建新的遗传连锁图谱。4)结合上述2个基于中国种质的遗传连锁图谱信息, 构建一张豌豆整合SSR遗传连锁图谱。

1 材料与方法

1.1 作图群体

本研究利用基于中国豌豆种质为亲本的2个F₂群体进行遗传连锁作图。群体1 (PSP1)与本实验室之前的研究相同^[15], 来自母本G0003973 (耐寒)和父本G0005527 (不耐寒)之间的杂交, 由190个F₂个体组成。群体2 (PSP2)则是以母本W6-22600 (多小叶)与父本W6-15174 (无小叶)进行杂交, 由480个F₂个体组成。

1.2 SSR标记筛选

利用本实验室自主开发^[38]和文献获取^[39,40,41]的12,491个SSR标记(包括11,145个基因组SSR和1346个EST-SSR), 对PSP1的亲本及随机选择的4个F₂个体进行全基因组扫描, 筛选出多态性SSR标记用于遗传连锁作图。此外, 从以往研究中已发表的豌豆遗传连锁图谱中, 选择具有已知连锁群位置的125个SSR标记^[10,42-44], 利用PSP1和PSP2的亲本和4个随机选择的F₂个体来筛选锚定标记。

1.3 DNA提取和PCR扩增

在2个F₂群体种植当年, 收集每个F₂个体植株的嫩叶, 经液氮速冻后, 使用改良的CTAB方法^[45]提取基因组DNA。用NanoDrop 2000检测DNA浓度并稀释到工作液浓度50 ng μL^-1后, 于-20℃保存备用。

PCR扩增反应体系为10 μL, 包含1.5 μL基因组DNA (50 ng μL^-1)、5 μL 2×Taq PCR Master Mix (Genstar, 中国北京)、0.5 μL正向引物 (2 μmol L^-1)、0.5 μL反向引物 (2 μmol L^-1)和2.5 μL ddH₂O。PCR产物通过8%非变性聚丙烯酰胺凝胶电泳(PAGE)分离, 并通过0.1%硝酸银染色。根据片段大小记录等位基因状态, SSR标记状态编码如下: 与父本相同的带型记为“AA”, 与母本相同的带型记为“BB”; 具有双亲带型的记为“AB”; 缺失或无效的带型记为“-”。只有那些能够扩增出清晰条带并可以显示亲本多态性的SSR标记才被选择用于后续的基因分型。

1.4 遗传连锁图谱构建

分别对PSP1和PSP2的所有F₂个体进行基因分型, 并去除缺失数据超过20%的标记或个体。使用χ2分析来检测标记偏分离状况, 并使用Bonferroni校正对P = 0.05的显著性水平进行校正, 在进一步的遗传作图中排除显著偏分离的标记。利用Kosambi作图函数对2个群体构建遗传连锁图谱, LOD>2。在以往公布的豌豆遗传连锁图谱的基础上, 通过筛选得到的锚定标记对每个连锁群进行分组^[10,42-44]。然后, 利用共有标记将这2个群体的信息整合到一张遗传连锁图谱上。以上所有分析均利用QTL IciMapping V4.0软件完成^[46]。遗传连锁图谱和物理图谱利用MapChart V2.3软件进行可视化展示^[47]。然后, 本研究以新近发表的豌豆基因组为参考(Caméor genome build 1a) ^[20], 利用KnowPulse网站(https:// knowpulse.usask.ca/blast/nucleotide/nucleotide)的BL ASTn工具对50个共有标记的扩增片段序列进行比对, 参数选取默认参数, E-value设为1e^-3。

2 结果与分析

2.1 多态性标记筛选

利用本实验室自主开发^[38]和文献获取^[39,40,41]的12,491个SSR标记(包括11,145个基因组SSR和1346个EST-SSR), 对PSP1的亲本及随机选择的4个F₂个体进行全基因组扫描, 初步筛选出扩增条带清晰且在父母本间呈多态性差异的954个多态性SSR标记, 用于PSP1群体190个F₂个体的基因分型, 最终得到729个在190份F₂群体单株中有清晰条带的多态性标记, 用于后续遗传连锁图谱的构建。然后利用这729个多态性标记对PSP2的亲本及随机选择的4个F₂个体进行多态性检测, 最终在480个F₂个体中成功筛选了103个多态性标记用于PSP2的遗传连锁图谱构建。

此外, 本研究还利用125个已知遗传连锁群位置信息的可公开获得的SSR标记^[10,39-41]在2个群体中筛选锚定标记, 分别在PSP1和PSP2中鉴定出11个和17个锚定标记, 其中有3个标记为2个群体共有的锚定标记, 共计25个锚定标记, 这些标记可在后续遗传连锁图谱构建中用于分配连锁群(表1)。

Table 1
表1
表1筛选得到的25个锚定标记在以往发表豌豆遗传连锁图谱的分布
Table 1Distribution of the screened 25 anchor markers on previous genetic linkage map in pea

连锁群 Linkage group	锚定标记的数目 Number of anchor markers	锚定标记的名称 Name of anchor markers
I	5	AA67, AD147, AF016458, D21, PsAS2
II	3	AA332, AD83, D23
III	4	AA355, AD270, PSAJ3318, PSBLOX13.2
IV	4	AA430942, AA122, AA285, AA315
V	1	PSGAPA1
VI	5	AA335, AB71, AD160, AD60, PSGSR1
VII	3	AB65, AF004843, PSAB60

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2.2 遗传连锁图谱构建

获得所有样本的基因型数据后, 首先对所有标记进行数据完整性和偏分离检测。对于PSP1作图群体, 740个标记(包含11个锚定标记)中, 有43个标记缺失信息大于20%, 占总标记数的5.81%, 而有80个标记检测到偏分离现象, 占总标记数的10.81%。

因此, 排除这123个标记后, 共有617个标记用于后续的遗传连锁图谱分析。针对PSP1构建的遗传连锁图谱, 将603个标记分配到7条连锁群上。根据11个锚定标记, PSP1图谱的7条连锁群分别对应于以往发表的遗传连锁图谱的6条连锁群^[10,39-41], 有2条连锁群均对应于LGI, 而缺少对应于LGV的连锁群, 可能是由于缺乏LGV的锚定标记。此外, 该图谱全长5330.6 cM, 相邻标记之间的平均距离为8.9 cM。每条连锁群的长度从494.9 cM (LGII)到904.7 cM (LGIV)不等, 平均为761.5 cM; 每条连锁群的标记数目从62 (LGII)到113 (LGI-2)不等, 平均为86个标记(图1和表2)。

图1

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图1豌豆PSP1群体遗传连锁图谱

粗体代表锚定标记, 标记名称以“e”开头的为EST-SSR标记。
Fig. 1Genetic linkage map of PSP1 population in pea

The bold labels on the marker name represent anchor markers, and marker names started with “e” represent EST-SSR markers.

Table 2
表2
表2利用豌豆PSP1和PSP2群体构建的2个遗传连锁图谱的标记分布
Table 2Marker distribution on the two genetic linkage maps from PSP1 and PSP2 population in pea

连锁群 Linkage group	上图标记数目 Number of mapped markers		图谱长度 Map length (cM)		平均标记密度 Average marker density
连锁群 Linkage group	PSP1	PSP2	PSP1	PSP2	PSP1	PSP2
I	74	11	751.0	137.4	10.1	12.5
II	62	13	494.9	160.6	8.0	12.4
III	90	20	855.3	158.4	9.5	7.9
IV	97	14	904.7	119.0	9.3	8.5
I-2/V	113	19	822.5	173.3	7.3	9.1
VI	70	21	658.6	175.0	9.4	8.3
VII	97	20	843.6	203.4	8.7	10.2
平均Mean	86	17	761.5	161.0	8.9	9.8
总计Total	603	118	5330.6	1127.1	62.3	68.9

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对于PSP2作图群体来说, 120个标记(包含17个锚定标记)中, 仅有1个标记由于缺失大于20%而被排除在后续分析之外。剩余的119个标记被用于进一步的连锁作图分析, 结果发现上图的118个标记分布在7条连锁群上。根据17个锚定标记, 这7条连锁群刚好完全对应于以往发表的遗传连锁图谱的7个连锁群^[10,39-41]。此外, 该图谱的累积长度为1127.1 cM, 相邻标记之间的平均遗传距离为9.8 cM (图2和表2)。每条连锁群的长度从119.0 cM (LG IV)到203.4 cM (LGVII)不等, 平均为161.0 cM; 每条连锁群的标记数目从11 (LGI)到21 (LGVI)不等, 平均为17个标记(图2和表2)。

图2

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图2豌豆PSP2群体遗传连锁图谱

粗体代表锚定标记。
Fig. 2Genetic linkage map of PSP2 population in pea

The bold labels on the marker name represent anchor markers.

2.3 整合遗传连锁图谱构建

基于以上2个遗传连锁图谱, 通过两两比较共发现了53个共有标记, 每个连锁群上的共有标记数为3 (LGI)到14 (LGVII)不等(表3)。利用上述2个遗传连锁图上的53个共有标记, 我们构建了一张包含668个SSR标记的整合遗传连锁图谱(标记信息详见附表1), 分布在7条连锁群上, 累积长度为6592.6 cM, 相邻标记之间的平均距离为10.0 cM。每条连锁群的长度从682.7 cM (LGII)到1077.2 cM (LGIII)不等, 平均为941.8 cM。在这7条连锁群中, 分布在LGV的标记数量最多, 有125个标记, 同时标记密度也最低, 为8.1 cM; 另有3条连锁群包含的标记数也都超过了100, 分别是LGIV (104)、LGVII (103)和LGIII (102); 而标记数量最少的连锁群为LGII, 仅有68个标记, 累积长度也是最小的, 仅有682.7 cM (图3和表3)。

Table 3
表3
表3豌豆整合遗传连锁图谱上的标记分布
Table 3Markers distribution on the integrated genetic linkage map in pea

连锁群 Linkage group	共有标记数目 Number of bridge markers	上图标记数目 Number of mapped markers	图谱长度 Map length (cM)	平均标记密度 Average marker density
I	3	82	976.8	11.9
II	7	68	682.7	10.0
III	8	102	1077.2	10.6
IV	7	104	1058.9	10.2
V	7	125	1018.2	8.1
VI	7	84	801.8	9.5
VII	14	103	977.0	9.5
平均Mean	7.6	95	941.8	10.0
总计Total	53	668	6592.6	69.8

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图3

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图3豌豆PSP1和PSP2整合遗传连锁图谱

粗体代表锚定标记, 下画线代表共有标记, 粗体加下画线代表共有锚定标记。标记名称以“e”开头的为EST-SSR标记。
Fig. 3Genetic linkage map integrated with PSP1 and PSP2 information in pea

The bold, underlined and bold underlined labels on the marker name represent anchor markers, common markers and common anchor markers, respectively. Marker names started with “e” represent EST-SSR markers.

Supplementary table 1
附表1
附表1豌豆整合遗传连锁图谱上图标记信息
Supplementary table 1Information of SSR markers used to generate the integrated genetic linkage map of pea

序号 No.	标记名称 Marker name	标记类型 Marker type	正向引物序列 Forward primer sequence (5°-3°)	反向引物序列 Reverse primer sequence (5°-3°)	退火温度T_m (℃)	片段大小Band size (bp)	连锁群Linkage group
序号 No.	标记名称 Marker name	标记类型 Marker type	正向引物序列 Forward primer sequence (5°-3°)	反向引物序列 Reverse primer sequence (5°-3°)	退火温度T_m (℃)	片段大小Band size (bp)	整合图谱Integrated map	PSP1 G0003973× G0005527	PSP2 W6-22600×W6-15174)
1	27256	基因组SSR	CATTCCATTCCATACATCCATTT	TGAAGTTGAAGCAGCCATTG	60	153	I		I
2	26075	基因组SSR	CTATGGCACCATCTCTTGGAC	AACACAATGTATGTGGTGCAAAT	59	186	I		I
3	28142	基因组SSR	TGCAAGCATATTGCCTTTTC	TCAGTGGTTGCTAGCTGTTGA	59	176	I		I
4	26020	基因组SSR	CTCTCAAATTTGGGGTCCTC	TCATTGCGTCTCAACCTCAG	59	129	I		I
5	25334	基因组SSR	TGAAGATGTGACAAACAACAGAAA	TCCTTCTCTGTTCCCACCAC	60	140	I	I	I
6	23971	基因组SSR	GCGTGTTGATGCTGAAAGAA	TCAAAATTGGGGTGTGACAA	60	131	I		I
7	25922	基因组SSR	CCAAGGGAAAACCCCTTCTA	GTGTGAAAGCTTATTGTATATGTATCG	58	176	I		I
8	AD147	锚定标记	AGCCCAAGTTTCTTCTGAATCC	AAATTCGCAGAGCGTTTGTTAC	61	330	I	I-2	I
9	AA67	锚定标记	CCCATGTGAAATTCTCTTGAAGA	GCATTTCACTTGATGAAATTTCG	51	370	I		I
10	20229	基因组SSR	GCGAAGACTTCCGACCATTA	CACGGTCAAGGCCTACATTT	60	181	I	I	I
11	18339	基因组SSR	TGGTTGAACTGGAACGAGTG	TGAAATTGCAATGTAAGCATGA	59	137	I	I
12	e1154	EST-SSR	GCCATGCGACCATATTTACC	ATGGCCACAGAAACGAAAAC			I	I
13	e1111	EST-SSR	TTCTTCGTCGGAGGATGAGT	TTGAGAGGAGATTGGAGAAAAA			I	I
14	28832	基因组SSR	CTCACGCGTTGAAGATACCA	CACCGCCATTGTAGTACAGC			I	I
15	27227	基因组SSR	GCATTCCATGAATTGCATCT	TTGCTGATTTCTTACTTGTTGTCA	59	167	I	I
16	e734	EST-SSR	AGGCAGTGACTGAATCATCGT	AATGGCTTTGAGGCAGAGAG			I	I
17	18591	基因组SSR	AGGGCCGAATGCTAAGTGAT	TTTTGAACCCTGGAGGGAGT	61	155	I	I
18	21662	基因组SSR	GAACATTATCGGAATCAACAGC	CCACACAAAAATGAACAACACA	59	158	I	I
19	17158	基因组SSR	CTCCCGAGTCTTGGCTAATG	AGGCGCTCATAAACAGTTCC	59	175	I	I
20	e1171	EST-SSR	AGTCCCATCCCACGAAAAAT	CTCTTTCAAATCCCCCAACA			I	I
21	PEACPLHPPS	基因组SSR	GTGGCTGATCCTGTCAACAA	CAACAACCAAGAGCAAAGAAAA	52	120	I	I
22	e613	EST-SSR	CAATAATTTCACACACACCAAGAA	TCCAAAGAATCCTAAGAAACATGA			I	I
23	24312	基因组SSR	TGCCATATGCATTTCATGGT	AAGCCCCTTTTCATCTTCAA	59	210	I	I
24	e1013	EST-SSR	ACACCAACGATGACCCATTT	CCGATCCACAAACCGTTATT			I	I
25	29847	基因组SSR	TTGTTCCTACCGTGCTTTCA	TCTTATTGGCCTGCACATCTT			I	I
26	29428	基因组SSR	CCATCCACATCCTTCCAAGT	CACTCAACCCACGGAAAAGT			I	I
27	e700	EST-SSR	GGTTTCGGTTGATCATGGTA	GGAACTTTCTCTCGGGATCA			I	I
28	e1189	EST-SSR	TGGCAATTGCGATGATTAGA	TTCATCCGTTTCATGATGTTG			I	I
29	25805	基因组SSR	CCAACTTACTTTTGCTTATCTGGT	TGGGTCCATGACAAAGACAA	58	124	I	I
30	25096	基因组SSR	CAACATTGTTATCATCAAAACTCAA	GGCGACAATCGATCTCAAGT	58	192	I	I
31	28733	基因组SSR	TGGCCTAGGTTTTTGTGTCC	GCATCTCAAAAGGGCATTATTT	59	116	I	I
32	23546	基因组SSR	TCCACCTTGTTGCCCTAATC	TGAATGCTTCTCAGATACAAAATGA	60	155	I	I
33	22611	基因组SSR	TGCAAATGTGCAATGAATGA	GGCGGACATGAGAAGGAATA	60	187	I	I
34	27835	基因组SSR	CATCACTTGGGATTTCTTGAGAG	AGGGCAATGGTAATCAGCAC	60	138	I	I
35	27301	基因组SSR	TGTCGGAAATTAAGAGGTGGA	TGGAAAAGTAAGCGGTGAACA	60	119	I	I
36	e1226	EST-SSR	CACACCAGGTATCAATCTGTGAA	CGTTCCGCTTTTCACTCTCT			I	I
37	16881	基因组SSR	ATGGGCTTTAGGGGAAGAAA	AAAAGCAGCACATGGAGGAC	60	133	I	I
38	21678	基因组SSR	CCCTTCAGCAACAATCACTG	TGCCTCAGATTTGGAATGGT	59	151	I	I
39	20646	基因组SSR	TCTCACATGTTGTTATTTCTTTCTCA	TGATGTTCCCCAGATTTTCA	59	120	I	I
40	PsAS2	锚定标记	CTAATCACACGTTTAGGACCGG	CGAAATCCAAACCGAACCTAATCC	52	300	I	I
41	e888	EST-SSR	GGCTTCTCCATTTGTGGTTC	GCCAATGGAGGTTCTACAGC			I	I
42	18011	基因组SSR	GACCAACGACTTGGACATCA	GGTGAGTTCCTAAGATGAATCAGA	59	197	I	I
43	e811	EST-SSR	TTTTGTGGGTCTCTCTTCACC	CACCACACATGCAACACTCA			I	I
44	27246	基因组SSR	CCAGGTTAAAACGATGATTTTTG	ACTTTTCCCCTTGGTTGGAC	60	180	I	I
45	3023	基因组SSR	GGTGCAAAATTTGAGGTGCT	CACACACGACTACACACACTACG	59	153	I	I
46	27167	基因组SSR	GGCACAACTACAACCCACAA	GGTTCAGGAATGGGTTCAGA	59	194	I	I
47	e799	EST-SSR	TGCAGGCTTTAGAAGTTGTTCA	CTCAGCAGCCACAATTACACA			I	I
48	5897	基因组SSR	GGCAATAACTTAAGAGTACTAAGGAAA	AGGGTGTCGTCGTGTGTGTA	57	282	I	I
49	27361	基因组SSR	CTGAAACGGTTTGCATTGTG	TCCAACCACTTCTTAACAACCT	58	113	I	I
50	22209	基因组SSR	AATCCACAACCCCGTCAATA	CAAAAGAGACCTTCTTCCTCTCA	59	164	I	I
51	22259	基因组SSR	CATCATGGCTCAATCTCAACA	TTCCCAAATTCCTTCGTTCA	60	110	I	I
52	22406	基因组SSR	GGAAAGAGTTATGGCAATGGAC	TGGTGGTGGAGCTAAGTGTG	60	174	I	I
53	24555	基因组SSR	CGCTTATGTAGCCCCTTTTG	GGCCAAAGGAGATTTGTGTC	60	123	I	I
54	24499	基因组SSR	AAAACAAACAAAACCGCAATG	TAGCCATCACCAAAGCAACA	60	169	I	I
55	e624	EST-SSR	CCTTAGCAAGTTTGTCTTTGAGTG	TGCAATGACATGATGGAAGAA			I	I
56	29029	基因组SSR	TAGGAGAGCGAGGAGCAAAG	CCACCAAAAGCAAGAATGTG			I	I
57	e884	EST-SSR	TTCTTTCCGCCATGAGATTC	GAGAGCAAGGGTTTGGAACA			I	I
58	28438	基因组SSR	GGAATGACGAAGTAACCACCT	GATGCAAGTGCAACCTTTGA	58	185	I	I
59	17628	基因组SSR	GGTTTTGTTTGCCGTTGATT	CCACCCCCAAACTTCCTTAT	60	153	I	I
60	17605	基因组SSR	CGCCCTTCATCATCATCTTC	AGAGTCGGTCCCTCCAACAT	61	150	I	I
61	1356	基因组SSR	CACGTGCACATACACACTCTT	TGTGTCAGAGCATGTGTTCG	57	106	I	I
62	171	基因组SSR	CAAACACACACGCACACAAA	CGTGTGAGCGTGCATAAGT	58	70	I	I
63	28654	基因组SSR	AGCGACGTGAATATCACAATG	GTTATCGCGGCGTGTAAATC	59	134	I	I
64	e658	EST-SSR	TGGTTTCTCTGCCAAAACAG	TGATGAGTGGTGACGCAAAT			I	I
65	e562	EST-SSR	CAAGATGCTTCTGATTCAGTGTC	AGGATTTGAGCTTGGGAGGT			I	I
66	17989	基因组SSR	CAGAGCCGGAGTTCTGGATA	TTTGGTTGACATTAGCACATGA	59	195	I	I
67	26333	基因组SSR	AAACACACGACATGTTTCCTTTT	TCACTGCAATTCGTCGATGT	60	116	I	I
68	D21	锚定标记	TATTCTCCTCCAAAATTTCCTT	GTCAAAATTAGCCAAATTCCTC	51	200	I	I
69	18928	基因组SSR	TGAATGTGGAAAGGAGGAATG	AGGGTCACCACTTTGGAGAG	59	178	I	I
70	18529	基因组SSR	GAATGTGCGTCCAACATCCT	AGATTTTGATGCGGAAGAGC	59	151	I	I
71	1863	基因组SSR	GCACACGAATACAGTCACGC	GTGTGTTGACGTGCGAGTTT	60	118	I	I
72	e997	EST-SSR	GCCTGGAGTGTTGAAGAGGA	CCATCACAATTTCCCACACA			I	I
73	1752	基因组SSR	GCACGCACACGAATACAGTC	GACGTCGTGAGTTTGCATGT	60	115	I	I
74	AF016458	锚定标记	CACTCATAACATCAACTATCTTTC	CGAATCTTGGCCATGAGAGTTGC	54		I	I
75	23261	基因组SSR	CTGCTTTTGGGGTTTGGTTA	GCAATGCAACTCCTCAACAA	60	156	I	I	I
76	22699	基因组SSR	CAACATGCCATTCTGGCTAA	GCCGAAACCCCATGTAGAC	60	157	I	I
77	24895	基因组SSR	AAGAAAGTTGCGTTGGATGTG	GTTTTGTACCGCCCAACACT	60	148	I	I
78	24731	基因组SSR	AGAAAATGGCCCACGAATTA	TGCATTGCATTGTGTTTGTG	59	204	I	I
79	25106	基因组SSR	AAGGCCAAACAGAAAGGAGA	CAATGTCCAAGAAAGATCCAGTT	59	178	I	I
80	4083	基因组SSR	TGCAAACTCACACGTCAACA	GTGCGTGTGCGAAGTACG	60	191	I	I
81	1753	基因组SSR	GCACGCACACGAATACAGTC	TCGTAGTTTGCATTGTGCGT	60	115	I	I
82	25341	基因组SSR	AATGCTTCTTCCACGGTCAC	TTCGCTCGAGTTCGATTCTT	60	184	I	I
83	25454	基因组SSR	TTCCAAGCAAGCGTTGAAGT	TCAAGAGAGACTTTTCAAGAGGTT	58	204	II	II
84	29797	基因组SSR	TGTGATCAGGTGCTCCCATA	GCGACAAATTATGGCTATGC			II	II
85	26436	基因组SSR	TTGCCTTGCCAACTTTTAGG	CTTGCTTCTGCGCCATAAAT	60	195	II	II
86	e581	EST-SSR	CCTTGATGCCACAAATGAGA	TTGCCACTTTCTCAAAAACTCA			II	II
87	S217	基因组SSR	CACTCAACTCACCGACCTCA	GACGGATGGAAATTGGTGTC	52	1035	II	II
88	e956	EST-SSR	CGAGCGTGAGACTGTGATGT	TCCACCGGTTCAACTTCAAT			II	II
89	e344	EST-SSR	ATGCAACCGGCGCAGTAT	CCACCTTTTCCTCGCTTTTT	61	159	II	II
90	e967	EST-SSR	TGACACTTTCGTGTACTGTGTTTTT	TTCCAAAAGCCTCTCTTTCATC			II	II
91	20922	基因组SSR	AAAAGGAGAACACATTTTATAATAGCA	TGCTCTTAAAGGCGACAATG	58	146	II	II
92	29141	基因组SSR	TTCTTTCTGCTAGGAGCCACT	CAAAGCCATCACCCTACACA			II	II
93	24959	基因组SSR	ATCCTCACCGGTTTGATGAC	TGGAGAGTGATAGAGAAAAATTGTG	59	115	II	II
94	25059	基因组SSR	ATGGATTGCGGATAGCTCAA	CAGCAGTTGTTCGCAGGTAA	60	189	II	II
95	24301	基因组SSR	TTGTGTTTTCCGGAGAGGTC	TCCCTCCCAACCTTGAATTT	60	142	II	II	II
96	AA332	锚定标记	TGAAAATAAAGGCATGCAAATA	TGATTAGTCAACTTGTTGTGGA	51	255	II		II
97	29220	基因组SSR	GGGGCAGATTTGTGGTATTG	TTCTTCTTCCTCACGTCTTTCTTT			II		II
98	AD83	锚定标记	CACATGAGCGTGTGTATGGTAA	GGGATAAGAAGAGGGAGCAAAT	61	270	II		II
99	18272	基因组SSR	CCCCAACATTTCTCTAGGTAACA	TTCTTCGCAGCTCGGTAAGT	59	131	II	II	II
100	S42	基因组SSR	AGTTTCGGGTTCCTTGGAGT	GTTGGCGTAGAACGATCCAT	53	211	II	II
101	e198	EST-SSR	ACCATCACCACCAACAACAC	CTGCATCTGGAGAGGGAGAG	59	188	II	II
102	e14	EST-SSR	TCCGCAATGTTCTCTCGAAT	GGAGGTCTCCGCATTATCAA	60	188	II	II
103	21776	基因组SSR	AACGGATATGCATGGAGAGG	AAAACACGACCATCCTTTGTG	60	172	II	II
104	21939	基因组SSR	GGTCCTCAAGCACCACCTAA	TGGGCGTCACTACTTAACTTTT	58	114	II	II
105	28257	基因组SSR	AAGGGCTGACGGTCTAACTG	GAACTGACGGACGCTAGAGG	59	161	II	II	II
106	16237	基因组SSR	GCAAACGAAGCAGGCTTATC	TTGGCTGATCCTGAAACTGA	59	152	II	II
107	18260	基因组SSR	AACCTTGAAATGGAGGTACATGA	GACCATGATCGGATGTTGTG	60	129	II	II
108	21641	基因组SSR	CGATTTACCGTCCTTCATCA	ACGGTCTCCCATGTGTTTGT	59	166	II	II
109	3015	基因组SSR	ACACATGCACACCCCCTTC	TGCGTGTGTACGTGTGTACG	60	153	II	II
110	D23	锚定标记	ATGGTTGTCCCAGGATAGATAA	GAAAACATTGGAGAGTGGAGTA	51	170	II	II
111	e1031	EST-SSR	CAACACAAGAACTTTGCACCT	TTGATCCACCTGCATCATTG			II	II
112	e1215	EST-SSR	GAGACAGAAGACGGCGAAAG	TGCCAAGAGTCAGGAGATTG			II	II
113	16433	基因组SSR	CACCGCAAACATAGCAAAAA	TCTCATAGCTGCGAGGTTCA	60	127	II	II
114	e617	EST-SSR	GCCAAACGGCTTTAAAACTTC	TCGCTGTTGGAAAGAGAAGAA			II	II
115	20723	基因组SSR	CTCACTTCACGTGCGCTATC	GCAGGAGCAGCTTGATTTTC	60	152	II	II
116	16437	基因组SSR	TTGTTTTTGTTGTTCTTGTTGTTG	TTTTCGGGTTTTGCTTATGG	59	128	II	II
117	17531	基因组SSR	TGCAGGGGTGTGTGTTACAT	TGAACATGGTGAAATGGATTG	59	140	II	II
118	18237	基因组SSR	GGGATATGAGAAGGCGATACC	TGGTTGTAGGATGTGGGATTT	59	127	II	II
119	17754	基因组SSR	AGCAACGGGCAACCTTATAG	CCTTTTGTTTGGAAGCTCAA	58	169	II	II	II
120	e850	EST-SSR	TTTCTTCTCCCAAACTACCTCAT	ATGCATGAACCAACCCATCT			II	II
121	17713	基因组SSR	AAAAAGGGGAAAGCAGGAGA	TTGACTGTGAGGCTGGTTTG	60	164	II	II
122	21945	基因组SSR	TTCACGCTCATCGCTAAGAC	TTCGAATCCTCCCTTCTTGA	59	113	II	II
123	e478	EST-SSR	GCCACCAACCAATTCAACTT	TGGGTATTGGGAATGGAAAA			II	II
124	e36	EST-SSR	GCAGGGTGGGTATATCTGTGA	GTGGTCCAATTCCTTTTTGC	59	219	II	II
125	e26	EST-SSR	TTTTGTCCCCGCGTTTAATA	CATTCATGCCACAAAAATGG	60	155	II	II
126	C24	基因组SSR	GCTACTGGAGGAGGCTTTCA	GCCTTCTACACAACGGCTTC	52	162	II	II
127	28383	基因组SSR	TCGATTGTTATTGTGTTTCCTCTC	TGAGATCAAGTGGGGGAAAA	60	150	II	II
128	e770	EST-SSR	GGTTTGAAAGGACCCCTAGC	GTTACCGATGGCCATGAATC			II	II
129	143	基因组SSR	ACATGCACACGTACACGCA	AGTTGGCGTGCAGTAGAGGT	60	69	II	II	II
130	16534	基因组SSR	TTGCAAATATACCAATTCCAAAA	ATTGGAGCCTGGTGAAGACC	58	139	II		II
131	28055	基因组SSR	GCCAGCAATTTTAGCATTACG	TTAGCTCAGCCCGGTAAAAA	60	163	II		II
132	25792	基因组SSR	GACGGAAACGAAATCGAAGA	TCAAAATTCACGCACACGAT	60	110	II	II	II
133	30024	基因组SSR	AGAGTGCCATCCCTTCAATG	GAACGTTTGGTTGGAGGAGA			II		II
134	25769	基因组SSR	GCAGCAAGATGGTTGGTAGTT	GACGTCGTAGTCGCCATCTC	59	144	II	II	II
135	e929	EST-SSR	TGAGGAGGGAGATGGAGAAA	GAAGGCAAACCTACCAACCTC			II	II
136	30218	基因组SSR	GGACGTGTCCCACTCCTATG	GGAAGGATAAAAACGTTGCAATA			II	II
137	30416	基因组SSR	GACACATGGAGCCACAAAAA	GAATGGAGGGGAGAGATGAA			II	II
138	e577	EST-SSR	TGTCATTTCTTTTTAGTTCCTTTCAA	CCTTCGCTTGATTCTTCACC			II	II
139	16570	基因组SSR	CAAACACCAACCACCACAGT	AAGGGGAGACGAAGTGGAGT	59	143	II	II
140	27099	基因组SSR	GGTACACCCACCGATACACC	TCTAGACGCGGAGAGGGTAA	60	140	II	II
141	27112	基因组SSR	GCAACAAGATTTTGACGTTTTT	GACGCTACCAACCGCTTTAG	58	129	II	II
142	28687	基因组SSR	CACGGAAGGCCCTACTTACA	GTGGCGAGTAGAGCGTAAGG	60	189	II	II
143	28704	基因组SSR	TTCTGCAGTCAGCTTCAACTTC	TAGTCACGGAAGCGATTCAA	59	147	II	II
144	22690	基因组SSR	GGTTCATCTGCACCCAAGTT	GGCAACTCTCTCACACACACA	60	138	II	II
145	24812	基因组SSR	GACCAAACCACCTCACAGATG	TGGCTCCTTTCTCATTTCTAACA	60	140	II	II
146	23154	基因组SSR	AAGACGAGGTGGCATGGTAG	GAGAATGCATGCTTCAATCAA	59	183	II	II
147	28785	基因组SSR	GATCCACCCAATTCCCTTTT	TGTATTGCAGCCGCTTTATG	60	208	II	II
148	19680	基因组SSR	GCCAACCCAACAATCTCAAC	CATTGGAACCAGATCGAACC	60	148	II	II
149	22052	基因组SSR	GTTACCGATGGCCATGAATC	AGCGAGTGAAGAGGGAAGTG	60	147	II	II
150	1078	基因组SSR	CACACGCACAGACACACGTA	GTGTGCGTATGCGTTACTGC	60	99	II	II
151	29138	基因组SSR	GCAGATTGAAACCAAAACGAC	TCGCAACCTGCACTTTCTTA			III	III
152	e763	EST-SSR	ATGCTTTGATGGGCTCAACT	TCCCAAACATGCTAGCAAAA			III	III
153	19532	基因组SSR	CCATGTTTGAATTCGGAGGA	GCGCGATGATTCAAGGTTTA	61	165	III	III
154	19656	基因组SSR	CCAACGTTGTTGTTTTAGTGG	CCAGAGTCGTGGAGCCCTAT	58	169	III	III
155	27876	基因组SSR	TGTTGTTGCCCATCAATCAT	AATCACACGAGGGATTGGAC	60	149	III	III
156	18323	基因组SSR	CAGACAATGGCAATTATTTGGTAA	CTGCTGTTGCTTCGATTTCA	60	136	III	III
157	21881	基因组SSR	CCATTCCCAACAATTTCCAC	GTGAGGTCCGGTTCTACAGG	60	119	III	III
158	30088	基因组SSR	TACTGGATCCGGATGAGGAC	TCGCATCAAAGCAAAAACCT			III	III
159	e605	EST-SSR	AGCACTTGCTACGGCAATTT	AAACCTAGAAATAACGATGCAAAA			III	III
160	24265	基因组SSR	GTTTGCGGCCAAACAATATC	TTTGCATGAGTGCACCTCTC	60	181	III	III
161	29627	基因组SSR	AGAAGACAACGACCGAGTGG	AACCGCATAACCGCAATTTA			III	III
162	27019	基因组SSR	GCAGTTTCCACACTTTAAGTCCA	TGGGTGTGTTAACAGGGTGA	60	155	III	III
163	e1208	EST-SSR	AACCATTGCGCGTCTTTTAC	AGACCACCGCCATAACATTC			III	III
164	28130	基因组SSR	CGAATTTGGTTGCGACGAC	TCGCGCCTCTTTAGGAATAA	60	208	III	III
165	27378	基因组SSR	GCCAATTATTCCCTCCAGGT	TTCGAAGGTTCTCCATCACC	60	147	III	III
166	28180	基因组SSR	TCGTTCATTTAACTTCGTGAGGTA	AGAGAGTTCTCCGCCAAACA	60	201	III	III
167	24903	基因组SSR	TGGTGTCAACTTTTTGATGTTCA	GACAAGTTGCTTTTGCTCCAC	60	147	III	III
168	26850	基因组SSR	TCACAGACAGTACACAAAGTTTTCTT	GCGAGGGAGAACAGAAACAG	59	160	III	III	III
169	24521	基因组SSR	AGGGAACCCCCAATTGACTA	CCAGAAACTGGGGTTGTGTT	60	208	III		III
170	23311	基因组SSR	TTTCAGAATGGTGCAGGGTAT	AGGATCTCAGTATACATGCGTAAA	57	168	III		III
171	24540	基因组SSR	CTCCCTCATGAGTCGTGACC	ATCAAAGGGGGAAGGTGAAG	60	143	III		III
172	26575	基因组SSR	GAAAATAAACAGTTGGCAACAAAA	CCACTCCAAACCCTTCAGGT	59	205	III		III
173	21726	基因组SSR	GGTGATGGAGAAAAGGGTGA	TGCATGCAGTCAAATCAAAA	59	157	III	III	III
174	23518	基因组SSR	CAAGGACGACGACAACAACA	GTGCCGACGTTCAAGAAAAT	60	202	III	III
175	e632	EST-SSR	AAACCTCTCTACAGCACCAACAC	GGGAGAGATTGTTTGAAGTATAGAA			III	III
176	28406	基因组SSR	CCGATTGTGCAGCAAGAGTA	ACGATGCACATGCAGAATTT	59	129	III	III
177	27038	基因组SSR	CGTCTACCTCCGACGATAGC	TTCGCCAGATATATACAATAAAAAGA	58	187	III	III
178	25151	基因组SSR	GCCTTCGAGGCATCCTAAT	TGGAACCATAAGATTGGTGAA	58	157	III	III	III
179	27270	基因组SSR	GGACCATTACCCTCCCATTT	TTCCTTCCGTTTTGCAGTCT	60	194	III	III
180	28194	基因组SSR	TGGGGTCTTAAAGTTGTGACTTC	CCGAAGGTGGGATGAAACTA	60	148	III	III
181	28645	基因组SSR	CTGAAATCGGAGTGGTCACA	GGTGAAGCCCTTAGCTACCA	59	188	III	III
182	17193	基因组SSR	CACAGCCATACCCAAGTTACAA	GGTTGCGAGGGATGAGAATA	60	178	III	III
183	e814	EST-SSR	GGTTGGTCCAATCCAACATC	AGAACGAACACACACGAAACA			III	III
184	e494	EST-SSR	GACCCGTCTGGACTGGTAAA	TTGAAAGATGCGGAGTGATG			III	III
185	29436	基因组SSR	TCTAGCAGCATTGGGGAAAC	TAATGAGGGGAAGGGGATTT			III	III
186	27710	基因组SSR	AACCACAGAAAACTGCCAAGT	TGAGAACCAAAAGCAGGTCA	59	110	III	III
187	29074	基因组SSR	TCGTCGAATGGTTGAAGAGA	TTCGCAAGTGAAGGAAAAAGA			III	III
188	29307	基因组SSR	GCGATTCCAGATGTCAGGTT	CGTCTCCCTACCAGCAAAAA			III	III
189	24534	基因组SSR	GTGGTGTGAAAGGGGTTTTG	CTTGCATTGGATTCCCTTTG	60	206	III	III
190	22543	基因组SSR	TTTCACGTACGTTCCCAACTC	CCAGATCAACCACCTAACTTCA	59	160	III	III
191	23262	基因组SSR	GGTGACGGAGGTGATAGAGG	TAGCAAATGCAAACCCAACA	60	183	III	III	III
192	24701	基因组SSR	TATGCTGGAGTGTGGAGTGG	TCAATCAATTCAACGGTACAGA	58	111	III	III
193	24784	基因组SSR	TTTAGACGGCCTTCGTTAGTG	CTGAGCCTAAAGGGCTGAAA	59	115	III	III
194	27254	基因组SSR	GAAGGCCTCTAACGGTGAAA	AATCAAACAGAGGCCACCAG	59	122	III	III
195	22754	基因组SSR	GGAACGACAACACGAACCTC	GACACGTTATGCGCACACTC	60	161	III	III	III
196	26131	基因组SSR	GGAAACGGTGGAAGATGAAA	TTGGCAAAAGGGATGAGAAG	60	175	III		III
197	24805	基因组SSR	TGCAGCAGATCAACCAAAAC	TTTTGAACTAAGGTGGTCTCAATC	59	144	III		III
198	18135	基因组SSR	CTTCAACCAACTGCGAGTGA	TCATTTGAGTTTTGCCATGTTC	60	120	III	III	III
199	PSAJ3318	锚定标记	CAGTGGTGACAGCAGGGCCAAG	CCTACATGGTGTACGTAGACAC	58		III		III
200	29634	基因组SSR	CAATTAACAAACGCAGCCTTA	TTAGCCCGTGGATTTTCAAC			III		III
201	26866	基因组SSR	CGATACATTAAGGGCGGAAC	TGACTCATTCGCATTTGGAGT	59	186	III		III
202	23757	基因组SSR	TGAAAGAGGGGAATTGAGAGA	TCAGGTTACAAGCCCGAGAT	59	187	III		III
203	AA355	锚定标记	AGAAAAATTCTAGCATGATACTG	GGAAATATAACCTCAATAACACA	51	180	III	III	III
204	23899	基因组SSR	CCCCATCCTTGTGAACAAAT	ACGGTGTTTTGGTGGTGAAT	60	151	III		III
205	AD270	锚定标记	CTCATCTGATGCGTTGGATTAG	AGGTTGGATTTGTTGTTTGTTG	51	290	III		III
206	20075	基因组SSR	GCCAGTCCCTTGAGTTAGGA	TGTTTCACGTGTTCCCCATA	59	122	III	III	III
207	e696	EST-SSR	CGCTATCACTCTCTCTTTCTCTTTT	ATCGGAGGACGAGGTCTTTT			III	III
208	e1021	EST-SSR	AAAAAGCTGAAACATTCAGACC	GGTCCATTCATTCTGCAGTG			III	III
209	e673	EST-SSR	GGAAGACGGAGTGGTGGTAA	GTCGTCGTGGTGCTTCTCTC			III	III
210	16524	基因组SSR	CCAGAGGATGTGAACCAGGTA	TTCAACCAAGCTGAACCCTTA	60	138	III	III
211	4826	基因组SSR	AACATGCGTCTGTCGTGTCT	TAGTGGGTGTGCGTGTGAGT	59	220	III	III
212	27093	基因组SSR	TCGTCATTCTCCCTCCAATC	TATGATGTCCACGCGTTTTT	59	128	III	III
213	19688	基因组SSR	GAATCGGGTCGCTGAGATAG	ACCTCCACCGTACCATTCAA	60	133	III	III
214	e998	EST-SSR	GGCAGACTGGTCTCAACTCC	ATCATCGTTGGTGGAATCGT			III	III
215	2875	基因组SSR	TATTAGCACCCCTCACGTCC	TTTCCCCTTCCTTCCAATCT	60	148	III	III
216	30442	基因组SSR	CCCACTCCATCAGTCTCTCC	AGAGGACCGGTGATGTGTTC			III	III
217	e446	EST-SSR	AACCAGAGATGAGTGGAAAAATG	GCCAACACCAGAGTTTGAATC			III	III
218	27160	基因组SSR	CTGCAGTTGCGTGTAGAGGA	TTGAATGATGATATAAATGCAATGAC	59	135	III	III
219	e472	EST-SSR	TCCATCACCAGGCATAGGTC	GCCGGTAGTGAGAAGGATTG			III	III
220	864	基因组SSR	CACATACATGCAATCAAGCG	GTGTGTTACGTGCGTGTGTG	59	92	III	III
221	24861	基因组SSR	TTCACAACCCCTCTCACTACA	TGGAGGGATGGTTTACAATGA	58	183	III	III
222	30382	基因组SSR	TTGGTGAGGCCTTGATTTTT	GCCAGTGGGGATTAGAGACC			III	III
223	17773	基因组SSR	TTCCACACGAGGCTATTTTC	TGCAAAAGCGACATCTTGAC	58	170	III	III
224	18363	基因组SSR	CATGCATGGAGTTGGAAGAG	GTCCCAAAATGCAGCCAATA	59	139	III	III
225	29125	基因组SSR	TCAGAGGTGTCATCGGTCTG	TTTCAAATAAGTTTTGAACAAAGTGT			III	III
226	e999	EST-SSR	GTTTAGGAGCCTTGGGGTGT	TCCAAACTCCGGCTTCTCTA			III	III
227	3647	基因组SSR	GGGGTCTTACAACACACGCT	AGGCAGAGGTGTGAGCATCT	60	176	III	III
228	6144	基因组SSR	CAGAAAAGGAAGCAAGGTGC	GTCGCCCTCGATTCTCATAC	60	297	III	III
229	4535	基因组SSR	CAGAAAAGGAAGCAAGGTGC	TGTGTGTACGTGTTCACCCTT	59	206	III	III
230	3374	基因组SSR	GGGGTCTTACAACACACGCT	TGAGCTAATCTCTCCGGGAA	60	165	III	III
231	2169	基因组SSR	GTTGTGTGTGTGCGTGTTCA	GGGGTCTTACAACACACGCT	60	126	III	III
232	1935	基因组SSR	GGGGTCTTACAACACACGCT	TGTGTGCGTGTTCACCTTTT	60	120	III	III
233	2303	基因组SSR	GGGGTCTTACAACACACGCT	TGTTGTGTGTGTGCGTGTTC	60	130	III	III
234	4653	基因组SSR	CAGAAAAGGAAGCAAGGTGC	TGTGTGCGTGTTCTACCCTT	59	211	III	III
235	5347	基因组SSR	CAGAAAAGGAAGCAAGGTGC	AGCATCTCTCCGGTAACCCT	60	248	III	III
236	2008	基因组SSR	GGGGTCTTACAACACACGCT	TGTGTGTGCGTGTTCTACCTT	59	122	III	III
237	5412	基因组SSR	CAGAAAAGGAAGCAAGGTGC	GTGAGCAATCTCTCCGGGTA	60	252	III	III
238	2051	基因组SSR	GGGGTCTTACAACACACGCT	TGTGTGTGTCGTGTTCTACCTTT	60	123	III	III
239	5400	基因组SSR	CAGAAAAGGAAGCAAGGTGC	GTGAGCAATCTCTCCGGAAC	60	251	III	III
240	5348	基因组SSR	CAGAAAAGGAAGCAAGGTGC	AGCAATCTCTCCGGTACCCT	60	248	III	III
241	4670	基因组SSR	CAGAAAAGGAAGCAAGGTGC	TGTGTGTGTCGTGTTCTACCC	60	212	III	III
242	5540	基因组SSR	CAGAAAAGGAAGCAAGGTGC	AGGCAGAGGTTGTGAGCAAT	60	260	III	III
243	21506	基因组SSR	ATAGGGGGAGCAGGACCTAA	TTTGACTTGTGGAAAGGAAGTT	58	151	III	III
244	25416	基因组SSR	CGCCCAATTGGATTATGATT	TGCTCAATGCACACTTACTAGC	58	145	III	III
245	e1197	EST-SSR	TCCAACCGTTAAACACTCTTCA	GCATGAAGGGCTCTGAGTTC			III	III
246	PSBLOX13.2	锚定标记	CTGCTATGCTATGTTTCACATC	CTTTGCTTGCAACTTAGTAACAG	56		III	III
247	18562	基因组SSR	TTCTTCTGCTGCTGCTCAAA	AAAACAAAAACCACAACCAAAAA	60	154	III	III
248	21610	基因组SSR	CGATTGATGCCGTGTCTAAG	TTTCAAGTTTCTTCTAGATTTTGTCA	58	114	III	III
249	27991	基因组SSR	AATACAGCTGGACCCCACAC	GCAGGCCATTTCATTTCATT	60	121	III	III
250	22724	基因组SSR	CCCAAGAAGAAGGATGGTGA	GAGCATTCTGGTGCTGTTGA	60	152	III	III
251	26028	基因组SSR	CGGCGAGATTTATTGACGAT	CAACGTGGCAAGCAAGTAGA	60	186	III	III
252	4397	基因组SSR	TAATCTCTCCGGGAACCCTT	TGATATGATGCCATGAGGGA	60	201	III	III
253	20084	基因组SSR	CTCCCTCCCGAATGTAATCA	CGGGACGATCAACTTTGTCT	60	111	IV	IV
254	16758	基因组SSR	CCCTTCAACAAAGCCTAACG	AGGGTGCGAAGGAGGTTAGT	60	115	IV	IV
255	21712	基因组SSR	CGGCGGGTTTGATAGAGTTA	CTTCACCCTTGCAACAAACA	60	166	IV	IV
256	21774	基因组SSR	GCAAGTTCCCAATCGTCCTA	TCAAAAGCAAGGTCCCATTC	60	123	IV	IV
257	e671	EST-SSR	TTGCCTCATTTATCATTCTCTTATG	CAAAAGGTTATCTAGCTACGACTTGA			IV	IV
258	29964	基因组SSR	CAATTCATGACGAAATTGACAAA	CATGGAGATGGAGAGTTCAAAA			IV	IV
259	e996	EST-SSR	GCCGGTAAACGATCCATCTA	TGCAGCCACACTCCTTTACA			IV	IV
260	19632	基因组SSR	AATGTAATTAACCCACGAAGTTG	TGCCAAAGCTCTCTCATCCT	57	201	IV	IV
261	21547	基因组SSR	TGAAAGCCTCAAAGCAACAA	CCATGGCATGTGCTAGTGTAG	59	156	IV	IV
262	21250	基因组SSR	GTGCAATTTTCACACAGTGG	ACGAAGGTTGGAGCATGATT	58	144	IV	IV
263	22067	基因组SSR	TATGCTCAGAGGGGCATAGG	TTACGACGATGAGCGACTTG	60	158	IV	IV
264	18533	基因组SSR	TCCAAAATGCGTGTCATCAT	TGACCGACACATTCATCTTCA	60	151	IV	IV
265	26592	基因组SSR	TCCGATCCTGGTAAAGTGGT	TCCAAAATGCGTGTCATCAT	59	174	IV	IV
266	26369	基因组SSR	GCTGAAACGTGGGAAACATT	TGGTTAGTGTTTGAAGGGTCTG	59	206	IV	IV
267	25046	基因组SSR	TCCTTTGTCAGTGGGAATTTTT	AGGATCATGGTTGTCGAGTTG	60	173	IV	IV
268	24606	基因组SSR	CCGAACAAGATGAACCACCT	AACAACATCGTGTGTGTTTGTCT	59	208	IV	IV
269	23750	基因组SSR	CTCGTTGTACAATCCAGATGAA	CACCGTCCACCTTCTCATTT	58	209	IV	IV
270	22599	基因组SSR	GGGTGTGAGGCAGTTGAAAT	AAATACCGAACCGAACCACTT	60	141	IV	IV
271	24038	基因组SSR	GCCCCCACTTTTTCAACA	CTCCTGACACAAGGCCCTAC	59	145	IV	IV
272	18438	基因组SSR	GATTGAGCCGTGCCAATATC	GATCCCACCCTAGAGGAAAAA	59	145	IV	IV
273	22352	基因组SSR	CCAACATCTTCCTCATCACCT	TGAGAGTCGCAGTCGGATAA	59	129	IV	IV
274	16512	基因组SSR	TAAGCCCGACGCTTCTATTC	GTGCCTCAGTTTCCGTTTGT	59	136	IV	IV
275	AA430942	锚定标记	CTGGAATTCTTGCGGTTTAAC	CGTTTTGGTTACGATCGAGCTA	54		IV	IV
276	17219	基因组SSR	TCATGTGCATGTGATGAAGAAA	GGTGTACCCATGTGCCATTT	60	181	IV	IV
277	28178	基因组SSR	TCAACCCATACTCTTGGAATCA	CCGGAGATTCCACAAATAACA	59	139	IV	IV
278	28085	基因组SSR	TGCTTGCAACGTTTCTTTCTT	ACCCCTCCCTGAAAGGAGTA	60	142	IV	IV
279	e538	EST-SSR	CTTCCCTTTCTTCCCTTTCA	GTTCGGAAGGATCGATTTGA			IV	IV
280	28374	基因组SSR	TCCACGGTCTTGCTATGTGT	CTGGTTGCACATCAGGGTAG	59	187	IV	IV
281	e578	EST-SSR	AGCAGCTCATATTCTCTGTCCA	GCAGAAGCAGGATCTAGGGTAG			IV	IV
282	e656	EST-SSR	AGCAGCTCATATTCTCTGTCCA	AGCAGAAGCAGAAGCAGGAT			IV	IV
283	e650	EST-SSR	GCAGCTCATATTCTCTGTCCA	AGCAGAAGCAGAAGCAGGAT			IV	IV
284	29955	基因组SSR	TCAAGTGCATTGGGAGAGACT	AAAAACCGACCCATAATCAATTT			IV	IV
285	e1073	EST-SSR	CTTGTTTCGCTCGGTACCTC	CTATTGCAGGCAGTCCTGGT			IV	IV
286	24844	基因组SSR	TTTCGTTTTCCCATTCTGGT	CCCCCTTACACACGAATCTG	59	121	IV	IV
287	26018	基因组SSR	TTGAGCTGCTCGCTGTAAGA	CACCAAACTGTTTCTTCACACA	59	146	IV	IV
288	26179	基因组SSR	GGAAGGTGAAATTCCGTTGA	AGCAAGTTGGTTCGGAAAAA	60	166	IV	IV
289	24485	基因组SSR	GTGTCAAAGCTCGCCCTAAT	ATTGGTACACTTGCCGAGAA	58	127	IV	IV
290	e1130	EST-SSR	GCTGCTACCGTGGATTGTCT	TTGAAGAGGTGTGGTGTGGA			IV	IV
291	e1027	EST-SSR	GTCTCGGTCCGAACCATTTA	TCTTCTGATCAACAAAAGTAACAACA			IV	IV
292	S5	基因组SSR	TGTGGGGCTTGTTACACTGA	AGCTACCATAACAGACAAAACC	54	205	IV	IV
293	22276	基因组SSR	ATGCGGCATTTTGCTTTATC	TTGGTCTGCAAATCGAAACA	60	127	IV	IV
294	29630	基因组SSR	CCTACCTTAAGTCGCCCATGT	TCCAAACATAGGCTTCGTCTC			IV	IV	IV
295	AA285	锚定标记	TCGCCTAATCTAGATGAGAATA	CTTAACATTTTAGGTCTTGGAG	51	248	IV		IV
296	28304	基因组SSR	TTTTCAGCTGATCGGATATCTACA	GGCAGCATCTTGAAAATCGT	60	132	IV	IV	IV
297	e535	EST-SSR	AAAAACCAAGCACACCCATAA	GCAAGACACAGCACAAAAACA			IV	IV
298	27384	基因组SSR	TTGTGCCAACAAAAATCACAA	CAGAATGCCGTTCACTTTTCT	59	115	IV	IV
299	e561	EST-SSR	TCCGATCTTGCTTCTGAATCT	TTCATCAACCCAGACGCATA			IV	IV
300	B83	基因组SSR	CCCTCTCCCATCTCATCTCA	AAAGAAAGTAGAGATCCAGCACTGA	54	205	IV	IV
301	24635	基因组SSR	CCCTCTCCCATCTCATCTCA	AGAGATCCCAGCACTGATTG	58	152	IV	IV
302	23788	基因组SSR	TGGAGAAATTATGAGAATGTTCAATG	GCACGTCGACACACACAAAC	61	184	IV	IV
303	24392	基因组SSR	GCGGAAACAGGAGAGAGAGA	GCACGTGCTCCATCATAGAC	59	121	IV	IV
304	e828	EST-SSR	AACCCCATTTTCAATATTTTTCA	GGATTGTTCTCCGCATCTTC			IV	IV
305	18358	基因组SSR	CCTGAACCGATTTTGGTGAT	ATTCCGCCCTCTTTCACTTC	60	138	IV	IV
306	27275	基因组SSR	TGCTCACATTAACCAAAAGCAC	TGGATGGGTATGTCCCATTT	60	147	IV	IV	IV
307	27252	基因组SSR	TAATGCCGACTGTGTGCTGT	GCAATTCAGCAAAAAGGAGAA	59	111	IV		IV
308	AA122	锚定标记	GGGTCTGCATAAGTAGAAGCCA	AAGGTGTTTCCCCTAGACATCA	61	190	IV		IV
309	28244	基因组SSR	TGGGAGAGGGGATAACTGAA	CATGTTGTTTGGTGCGTTTC	59	182	IV		IV
310	19487	基因组SSR	CCACCTGCTCAATTCCAAAT	GGCGAAGCGAATCTAACATC	60	178	IV	IV	IV
311	22506	基因组SSR	CGAAACATGCACAACCATTT	TGAACGTTCTGACCCAGATG	59	208	IV	IV
312	22693	基因组SSR	CGACAACAACAACCACATCA	CTCCATCGAACGAAAGGAAC	59	148	IV	IV
313	24465	基因组SSR	TCAAGCAGAAGAGTCGACCA	TAGCTATGTTCCCGCCAAAT	60	161	IV	IV
314	25164	基因组SSR	CCAAATACAAGCATTAATAGGGAGA	TGGTCGACTCTTCTGCTTGA	60	110	IV	IV
315	25419	基因组SSR	TGCAAGTCCTGATGCAAGTC	GCGATTCAGGATTGGCTTAC	60	140	IV	IV
316	27605	基因组SSR	AAATGAACGGAAACAGAAAGAA	ACATAGCACACGCAGCAAAC	58	155	IV	IV
317	29578	基因组SSR	CAAGTCATGAACGTCTCAAAAGA	TGGACGCGTTTTAAAGTTCC			IV	IV
318	27854	基因组SSR	TCCTTCATCAAAACGCAACA	ATTGACGTTCAAGCGGGTAG	60	138	IV	IV
319	e155	EST-SSR	TTTCTCGTTGCACTCATCCA	TCGGTTGTCGTTTCTTGTTG	60	174	IV	IV
320	21866	基因组SSR	GCAGCCTTCAAATCCTCTTC	AAAACGCGCTTACGCTTCTA	59	139	IV	IV
321	29760	基因组SSR	TGTGCCTCAGAGATGTTCAAA	AGAGGTGGTGCGGTGACTAT			IV	IV
322	18049	基因组SSR	ACCCCTCTTTGCTAGGGTGA	ACCACACATCTCGCACACAT	60	202	IV	IV
323	e342	EST-SSR	CACAACAACCCCTCCAAAAC	TTTGGATTTTCGCTTGGGTA	60	189	IV	IV
324	e546	EST-SSR	TGACAGTGAGTGAGTGGCTTCT	TTGCGGGTGAAAAGAAAAAG			IV	IV
325	21805	基因组SSR	TGGGAATGTGAAGTGGTGAA	TGTGGTGTGGTTGGTTTCTG	60	128	IV	IV
326	21622	基因组SSR	ACAGCATGAAATGCGTGAAA	TCGTCATCCCAACTTCATCA	60	139	IV	IV	IV
327	AA315	锚定标记	AGTGGGAAGTAAAAGGTGTAG	TTTCACTAGATGATATTTCGTT	51		IV		IV
328	79	基因组SSR	GCTCAGTCAGCCCGTCATA	GTGCGTGTGTGCGTGTGT	60	66	IV		IV
329	17066	基因组SSR	CTCTCCCCCACACCTGATAA	GAGGACCCAGTAGGGATCGT	60	159	IV		IV
330	24036	基因组SSR	GAAGGACCAAATCAATTCTCTAAA	ACCGACGTCAACGACTGATA	58	196	IV	IV	IV
331	24423	基因组SSR	CATCCCACTCTAACCGCACT	GCATAATCGGCTCTCTCTCC	59	184	IV	IV
332	25717	基因组SSR	CGTGCATGCATGTGTATGTT	TCACCGATCAACACCAATTT	59	192	IV	IV
333	28434	基因组SSR	GTTTTCAATCGATCCGTCCA	TTCCACCGTCTTCTTCAACA	59	164	IV	IV
334	e1213	EST-SSR	TTGGTTTCCGGTTAAAATGA	CAATCCCATTCACACCACAA			IV	IV
335	e782	EST-SSR	CATTGAGTTTGAGGATGAGGA	CCCATAACCATATCTCACAGTTCA			IV	IV
336	e336	EST-SSR	CCCCAAACCATATCCCTACA	TTCCATTCCCAAACTCACTTG	60	170	IV	IV
337	21227	基因组SSR	CGGATTCAACAAGCAGAACA	CGAGAATGGAGGAAGAAGTTG	60	153	IV	IV
338	S236	基因组SSR	AAATGGCCGTTTTATGATCG	CGGAGCTGAACCTTCTGGTA	53	604	IV	IV
339	e182	EST-SSR	TGGTAACCCTAGCAATCATCA	CTCTTTGGCAACAACATCTCA	58	241	IV	IV
340	28173	基因组SSR	TGCATTGCTAATAACATTAGAACCAT	TTCCTTTTAAGCAAGGTGAGGT	59	197	IV	IV
341	29016	基因组SSR	TTTCAAAGGCAAGGCAAAAC	CACCTCGCAAAATTGGACTT			IV	IV
342	e273	EST-SSR	CAACAACTTCTACAGCAGCAA	GCAGTAGCATCTGGCTGTGA	57	152	IV	IV
343	e1121	EST-SSR	AACAACGGCAACAACAACAA	GTGGCCTTAGTCCCAAGAAA			IV	IV
344	e908	EST-SSR	TGCAGTGATGAAGTGGTTGA	CACTGCTCCATATCCCACAA			IV	IV
345	27057	基因组SSR	TGACCCTAGCAATTAGGATTTGA	ACCATGCCTCCAAAAACTTG	60	160	IV	IV
346	24907	基因组SSR	AAGCAATCCTAATCCATGTGTG	CATCCTTTCCGCCTTTGTTA	59	134	IV	IV	IV
347	22155	基因组SSR	ACCCGAGTCAGTCGCTTATG	AACACGGCTTCAATTTACGA	58	135	IV	IV
348	e878	EST-SSR	CGCATTTTCACTCCACACAC	CGTTCGGAACATCCAAGGT			IV	IV
349	o79	基因组SSR	TTGTCTTCACCACCTTAC	GATCATCAGCCAATAGTT	52	300	IV	IV
350	27068	基因组SSR	TTTCGGGCGTCAAATAATTC	GCCACACCTCCAAATGAGTT	60	152	IV	IV
351	20339	基因组SSR	CCTTCCGTGACCAAGAGAAA	GGTGGATGAGATGGATGAATG	60	146	IV	IV
352	e307	EST-SSR	CTTGTCAGCTTGGCATTCAA	GCGAGTTTCCATTCATCACA	60	177	IV	IV
353	e360	EST-SSR	CTTGCCCAAATTCAAGCTGT	CTCTATTACACAAATGCCAGTG	55	239	IV	IV
354	B110	基因组SSR	CCTCTTCAACGGTACGAGGA	TTGCAGAGAGACGAGAGAGAAA	56	208	IV	IV
355	27300	基因组SSR	CGGCAGTATTTGCAACAAGA	CCTCAAGGCCAGATGATTTT	59	159	IV	IV
356	e1029	EST-SSR	TCATTGCATGCCATTCTTTC	CGAAATAGAGAAAAGATAGAACCAG			IV	IV
357	29971	基因组SSR	CGAAATTGAATGCAGGAATG	CAACCTCCAAACTCCAAACAA			V		V
358	28387	基因组SSR	GGCTCCATATCATGTTTCTATGC	AAAAGGAGGGAACATGGAAGA	60	201	V	I-2	V
359	22059	基因组SSR	ATCTTCCGCAACAACACACA	ACGTGAAACGGCACAGTATT	58	203	V	I-2
360	27937	基因组SSR	ACAAGGCATGGTATGGTGGA	TGAACACAAATTGCAGCCTAA	59	124	V	I-2
361	S220	基因组SSR	AGCTCTTTCTTCCACCACCA	CAGGTTCCAGCTGAGAGGAG	55	1001	V	I-2
362	e59	EST-SSR	GAGGGTTTCCCGACTTCATT	TAAAGGTTTTCGCCACCATC	60	154	V	I-2
363	e723	EST-SSR	GGGGGTGTCTTACGTTGATG	CCCCAAAACCAGCTGAACTA			V	I-2
364	29094	基因组SSR	CCCCAAAACCAGCTGAACTA	GGGGGTGTCTTACGTTGATG			V	I-2
365	18391	基因组SSR	CCATCCTCCACGTGTCTCTT	TCGCATATCCAAATGCAAAC	60	142	V	I-2
366	e121	EST-SSR	AGCTCCATTTTGGAGTTTGT	CCTGAACCTGATTATAGCCAAGA	56	211	V	I-2
367	e619	EST-SSR	AAGTCTCTCATACCTAACCAACCAC	GCAGCCAAATTTGAGGAAGA			V	I-2
368	20702	基因组SSR	GTTCTCCATCGCCTTCTTTC	TGTGTTATGCCGAGCTTTTG	59	151	V	I-2
369	e611	EST-SSR	CCACAACCCCCTCTCTCTC	CTGCGAATTCGGAAAGAAAC			V	I-2
370	e1060	EST-SSR	AGAAGTTTTGTTGGTGCAAAGA	TGCTCATTTCTTTACCTTTCTTGA			V	I-2
371	e1233	EST-SSR	CGTTCCTTGTCTCTCCTCAAA	ATTCCCAACATGCACCATTT			V	I-2
372	22184	基因组SSR	GGGCGAAAACAACTTCCATA	CCTGGATGCTCCCAAAATAA	60	149	V	I-2
373	26117	基因组SSR	CATCGGGCGAGATAACAAAT	TTCCAAGCCTCACTTTCTCC	59	202	V	I-2
374	26857	基因组SSR	TGCTACAAGTCTAAATACAACACTCTT	CGGGAAGAGAATGATGAGGA	57	133	V	I-2
375	24882	基因组SSR	TTTCTGGAACCTCGCAAAAC	TTGCCTCAATTTGGAGACCT	60	173	V	I-2
376	23829	基因组SSR	CGCTCGGCCATGTAACTTAT	GGAAATGGGACTGAAACTGG	59	199	V	I-2	V
377	24112	基因组SSR	TTGATCATCCTCTCGCTTTT	TGTTGTCGTCATCAAAACACAG	57	187	V	I-2
378	23525	基因组SSR	TGTGCTTTTCTCTTGGCTTCT	CCAGAGGAACCACAAGGTGT	59	125	V	I-2
379	24512	基因组SSR	AAGCGTACGTGGCAAGAAAT	TCCCTGGGAGAGATGAAAGA	60	171	V	I-2
380	24236	基因组SSR	CAAACCTTCTTTATTTCCATTTCA	ACTTCTGGTCCACGCAAAAC	59	152	V	I-2
381	e738	EST-SSR	CCAATGGACTAGGTGGTGGA	TGATGGATGGGGTGATCATA			V	I-2
382	25394	基因组SSR	AATGGGTTTTGCTACGTGGT	GGGTGAGTGGAGAAAGCACT	59	142	V	I-2
383	25076	基因组SSR	GCTTGCAAGTGTGCGTGTAT	CCAGCCAAATGCACAATAAA	60	181	V	I-2
384	25618	基因组SSR	TTCCATCGTGAACCTTCCTC	CACACGACTTGCAATGTTCC	60	204	V	I-2	V
385	e122	EST-SSR	TCCACCGACATCTCTTCTCA	AGGTGGTGGTTGTTGTTGGT	59	180	V	I-2
386	19657	基因组SSR	TCCAAACCCTAGTTAGAGAAAGAA	AGCACCATCATCGTTCATCA	58	188	V	I-2
387	28929	基因组SSR	GGACTTTTGCGGGTATGAAA	TGTCTCTTTAGATTCGTTCCAAAA			V	I-2
388	e5	EST-SSR	ATTAGGGCCGGATAATTTGG	TCCTCAGCAGCTGTCTCAAA	60	162	V	I-2
389	25089	基因组SSR	ATTCTTGTTGGCGAAACACC	TTGCATTACCCAAAGCTCCT	60	185	V	I-2
390	24228	基因组SSR	GCAAATTTTCGTTAAATGGATGT	GACAACCTGGAGACGCATTC	59	134	V	I-2
391	26521	基因组SSR	TGTCTAAGGGTGACAAAAGATCA	TGAACCCGCTCTTCCTTACT	59	159	V	I-2
392	25851	基因组SSR	AGGCAACACGAGGACGAATA	CGACGGAATTGAAAAACAAAA	60	152	V	I-2
393	25986	基因组SSR	CAATAGGCCGCGTAAGAAAA	TTGCCATCGATTTGATTTGA	60	158	V	I-2	V
394	22848	基因组SSR	GTGGTGGAAGAGCGTTTGAT	CATGGTGCGTTAACCCAGTT	60	190	V	I-2
395	22829	基因组SSR	TAGAAGGTTGCCTTGGGTGT	GCCCACCAAAGAAATCAAAA	60	114	V	I-2
396	24850	基因组SSR	TGGCACACATCTTCAATACAAA	GCACAACCGTTTTTGGTTCT	59	137	V	I-2
397	26012	基因组SSR	TGGCCCTCAACCTTGTATGT	GCAACACAGAACAAAGCACAA	60	138	V	I-2
398	29460	基因组SSR	TTCCTGACGCGGACATTAAC	GGAAATTCGGCAAGGACTTA			V	I-2
399	e599	EST-SSR	GCAATTTTCTCACTCCATCTCC	AAAGGAAAGCAACTCGGTGA			V	I-2
400	29839	基因组SSR	GAACCTCGTTTTTGCATCCT	AATGATAGGGGTTGCCACAT			V	I-2
401	21936	基因组SSR	TGTTGTTTGTTTGGTTGAGGA	CGTTGGCAAACATCATTATCA	59	187	V	I-2
402	22325	基因组SSR	TGCGAGGGATGAGTTTCTTT	TGTGTGGCCAAATCGAAGTA	60	187	V	I-2	V
403	25721	基因组SSR	AAAAGGAAGCAACTCGGTGA	GCAATTTTCTCACTCCATCTCC	60	168	V		V
404	16452	基因组SSR	CGATGGTTGCTGTTGTGAGA	ACCCCAAACAAACACCAATG	61	129	V		V
405	24388	基因组SSR	GGTCTGGGTCTTTGGCCTAT	AGCATTGCAACGAAGGTTTC	60	126	V		V
406	23759	基因组SSR	GGGGTGACAGTGTAGGGTTTT	TAGGCACACGCTTTCATGTT	60	157	V	I-2	V
407	23282	基因组SSR	TGGTGATGCATGATCATTTAGA	TACAACCCCACCCTGATTGT	59	152	V		V
408	25755	基因组SSR	TTTTCCAACTAAGGTTGTTTCTTTC	CAAAAGGAGGAGGCTGAAGA	59	150	V		V
409	28889	基因组SSR	ATTTGTGGTGCAAGCCTTCT	AAAATTGTACATGGACTCCTTTCTC			V		V
410	27456	基因组SSR	GCACATCCCATTTTTCCAGT	TTCATTACTTTGATAGTGTTCACAAA	57	120	V		V
411	27661	基因组SSR	ATTCATCTCTTTTCCTAAACAAAAAT	CACACCTCCACGTTCATCTC	57	152	V		V
412	29062	基因组SSR	TCACCATCGTGAGCAAGTTC	GGATGTTACGCCCACAATG			V		V
413	29741	基因组SSR	GCAAAAAGCATTGTCCATTTC	GCCTAATCTACAAACGGCTGAG			V		V
414	PSGAPA1	锚定标记	GACATTGTTGCCAATAACTGG	GGTTCTGTTCTCAATACAAG	51		V		V
415	16914	基因组SSR	AACCTCGAGCAACAACAGGT	TTAGGTTGGCGTTTTTGGTC	60	149	V	I-2	V
416	e691	EST-SSR	GATTTAATGCGCGGTTGATG	TGAGTGAAATCATGGGTGGA			V	I-2
417	28482	基因组SSR	CCGACACACTCCTCAACAAA	TCATCAGGATGAGGACACTCC	60	155	V	I-2
418	29694	基因组SSR	GAGTGCCTGATCCAAGAGGA	CTCTAAAGGGTGGCAACGAC			V	I-2
419	28277	基因组SSR	AACACAAGCGCGTTAGTTGA	GACCAGAGTCGAAGCGAAAC	60	183	V	I-2
420	29430	基因组SSR	TGCGATTTTTCAGTGAGGTG	AACGCAGGTGATGAGCCTAT			V	I-2
421	27008	基因组SSR	CGAGCAACAGACTGCAAAAA	GCCAACTTTCAATGTTTGACATA	59	132	V	I-2
422	16549	基因组SSR	CAATGAGATGCTGGCGATAA	GTTCGGTGTTGTGGGTTTTT	60	140	V	I-2
423	e506	EST-SSR	CCCCTTTATCCCCCTATTTC	CCTCAACACCAATGAACCAC			V	I-2
424	e938	EST-SSR	CTCCTCCTCTGATCCCTTCA	AAATTTCGATCAGGGGTTCC			V	I-2
425	e625	EST-SSR	GCTCCAATGGCTTCCTAACA	AACAAGGGGCAATCACAATC			V	I-2
426	e800	EST-SSR	AATCGCCAAAGGGTTTGTTT	CGCTTTGGTTCTAGCAGGAT			V	I-2
427	e1095	EST-SSR	TATCCATTGCCAGCAGCATA	AATCGCCAAAGGGTTTGTTT			V	I-2
428	157	基因组SSR	CACATCGACAGAGACATACGA	GTGTGTGTGATGTGTGTGGTG	57	70	V	I-2
429	21491	基因组SSR	ACACGGGATCGAGCTTTAGA	TCCTTTCCTCTAACTTCTTCCTTCT	59	175	V	I-2
430	3644	基因组SSR	CACACGCAGAATCACACGTA	CGTGTGTGTGCATGTAATCG	59	176	V	I-2
431	3790	基因组SSR	GCGTGTGTACATGTGTGTGC	GCGACTTGCACAAGCAGA	60	182	V	I-2
432	4696	基因组SSR	TGCATGTGCATGTAATCGTG	TCACACGCACGTACAAATCA	60	213	V	I-2
433	5849	基因组SSR	ACATTCACACACACACGCAA	AAGCTGTGTGCACGTGAGTT	60	278	V	I-2
434	3919	基因组SSR	GCACACGCAAACTCACAAGT	TGCGTGTGTGCATTTGTTTA	60	186	V	I-2
435	919	基因组SSR	GCACACGCAAACTCACACTT	GCGCATGTGCATTCGTGT	60	94	V	I-2
436	4190	基因组SSR	ACACGCATGCACGATTACAT	TGTCCGTGTACGAGCTTTTG	60	194	V	I-2
437	4013	基因组SSR	ACACGCATGCACGATTACAT	GTACGAGCTTTTGTCACGCA	60	189	V	I-2
438	3772	基因组SSR	CATGCACGATTACATGCACA	CGTAGCTTTGTCACGCATGT	60	181	V	I-2
439	4156	基因组SSR	ACACGCATGCACGATTACAT	GTGTCGTGTACGAGCTTTGC	60	193	V	I-2
440	3835	基因组SSR	ACACGCATGCACGATTACAT	GCTTTGTCACGCGATGTGTA	60	183	V	I-2
441	4427	基因组SSR	ACACGCATGCACGATTACAT	CGTACGTGTCCGTGTACGAG	60	202	V	I-2
442	3955	基因组SSR	ACACGCATGCACGATTACAT	ACGTAGCTTTTGTCACGCAT	58	187	V	I-2
443	4014	基因组SSR	ACACGCATGCACGATTACAT	GTGTACGAGCTTTGTCACGC	60	189	V	I-2
444	3567	基因组SSR	TGCATATGTGTGTGTCTGCG	ATACACATGCGTGCAAAAGC	60	173	V	I-2
445	4274	基因组SSR	CACACAGAAGCACACGCC	GCATGTGTGTGTGCGATGTA	60	197	V	I-2
446	4043	基因组SSR	ACACGCATGCACGATTACAT	CGTGTACGTAGCTTTGCACG	60	190	V	I-2
447	3695	基因组SSR	ACACGCATGCACGATTACAT	TTGTCACGCATGTGTATGTGTT	60	178	V	I-2
448	3771	基因组SSR	ACACGCATGCACGATTACAT	GCTTTGTCACGCATGTGTATG	60	181	V	I-2
449	1683	基因组SSR	GTGCGCATGGTGCATATAAA	CATATACACACACGCACGCA	60	113	V	I-2
450	3807	基因组SSR	ACACGCATGCACGATTACAT	GCTTTGCACGCATGTGTAGT	60	182	V	I-2
451	4114	基因组SSR	ATACACGCATGGCACGATTA	GTACGAGCTTTTGTCACGCA	60	192	V	I-2
452	3600	基因组SSR	CACATGCACACGCACACTTA	TGTCCGTGTACGAGCTTTTG	60	174	V	I-2
453	4080	基因组SSR	ACACGCATGCACGATTACAT	TGTACGAGCTTTTGTCACGC	60	191	V	I-2
454	3770	基因组SSR	ACACGCATGCACGATTACAT	AGCTTTGCACGCATGTGTAT	59	181	V	I-2
455	3808	基因组SSR	ACACGCATGCACGATTACAT	GCTTTTGCACGCATGTGTAT	60	182	V	I-2
456	3921	基因组SSR	ACACGCATGCACGATTACAT	TACGAGCTTTTGTCACGCAC	60	186	V	I-2
457	3377	基因组SSR	CACATGCACACGCACACTTA	CGTAGCTTTGTCACGCATGT	60	165	V	I-2
458	3597	基因组SSR	CGGGTTCACGTATGTGTGTT	ACGCGTATATTCACACGCAC	59	174	V	I-2
459	3923	基因组SSR	TACACGCATGCACGATTACA	CGTAGCTTTGTCACGCATGT	60	186	V	I-2
460	4825	基因组SSR	GCATGTGCATTTAATCGTGC	TGCACACGTACACACAAATCA	60	220	V	I-2
461	3298	基因组SSR	TATGCGTGTGTGTGCTTGTG	CACACATACACGTGTGAACCC	60	162	V	I-2
462	3181	基因组SSR	CACATACACATGCGTGCAAA	CGTGTGGTCATGTACGTGTG	59	158	V	I-2
463	24331	基因组SSR	AATGGCGCACTTCACTTTCT	CCGTTAACGCCTAGCTCAAG	60	137	V	I-2
464	25387	基因组SSR	GGCTCATGCATCTACCACCT	ATCCCGACGTTCACATTTTC	60	170	V	I-2
465	24186	基因组SSR	GGTGGATCCTCCTTTTGTCA	TCCCAATCACCACTTCTTCA	59	167	V	I-2
466	19075	基因组SSR	CACGAGTACAACATGGAGTGAAG	CAAGCTCAACCTCCTCATACC	59	187	V	I-2
467	e531	EST-SSR	CACCTCCACCCTTTCACCT	CTGGAGGTGGGAGATTGTCT			V	I-2
468	28857	基因组SSR	CCGAAATGTTCCGAAGAGAG	TTTCAATTCAATGCCGAAAT			V	I-2
469	27046	基因组SSR	AAAGAAGGGGATGCGAGAAG	GCTCAAGTCAGTCGGACCAC	61	145	V	I-2
470	30087	基因组SSR	CGCATACACTGAGGTAACACC	AATACACCGGAAGAGGACCA			V	I-2
471	o65	基因组SSR	ACCGCAACAACAGGATAAT	TGAGGTGAAATCGGAAGAC	53	143	V	I-2
472	e1035	EST-SSR	TTTTGCACCCCCTTATGTCT	CCACAAAACTCGGGTGAAAT			V	I-2
473	19252	基因组SSR	CAATATTGATCGGAATTTGTTTC	TGCGGTTTGATTGAGTTTGA	58	199	V	I-2
474	29379	基因组SSR	AGGCACGTTGGTGCTAGACT	CCTCAATGATCCCAAAGCAC			V	I-2
475	e178	EST-SSR	CGAAGAAGATCAAAATCACCAC	AGCTTCAGGGGTTTCTTTCC	59	195	V	I-2
476	e135	EST-SSR	GTTCGTTCGTTCGTTCCTTG	TCTGTCTGGAAATGAAATGG	56	219	V	I-2
477	AD147	锚定标记	AGCCCAAGTTTCTTCTGAATCC	AAATTCGCAGAGCGTTTGTTAC	61	330	V	I-2	I
478	e921	EST-SSR	AAGGGGTGATCAAGCATCAA	TTGAGGGAACATGAAGAAATCA			V	I-2
479	e915	EST-SSR	GTGGACTCGGATTGGGACTA	GCATCGACGACGAAGAAGAC			V	I-2
480	111	基因组SSR	CGCACAGCAACACACACAT	GCAGTTAGTGCGTGCGTG	60	68	V	I-2
481	22913	基因组SSR	TCCAACAAACTCAGCCACAG	CAATGGTGGTGGTGCTCATA	60	175	V	I-2
482	23383	基因组SSR	TGGAGAAATTGGTGGTGACA	TGCAACCATGTTCTTGTTCC	60	141	VI	VI
483	23611	基因组SSR	TGCAAATGTGCAATGAATGA	GGCGGACATGAGAAGGAATA	60	187	VI	VI
484	25953	基因组SSR	GGCCACAACCGTGATGAG	GGATCCAAGACCGAGACAAC	60	117	VI	VI
485	24326	基因组SSR	AAAACGAGAGGCTCGAAACA	ACTAAAACCTCGCGCATCAC	60	185	VI	VI	VI
486	17422	基因组SSR	ACCACAAATGCTTCCGCTTA	GTTGTTGTTGCTGCTGCTGT	60	200	VI	VI
487	22197	基因组SSR	GTTGTTGTTGCTGCTGCTGT	ACCACAAATGCTTCCGCTTA	60	200	VI	VI
488	29622	基因组SSR	AACTTCTGCAGTGGCATGTG	CAAAACAACCTATAAGGATGGAAAA			VI	VI
489	29331	基因组SSR	GGGTGGACCGAATATTTCAA	CGTCACCTCTACCGAAGCTC			VI	VI
490	e124	EST-SSR	GCTTCTGAACCAAGCACACA	AACAATCCCATGTATCAGCAAC	59	235	VI	VI
491	e1097	EST-SSR	CCCTTCTCATGGGGAATGAT	TAGTCCATGGAAGCGGAAAA			VI	VI
492	e1105	EST-SSR	TAGTCCATGGAAGCGGAAAA	CCCTTCTCATGGGGAATGAT			VI	VI
493	27153	基因组SSR	GGGAGCGATGCACATAGTATT	GCCCTACAACGAGTGACACA	59	134	VI	VI
494	26929	基因组SSR	CACATTCACGACGAGGACAG	GCACACTGTAAGCACTTTTCTCA	60	142	VI	VI
495	e853	EST-SSR	CTTCCCGGGTAAGAACAACA	GCTATGGTTCAGGCGTTTCT			VI	VI
496	23637	基因组SSR	AAGAGGCTCGTGACCCAATA	TGCATTGCATCCTTCAAGAG	60	192	VI	VI
497	AD160	锚定标记	ACCAGTCAAATGGTTAGAAAGT	GAATGGAAAAGAGAATCAAGTT	51	190	VI	VI	VI
498	23578	基因组SSR	AAGGAAGGTGGTGTGGAATG	CAATATTACTCAGCCATTAATTAACCT	58	205	VI		VI
499	PSGSR1	锚定标记	TGAAACCACCATTCTCTGGA	AAGACCCCACTTGAAAATTACTTC	58		VI		VI
500	25075	基因组SSR	CATCACTCACTCGCCAAAAA	AACCATCTTTGCCAGGTACG	60	192	VI	VI	VI
501	e709	EST-SSR	TGTGCAACCGAGATTGGTAA	CGCCAAAAATACTGATTCACTTC			VI	VI
502	27176	基因组SSR	TGCCTTCAGGTTTTCAAGGT	TGATGAAAGCAATTTTCATGACTT	60	147	VI	VI
503	29246	基因组SSR	CCCTTGCTTGGGTAAGAAATC	GTGCCGGGTATGTATCTGGT			VI	VI	VI
504	4816	基因组SSR	CGTCATCATTGTTCGTCATTCT	GGTCGTAGGGTGTGTCGTCT	60	219	VI	VI
505	2089	基因组SSR	ACACACGTCACACACACGTC	TGATGTGTATGCGTGATGGA	59	124	VI	VI
506	28621	基因组SSR	CGTTTTCACATTCGCTAACC	TGGAGAAAGGTTTCCTGATGA	58	171	VI	VI
507	27428	基因组SSR	GCACGCCTGACTTCTTCTTT	AAATGGATTGCGACGTGATT	60	174	VI	VI
508	24560	基因组SSR	GATAAAGGCAGCGACAGAGG	AATGAAGTGCAAGCCCAAAT	60	202	VI	VI
509	23737	基因组SSR	CGTGCAACCATAGCAAGAGA	AAACCGCTCAAGCTCAGGTA	60	182	VI	VI
510	26025	基因组SSR	ATGCACTCAAAGGCCATCAT	CACTTGCAGAGCGAGAGAAA	59	113	VI	VI
511	24906	基因组SSR	TCGAGTCAATCGCTCAGAAC	TGCCCAGATGTCATAAGGTG	59	134	VI	VI
512	16445	基因组SSR	TCAAACCGCTGAAAAACAAA	GCGGTGGGAGGGAGATAC	59	128	VI	VI
513	16397	基因组SSR	AGGGCCAGGTTTATTTCCAC	TTTCCCAATGGCAAGTTAGC	60	123	VI	VI
514	e715	EST-SSR	CGTTGAAACAGCGATTCTGA	TTTCTTCAATACCTCAATGGTTC			VI	VI
515	28153	基因组SSR	TGGGTTGTCGTGTTGTTGTT	AACACTCCCAACTCCATTTTT	58	183	VI	VI
516	27491	基因组SSR	TCCTAACCAACCAATAACACGAT	TTGAGGATTTCGGTGACCTC	60	180	VI	VI
517	C58	基因组SSR	TCACGTGCTTGTCGTTCTTC	TAAGAAACCGCCATGGAT TT			VI	VI
518	B117	基因组SSR	ACATCAGGGAAGAACGCATC	GAGGGTGAAGACCAGCTTTG			VI	VI
519	30379	基因组SSR	TGTTGGCAGGAAACTCTTCA	AGCCACAAATTTCGTTGTGTT			VI	VI
520	24575	基因组SSR	TTGTGAGCACATTGGAGGAG	GGATTGTGTTGGTTAGAGAAAGAGA	60	161	VI	VI
521	23468	基因组SSR	CGGCAGCATCTACACAAGAG	ACGTTGAAGACTCCGTCACC	60	171	VI	VI
522	29440	基因组SSR	TGTTCCCCTTTAATTTTCATCCT	AAGAAGCCGTCACGAAATGT			VI	VI
523	30004	基因组SSR	TCTTTGCGGATATGCATTTTA	CGGGTGAGGACTGAAAACTC			VI	VI
524	2614	基因组SSR	ATGTGTGTGCGTGTGTGTTG	GATTGTTATGTGCTGCGTGG	60	140	VI	VI
525	3494	基因组SSR	GCACCGCTCTGACACTCATA	TGAGAGTGGAGTGGCTGAAG	59	170	VI	VI
526	3244	基因组SSR	CTTCCCCTCGCAATTTATGA	ATGTGTGTGCGTGTGTGTTG	60	160	VI	VI
527	4583	基因组SSR	ACACCATTGCACCATTCTGA	TGCGTGTGTTGTGAGTGAGA	60	208	VI	VI
528	4581	基因组SSR	ACACCATTGCACCATTCTGA	GTGCGTGTGTGTTGTGAGTG	60	208	VI	VI
529	4629	基因组SSR	ACACCATTGCACCATTCTGA	GTGCGTGTGTGTGTGAGTGA	60	210	VI	VI
530	4811	基因组SSR	ATGTGTGTGCGTGTGTGTTG	ACACCATTGCACCATTCTGA	60	219	VI	VI
531	e93	EST-SSR	ATGGCCTTTGCAATTACAGG	GCTGATGTTGGCCAAGGTAT	60	203	VI	VI
532	e771	EST-SSR	TCCGGCAAGATATTGGAAAA	CTGCAGAGGCTGTCACTCAA			VI	VI
533	e1109	EST-SSR	TCCGGCAAGATATTGGAAAA	GCTTGGATCGCAGGAAAATA			VI	VI
534	20896	基因组SSR	TGATGACCCTGCAAATTCAA	TGCACCACTGTCAGGTGATT	60	131	VI	VI
535	e596	EST-SSR	TCCCTCATTCTCCCTTTTCA	GACGGCGCTGATGATAGACT			VI	VI
536	e1098	EST-SSR	AGAGGACGTGTTGCTGTGTG	CACAGAATTGGCAGAAACAGAG			VI	VI
537	27272	基因组SSR	TGTAGCGGCACACTTTGAGA	GATCTCTGCCACCCATCTTC	60	190	VI	VI
538	28790	基因组SSR	GCTGTGGGGGTTTAATCAGA	CCGCAATCCTTCAAGAACTC	60	133	VI	VI
539	29200	基因组SSR	ATGCTGATGAAATGCGAATG	CATCTGTACCCGGACCTTTG			VI	VI
540	27844	基因组SSR	GCTTCAAGCTACCAAGTGGA	CCTCACGGGCTCTACCATAC	58	118	VI	VI
541	29877	基因组SSR	GTCGTGGGGAAAAGGTATCA	GGTACGACAACCCTACCTTTG			VI	VI
542	e975	EST-SSR	AGCAGCTCCTACTCCTTCTCC	GCGCAAATCCTATTCCAAAG			VI	VI
543	e940	EST-SSR	GCGCAAATCCTATTCCAAAG	ATTTCAGCAGCTCCTACTCCT			VI	VI
544	19691	基因组SSR	TTGTAAGACCGACTCGTCCA	CGGTCTGAGGTTGTTGTGAA	59	146	VI	VI
545	16588	基因组SSR	CGGTCTGAGGTTGTTGTGAA	TTGTAAGACCGACTCGTCCA	59	146	VI	VI
546	25488	基因组SSR	TGAAGAATGAGCTTCAATTTTTGT	GGGTGCAATCATGAGTGTTG	59	145	VI	VI
547	26625	基因组SSR	GCTCCATCACGGTGAGTTTT	TCCCACTTTCACGATGTTCA	60	191	VI	VI
548	25280	基因组SSR	CTCTCTGCCCACTGCTCTG	TCTCACGTTGGGATGCTAAA	59	122	VI	VI
549	25711	基因组SSR	AAGGTTTTGAAATAAATGAAGTTTG	TGAAAGCCCACTTGATCTTC	57	188	VI	VI
550	25888	基因组SSR	AAGGGGGAGAGAGGTGGTTA	TCGCCTTTTCTTTCTTCTTCA	59	164	VI	VI	VI
551	AA335	锚定标记	ACGCACACGCTTAGATAGAAAT	ATCCACCATAAGTTTTGGCATA	61	220	VI		VI
552	23568	基因组SSR	TCCCTCATTCTCCCTTTTCA	GACGGCGCTGATGATAGACT	60	127	VI	VI	VI
553	23081	基因组SSR	ACCCTTGCTTTGCCACATAA	TGTTGCTCTTTTGTTGAGTTGA	59	149	VI	VI
554	23431	基因组SSR	GCAACAACAGCAACCTCTGA	TGAAGGTGGAACTTGGTTTTG	60	145	VI	VI	VI
555	27435	基因组SSR	TTGATGCTCTTCTTCCATTCAA	CATACAAAACACACAAAAAGGATTG	60	171	VI		VI
556	16410	基因组SSR	AAGGTCATGCTTCTTCATCTCT	GGGTGAGGTGTTATGGCACT	57	125	VI		VI
557	28516	基因组SSR	CCAAAATTCATGCATGGTACG	TCCAGTGGCTCATAGAGGAGA	60	169	VI		VI
558	26140	基因组SSR	TTGTGTGCAAACCACCTAGC	GATTGCATCACACGGTCAAG	60	200	VI		VI
559	23309	基因组SSR	GAAGATGGCAACGTGTCAAA	AACTCATCCTCCACCACCAC	60	128	VI		VI
560	29872	基因组SSR	TCCACTTCCACCCACAAAAT	GCAATGGAGGTTTTGCTTTT			VI		VI
561	26514	基因组SSR	TGGGCACAAGTCTGTGAGAA	TTGGGTTGAGGTGTTTAGGTG	60	153	VI		VI
562	AD60	锚定标记	CTGAAGCACTTTTGACAACTAC	ATCATATAGCGACGAATACACC	51	216	VI		VI
563	AB71	锚定标记	CCAACCATTTGTGAGTTCCCTT	TTCGTCGAACCACGAGAATAGA	61	145	VI		VI
564	23949	基因组SSR	TCGGTCAATTTTCACGTAGC	GCAGAGAATGAAAGAAATATAAAGAAA	58	142	VI		VI
565	22903	基因组SSR	TGCTTGCCAGAATAAAAGTCC	CCTCTAGGGTTTCGGGTCTC	59	181	VI		VI
566	27055	基因组SSR	TCTGCAAACTCCCAACACTG	GGTGGTTTGGTTGCAATCTT	60	206	VII	VII
567	e282	EST-SSR	GGCAAGCATAAAAGGGACAC	TTCATCCAAGAACCCTCGAC	60	185	VII	VII
568	e144	EST-SSR	TCCATTTCCCGAGTTTATCG	AACACGAATATGCAACAAGC	56	151	VII	VII
569	S244	基因组SSR	TTTAGCACAGAACAGCGTAGT	TAACGCCCTTGAGAATTTCG	53	606	VII	VII
570	21399	基因组SSR	TGATTCTAGTTCATTTCACAAACACA	CGTCCTGCACCTAGCTTCTT	60	153	VII	VII	VII
571	26099	基因组SSR	GCTCTTCGTAACGCTCACAA	TGACGACGGAGACTGAGTTG	59	201	VII		VII
572	19979	基因组SSR	CACCAGAAAATTTGTTATCAAAAAGA	GCACCCTGGGAAATTACAAA	60	161	VII	VII	VII
573	24547	基因组SSR	CGGCAGAATTAGGGTTTTGA	TCAATTCCGAACCACCTTTC	60	198	VII	VII
574	e968	EST-SSR	ACCGCTTGAACTCCAAACAA	GAAGGTAACAACGCCGAGAA			VII	VII
575	18013	基因组SSR	TCAATTCCGAACCACCTTTC	CGGCAGAATTAGGGTTTTGA	60	198	VII	VII
576	S144	基因组SSR	TTTTCTCACCGCGCTTATTT	AACAACCACCGAAGACGAAG	54	235	VII	VII
577	20828	基因组SSR	CATGGATCCCAACAGAAACA	TGGTTTTTACCCGAGACTGG	59	144	VII	VII
578	e876	EST-SSR	GCAGATTGACTGCTCATGATGT	AGCTGATTATTGGGCACCTG			VII	VII
579	21405	基因组SSR	TCCACCATCTAATCCCCTCTT	AGCTGATTATTGGGCACCTG	60	147	VII	VII	VII
580	18103	基因组SSR	CATGTGCATGTGCAAGTACG	CTTTAAAATGCCCGGACAAA	60	209	VII		VII
581	17431	基因组SSR	TTCACAATTCACCACCAATCA	CCAACGTCAGGTACGATTCA	60	201	VII	VII	VII
582	17666	基因组SSR	GTGCATTGGCTCGTACTCAA	TCCACAATATAGCCCAGACCA	60	158	VII	VII	VII
583	S85	基因组SSR	TTCCAACCATGGAAGCTTTT	TTCTTCGTCGGGTACAGTGA	54	188	VII	VII
584	e255	EST-SSR	TGGAGAAGGGCAAAATATCG	TCTCAACACCACATCAAGGAA	59	249	VII	VII
585	25965	基因组SSR	TGATTCGTAGACCCCACACA	AAGGTTAATGTCTTCTTTTTGAAGTT	57	150	VII	VII	VII
586	17384	基因组SSR	AGTAGCGGTGTGTGGTTGTG	GGGAAGAAAAAGGTTGGAAGA	60	197	VII	VII	VII
587	30253	基因组SSR	CTACGTTTGGCCCTTGTGTT	GGCCCTAAATCTAAAATGAACAA			VII	VII
588	B179	基因组SSR	ACCGACGCTTCAATGGTATT	TTCATCTCCGACCCTACACC	55	267	VII	VII
589	e499	EST-SSR	GACTCCACGCACAGAACTGA	GGAGAGGGGAGTGAATGTGA			VII	VII
590	e544	EST-SSR	CACTACACAAGAAGCAAAGAAAAA	ATTCCTTTCCGGTCCATTTC			VII	VII
591	e1002	EST-SSR	CCACGCGTGACAAGTAAAGA	ATTCCTTTCCGGTCCATTTC			VII	VII
592	21420	基因组SSR	GGTGGTCGACGTATCGAAGT	TCAATGTTGTTGCGCTTACAT	59	142	VII	VII
593	18255	基因组SSR	TGTCAAATCCAATAAAAACACACA	TTTGTGCACACCGTCAATTT	59	129	VII	VII
594	e643	EST-SSR	CACCACCACTCTCACACCAT	TGCATTGCGAGAGTAAGACAA			VII	VII
595	e825	EST-SSR	AGTTTCCGCCATCAACATTT	CCTACCTGCATTCACAACCAT			VII	VII
596	17225	基因组SSR	GTTGCAAGCTGCTACCATCA	AGACGGATCCAACAATCTCC	59	182	VII	VII
597	17593	基因组SSR	CATCCTCCTCCTCCATACCA	TCATCATCAATGCAAAGGACA	60	148	VII	VII
598	24407	基因组SSR	GGGATCAAAGCAACCCTTTT	CATGGCAAGGAAGACCGTAG	60	208	VII	VII
599	24810	基因组SSR	TGAGCGAGGTAGGAAGAACC	CCTCTACAGTGGCCCTCTCA	59	132	VII	VII
600	25354	基因组SSR	ACCCTTGGGGCTTACAATTC	GACGTGGCTGGACATAACAA	60	184	VII	VII
601	25799	基因组SSR	CTGCAGAAGGCCCTGTTCTA	GATTCTTCATTCTCAACACACATTG	60	137	VII	VII
602	25430	基因组SSR	GCACTTGACGATGCATTTGA	GGGAAAGGGAACGATTTCAT	60	210	VII	VII
603	23294	基因组SSR	GGAGGAGGATGACGATGAAA	AGGGCTACCGGACTGAAACT	60	172	VII	VII
604	24142	基因组SSR	GCAGCCATGGTTGATTGATT	TCAAGAACATTACTTTTTCCCTCT	58	194	VII	VII
605	27051	基因组SSR	TGGTGGTGGAGAGTGATTGA	TCTGGTGGTACTTCCTCCAAAT	60	208	VII	VII
606	e1114	EST-SSR	GTGCGGCTTCATTTTCAACT	TTCTCAACTGGTTGGTTCCATA			VII	VII
607	e1209	EST-SSR	AGTGCGGCTTCATTTTCAAC	TCAATTTGATCCATGCAGTAGG			VII	VII
608	AF004843	锚定标记	CCATTTCTGGTTATGAAACCG	CTGTTCCTCATTTTCAGTGGG	54		VII	VII
609	23606	基因组SSR	GGGTTTGCCTCTTTTTCTCTC	ATCGTCAAAACTGCCCAAAC	59	113	VII	VII
610	21345	基因组SSR	TGCAATGCATGTTGATACGTC	GCAAAAACTCAAACTCAAACTCAA	60	152	VII	VII
611	27583	基因组SSR	TGCACAGAGGATGGTTCTCA	TGGATTGAGCCTCTTGTCCT	60	110	VII	VII
612	25610	基因组SSR	TTTGGTCGTTGCCAAACTAA	AGGACATACCGAGCCAGATG	59	133	VII	VII
613	26656	基因组SSR	AGAAGAGCGTCGGGAAGAGT	CCATGACGGAAAACAACCTT	60	188	VII	VII
614	25728	基因组SSR	TGGACAAATCTCGTGCAAAG	CCAACTTCCCATCTCCAACA	60	164	VII	VII
615	24471	基因组SSR	CAACACAACCTCCTCCAGGT	TAAGCCATTCCCACCTTCTG	60	121	VII	VII
616	24734	基因组SSR	CGCATGAGAGGATAATGATGAA	CCATAGCTTTCCCGAATCAG	60	183	VII	VII
617	24588	基因组SSR	AAATAGATGAGAAAGAGAGAGATTACG	CGCACTTCCATTCACATGAT	57	137	VII	VII
618	24602	基因组SSR	TGAGTGGGCGTGTGATTTAG	TTGCACTGTCGCATTTGAGT	60	118	VII	VII
619	24503	基因组SSR	AAAGGAGATCACCTATGAGAGAGAA	AATTGATTTGGGATCTTGGATA	57	154	VII	VII	VII
620	28097	基因组SSR	CAAAATCGGCAGGATTTACAA	TTTGCCTATGAACCTAAAACCAT	59	124	VII		VII
621	24814	基因组SSR	GTGCAACCAGAGCATGTTTC	CATCGACTGTGGAGACATTGA	59	152	VII		VII
622	AB65	锚定标记	CTCGTATCCAAAGATTCGTAGA	AGGGTTAATCGGAGTTTTATGA	51		VII		VII
623	29468	基因组SSR	AGTACTCTCGCCGCATCAGT	CACCCCACTTGAGCATATCA			VII	VII	VII
624	28967	基因组SSR	CTCGTCCTCATCGAAAAGCTA	GTGAACAACGCAAGGGTTTC			VII	VII
625	29500	基因组SSR	TGGGAATTGACGAAGAGTGTT	ATGGAGAGGGTTGCTGACAT			VII	VII
626	e46	EST-SSR	AGGAGGGAGTGGTGGAGATT	CATCACGTGCTTGTGCTTCT	60	227	VII	VII
627	e1173	EST-SSR	TGATTCCGAATGGGAAACTT	AATCCGCAAACACATCAACA			VII	VII
628	e1276	EST-SSR	TGAAACAATAGTGCTTTGTTGAAACT	TTTTCTCGTCTGCGTGTGAC			VII	VII
629	22060	基因组SSR	CCGCCTTAGGAAGCCTAACT	CGGCTTGATAATTTGGTGCT	60	181	VII	VII
630	e628	EST-SSR	GCCACCCTGTTTCTGCTAAC	CTTGTGGATCGGTTCGTGAT			VII	VII
631	e524	EST-SSR	TCACACCATAGAGAATAACAACAACA	TCTGAAGCCATCTCCATTCTC			VII	VII
632	28608	基因组SSR	TCATGATTTCAAATTTCTTTCACAA	CGCCGTTGGGTAATTGTAAC	60	136	VII	VII
633	22101	基因组SSR	ACCAATCCAGACGCAAATTC	GGGGACAGTGACGAGAACAT	60	156	VII	VII
634	19460	基因组SSR	CAATCCAAACGCAAATCTAACA	AATTGCAAGCCCTACACACA	59	187	VII	VII
635	17910	基因组SSR	CATGCCTGCTTCCTTCTGTT	TTGCAATTTCAAGCCTTCAC	59	187	VII	VII	VII
636	27398	基因组SSR	GAACCACATTGGGGATTCTG	AACCTGCAAAAGCCATAAGC	59	180	VII	VII
637	30165	基因组SSR	CCTTTTTACCCCTCCCTCAG	TCAAATGCAAAGGGAAACAA			VII	VII
638	27755	基因组SSR	TGCATTTGAGCTAGTGACATTCT	AAAAACCAACCCAACCACTT	58	163	VII	VII
639	e1193	EST-SSR	GTTGGCGAGGAATGATTGTT	TCACACACTCTGCCATTTCA			VII	VII
640	PSAB60	基因组SSR	AATTAATGCCAATCCTAAGGTATT	GGTTGCACTATTTTCGTTCTC	59		VII	VII
641	28054	基因组SSR	AGCAAAGTCACCAGCTGTTTT	TCCTGTTCTAAAACAAAAACAAGAGA	59	113	VII	VII
642	30124	基因组SSR	GACTATGTGTGTGCATGAATTTGA	GGCCATCTCGTTTCAGTACC			VII	VII
643	30304	基因组SSR	CACGCACAGACAGATCATCA	GAAGTTGGGGATGGGAAGAT			VII	VII
644	28108	基因组SSR	CGACAATGTTGCCAGCTATC	TTTTAGGATTTTATCGACGTTTTTC	59	119	VII	VII
645	e291	EST-SSR	CAAGCGTCGAAGATGAACAA	GCTGGCTGCAAAAGTTTACC	60	205	VII	VII
646	e292	EST-SSR	GCACATGAAAAATGCCAAAG	CTGTTGCTGTTGGTGGTGAG	59	220	VII	VII
647	PS11824	基因组SSR	ACCACCACCACCGAGAAGAT	TTTGTGGCAATGGAGAAACA	52	210	VII	VII
648	e527	EST-SSR	TTGAAGCAGTGGCAGAGTTG	TCTCAATGAAACATAAGAATGACCTT			VII	VII
649	e1202	EST-SSR	TGCATGGTTATGATGCTTGA	TCACACACCCCTTCAATTATTC			VII	VII
650	23521	基因组SSR	CGCCAATTCCTTTTCCCTAC	AGAACTCACAGGCGATGGTC	60	112	VII	VII
651	24652	基因组SSR	GAGAAAGCGGCTGCTTAGAA	GCTGTCACCGAGAATGATGA	60	190	VII	VII
652	25241	基因组SSR	TGCAAGCAAGCAAAATTGAA	TGCATGCCCTTTATTTCTTTG	60	142	VII	VII
653	24944	基因组SSR	TCAACCGGAATCTGGAAAAC	ATGGCGCAATCCTAGTGAAC	60	193	VII	VII
654	22896	基因组SSR	ATGTACGCCATGCAGTCAAA	ACAAGATGGGCGTCGAATAC	60	151	VII	VII
655	24398	基因组SSR	TTCGATGCATGAATGACAAA	ATCGGCGGAGACTAAGATCA	59	142	VII	VII
656	20817	基因组SSR	GGCGTAGAGGGCTAAACCTT	TTCCCGAGTCCTAACTTTCTTG	60	135	VII	VII
657	19526	基因组SSR	ACTCCTGGACACCCTGAGAA	CTGACCAAGGGGACCTGTAA	60	198	VII	VII
658	29620	基因组SSR	CCACTGAAGGCTCCTGAACT	AGCGATCACCGATAGTGTCC			VII	VII	VII
659	e1220	EST-SSR	ATGGTGGTGGAGGTGTGATT	CATCGCCAAATGGATCTTCT			VII	VII
660	17980	基因组SSR	CAATTCACAACGTTCCACTCA	TTTTCGTGAAATTGAAATGACC	59	194	VII	VII	VII
661	29248	基因组SSR	CAACAATGTGCATGGAAAAA	CTCGCATTGCGTAACGATAA			VII	VII	VII
662	28595	基因组SSR	CCGGTTCATCGATAAATGTGA	TCTCAAACCCACCAACAACA	60	199	VII	VII
663	2629	基因组SSR	CAAAACACATACGCACACACA	CCCGTCATGATGTCATGTAAA	59	141	VII	VII
664	e123	EST-SSR	GAGCACAACTTTGCAAGCAG	ACACGTCATTTCAAACCACT	55	198	VII	VII
665	e1249	EST-SSR	CCCGTTTCAAATTAGAACGATAA	CACGGTTCGGCATTTATTTC			VII	VII
666	23536	基因组SSR	GACGTTGATTGGCCTGTTTT	TGACCATGACATGCCTGTTT	60	180	VII	VII
667	21796	基因组SSR	TCTTCGCTGGGAAGTTGAGT	GGAAGCGATGTCGTTTCATT	60	210	VII	VII	VII
668	28261	基因组SSR	CTCTCCCCATGGAGAACAAC	AACAGCTGAAATTGGCGTAGA	60	131	VII		VII

粗体代表锚定标记,下划线代表共有标记,粗体加下划线代表共有锚定标记。标记名称以“e”开头的为EST-SSR标记。
The bold, underlined and bold underlined labels represent anchor markers, common markers and common anchor markers, respectively. Marker names that start with “e” represent EST-SSR markers.

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将我们的整合图谱与以往的遗传图谱相比较, 结果发现, 本研究中的7条连锁群能够与以往遗传连锁图谱中7条连锁群相对应^[10,42-44]。通过比较整合遗传图谱和2个单独的遗传图谱之间共有标记的顺序和位置, 结果发现, LGIV和LGVII这2个连锁群在不同图谱中的标记顺序存在较高的共线性, 而其他连锁群则由于作图群体差异而观察到了一些倒位和标记重排的现象。此外, 在整合图谱的LGV上, 同时包含2个锚定标记, 一个是在以往图谱中定位于LGV的PSGAPA1, 另一个是在以往图谱中定位于LGI的锚定标记AD147, 因而导致LGV的定义存疑(附图1)。

附图1

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附图1整合图谱和2个单独图谱及其与豌豆参考基因组物理图谱之间共有标记比较(A–G)

图谱左侧数字为标记位置信息，而右侧为标记名称。遗传图谱包括PSP1、PSP2和整合图谱的图距单位为cM，参考基因组(Reference genome)的物理图谱图距单位为MB。图谱标记名称红色代表锚定标记，标记名称粗体加下划线代表共有标记。标记名称以“e”开头的为EST-SSR标记。
Supplementary Fig. 1Comparison of the common markers among the integrated map and the two individual maps as well as the physical map of the reference genome (A–G)

Numbers on the left of the map and labels on the right of the map represent the position and name of markers, respectively. The map distance unit of genetic maps including PSP1, PSP2 and integrated map is cM, while the map distance unit of physical map of the reference genome is MB. The red and underline labels represent anchor markers and common markers, respectively. Marker names that start with “e” represent EST-SSR markers.

本研究以新近发表的豌豆基因组为参考(Caméor genome build 1a)^[20], 对50个共有标记的扩增片段序列进行了BLAST比对分析, 从而对本文得到的遗传图谱和豌豆的物理图谱相比较(附表2)。发现有45个标记被成功比对到豌豆的7条染色体上, 上述有争议的1条连锁群(LGI-2/LGV)被证明为LGV。其中30个标记具有唯一比对位置, 15个标记比对到2个以上的位置; 剩下的5个标记比对到了未挂载到染色体的组装支架上。此外, 还比较了3个遗传图谱和物理图谱的标记顺序(附图1), 发现chr2LG1与LGI-PSP2高度一致, chr6LG2与LGII-PSP1高度一致, chr5LG3与LGIII-Integrated map基本一致, chr4LG4与LGIV- PSP1、LGIV-PSP2和LGIV-Integrated map基本一致, chr3LG5与LGV-PSP2和LGV-Integrated map基本一致, chr1LG6与LGVI-PSP2和LGVI-Integrated map基本一致, chr7LG7与LGIV-PSP1、LGIV-PSP2和LGIV-Integrated map一致率较高。

Supplementary table 2
附表2
附表2豌豆遗传连锁图谱上共有标记BLAST比对结果
Supplementary table 2BLAST results of common markers on genetic linkage maps of pea

标记名称Marker name	连锁群Linkage group			染色体/连锁群 Chromosome/LG	比对条数 Number of blast results	最小E-value The minimal of E-value	最高比对率 The maximal identifies (%)	起始位置 Start position	终止为止 End position	比对长度Mapped length
标记名称Marker name	整合图谱 Integrated map	PSP1 G0003973 × G0005527	PSP2 W6-22600×W6-15174)	染色体/连锁群 Chromosome/LG	比对条数 Number of blast results	最小E-value The minimal of E-value	最高比对率 The maximal identifies (%)	起始位置 Start position	终止为止 End position	比对长度Mapped length
20229	I	I	I	chr2LG1	20	6.7E-94	100	21,564,671	21,564,851	180
25334	I	I	I	chr2LG1	4	2.7E-70	100	329,941,758	329,941,897	139
23261	I	I	I	chr2LG1	1	1.7E-79	100	410,201,631	410,201,786	155
24301	II	II	II	chr6LG2	1	3.9E-58	96	44,747,008	44,747,143	135
28257	II	II	II	chr6LG2	1	3.5E-55	93	89,359,789	89,359,941	152
17754	II	II	II	chr6LG2	1	3.0E-85	99	284,992,981	284,993,149	168
25769	II	II	II	chr6LG2	17	3.9E-58	95	379,342,213	379,342,360	147
25792	II	II	II	chr6LG2	3	6.5E-121	93	430,274,319	430,274,608	289
26850	III	III	III	chr5LG3	1	4.5E-80	99	95,945,216	95,945,375	159
21726	III	III	III	chr5LG3	2	4.4E-80	100	105,392,839	105,392,995	156
25151	III	III	III	chr5LG3	31	4.4E-80	100	128,426,677	128,426,833	156
18135	III	III	III	chr5LG3	1	8.3E-59	100	157,545,697	157,545,816	119
22754	III	III	III	chr5LG3	3	2.2E-82	100	178,517,012	178,517,172	160
20075	III	III	III	chr5LG3	1	6.6E-45	95	255,763,443	255,763,559	116
29630	IV	IV	IV	chr4LG4	1	2.3E-97	100	77,755,735	77,755,921	186
28304	IV	IV	IV	chr4LG4	4	5.8E-64	99	77,763,197	77,763,328	131
27275	IV	IV	IV	chr4LG4	5	1.1E-58	94	144,374,211	144,374,362	151
24036	IV	IV	IV	chr4LG4	1	1.5E-102	100	298,729,278	298,729,473	195
19487	IV	IV	IV	chr4LG4	2	3.6E-92	100	329,433,230	329,433,407	177
21622	IV	IV	IV	chr4LG4	1	1.0E-69	100	411,138,053	411,138,191	138
24907	IV	IV	IV	chr4LG4	1	7.5E-67	100	441,578,306	441,578,439	133
23829	V	V	V	chr3LG5	4	2.8E-104	100	40,897,218	40,897,416	198
25618	V	V	V	chr3LG5	10	8.7E-101	99	91,875,171	91,875,372	201
25986	V	V	V	chr3LG5	1	9.0E-56	91	181,332,864	181,333,028	164
16914	V	V	V	chr3LG5	1	2.2E-45	89	191,073,596	191,073,747	151
22325	V	V	V	chr3LG5	1	5.4E-91	98	221,771,656	221,771,845	189
23759	V	V	V	chr3LG5	1	4.4E-80	100	276,140,552	276,140,708	156
24326	VI	VI	VI	chr1LG6	1	3.3E-96	100	30,662,379	30,662,563	184
25075	VI	VI	VI	chr1LG6	1	1.0E-92	98	46,237,147	46,237,342	195
29246	VI	VI	VI	chr1LG6	2	5.0E-72	100	63,898,496	63,898,638	142
23568	VI	VI	VI	chr1LG6	1	7.9E-63	100	339,601,745	339,601,871	126
25888	VI	VI	VI	chr1LG6	1	4.1E-84	100	342,366,936	342,367,099	163
23431	VI	VI	VI	chr1LG6	1	3.5E-73	100	359,154,849	359,154,993	144
21405	VII	VII	VII	chr7LG7	1	4.0E-47	90	23,151,033	23,151,166	133
17431	VII	VII	VII	chr7LG7	2	7.5E-94	98	28,469,955	28,470,153	198
19979	VII	VII	VII	chr7LG7	1	8.3E-71	97	33,247,383	33,247,539	156
17666	VII	VII	VII	chr7LG7	1	8.1E-71	97	33,247,466	33,247,619	153
25965	VII	VII	VII	chr7LG7	1	7.5E-49	89	79,270,351	79,270,512	161
17384	VII	VII	VII	chr7LG7	1	4.0E-103	100	86,051,350	86,051,546	196
29468	VII	VII	VII	chr7LG7	1	3.9E-58	95	196,186,432	196,186,576	144
24503	VII	VII	VII	chr7LG7	4	2.6E-52	91	201,461,481	201,461,642	161
17910	VII	VII	VII	chr7LG7	1	2.0E-68	97	329,043,834	329,043,990	156
17980	VII	VII	VII	chr7LG7	1	1.9E-94	98	474,504,396	474,504,591	195
29248	VII	VII	VII	chr7LG7	1	7.2E-75	99	481,640,880	481,641,037	157
21796	VII	VII	VII	chr7LG7	1	2.7E-97	97	488,671,955	488,672,158	203
18272	II	II	II	scaffold00024	1	9.2E-59	98	43,943	44,076	133
23262	III	III	III	scaffold03071	1	4.7E-95	100	151,557	151,739	182
28387	V	V	V	scaffold03509	2	2.0E-105	100	40,828	41,028	200
21399	VII	VII	VII	scaffold00916	1	8.8E-67	96	9,846	10,002	156
29620	VII	VII	VII	scaffold01087	12	1.1E-54	96	47,702	47,832	130

遗传图谱包括PSP1、PSP2和整合图谱的图距单位为cM,参考基因组(Reference genome)物理图谱图距单位为MB。图谱标记名称粗体加下划线代表共有标记。
The map distance unit of genetic maps including PSP1, PSP2 and Integrated map is cM, while the map distance unit of physical map of the reference genome is MB. The underline labels represent common markers. Marker names that start with “e” represent EST-SSR markers.

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3 讨论

3.1 SSR标记筛选

由于具有多态性高、多等位基因、共显性、可重复、可转移和基因组覆盖度高等特性, SSR标记包括基因组SSR和基于表达序列标签的EST-SSR已被广泛应用于种质资源鉴定评价、遗传变异检测、遗传进化分析、遗传连锁图谱构建、功能基因定位以及标记辅助育种等诸多领域^[21,48-50]。通过对基因组SSR和EST-SSR的应用范围进行比较发现, 基因组SSR因为多态性较高更适合于区分遗传关系比较近的基因型, 并且在指纹图谱构建和种质资源鉴定研究中更具优势; 而从基因或表达序列标签数据生成的EST-SSR标记则有利于获得更多基因的相关信息, 在重要农艺性状的标记辅助选择中具有更大优势, 同时在近缘物种之间具有更高的可转移性^[21,51]。以往的研究已经成功开发了一些SSR标记^{[10-12,38,44]}, 并用于遗传多样性评估^[1,33,52-55], 遗传连锁图谱构建^[10,15], 以及标记性状关联分析^[32,56]等。然而, 豌豆作为具有重要经济价值和显著生态优势的食用豆类作物之一, 目前具有明确定位信息同时可公开获取的基因组SSR和EST-SSR标记数量还相对有限, 这极大地阻碍了豌豆的分子遗传研究和标记辅助育种。在本研究中, 我们利用大规模筛选从本实验室开发的12,491个SSR标记中得到了729个多态性SSR标记, 同时从125个已发表的具有明确遗传图谱定位信息的标记中筛选了25个锚定标记, 分别用于2个基于中国豌豆种质F₂群体的遗传连锁图谱构建和整合。在最终得到的整合遗传图谱中, 上图标记达到668个, 包括509个基因组SSR、134个EST-SSR和25个锚定标记, 分布在7条连锁群上, 对应于豌豆的7条染色体, 相关标记信息详见附表1。这些已定位的SSR标记将为豌豆种质资源鉴定、遗传关系分析、分子遗传作图以及标记辅助选择提供宝贵资源。

3.2 遗传连锁图谱构建

高密度遗传连锁图谱是进行遗传研究和分子育种的有力工具。针对豌豆的遗传连锁作图已经有很长的研究历史, 前人利用不同的分子标记和不同的作图群体已经构建了许多豌豆遗传连锁图谱^[31]。随着二代测序技术的发展, 基因组SSR、EST-SSR和SNP标记的高通量开发为豌豆的分子作图奠定了重要基础^[14-18,57]。最近, 有****还利用SRAP、SSR和SNP这3种标记基于F₂群体构建了一张豌豆的遗传连锁图谱, 包含128个遗传标记, 分布在9个连锁群上, 然后他们利用9个SSR标记作为锚定标记, 将其中的6个连锁群与以往发表的遗传连锁图谱中的连锁群对应起来^[58]。尽管在豌豆中已经构建了超过50张的遗传连锁图谱, 然而目前国际上还没有基于SSR标记构建的标记数目达500个以上的图谱, 因此豌豆SSR遗传连锁图谱无论在标记数目和密度上均有待完善^[10,15]。此外, 前人的研究已经证明中国豌豆种质资源具有独特的遗传背景^[1,32-33], 然而基于中国种质的豌豆遗传连锁图谱很少。过去的一项研究基于中国种质构建的豌豆遗传连锁图谱总长1518 cM, 仅包含157个SSR标记, 标记间平均距离为9.7 cM, 而且分布在11条连锁群上, 与豌豆的单倍体染色体数并不一致(2n = 2x = 14)^[15]。在本研究中, 我们首先基于与以往研究^[15]相同的作图群体(PSP1), 筛选了大量多态性SSR标记, 对以往的图谱进行加密。加密后的图谱累计长度扩展到5330.6 cM, 标记数目增加到603个, 标记间平均距离缩小为8.9 cM, 所有标记分布在7个连锁群上(图1和表2), 其中6条与以往发表的豌豆遗传图谱一一对应。此外, 我们还利用一个新的大样本作图群体(PSP2)构建了一张新的豌豆遗传连锁图谱, 并通过17个锚定标记将总共118个标记定位在与以前发表的豌豆遗传图谱完全一致的7条连锁群上(图2和表2)^[10]。本研究基于中国豌豆种质构建的2个作图群体代表了与国外豌豆群体具有显著差异的基因组背景, 得到了2个SSR遗传连锁图谱, 与以往的研究相比^[15], PSP1图谱在标记数目和密度上具有明显提高, 而PSP2在连锁群装配方面则具有明显改善, 这些图谱将为中国豌豆的标记辅助育种提供有力工具。

3.3 整合遗传图谱

整合图谱凭借较高的标记数目和密度以及更完整的基因组覆盖范围等优势^[34,35], 在许多作物包括豌豆中均有应用。过去几十年, 在豌豆遗传连锁作图悠久的研究历史中, 前人已经通过整合来自多个作图群体的信息而获得了很多复合遗传连锁图谱^{[10,16,31,36-37,42,59-61]}。2015年, 法国科学家利用13.2K的基因芯片对12个豌豆RIL群体进行基因分型, 同时整合了以往图谱中的2277个标记, 构建了一个包含15,079个标记和7条连锁群的高密度一致性连锁图谱, 该图谱全长794.9 cM, 平均标记间距离0.24 cM, 为评估豌豆基因组结构、基因含量和进化历史以及候选基因鉴定提供了有力工具^[16]。然而上述一致性图谱包含的EST-SSR和基因组SSR标记仅有187个, 为了在豌豆遗传连锁图谱上积累最大数量的SSR标记, 本研究通过53个共有标记将2个F₂作图群体的数据合并构建了一个整合遗传连锁图谱, 覆盖范围为6592.6 cM, 包括668个SSR标记(509 基因组SSR、134 EST-SSR和25个锚定标记), 分布在7条连锁群上(图3和表3)。值得注意的是, 遗传连锁图谱的长度随标记数目的增多而延伸, 这种现象在包括豌豆在内的多个物种中均有发现, 有人推测可能与重组事件和偏分离有关^{[36-37,62-64]}。此外, 有****认为延伸的图谱长度对图谱上标记的映射顺序几乎没有影响^[61]。过去的研究发现, 由于多等位标记、数据缺失、偏分离和染色体重排等原因, 在比较不同杂交群体获得的遗传连锁图谱时, 会出现标记定位和顺序的不一致^{[10,36-37,65]}。在本研究中, 通过对2个单独的遗传连锁图谱之间53个共有标记的比较发现, 基于2个群体构建的遗传图谱均可组装到7条连锁群上, 与以往研究中的7个连锁群具有较好的对应关系, 然而PSP1图谱的LGI-2 (包含LGI锚定标记AD147)对应于PSP2图谱的LGV (包含LGV锚定标记PSGAPA1), 导致整合图谱中LGV的不确定性(附图1)。但是由于PSP1图谱中7个连锁群有6个与以往发表的图谱一一对应, 仅剩1个连锁群(LGI-2)因为缺乏额外的锚定标记无法与以往发表的LGV相对应; 同时PSP1图谱中的LGI-2与PSP2图谱中的LGV之间存在7个共有标记, 基于上述两点原因, 我们推测LGI-2应该对应于以往发表图谱中的LGV。然后通过对50个共有标记的扩增片段序列的BLAST比对分析, 对3个遗传图谱和物理图谱进行共线性比较, 结果支持有争议的一条连锁群确实对应于以往遗传图谱中的LGV (附表2), 而该连锁群上的锚定标记AD147可能是异位所致。另一方面, 单独图谱和整合图谱共线性比较发现, 不同图谱在LGIV和LGVII这2个连锁群上的标记顺序具有高度一致性, 而在其他连锁群上则观察到标记的颠倒和错位(附图1), 推测这种不一致性可能是由不同作图群体中发生的染色体重排引起的^[10,36-37]。不同遗传图谱与物理图谱的共线性比较得到了类似的结果(附图1和附表2), 物理图谱的chr4LG4和chr7LG7与3个遗传图谱在LGIV和LGVII这2个连锁群上的标记顺序具有较高的一致性, 说明这2条连锁群或染色体在不同群体的亲本间并未发生明显结构变异; 物理图谱中的chr2LG1、chr3LG5和chr1LG6与PSP2遗传图谱的LGI、LGV和LGVI基本一致, 而与PSP1遗传图谱的标记顺序全部或部分相反, 说明PSP1群体的亲本在这3条连锁群或染色体上可能发生了倒位; 物理图谱中的chr6LG2与PSP1遗传图谱LGII的标记顺序基本一致, 而与PSP2遗传图谱的标记顺序全部相反, 说明PSP2群体的亲本在这条连锁群或染色体上可能发生了倒位; 物理图谱中的chr5LG3与LGIII-Composite map的标记顺序一致, 而与PSP1和PSP2遗传图谱的标记顺序相反, 可能是参考基因组的豌豆基因型在该条染色体上发生了倒位或者错误组装。

4 结论

利用2个基于中国豌豆种质的F₂群体, 通过QTL IciMapping V4.0软件, 整合得到一张包含7条连锁群、668个SSR标记、总遗传距离为6592.6 cM、平均标记密度为10 cM的豌豆遗传连锁图谱。本研究将为豌豆的分子遗传研究和标记辅助育种提供有力工具。

附表和附图 请见网络版: 1) 本刊网站http://zwxb. chinacrops.org/; 2) 中国知网http://www.cnki.net/; 3) 万方数据http://c.wanfangdata.com.cn/Periodical-zuow xb.aspx。

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Plant J, 2015,84:1257-1273.

DOI:10.1111/tpj.13070 URLPMID:26590015 [本文引用: 4]

Single nucleotide polymorphism (SNP) arrays represent important genotyping tools for innovative strategies in both basic research and applied breeding. Pea is an important food, feed and sustainable crop with a large (about 4.45 Gbp) but not yet available genome sequence. In the present study, 12 pea recombinant inbred line populations were genotyped using the newly developed GenoPea 13.2K SNP Array. Individual and consensus genetic maps were built providing insights into the structure and organization of the pea genome. Largely collinear genetic maps of 3918-8503 SNPs were obtained from all mapping populations, and only two of these exhibited putative chromosomal rearrangement signatures. Similar distortion patterns in different populations were noted. A total of 12 802 transcript-derived SNP markers placed on a 15 079-marker high-density, high-resolution consensus map allowed the identification of ohnologue-rich regions within the pea genome and the localization of local duplicates. Dense syntenic networks with sequenced legume genomes were further established, paving the way for the identification of the molecular bases of important agronomic traits segregating in the mapping populations. The information gained on the structure and organization of the genome from this research will undoubtedly contribute to the understanding of the evolution of the pea genome and to its assembly. The GenoPea 13.2K SNP Array and individual and consensus genetic maps are valuable genomic tools for plant scientists to strengthen pea as a model for genetics and physiology and enhance breeding.

[17]

Boutet

, Carvalho

S A

, Falque

, Peterlongo

, Lhuillier

, Bouchez

, Lavaud

, Pilet-Nayel

M L

, Riviere

, Baranger

. SNP discovery and genetic mapping using genotyping by sequencing of whole genome genomic DNA from a pea RIL population
BMC Genomics, 2016,17:121.

DOI:10.1186/s12864-016-2447-2 URLPMID:26892170 [本文引用: 1]

BACKGROUND: Progress in genetics and breeding in pea still suffers from the limited availability of molecular resources. SNP markers that can be identified through affordable sequencing processes, without the need for prior genome reduction or a reference genome to assemble sequencing data would allow the discovery and genetic mapping of thousands of molecular markers. Such an approach could significantly speed up genetic studies and marker assisted breeding for non-model species. RESULTS: A total of 419,024 SNPs were discovered using HiSeq whole genome sequencing of four pea lines, followed by direct identification of SNP markers without assembly using the discoSnp tool. Subsequent filtering led to the identification of 131,850 highly designable SNPs, polymorphic between at least two of the four pea lines. A subset of 64,754 SNPs was called and genotyped by short read sequencing on a subpopulation of 48 RILs from the cross 'Baccara' x 'PI180693'. This data was used to construct a WGGBS-derived pea genetic map comprising 64,263 markers. This map is collinear with previous pea consensus maps and therefore with the Medicago truncatula genome. Sequencing of four additional pea lines showed that 33 % to 64 % of the mapped SNPs, depending on the pairs of lines considered, are polymorphic and can therefore be useful in other crosses. The subsequent genotyping of a subset of 1000 SNPs, chosen for their mapping positions using a KASP assay, showed that almost all generated SNPs are highly designable and that most (95 %) deliver highly qualitative genotyping results. Using rather low sequencing coverages in SNP discovery and in SNP inferring did not hinder the identification of hundreds of thousands of high quality SNPs. CONCLUSIONS: The development and optimization of appropriate tools in SNP discovery and genetic mapping have allowed us to make available a massive new genomic resource in pea. It will be useful for both fine mapping within chosen QTL confidence intervals and marker assisted breeding for important traits in pea improvement.

[18]

, Coyne

C J

, Grusak

M A

, Mazourek

, Cheng

, Main

, McGee

R J

. Genome-wide SNP identification, linkage map construction and QTL mapping for seed mineral concentrations and contents in pea (Pisum sativum L.)
BMC Plant Biol, 2017,17:43.

DOI:10.1186/s12870-016-0956-4 URLPMID:28193168 [本文引用: 2]

BACKGROUND: Marker-assisted breeding is now routinely used in major crops to facilitate more efficient cultivar improvement. This has been significantly enabled by the use of next-generation sequencing technology to identify loci and markers associated with traits of interest. While rich in a range of nutritional components, such as protein, mineral nutrients, carbohydrates and several vitamins, pea (Pisum sativum L.), one of the oldest domesticated crops in the world, remains behind many other crops in the availability of genomic and genetic resources. To further improve mineral nutrient levels in pea seeds requires the development of genome-wide tools. The objectives of this research were to develop these tools by: identifying genome-wide single nucleotide polymorphisms (SNPs) using genotyping by sequencing (GBS); constructing a high-density linkage map and comparative maps with other legumes, and identifying quantitative trait loci (QTL) for levels of boron, calcium, iron, potassium, magnesium, manganese, molybdenum, phosphorous, sulfur, and zinc in the seed, as well as for seed weight. RESULTS: In this study, 1609 high quality SNPs were found to be polymorphic between 'Kiflica' and 'Aragorn', two parents of an F6-derived recombinant inbred line (RIL) population. Mapping 1683 markers including 75 previously published markers and 1608 SNPs developed from the present study generated a linkage map of size 1310.1 cM. Comparative mapping with other legumes demonstrated that the highest level of synteny was observed between pea and the genome of Medicago truncatula. QTL analysis of the RIL population across two locations revealed at least one QTL for each of the mineral nutrient traits. In total, 46 seed mineral concentration QTLs, 37 seed mineral content QTLs, and 6 seed weight QTLs were discovered. The QTLs explained from 2.4% to 43.3% of the phenotypic variance. CONCLUSION: The genome-wide SNPs and the genetic linkage map developed in this study permitted QTL identification for pea seed mineral nutrients that will serve as important resources to enable marker-assisted selection (MAS) for nutritional quality traits in pea breeding programs.

[19]

Barilli

, Cobos

M J

, Carrillo

, Kilian

, Carling

, Rubiales

. A high-density integrated DArTseq SNP-based genetic map of Pisum fulvum and identification of QTLs controlling rust resistance
Front Plant Sci, 2018,9:167.

DOI:10.3389/fpls.2018.00167 URLPMID:29497430 [本文引用: 1]

Pisum fulvum, a wild relative of pea is an important source of allelic diversity to improve the genetic resistance of cultivated species against fungal diseases of economic importance like the pea rust caused by Uromyces pisi. To unravel the genetic control underlying resistance to this fungal disease, a recombinant inbred line (RIL) population was generated from a cross between two P. fulvum accessions, IFPI3260 and IFPI3251, and genotyped using Diversity Arrays Technology. A total of 9,569 high-quality DArT-Seq and 8,514 SNPs markers were generated. Finally, a total of 12,058 markers were assembled into seven linkage groups, equivalent to the number of haploid chromosomes of P. fulvum and P. sativum. The newly constructed integrated genetic linkage map of P. fulvum covered an accumulated distance of 1,877.45 cM, an average density of 1.19 markers cM(-1) and an average distance between adjacent markers of 1.85 cM. The composite interval mapping revealed three QTLs distributed over two linkage groups that were associated with the percentage of rust disease severity (DS%). QTLs UpDSII and UpDSIV were located in the LGs II and IV respectively and were consistently identified both in adult plants over 3 years at the field (Cordoba, Spain) and in seedling plants under controlled conditions. Whenever they were detected, their contribution to the total phenotypic variance varied between 19.8 and 29.2. A third QTL (UpDSIV.2) was also located in the LGIVand was environmentally specific as was only detected for DS % in seedlings under controlled conditions. It accounted more than 14% of the phenotypic variation studied. Taking together the data obtained in the study, it could be concluded that the expression of resistance to fungal diseases in P. fulvum originates from the resistant parent IFPI3260.

[20]

Kreplak

, Madoui

M A

, Capal

, Novak

, Labadie

, Aubert

, Bayer

P E

, Gali

K K

, Syme

R A

, Main

, Klein

, Berard

, Vrbova

, Fournier

, d’Agata

, Belser

, Berrabah

, Toegelova

, Milec

, Vrana

, Lee

, Kougbeadjo

, Terezol

, Huneau

, Turo

C J

, Mohellibi

, Neumann

, Falque

, Gallardo

, McGee

, Tar’an

, Bendahmane

, Aury

J M

, Batley

, Le

Paslier M C

, Ellis

, Warkentin

T D

, Coyne

C J

, Salse

, Edwards

, Lichtenzveig

, Macas

, Dolezel

, Wincker

, Burstin

. A reference genome for pea provides insight into legume genome evolution
Nat Genet, 2019,51:1411-1422.

DOI:10.1038/s41588-019-0480-1 URLPMID:31477930 [本文引用: 4]

We report the first annotated chromosome-level reference genome assembly for pea, Gregor Mendel's original genetic model. Phylogenetics and paleogenomics show genomic rearrangements across legumes and suggest a major role for repetitive elements in pea genome evolution. Compared to other sequenced Leguminosae genomes, the pea genome shows intense gene dynamics, most likely associated with genome size expansion when the Fabeae diverged from its sister tribes. During Pisum evolution, translocation and transposition differentially occurred across lineages. This reference sequence will accelerate our understanding of the molecular basis of agronomically important traits and support crop improvement.

[21]

Kalia

R K

, Rai

M K

, Kalia

, Singh

, Dhawan

A K

. Microsatellite markers: an overview of the recent progress in plants
Euphytica, 2011,177:309-334.

DOI:10.1007/s10681-010-0286-9 URL [本文引用: 4]

In recent years, molecular markers have been utilized for a variety of applications including examination of genetic relationships between individuals, mapping of useful genes, construction of linkage maps, marker assisted selections and backcrosses, population genetics and phylogenetic studies. Among the available molecular markers, microsatellites or simple sequence repeats (SSRs) which are tandem repeats of one to six nucleotide long DNA motifs, have gained considerable importance in plant genetics and breeding owing to many desirable genetic attributes including hypervariability, multiallelic nature, codominant inheritance, reproducibility, relative abundance, extensive genome coverage including organellar genomes, chromosome specific location and amenability to automation and high throughput genotyping. High degree of allelic variation revealed by microsatellite markers results from variation in number of repeat-motifs at a locus caused by replication slippage and/or unequal crossing-over during meiosis. In spite of limited understanding of the functions of the SSR motifs within the plant genes, SSRs are being widely utilized in plant genome analysis. Microsatellites can be developed directly from genomic DNA libraries or from libraries enriched for specific microsatellites. Alternatively, microsatellites can also be found by searching public databases such as GenBank and EMBL or through cross-species transferability. At present, EST databases are an important source of candidate genes, as these can generate markers directly associated with a trait of interest and may be transferable in close relative genera. A large number of SSR based techniques have been developed and a quantum of literature has accumulated regarding the applicability of SSRs in plant genetics and genomics. In this review we discuss the recent developments (last 4-5 years) made in plant genetics using SSR markers.

[22]

Vieira

M L C

, Santini

, Diniz

A L

, Munhoz

C D

. Microsatellite markers: what they mean and why they are so useful
Genet Mol Biol, 2016,39:312-328.

DOI:10.1590/1678-4685-GMB-2016-0027 URLPMID:27561112 [本文引用: 1]

Microsatellites or Single Sequence Repeats (SSRs) are extensively employed in plant genetics studies, using both low and high throughput genotyping approaches. Motivated by the importance of these sequences over the last decades this review aims to address some theoretical aspects of SSRs, including definition, characterization and biological function. The methodologies for the development of SSR loci, genotyping and their applications as molecular markers are also reviewed. Finally, two data surveys are presented. The first was conducted using the main database of Web of Science, prospecting for articles published over the period from 2010 to 2015, resulting in approximately 930 records. The second survey was focused on papers that aimed at SSR marker development, published in the American Journal of Botany's Primer Notes and Protocols in Plant Sciences (over 2013 up to 2015), resulting in a total of 87 publications. This scenario confirms the current relevance of SSRs and indicates their continuous utilization in plant science.

[23]

Ali

, Pan

Y B

, Wang

Q N

, Wang

J D

, Chen

J L

, Gao

S J

. Genetic diversity and population structure analysis of Saccharum and Erianthus genera using microsatellite (SSR) markers
Sci Rep, 2019,9:10.

DOI:10.1038/s41598-018-36877-0 URLPMID:30626881 [本文引用: 1]

=0.9 for Ceriodaphnoa dubia, Asellus aquaticus, Daphnia magna, Daphnia pulex; r(2) >=0.8 for Hyalella azteca, Chironomus spec. larvae and Culex spec. larvae) to convert size measured on the spheroid counter to traditional, microscope based, length measurements, which follow the longest orientation of the body. Finally, we demonstrate semi-automated measurement of growth curves of individual daphnids (C. dubia and D. magna) over time and find that the quality of individual growth curves varies, partly due to methodological reasons. Nevertheless, this novel method could be adopted to other species and represents a step change in experimental throughput for measuring organisms' shape, size and growth curves. It is also a significant qualitative improvement by enabling high-throughput assessment of inter-individual variation of growth.]]>

[24]

Hao

, Zhang

G S

, Lu

D Y

, Hu

J J

, Jia

H X

. Analysis of the genetic diversity and population structure of Salix psammophila based on phenotypic traits and simple sequence repeat markers
PeerJ, 2019,7:e6419.

DOI:10.7717/peerj.6419 URLPMID:30805247 [本文引用: 1]

Salix psammophila (desert willow) is a shrub endemic to the Kubuqi Desert and the Mu Us Desert, China, that plays an important role in maintaining local ecosystems and can be used as a biomass feedstock for biofuels and bioenergy. However, the lack of information on phenotypic traits and molecular markers for this species limits the study of genetic diversity and population structure. In this study, nine phenotypic traits were analyzed to assess the morphological diversity and variation. The mean coefficient of variation of 17 populations ranged from 18.35% (branch angle (BA)) to 38.52% (leaf area (LA)). Unweighted pair-group method with arithmetic mean analysis of nine phenotypic traits of S. psammophila showed the same results, with the 17 populations clustering into five groups. We selected 491 genets of the 17 populations to analyze genetic diversity and population structure based on simple sequence repeat (SSR) markers. Analysis of molecular variance (AMOVA) revealed that most of the genetic variance (95%) was within populations, whereas only a small portion (5%) was among populations. Moreover, using the animal model with SSR-based relatedness estimated of S. psammophila, we found relatively moderate heritability values for phenotypic traits, suggesting that most of trait variation were caused by environmental or developmental variation. Principal coordinate and phylogenetic analyses based on SSR data revealed that populations P1, P2, P9, P16, and P17 were separated from the others. The results showed that the marginal populations located in the northeastern and southwestern had lower genetic diversity, which may be related to the direction of wind. These results provide a theoretical basis for germplasm management and genetic improvement of desert willow.

[25]

Choi

J K

, Sa

K J

, Park

D H

, Lim

S E

, Ryu

S H

, Park

J Y

, Park

K J

, Rhee

H I

, Lee

J K

. Construction of genetic linkage map and identification of QTLs related to agronomic traits in DH population of maize (Zea mays L.) using SSR markers
Genes Genomics, 2019,41:667-678.

DOI:10.1007/s13258-019-00813-x URLPMID:30953340 [本文引用: 1]

BACKGROUND: In this study, we used phenotypic and genetic analysis to investigate Double haploid (DH) lines derived from normal corn parents (HF1 and 11S6169). DH technology offers an array of advantages in maize genetics and breeding as follows: first, it significantly shortens the breeding cycle by development of completely homozygous lines in two or three generations; and second, it simplifies logistics, including requiring less time, labor, and financial resources for developing new DH lines compared with the conventional RIL population development process. OBJECTIVES: In our study, we constructed a maize genetic linkage map using SSR markers and a DH population derived from a cross of normal corn (HF1) and normal corn (11S6169). METHODS: The DH population used in this study was developed by the following methods: we crossed normal corn (HF1) and normal corn (11S6169), which are parent lines of a normal corn cultivar, in 2014; and the next year, the F1 hybrids were crossed with a tropicalized haploid inducer line (TAIL), which is homozygous for the dominant marker gene R1-nj (Nanda and Chase in Crop Sci 6:213-215, 1966), and we harvested seeds of the haploid lines. RESULTS: A total of 200 SSR markers were assigned to 10 linkage groups that spanned 1145.4 cM with an average genetic distance between markers of 5.7 cM. 68 SSR markers showed Mendelian segregation ratios in the DH population at a 5% significance threshold. A total of 15 quantitative trait loci (QTLs) for plant height (PH), ear height (EH), ear height ratio (ER), leaf length (LL), ear length (EL), set ear length (SEL), set ear ratio (SER), ear width (EW), 100 kernel weight (100 KW), and cob color (CC) were found in the 121 lines in the DH population. CONCLUSION: The results of this study may help to improve the detection and characterization of agronomic traits and provide great opportunities for maize breeders and researchers using a DH population in maize breeding programs.

[26]

Yang

, Jiang

J Y

, Zhang

H Y

, Liu

, Strelkov

, Hwang

S F

, Chang

K F

, Yang

, Miao

Y M

, He

Y H

, Zong

X X

. Density enhancement of a faba bean genetic linkage map (Vicia faba) based on simple sequence repeats markers
Plant Breed, 2019,138:207-215.

DOI:10.1111/pbr.2019.138.issue-2 URL [本文引用: 1]

[27]

Anjani

, Ponukumatla

, Mishra

, Ravulapalli

D P

. Identification of simple-sequence-repeat markers linked to Fusarium wilt (Fusarium oxysporum f. sp carthami) resistance and marker- assisted selection for wilt resistance in safflower (Carthamus tinctorius L.) interspecific offsprings
Plant Breed, 2018,137:895-902.

DOI:10.1111/pbr.2018.137.issue-6 URL [本文引用: 1]

[28]

Swathi

, Rani

C V D

, Md

, Madhav

M S

, Vanisree

, Anuradha

, Kumar

N R

, Kumar

N A P

, Kumari

K A

, Bhogadhi

, Ramprasad

, Sravanthi

, Raju

S K

, Bhuvaneswari

, Rajan

C P D

, Jagadeeswar

. Marker-assisted introgression of the major bacterial blight resistance genes, Xa21 and xa13, and blast resistance gene, Pi54, into the popular rice variety, JGL1798
Mol Breed, 2019,39:12.

DOI:10.1007/s11032-018-0919-6 URL [本文引用: 1]

[29]

Kumar

, Shikha

, Kumari

, Choudhary

B K

, Kumar

, Singh

I S

. SSR-based DNA Fingerprinting and diversity assessment among Indian germplasm of Euryale ferox: an aquatic underutilized and neglected food crop
Appl Biochem Biotechnol, 2018,185:34-41.

DOI:10.1007/s12010-017-2643-9 URLPMID:29082475 [本文引用: 1]

Euryale ferox is native to Southeast Asia and China, and it is one of the important aquatic food crops propagated mostly in eastern part of India. The aim of the present study was to characterize and evaluate the genetic diversity of ex situ collections of E. ferox germplasm from different geographical states of India using microsatellite (simple sequence repeats (SSRs)) markers. Ten SSR markers were analyzed to assess DNA fingerprinting and genetic diversity of 16 cultivated germplasm of E. ferox. Total 37 polymorphic alleles were recorded with an average of 3.7 allele frequency per primer. The polymorphic information content value varied from 0.204 to 0.735 with mean of 0.448. A high range of heterozygosity (Ho 0.228; He 0.512) was detected in the present study. The neighbor-joining (N-J) tree and the principle coordinate analysis showed that the germplasm divided in to three main clusters. The results of the present investigation comply that SSR markers are effective for computing genetic assessment of genetic diversity and similarity with classifying cultivated varieties of E. ferox. Evaluation of genetic diversity among Indian E. ferox germplasm could provide useful information for genetic improvement.

[30]

Siew

G Y

, Ng

W L

, Tan

S W

, Alitheen

N B

, Tan

S G

, Yeap

S K

. Genetic variation and DNA fingerprinting of durian types in Malaysia using simple sequence repeat (SSR) markers
PeerJ, 2018,6:e4266.

DOI:10.7717/peerj.4266 URLPMID:29511604 [本文引用: 1]

Durian (Durio zibethinus) is one of the most popular tropical fruits in Asia. To date, 126 durian types have been registered with the Department of Agriculture in Malaysia based on phenotypic characteristics. Classification based on morphology is convenient, easy, and fast but it suffers from phenotypic plasticity as a direct result of environmental factors and age. To overcome the limitation of morphological classification, there is a need to carry out genetic characterization of the various durian types. Such data is important for the evaluation and management of durian genetic resources in producing countries. In this study, simple sequence repeat (SSR) markers were used to study the genetic variation in 27 durian types from the germplasm collection of Universiti Putra Malaysia. Based on DNA sequences deposited in Genbank, seven pairs of primers were successfully designed to amplify SSR regions in the durian DNA samples. High levels of variation among the 27 durian types were observed (expected heterozygosity, HE = 0.35). The DNA fingerprinting power of SSR markers revealed by the combined probability of identity (PI) of all loci was 2.3x10(-3). Unique DNA fingerprints were generated for 21 out of 27 durian types using five polymorphic SSR markers (the other two SSR markers were monomorphic). We further tested the utility of these markers by evaluating the clonal status of shared durian types from different germplasm collection sites, and found that some were not clones. The findings in this preliminary study not only shows the feasibility of using SSR markers for DNA fingerprinting of durian types, but also challenges the current classification of durian types, e.g., on whether the different types should be called

[31]

Tayeh

, Aubert

, Pilet Nayel

M L

, Lejeune Henaut

, Warkentin

T D

, Burstin

. Genomic tools in pea breeding programs: Status and perspectives
Front Plant Sci, 2015,6:1037.

DOI:10.3389/fpls.2015.01037 URLPMID:26640470 [本文引用: 3]

Pea (Pisum sativum L.) is an annual cool-season legume and one of the oldest domesticated crops. Dry pea seeds contain 22-25% protein, complex starch and fiber constituents, and a rich array of vitamins, minerals, and phytochemicals which make them a valuable source for human consumption and livestock feed. Dry pea ranks third to common bean and chickpea as the most widely grown pulse in the world with more than 11 million tons produced in 2013. Pea breeding has achieved great success since the time of Mendel's experiments in the mid-1800s. However, several traits still require significant improvement for better yield stability in a larger growing area. Key breeding objectives in pea include improving biotic and abiotic stress resistance and enhancing yield components and seed quality. Taking advantage of the diversity present in the pea genepool, many mapping populations have been constructed in the last decades and efforts have been deployed to identify loci involved in the control of target traits and further introgress them into elite breeding materials. Pea now benefits from next-generation sequencing and high-throughput genotyping technologies that are paving the way for genome-wide association studies and genomic selection approaches. This review covers the significant development and deployment of genomic tools for pea breeding in recent years. Future prospects are discussed especially in light of current progress toward deciphering the pea genome.

[32]

Liu

, Fang

, Yang

, Zhang

X Y

, Hu

J G

, Zhang

H Y

, Han

W L

, Hua

Z K

, Hao

J J

, Zong

X X

. Marker-trait association analysis of frost tolerance of 672 worldwide pea (Pisum sativum L.) collections
Sci Rep, 2017,7:5919.

DOI:10.1038/s41598-017-06222-y URLPMID:28724947 [本文引用: 3]

Frost stress is one of the major abiotic stresses causing seedling death and yield reduction in winter pea. To improve the frost tolerance of pea, field evaluation of frost tolerance was conducted on 672 diverse pea accessions at three locations in Northern China in three growing seasons from 2013 to 2016 and marker-trait association analysis of frost tolerance were performed with 267 informative SSR markers in this study. Sixteen accessions were identified as the most winter-hardy for their ability to survive in all nine field experiments with a mean survival rate of 0.57, ranging from 0.41 to 0.75. Population structure analysis revealed a structured population of two sub-populations plus some admixtures in the 672 accessions. Association analysis detected seven markers that repeatedly had associations with frost tolerance in at least two different environments with two different statistical models. One of the markers is the functional marker EST1109 on LG VI which was predicted to co-localize with a gene involved in the metabolism of glycoproteins in response to chilling stress and may provide a novel mechanism of frost tolerance in pea. These winter-hardy germplasms and frost tolerance associated markers will play a vital role in marker-assisted breeding for winter-hardy pea cultivar.

[33]

X B

, Li

N N

, Hao

J J

, Hu

J G

, Zhang

X Y

, Blair

M W

. Genetic diversity of Chinese and global pea (Pisum sativum L.) collections
Crop Sci, 2017,57:1-11.

DOI:10.2135/cropsci2015.07.0415 URL [本文引用: 3]

[34]

Milczarski

, Bolibok Br?goszewska

, My?ków

, Stoja?owski

, Heller Uszyńska

, Góralska

, Br?goszewski

, Uszyński

, Kilian

, Rakoczy Trojanowska

. A high density consensus map of rye (Secale cereale L.) based on DArT markers
PLoS One, 2011,6:e28495.

DOI:10.1371/journal.pone.0028495 URLPMID:22163026 [本文引用: 2]

BACKGROUND: Rye (Secale cereale L.) is an economically important crop, exhibiting unique features such as outstanding resistance to biotic and abiotic stresses and high nutrient use efficiency. This species presents a challenge to geneticists and breeders due to its large genome containing a high proportion of repetitive sequences, self incompatibility, severe inbreeding depression and tissue culture recalcitrance. The genomic resources currently available for rye are underdeveloped in comparison with other crops of similar economic importance. The aim of this study was to create a highly saturated, multilocus linkage map of rye via consensus mapping, based on Diversity Arrays Technology (DArT) markers. METHODOLOGY/PRINCIPAL FINDINGS: Recombinant inbred lines (RILs) from 5 populations (564 in total) were genotyped using DArT markers and subjected to linkage analysis using Join Map 4.0 and Multipoint Consensus 2.2 software. A consensus map was constructed using a total of 9703 segregating markers. The average chromosome map length ranged from 199.9 cM (2R) to 251.4 cM (4R) and the average map density was 1.1 cM. The integrated map comprised 4048 loci with the number of markers per chromosome ranging from 454 for 7R to 805 for 4R. In comparison with previously published studies on rye, this represents an eight-fold increase in the number of loci placed on a consensus map and a more than two-fold increase in the number of genetically mapped DArT markers. CONCLUSIONS/SIGNIFICANCE: Through the careful choice of marker type, mapping populations and the use of software packages implementing powerful algorithms for map order optimization, we produced a valuable resource for rye and triticale genomics and breeding, which provides an excellent starting point for more in-depth studies on rye genome organization.

[35]

Blenda

, Fang

D D

, Rami

J F

, Garsmeur

, Luo

, Lacape

J M

. A high density consensus genetic map of tetraploid cotton that integrates multiple component maps through molecular marker redundancy check
PLoS One, 2012,7:e45739.

DOI:10.1371/journal.pone.0045739 URLPMID:23029214 [本文引用: 2]

A consensus genetic map of tetraploid cotton was constructed using six high-density maps and after the integration of a sequence-based marker redundancy check. Public cotton SSR libraries (17,343 markers) were curated for sequence redundancy using 90% as a similarity cutoff. As a result, 20% of the markers (3,410) could be considered as redundant with some other markers. The marker redundancy information had been a crucial part of the map integration process, in which the six most informative interspecific Gossypium hirsutumxG. barbadense genetic maps were used for assembling a high density consensus (HDC) map for tetraploid cotton. With redundant markers being removed, the HDC map could be constructed thanks to the sufficient number of collinear non-redundant markers in common between the component maps. The HDC map consists of 8,254 loci, originating from 6,669 markers, and spans 4,070 cM, with an average of 2 loci per cM. The HDC map presents a high rate of locus duplications, as 1,292 markers among the 6,669 were mapped in more than one locus. Two thirds of the duplications are bridging homoeologous A(T) and D(T) chromosomes constitutive of allopolyploid cotton genome, with an average of 64 duplications per A(T)/D(T) chromosome pair. Sequences of 4,744 mapped markers were used for a mutual blast alignment (BBMH) with the 13 major scaffolds of the recently released Gossypium raimondii genome indicating high level of homology between the diploid D genome and the tetraploid cotton genetic map, with only a few minor possible structural rearrangements. Overall, the HDC map will serve as a valuable resource for trait QTL comparative mapping, map-based cloning of important genes, and better understanding of the genome structure and evolution of tetraploid cotton.

[36]

Sudheesh

, Lombardi

, Leonforte

, Cogan

N O I

, Materne

, Forster

J W

, Kaur

. Consensus genetic map construction for field pea (Pisum sativum L.), trait dissection of biotic and abiotic stress tolerance and development of a diagnostic marker for the er1 powdery mildew resistance gene
Plant Mol Biol Rep, 2015,33:1391-1403.

DOI:10.1007/s11105-014-0837-7 URL [本文引用: 5]

[37]

Sudheesh

, Rodda

, Kennedy

, Verma

, Leonforte

, Cogan

N O I

, Materne

, Forster

J W

, Kaur

. Construction of an integrated linkage map and trait dissection for bacterial blight resistance in field pea (Pisum sativum L.)
Mol Breed, 2015,35:185.

DOI:10.1007/s11032-015-0376-4 URL [本文引用: 5]

[38]

Yang

, Fang

, Zhang

X Y

, Hu

J G

, Bao

S Y

, Hao

J J

, Li

, He

Y H

, Jiang

J Y

, Wang

, Tian

, Zong

X X

. High-throughput development of SSR markers from pea (Pisum sativum L.) based on next generation sequencing of a purified Chinese commercial variety
PLoS One, 2015,10:e0139775.

DOI:10.1371/journal.pone.0139775 URLPMID:26440522 [本文引用: 3]

Pea (Pisum sativum L.) is an important food legume globally, and is the plant species that J.G. Mendel used to lay the foundation of modern genetics. However, genomics resources of pea are limited comparing to other crop species. Application of marker assisted selection (MAS) in pea breeding has lagged behind many other crops. Development of a large number of novel and reliable SSR (simple sequence repeat) or microsatellite markers will help both basic and applied genomics research of this crop. The Illumina HiSeq 2500 System was used to uncover 8,899 putative SSR containing sequences, and 3,275 non-redundant primers were designed to amplify these SSRs. Among the 1,644 SSRs that were randomly selected for primer validation, 841 yielded reliable amplifications of detectable polymorphisms among 24 genotypes of cultivated pea (Pisum sativum L.) and wild relatives (P. fulvum Sm.) originated from diverse geographical locations. The dataset indicated that the allele number per locus ranged from 2 to 10, and that the polymorphism information content (PIC) ranged from 0.08 to 0.82 with an average of 0.38. These 1,644 novel SSR markers were also tested for polymorphism between genotypes G0003973 and G0005527. Finally, 33 polymorphic SSR markers were anchored on the genetic linkage map of G0003973 x G0005527 F2 population.

[39]

Kwon

S J

, Brown

A F

, Hu

, McGee

, Watt

, Kisha

, Timmerman-Vaughan

, Grusak

, McPhee

K E

, Coyne

C J

. Genetic diversity, population structure and genome-wide marker-trait association analysis emphasizing seed nutrients of the USDA pea (Pisum sativum L.) core collection
Genes Genomics, 2012,34:305-320.

DOI:10.1007/s13258-011-0213-z URL [本文引用: 5]

[40]

Kaur

S J

, Pembleton

L W

, Cogan

N O

, Savin

K W

, Leonforte

, Paull

, Materne

, Forster

J W

. Transcriptome sequencing of field pea and faba bean for discovery and validation of SSR genetic markers
BMC Genomics, 2012,13:104.

DOI:10.1186/1471-2164-13-104 URLPMID:22433453 [本文引用: 2]

BACKGROUND: Field pea (Pisum sativum L.) and faba bean (Vicia faba L.) are cool-season grain legume species that provide rich sources of food for humans and fodder for livestock. To date, both species have been relative 'genomic orphans' due to limited availability of genetic and genomic information. A significant enrichment of genomic resources is consequently required in order to understand the genetic architecture of important agronomic traits, and to support germplasm enhancement, genetic diversity, population structure and demographic studies. RESULTS: cDNA samples obtained from various tissue types of specific field pea and faba bean genotypes were sequenced using 454 Roche GS FLX Titanium technology. A total of 720,324 and 304,680 reads for field pea and faba bean, respectively, were de novo assembled to generate sets of 70,682 and 60,440 unigenes. Consensus sequences were compared against the genome of the model legume species Medicago truncatula Gaertn., as well as that of the more distantly related, but better-characterised genome of Arabidopsis thaliana L.. In comparison to M. truncatula coding sequences, 11,737 and 10,179 unique hits were obtained from field pea and faba bean. Totals of 22,057 field pea and 18,052 faba bean unigenes were subsequently annotated from GenBank. Comparison to the genome of soybean (Glycine max L.) resulted in 19,451 unique hits for field pea and 16,497 unique hits for faba bean, corresponding to c. 35% and 30% of the known gene space, respectively. Simple sequence repeat (SSR)-containing expressed sequence tags (ESTs) were identified from consensus sequences, and totals of 2,397 and 802 primer pairs were designed for field pea and faba bean. Subsets of 96 EST-SSR markers were screened for validation across modest panels of field pea and faba bean cultivars, as well as related non-domesticated species. For field pea, 86 primer pairs successfully obtained amplification products from one or more template genotypes, of which 59% revealed polymorphism between 6 genotypes. In the case of faba bean, 81 primer pairs displayed successful amplification, of which 48% detected polymorphism. CONCLUSIONS: The generation of EST datasets for field pea and faba bean has permitted effective unigene identification and functional sequence annotation. EST-SSR loci were detected at incidences of 14-17%, permitting design of comprehensive sets of primer pairs. The subsets from these primer pairs proved highly useful for polymorphism detection within Pisum and Vicia germplasm.

[41]

S C

, Gong

Y M

, Mao

W H

, Hu

Q Z

, Zhang

G W

, Fu

, Xian

Q Q

. Development and characterization of 41 novel EST-SSR markers for Pisum sativum (Leguminosae)
Am J Bot, 2012,99:E149-E153.

DOI:10.3732/ajb.1100445 URL [本文引用: 5]

Methods and Results: Forty-one novel EST-SSR primers were developed and characterized for size polymorphism in 32 Pisum sativum individuals from four populations from China. In each population, the number of alleles per locus ranged from one to seven, with observed heterozygosity and expected heterozygosity ranging from 0 to 0.8889 and 0 to 0.8400, respectively. Furthermore, 53.7% of these markers could be transferred to the related species, Vicia faba.Conclusions: The developed markers have potential for application in the study of genetic diversity, germplasm appraisal, and marker-assisted breeding in pea and other legume species.]]>

[42]

Bordat

, Savois

, Nicolas

, Salse

, Chauveau

, Bourgeois

, Potier

, Houtin

, Rond

, Murat

, Marget

, Aubert

, Burstin

. Translational genomics in legumes allowed placing in silico 5460 unigenes on the pea functional map and identified candidate genes in Pisum sativum L
G3: Genes Genom Genet 2011,1:93-103.

[本文引用: 4]

[43]

顾竟, 李玲, 宗绪晓, 王海飞, 关建平, 杨涛. 豌豆种质表型性状SSR标记关联分析
植物遗传资源学报, 2011,12:833-839.

, Li

, Zong

X X

, Wang

H F

, Guan

J P

, Yang

. Association analysis between morphological traits of pea and its polymorphic SSR markers
J Plant Genet Resour, 2011,12:833-839 (in Chinese with English abstract).

[44]

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, Aubert

, Barranger

. Microsatellite polymorphism in Pisum sativum
Plant Breed, 2001,120:311.

DOI:10.1046/j.1439-0523.2001.00608.x URL [本文引用: 4]

[45]

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S L

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J B

. A plant DNA minipreparation: version II
Plant Mol Biol Rep, 1983,1:19-21.

DOI:10.1007/BF02712670 URL [本文引用: 1]

[46]

Meng

, Li

H H

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L Y

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J K

. QTL IciMapping: integrated software for genetic linkage map construction and quantitative trait locus mapping in biparental populations
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DOI:10.1016/j.cj.2015.01.001 URL [本文引用: 1]

[47]

Voorrips

R E

. MapChart: software for the graphical presentation of linkage maps and QTLs
J Hered, 2002,93:77-78.

DOI:10.1093/jhered/93.1.77 URLPMID:12011185 [本文引用: 1]

[48]

Parida

S K

, Kalia

S K

, Kaul

, Dalal

, Hemaprabha

, Selvi

, Pandit

, Singh

, Gaikwad

, Sharma

T R

, Srivastava

P S

, Singh

N K

, Mohapatra

. Informative genomic microsatellite markers for efficient genotyping applications in sugarcane
Theor Appl Genet, 2009,118:327-338.

DOI:10.1007/s00122-008-0902-4 URL [本文引用: 1]

Genomic microsatellite markers are capable of revealing high degree of polymorphism. Sugarcane (Saccharum sp.), having a complex polyploid genome requires more number of such informative markers for various applications in genetics and breeding. With the objective of generating a large set of microsatellite markers designated as Sugarcane Enriched Genomic MicroSatellite (SEGMS), 6,318 clones from genomic libraries of two hybrid sugarcane cultivars enriched with 18 different microsatellite repeat-motifs were sequenced to generate 4.16Mb high-quality sequences. Microsatellites were identified in 1,261 of the 5,742 non-redundant clones that accounted for 22% enrichment of the libraries. Retro-transposon association was observed for 23.1% of the identified microsatellites. The utility of the microsatellite containing genomic sequences were demonstrated by higher primer designing potential (90%) and PCR amplification efficiency (87.4%). A total of 1,315 markers including 567 class I microsatellite markers were designed and placed in the public domain for unrestricted use. The level of polymorphism detected by these markers among sugarcane species, genera, and varieties was 88.6%, while cross-transferability rate was 93.2% within Saccharum complex and 25% to cereals. Cloning and sequencing of size variant amplicons revealed that the variation in the number of repeat-units was the main source of SEGMS fragment length polymorphism. High level of polymorphism and wide range of genetic diversity (0.16–0.82 with an average of 0.44) assayed with the SEGMS markers suggested their usefulness in various genotyping applications in sugarcane.]]>

[49]

Kamaluddin

, Khan

M A

, Kiran

, Ali

, Abdin

M Z

, Zargar

M Y

, Ahmad

, Sofi

P A

, Gulzar

. Molecular markers and marker-assisted selection in crop plants
In: Abdin M Z, Kiran U, Kamaluddin, Ali A, eds. Plant Biotechnology: Principles and Applications. Singapore: Springer Singapore, 2017. pp 295-328.

[50]

Nadeem

M A

, Nawaz

M A

, Shahid

M Q

, Do?an

, Comertpay

, Y?ld?z

, Hatipo?lu

, Ahmad

, Alsaleh

, Labhane

, ?zkan

, Chung

, Baloch

F S

. DNA molecular markers in plant breeding: current status and recent advancements in genomic selection and genome editing
Biotechnol Biotechnol Equip, 2018,32:261-285.

DOI:10.1080/13102818.2017.1400401 URL [本文引用: 1]

[51]

Varshney

R K

, Graner

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M E

. Genic microsatellite markers in plants: features and applications
Trends Biotechnol, 2005,23:48-55.

DOI:10.1016/j.tibtech.2004.11.005 URLPMID:15629858 [本文引用: 1]

Expressed sequence tag (EST) projects have generated a vast amount of publicly available sequence data from plant species; these data can be mined for simple sequence repeats (SSRs). These SSRs are useful as molecular markers because their development is inexpensive, they represent transcribed genes and a putative function can often be deduced by a homology search. Because they are derived from transcripts, they are useful for assaying the functional diversity in natural populations or germplasm collections. These markers are valuable because of their higher level of transferability to related species, and they can often be used as anchor markers for comparative mapping and evolutionary studies. They have been developed and mapped in several crop species and could prove useful for marker-assisted selection, especially when the markers reside in the genes responsible for a phenotypic trait. Applications and potential uses of EST-SSRs in plant genetics and breeding are discussed.

[52]

Smykal

, Hybl

, Corander

, Jarkovsky

, Flavell

A J

, Griga

. Genetic diversity and population structure of pea (Pisum sativum L.) varieties derived from combined retrotransposon, microsatellite and morphological marker analysis
Theor Appl Genet, 2008,117:413-424.

DOI:10.1007/s00122-008-0785-4 URL [本文引用: 1]

One hundred and sixty-four accessions representing Czech and Slovak pea (Pisum sativum L.) varieties bred over the last 50years were evaluated for genetic diversity using morphological, simple sequence repeat (SSR) and retrotransposon-based insertion polymorphism (RBIP) markers. Polymorphic information content (PIC) values of 10 SSR loci and 31 RBIP markers were on average high at 0.89 and 0.73, respectively. The silhouette method after the Ward clustering produced the most probable cluster estimate, identifying nine clusters from molecular data and five to seven clusters from morphological characters. Principal component analysis of nine qualitative and eight quantitative morphological parameters explain over 90 and 93% of total variability, respectively, in the first three axes. Multidimensional scaling of molecular data revealed a continuous structure for the set. To enable integration and evaluation of all data types, a Bayesian method for clustering was applied. Three clusters identified using morphology data, with clear separation of fodder, dry seed and afila types, were resolved by DNA data into 17, 12 and five sub-clusters, respectively. A core collection of 34 samples was derived from the complete collection by BAPS Bayesian analysis. Values for average gene diversity and allelic richness for molecular marker loci and diversity indexes of phenotypic data were found to be similar between the two collections, showing that this is a useful approach for representative core selection.]]>

[53]

宗绪晓, 关建平, 王述民, 刘庆昌. 中国豌豆地方品种SSR标记遗传多样性分析
作物学报, 2008,34:1330-1338.

DOI:10.3724/SP.J.1006.2008.01330 URL

利用21对豌豆多态性SSR引物, 对来自全国春、秋播区19省区市的1 221份豌豆地方品种进行遗传多样性分析, 共扩增出104条多态性带, 每对引物平均扩增出4.95个等位变异, 其中有效等位变异占62.52%。省份间SSR等位变异分布均匀, 但是省份间有效等位变异数、Shannon’s信息指数(I)差异明显, 省籍资源群间遗传多样性差异显著。遗传多样性以内蒙古资源群最高, 甘肃、四川、云南和西藏等资源群其次, 辽宁资源群最低。PCA三维空间聚类图揭示, 我国豌豆地方品种资源分化成3个基因库, 基因库I主要由春播区的内蒙古、陕西资源构成, 基因库II主要由秋播区最北端的河南资源构成, 基因库III主要由除上述省份之外的其他省区市的资源构成。UPGMA聚类分析表明, 不同省份资源群间的遗传距离变化范围为5.159~27.586, 中国豌豆地方资源据此聚类成2个组群8个亚组群, 与3个基因库的聚类结果相呼应。聚类结果显示, 我国豌豆地方品种资源群间遗传距离与其来源地生态环境相关联。

Zong

X X

, Guan

J P

, Wang

S M

, Liu

Q C

. Genetic diversity among Chinese pea (Pisum sativum L.) landraces revealed by SSR markers
Acta Agron Sin, 2008,34:1330-1338 (in Chinese with English abstract).

[54]

Zong

X X

, Ford

, Redden

R R

, Guan

J P

, Wang

S M

. Identification and analysis of genetic diversity structure within Pisum genus based on microsatellite markers
Agric Sci China, 2009,8:257-267.

DOI:10.1016/S1671-2927(08)60208-4 URL

[55]

宗绪晓, 关建平, 王述民, 刘庆昌, Redden R R, Ford R. 国外栽培豌豆遗传多样性分析及核心种质构建
作物学报, 2008,34:1518-1528.

DOI:10.3724/SP.J.1006.2008.01518 URL [本文引用: 1]

Pisum sativum L.)进行遗传多样性分析与核心种质构建。共扩增出109条多态性带, 每对引物平均扩增出5.19个等位变异。SSR等位变异在各大洲间分布不均匀, 有效等位变异数、Shannon’s信息指数(I)洲际间差异明显。各大洲资源群间遗传多样性差异显著, 其中亚洲最高(I = 1.1753), 欧洲其次(I = 1.1387), 俄罗斯联邦(I = 1.0285)、美洲(I = 1.0196)、非洲(I = 0.9254)、大洋洲(I = 0.8608)依次降低。利用Popgene 1.32软件, 依豌豆栽培资源洲际间Nei78遗传距离可聚类成2个组群和4个亚组群; 基于Structure 2.2软件分析, 国外栽培豌豆资源实际由3大类群组成, 并与Popgene 1.32聚类结果呼应得较好。上述两种分析方法均表明, 国外栽培豌豆类群的遗传多样性与其地理分布相关。设计并实践了一套基于Structure分析的科学可靠、逻辑性强的核心种质构建标准化方案, 并依此构建了一套以6.57%的资源(48份)涵盖总体84.4%等位变异的国外栽培豌豆核心种质。]]>

Zong

X X

, Guan

J P

, Wang

S M

, Liu

Q C

, Redden

R R

, Ford

. Genetic diversity and core collection of alien Pisum sativum L. germplasm
Acta Agron Sin, 2008,34:1518-1528 (in Chinese with English abstract).

[本文引用: 1]

[56]

Prakash

, Kumar

, Choudhary

V K

, Singh

C M

. Molecular assessment of genetic divergence in pea genotypes using microsatellite markers
Legume Res, 2016,39:183-188.

[本文引用: 1]

[57]

Duarte

, Riviere

, Baranger

, Aubert

, Burstin

, Cornet

, Lavaud

, Lejeune Henaut

, Martinant

J P

, Pichon

J P

, Pilet Nayel

M L

, Boutet

. Transcriptome sequencing for high throughput SNP development and genetic mapping in pea
BMC Genom, 2014,15:126.

DOI:10.1186/1471-2164-15-126 URL [本文引用: 1]

[58]

Guindon

M F

, Martin

, Cravero

, Gali

K K

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T D

, Cointry

. Linkage map development by GBS, SSR, and SRAP techniques and yield-related QTLs in pea
Mol Breed, 2019,39:54.

DOI:10.1007/s11032-019-0949-8 URL [本文引用: 1]

[59]

Aubert

, Morin

, Jacquin

, Loridon

, Quillet

M C

, Petit

, Rameau

, Lejeune Henaut

, Huguet

, Burstin

. Functional mapping in pea, as an aid to the candidate gene selection and for investigating synteny with the model legume Medicago truncatula
Theor Appl Genet, 2006,112:1024-1041.

DOI:10.1007/s00122-005-0205-y URL [本文引用: 1]

The identification of the molecular polymorphisms giving rise to phenotypic trait variability—both quantitative and qualitative—is a major goal of the present agronomic research. Various approaches such as positional cloning or transposon tagging, as well as the candidate gene strategy have been used to discover the genes underlying this variation in plants. The construction of functional maps, i.e. composed of genes of known function, is an important component of the candidate gene approach. In the present paper we report the development of 63 single nucleotide polymorphism markers and 15 single-stranded conformation polymorphism markers for genes encoding enzymes mainly involved in primary metabolism, and their genetic mapping on a composite map using two pea recombinant inbred line populations. The complete genetic map covers 1,458cM and comprises 363 loci, including a total of 111 gene-anchored markers: 77 gene-anchored markers described in this study, 7 microsatellites located in gene sequences, 16 flowering time genes, the Tri gene, 5 morphological markers, and 5 other genes. The mean spacing between adjacent markers is 4cM and 90% of the markers are closer than 10cM to their neighbours. We also report the genetic mapping of 21 of these genes in Medicago truncatula and add 41 new links between the pea and M. truncatula maps. We discuss the use of this new composite functional map for future candidate gene approaches in pea.]]>

[60]

Duarte

, Riviere

, Baranger

, Aubert

, Burstin

, Cornet

, Lavaud

, Lejeune Henaut

, Martinant

J P

, Pichon

J P

, Pilet Nayel

M L

, Boutet

. Transcriptome sequencing for high throughput SNP development and genetic mapping in pea
BMC Genom, 2014,15:126.

DOI:10.1186/1471-2164-15-126 URL

[61]

Sindhu

, Ramsay

, Sanderson

L A

, Stonehouse

, Li

, Condie

, Shunmugam

A S K

, Liu

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A B

, Diapari

, Burstin

, Aubert

, Tar’an

, Bett

K E

, Warkentin

T D

, Sharpe

A G

. Gene-based SNP discovery and genetic mapping in pea
Theor Appl Genet, 2014,127:2225-2241.

DOI:10.1007/s00122-014-2375-y URL [本文引用: 2]

Sybenga

. Recombination and chiasmata: few but intriguing discrepancies
Genome, 1996,39:473-484.

DOI:10.1139/g96-061 URLPMID:18469909 [本文引用: 1]

The paradigm that meiotic recombination and chiasmata have the same basis has been challenged, primarily for plants. High resolution genetic mapping frequently results in maps with lengths far exceeding those based on chiasma counts. In addition, recombination between specific homoeologous chromosomes derived from interspecific hybrids is sometimes much higher than can be explained by meiotic chiasma frequencies. However, almost the entire discrepancy disappears when proper care is taken of map inflation resulting from the shortcomings of the mapping algorithm and classification errors, the use of dissimilar material, and the difficulty of accurately counting chiasmata. Still, some exchanges, especially of short interstitial segments, cannot readily be explained by normal meiotic behaviour. Aberrant meiotic processes involving segment replacement or insertion can probably be excluded. Some cases of unusual recombination are somatic, possibly premeiotic exchange. For other cases, local relaxation of chiasma interference caused by small interruptions of homology disturbing synaptonemal complex formation is proposed as the cause. It would be accompanied by a preference for compensating exchanges (negative chromatid interference) resulting from asymmetry of the pairing chromatid pairs, so that one side of each pair preferentially participates in pairing. Over longer distances, the pairing face may switch, causing the normal random chromatid participation in double exchanges and the relatively low frequency of short interstitial exchanges. Key words : recombination frequency, map length, chiasmata, discrepancy, chromatid interference.

[63]

Knox

M R

, Ellis

T H N

. Excess heterozygosity contributes to genetic map expansion in pea recombinant inbred populations
Genetics, 2002,162:861-873.

URLPMID:12399396

Several plant genetic maps presented in the literature are longer than expected from cytogenetic data. Here we compare F(2) and RI maps derived from a cross between the same two parental lines and show that excess heterozygosity contributes to map inflation. These maps have been constructed using a common set of dominant markers. Although not generally regarded as informative for F(2) mapping, these allowed rapid map construction, and the resulting data analysis has provided information not otherwise obvious when examining a population from only one generation. Segregation distortion, a common feature of most populations and marker systems, found in the F(2) but not the RI, has identified excess heterozygosity. A few markers with a deficiency of heterozygotes were found to map to linkage group V (chromosome 3), which is known to form rod bivalents in this cross. Although the final map length was longer for the F(2) population, the mapped order of markers was generally the same in the F(2) and RI maps. The data presented in this analysis reconcile much of the inconsistency between map length estimates from chiasma counts and genetic data.

[64]

Truong

S K

, McCormick

R F

, Morishige

D T

, Mullet

J E

. Resolution of genetic map expansion caused by excess heterozygosity in plant recombinant inbred populations
G3: Genes Genom Genet, 2014,4:1963-1969.

[本文引用: 1]

[65]

Ellis

T H

, Turner

, Hellens

R P

, Lee

, Harker

C L

, Enard

, Domoney

, Davies

D R

. Linkage maps in pea
Genetics, 1992,130:649-663.

URLPMID:1551583 [本文引用: 1]

We have analyzed segregation patterns of markers among the late generation progeny of several crosses of pea. From the patterns of association of these markers we have deduced linkage orders. Salient features of these linkages are discussed, as is the relationship between the data presented here and previously published genetic and cytogenetic data.