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刘章雄, 孙石, 李卫东, 陈立军, 常汝镇, 邱丽娟. 1983—2010年北京市大豆育成品种的亲缘关系分析 . 作物学报, 2013, 39(9): 1693-1700[LIU Zhang-Xiong, SUN Shi, LI Wei-Dong, CHEN Li-Jun, CHANG Ru-Zhen, QIU Li-Juan. Analysis of Parental Relationship for Soybean Cultivars Released from 1983 to 2010 in Beijing. Acta Agronomica Sinica, 2013, 39(9): 1693-1700] 复制到剪切板

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1983—2010年北京市大豆育成品种的亲缘关系分析

刘章雄¹, 孙石¹, 李卫东², 陈立军³, 常汝镇¹, 邱丽娟^1,^*

¹农作物基因资源与基因改良国家重大科学工程 / 农业部作物种质资源利用重点开放实验室 / 中国农业科学院作物科学研究所, 北京 100081

²河南省农业科学院经济作物研究所, 河南郑州 450002

³北京市种子管理站, 北京 100088

^* 通讯作者(Corresponding author)：邱丽娟, E-mail:qiulijuan@caas.cn, Tel: 010-82105843

基金:本研究由农业部保种项目(NB2010-2130135-25-05)和国家科技支撑项目(2011BAD35B06-2-9)资助。

摘要

追溯北京市1983—2010年90个大豆品种的系谱信息, 计算其亲本间及品种组合间亲缘系数(coefficient of parentage, COP)并据此聚类, 旨在探讨北京市育成大豆品种的遗传多样性特点。结果表明, 19个品种的父母本间存在亲缘关系, 占品种总数的21.11%; 90个品种间共组成4005对组合, 其中63.62%存在亲缘关系, COP平均值为0.031, 表明大部分品种亲缘关系相对较远。随着时间的推移, 品种间亲缘系数呈下降趋势, 但从2005年至2010年亲缘系数值增大。除12个品种各自独为一类外, 其余78个品种被聚成10类, 反映出不同年代的育种特征和品种演化过程; 中品661等9份种质是对北京市大豆品种培育贡献较大的骨干亲本。北京市品种总体的遗传背景丰富, 但随年代推移, 遗传多样性有逐渐降低趋势。

关键词: 大豆; 亲缘系数; 遗传多样性; 系谱; 演化

Analysis of Parental Relationship for Soybean Cultivars Released from 1983 to 2010 in Beijing

LIU Zhang-Xiong¹, SUN Shi¹, LI Wei-Dong², CHEN Li-Jun³, CHANG Ru-Zhen¹, QIU Li-Juan^1,^*

¹National Key Facility for Gene Resources and Genetic Improvement / Key Laboratory of Crop Germplasm Utilization, Ministry of Agriculture, Institute of Crop Sciences, Chinese Academy of Agricultural Science, Beijing 100081, China

²Institute of Industrial Crops Research, Henan Academy of Agricultural Sciences, Zhengzhou 450002, China

³Beijing Seed Station, Beijing 100088, China

Abstract

To explore the relationship and genetic diversity of soybean cultivars in Beijing and provide theoretical guidance for soybean breeding, we traced the pedigree information of 90 soybean cultivars released from 1983 to 2010 and calculated the coefficients of parentage (COP) between the parents and between cross combinations. The results showed that 19 of 90 cultivars had relationships in their parents. A total of 4005 cross combinations were made among 90 cultivars, and 63.62% of the combinations shared parental relationship, but their average COP value was only 0.031, showing that there existed abundant genetic diversity because there were not close relationship among most cultivars; COP value between cultivars reduced in the process of time, but increased from 2005 to 2010. Cluster analysis suggested that 78 cultivars could be divided into 10 groups, reflecting breeding characteristics in different times and evolution of cultivars. Nine accessions were found to be core parents which contributed more than the other parents. Therefore, the genetic basis of soybean cultivars in Beijing was abundant but genetic diversity became narrower in the process of time.

Keyword: Soybean; Coefficient of parentage; Genetic diversity; Pedigree; Evolution

1923—1995年, 中国利用348个祖先亲本培育出 651个大豆品种, 其祖先亲本仅占大豆品种资源总数的1%^[1]。目前广泛存在利用骨干亲本及其衍生系杂交选育新品种的倾向^[2], 另外, 长期对产量、抗性、品质等性状的连续定向选择, 有可能使品种的遗传基础日益狭窄, 遗传多样性下降^[3]。因此, 对以前育成品种进行遗传背景分析, 从遗传亲缘关系中筛选出可供育种或生产上利用的种质(或性状)越来越重要。

亲缘系数(coefficient of parentage, COP)是衡量个体之间亲缘关系远近或共占某一特定基因概率的指标。COP的计算依赖于个体的系谱信息, 不受环境因素影响, 既可以研究单个品种的遗传信息, 还可以预测品种间的遗传背景丰富程度, 因此, 对于已知系谱信息的自花授粉植物, COP分析是一种简便的研究遗传多样性的方法^[4,5], 并已应用于玉米^[6]、小麦^{[7,8,9,10,11]}、燕麦^[4]、甘蔗^[12,13]、水稻^[14]和大豆^{[15,16,17,18]}等作物。在大豆方面, Sneller^[15]和Helms等^[16]对美国大豆育成品种、Cui等^[17]对中国1923—1995大豆育成品种、Zhou等^[18]对日本1950—1988年86个大豆育成品种进行了COP分析, Helms等^[16]及Zhou等^[18]指出, 亲缘关系较近的亲本杂交后代的遗传变异要小于亲缘关系较远的亲本杂交后代, 在亲本选配时要选择COP值较小的亲本组合。

位于北京市的大豆育种单位主要针对黄淮海地区夏大豆生产需要选育丰产、抗病、优质的大豆品种, 并兼顾北方春大豆生产对品种的需要, 所育成品种遍及黄淮海各省市, 取得了较大的成绩。自1983年第一个大豆品种早熟3号审定以来, 至2010年已育成品种90个, 特别是2001—2010年育成品种58个, 占总数的64.44%, 平均每年审定品种约6个; 广适品种中黄13累计种植面积已达400万公顷, 2012年获国家科技进步一等奖; 高产品种中黄35于2010年在新疆创造了6088.4 kg hm^-2单产最高记录。为总结育种经验, 促进优异品种选育, 本文采用亲缘系数(COP)分析方法, 对1983—2010年北京市育成的90个大豆品种进行遗传多样性分析, 探讨其品种间的遗传关系和品种演化规律, 以为合理利用种质资源和选配亲本、制定育种方案、培育新品种提供理论依据。

1 材料与方法

1.1 试验材料

为了便于比较和分析不同年代育成品种的特点, 将北京市科研单位于1983—2010年育成的90个大豆品种(表1)分为4个时期, 1980—1989年为第1时期, 育成品种10个; 1990—1999年为第2时期, 育成品种18个; 2000—2010年育成品种较多, 分为2个时期, 其中2000—2004年为第3时期, 2005—2010年为第4时期, 分别育成品种36个和26个。

1.2 试验方法

追溯品种的系谱直至祖先亲本(指终极的地方品种或无法再进一步追溯其遗传来源的育成品系或品种, 国外引进种质视为祖先亲本)。系谱资料来源于崔章林等^[1]编著的《中国大豆育成品种系谱分析(1923—1995)》和李卫东主编的《黄淮海大豆种质改良》书稿(尚未正式出版), 2005年以后的系谱资料由育种单位专家提供。

参考Cox等^{[10,17,18,19]}的方法计算90个品种父母本间及成对组合的COP。计算原则如下: (1)一个品种分别从其双亲得到一半的基因; (2)所有祖先种、亲本及其后代品种都是纯合的; (3)最早的祖先品种(系)间的亲缘系数为0; (4)系选品种、自然突变和诱导突变材料与其祖先的亲缘系数为0.75; (5)同一亲本组合的后代两品种间的 COP=(0.75)²; (6)一个品种与其自身的 COP = 1; (7)含有部分相同亲本的旁系品种S和品种D间的亲缘系数 COP_SD＝Σ[(1/2) ⁿ], n代表S品种和D品种的共同亲本与品种S间的世代和品种 D间的世代数之和。用(1-COP)表示系谱遗传距离, 采用SAS9.1软件以欧氏距离的类平均法对COP (90×90)亲缘系数矩阵进行聚类分析。

2 结果与分析

2.1 供试品种父母本间亲缘系数分析

在供试的90个品种中, 7个由单一亲本系选或诱变育成, 64个父母本间的COP值为0, 19个父母本间的COP值为0.001~0.750, 平均值为0.097。在有亲缘关系的19个品种中, 除中黄53 (亲缘系数为0.750)与京豆1号(亲缘系数为0.563)的两亲本亲缘关系较近外, 其他品种亲本间亲缘关系均相对较远, 亲缘系数均小于0.089。可见, 北京市大豆品种除少数品种的父母本间有亲缘关系外, 其余均无亲缘关系, 表明北京市大部分育成品种遗传信息较为丰富。

2.2 品种间的亲缘系数和遗传关系

90份供试品种共组成4005对组合, 其中2548个组合存在亲缘关系, 占组合数的63.62%, 所有组合的COP值变化范围为0~0.781, COP总和为122.855, 平均为0.031, 表明北京市大部分大豆品种间存在亲缘关系, 但其遗传相似程度较低。所有组合中COP值最大的是0.781, 即京豆1号与早熟6号及京豆1号与早熟3号组合, 这是因为早熟3号与早熟6号为姊妹系, 而京豆1号为早熟3号与早熟6号杂交选育后代; COP次大值是0.750, 为中品661与科新7号组合, 这是因为科新7号为中品661化学诱变选育而成; COP第3大值为0.563, 很多姊妹系品种间的COP值为0.563, 如早熟3号与早熟6号等。

2.3 基于亲缘系数的品种聚类分析

对品种间COP值反映的遗传相似性进行聚类分析(图1), 除中黄26、中黄42、中黄24、中黄40、科丰37、豪彩1号、中黄8号、中黄35、中黄57、中黄29、顺豆92-51及中作引1号等12个品种各自单独成一类外, 其他品种共聚成10个类群(A、B、C、D、E、F、G、H、I和J)。从整体来看, 大多具有相同或相似亲本的品种聚在一起, 表明其亲缘关系近, 且大多同一年代的品种聚在一类, 体现出不同年代的育种特征。A群以早熟3号、早熟6号及其姊妹系及其以它们为主要亲本选育的品种组成; B群是以诱变30、诱变31 (组合为58-161×徐豆1号)及其姊妹系以及以它们为主要亲本选育的品种组成; C群是以品系遗-4和遗-2 (组合均为诱变30×Clack63)为主要亲本选育的品种组成; D群是以科新3号及以其为主要亲本选育的品种组成; E群是以早熟18及以其为主要亲本选育的品种组成; F群是以中品661及以其为主要亲本选育的品种组成; G群是以豫豆8号为主要亲本选育的品种组成; H群是以L81-4590的衍生品系Ti15176为主要亲本选育的品种组成; I群是以鲁豆4号为主要亲本选育的品种组成; I群包括2个品种, 中野1号和中野2号, 为姊妹系, 是利用一年生野生大豆为亲本选育而成。

表1 育成品种的来源、亲本组合及其审定时间 Table 1 Origin, parental combination, and released time of soybean cultivars

序号 No.	品种名称 Cultivar name	审定时间 Released time	组合 Combination
1	^※京豆1号 Jingdou 1	1990	早熟3号×早熟6号 Zaoshu 3×Zaoshu 6
2	^※顺豆92-51 Shundou 92-51	2002	墩子黄×红大豆 Dunzihuang×Hongdadou
3	^※豪彩1号 Haocai 1	2004	齐黄26×太空5号 Qihuang 26×Taikong 5
4	^※京黄1号 Jinghuang 1	2004	晋遗20×遗-4 Jinyi 20×Yi-4
5	^※京黄2号Jinghuang 2	2004	豫豆2号×早熟18 Yudou 2×Zaoshu 18
6	^＃早熟3号 Zaoshu 3	1983	科系8号×铁4117 Kexi 8×Tie 4117
7	^＃早熟6号 Zaoshu 6	1983	科系8号×铁4117 Kexi 8×Tie 4117
8	^＃早熟9号 Zaoshu 9	1983	科系8号×6810 Kexi 8×6810
9	^＃早熟15 Zaoshu 15	1983	科系8号×铁4117 Kexi 8×Tie 4117
10	^＃诱变30 Youbian 30	1983	58-161×徐豆1号 58-161×Xudou 1
11	^＃诱变31 Youbian 31	1983	58-161×徐豆1号 58-161×Xudou 1
12	^＃早熟14 Zaoshu 14	1987	科系8号×铁4117 Kexi 8×Tie 4117
13	^＃早熟17 Zaoshu 17	1989	耐阴黑豆×诱变31 Naiyinheidou×Youbian 31
14	^＃科丰6号 Kefeng 6	1989	7611-3-3×诱变30 7611-3-3×Youbian 30
15	^＃早熟18 Zaoshu 18	1992	7902×7821
16	^＃科丰34 Kefeng 34	1993	58-161×徐豆1号 58-161×Xudou 1
17	^＃科丰35 Kefeng 35	1993	7902×诱变16 7902×Youbian 16
18	^＃科新3号 Kexin 3	1995	豫豆2号 EMS诱变 Yudou 2 EMS mutagenesis
19	^＃科丰36 Kexin 36	1999	早熟18×7923 Zaoshu 18×7923
20	科新5号 Kexin 5	2000	鲁豆4号 EMS诱变 Ludou 4 EMS mutagenesis
21	^＃科丰14 Kefeng 14	2001	铁丰18×7913 Tiefeng 18×7913
22	^＃科丰53 Kefeng 53	2001	7409×诱变30 7409×Youbian 30
23	^＃科新6号 Kexin 6	2001	豫豆2号 EMS诱变Yudou 2 EMS mutagenesis
24	^＃科丰15 Kefeng 15	2002	8511×吉林20 8511×Jilin 20
25	^＃科丰37 Kefeng 37	2002	8033×密荚1号 8033×Mijia 1
26	^＃科新4号Kexin 4	2002	豫豆2号 EMS诱变Yudou 2 EMS mutagenesis
27	^＃科新7号 Kexin 7	2003	中品661 EMS和EI诱变 Zhongpin 661 EMS and EI mutagenesis
28	^＃科新8号 Kexin 8	2003	科9103 EMS诱变 Ke 9103 EMS EI mutagenesis
29	^＃科丰17 Kefeng 17	2004	8511×铁杂 8511×Tieza
30	中豆28 Zhongdou 28	1999	鲁豆4号×L83-4387 Ludou 4×L83-4387
31	中豆27 Zhongdou 27	2000	中豆19×L81-4590 Zhongdou 19×L81-4590
32	中黄1号Zhonghuang 1	1989	早熟6号×晋豆4号 Zaoshu 6×Jindou 4
33	中黄2号Zhonghuang 2	1990	7907-2-16 系选 7907-2-16 systemic selection
34	中黄3号Zhonghuang 3	1990	诱变30×7614 Youbian 30×7614
35	中黄4号Zhonghuang 4	1990	遗112×上海红芒早毛豆Yi 112×Shanghaihongmangzaomaodou
36	中黄5号Zhonghuang 5	1992	诱变30×7614 Youbian 30×7614
37	中黄7号Zhonghuang 7	1993	科丰6号×菏7308-1-2 Kefeng 6×He 7308-1-2
38	中黄6号Zhonghuang 6	1994	早熟6号×晋豆4号 Zaoshu 6×Jindou 4
39	中品661 Zhongpin 661	1994	William82⁶×Bufallo
40	中黄8号Zhonghuang 8	1995	大金元×兖黄1号 Dajinyuan×Yanhuang 1
41	中黄9号Zhonghuang 9	1996	科丰6号×菏7308-1-2 Kefeng 6×He 7308-1-2
42	中黄10号Zhonghuang 10	1996	文丰7号×鲁豆4号 Wenfeng 7×Ludou 4
43	中野1号Zhongye 1	1999	(察隅1号×野大豆ZYD3576)F₁×大湾大粒 (Cayu 1×Yedadou ZYD3576)F₁×Dawandali
44	中黄11 Zhonghuang 11	2000	Crowford×豫豆8号Crowford×Yudou 8
45	中黄12 Zhonghuang 12	2000	晋遗20×遗-4 Jinyi 20×Yi-4
46	中黄13 Zhonghuang 13	2001	豫豆8号×中作90052-76 Yudou 8×Zhongzuo 90052-76
47	中黄14 Zhonghuang 14	2001	Crowford×豫豆8号 Crowford×Yudou 8
48	中黄15 Zhonghuang 15	2001	中品661×早熟17 Zhongpin 661×Zaoshu 17

(续表1)

表1 育成品种的来源、亲本组合及其审定时间Table 1 Origin, parental combination, and released time of soybean cultivars

续表1 育成品种的来源、亲本组合及其审定时间 Table 1 Origin, parental combination, and released time of soybean cultivars

序号 No.	品种名称 Cultivar name	审定时间 Released time	组合 Combination
49	中黄17 Zhonghuang 17	2001	遗-2×Hobbit Yi-2×Hobbit
50	中黄18 Zhonghuang 18	2001	中品661×Century-2 Zhongpin 661×Century-2
51	中黄20 Zhonghuang 20	2001	遗-2 Yi-2×Hobbit
52	中野2号Zhongye 2	2001	(察隅1号×野大豆ZYD3576)F₁×大湾大粒 (Cayu 1×Yedadou ZYD3576)F₁×Dawandali
53	中品662 Zhongpin 662	2002	早5粒×鲁豆4号 Zao 5 li×Ludou 4
54	中黄16 Zhonghuang 16	2002	ti15176×Century-2.3
55	中黄22 Zhonghuang 22	2002	中品661×91-1 Zhongpin 661×91-1
56	中黄23 Zhonghuang 23	2002	杂抗F₆×鲁豆4号 Zakang F₆×Ludou 4
57	中作引1号Zhongzuoyin 1	2003	Fabio中系选 Fabio systemic selection
58	中黄19 Zhonghuang 19	2003	中品661×豫豆10号 Zhongpin 661×Yudou 10
59	中黄21 Zhonghuang 21	2003	中品661×91-1 Zhongpin 661×91-1
60	中黄24 Zhonghuang 24	2003	吉林21×(汾豆31×中豆19)F₁ Jilin 21×(Fendou 31×Zhongdou 19)F₁
61	中黄25 Zhonghuang 25	2003	中黄4号×诱变30 Zhonghuang 4×Youbian 30
62	中黄26 Zhonghuang 26	2003	单8×PI437654 Dan 8×PI437654
63	中黄27 Zhonghuang 27	2003	晋遗20×中作90052-76 Jinyi 20×Zhongzuo 90052-76
64	中黄28 Zhonghuang 28	2003	ti15176×Century-2.3
65	中黄29 Zhonghuang 29	2005	鲁豆861168×鲁豆11 Ludou 861168×Ludou 11
66	中黄31 Zhonghuang 31	2005	ti15176×Century-2.3
67	中黄33 Zhonghuang 33	2005	豫豆8号×晋遗20 Yudou 8×Jinyi 20
68	中黄30 Zhonghuang 30	2006	中品661×中黄14 Zhongpin 661×Zhonghuang 14
69	中黄34 Zhonghuang 34	2006	晋遗20×遗-4 Jinyi 20×Yi-4
70	中黄35 Zhonghuang 35	2006	(PI486355×郑8431)F₃×郑6062 (PI486355×Zheng 8431)F₃×Zheng 6062
71	中黄36 Zhonghuang 36	2006	遗-2×Hobbit Yi-2×Hobbit
72	中黄37 Zhonghuang 37	2006	95B020×早熟18 95B020×Zaoshu 18
73	中黄38 Zhonghuang 38	2006	遗-2×Hobbit Yi-2×Hobbit
74	中黄39 Zhonghuang 39	2006	中品661×中黄14 Zhongpin 661×Zhonghuang 14
75	中黄40 Zhonghuang 40	2007	晋豆6号×豫豆12 Jindou 6×Yudou 12
76	中黄41 Zhonghuang 41	2007	科丰14×科新3号 Kefeng 14×Kexin 3
77	中黄42 Zhonghuang 42	2007	诱处4号×锦豆33 Youchu 4×Jindou 33
78	中黄43 Zhonghuang 43	2008	冀豆7号×科新3号 Jidou 7×Kexin 3
79	中黄44 Zhonghuang 44	2009	科丰14×科新3号 Kefeng 14×Kexin 3
80	中黄45 Zhonghuang 45	2009	中黄21×WI995 Zhonghuang 21×WI995
81	中黄46 Zhonghuang 46	2009	ti15176×Century-2.3
82	中黄47 Zhonghuang 47	2009	D90×L81-4590
83	中黄48 Zhonghuang 48	2009	科丰14×科新3号 Kefeng 14×Kexin 3
84	中黄49 Zhonghuang 49	2009	科丰14×科新3号 Kefeng 14×Kexin 3
85	中黄50 Zhonghuang 50	2010	中黄13×中品661 Zhonghuang 13×Zhongpin 661
86	中黄52 Zhonghuang 52	2010	冀豆7号×早熟18 Jidou 7×Zaoshu 18
87	中黄53 Zhonghuang 53	2010	中作M17×(豫豆8号×D90) Zhongzuo M17×(Yudou 8×D90)
88	中黄55 Zhonghuang 55	2010	T200×早熟18 T200×Zaoshu 18
89	中黄56 Zhonghuang 56	2010	中品95-6051×DP3480 Zhongpin 95-6051×DP3480
90	中黄57 Zhonghuang 57	2010	Hartwig×晋1265 Hartwig×Jin 1265

此处审定时间表示品种的第一次审定时间, 因少数品种在不同时间内通过多个地区审定; 品种名称前标^※代表选育单位为其他单位, 标^＃为原中国科学院遗传所和现遗传与发育研究所, 未标注的为中国农业科学院。

The released time here is the first released time of soybean cultivars, among which a few cultivars are released by different districts in different years.^※ marked before the name of cultivars stands for that bred by other units, and^＃stands for that bred by former Institute of Genetics and Institute of Genetics and Development Biology of the Chinese Academy of Sciences; the others come from the Chinese Academy of Agricultural Sciences.

续表1 育成品种的来源、亲本组合及其审定时间Table 1 Origin, parental combination, and released time of soybean cultivars

图1 基于亲缘系数的品种聚类左侧“数字”同表1的品种序号, 如“12”代表早熟14。Fig. 1 Cultivars clustering on the basis of relationship coefficient Numbers on the left side correspond with the numbers of cultivar names given in Table 1. For example, “12” is Zaoshu 14.

从聚类图上可以分析北京市品种演化的过程, 20世纪80年代育成品种主要为原中国科学院遗传所选育的早熟3号、早熟6号及其姊妹系、诱变30、诱变31等, 部分品种如早熟3号和诱变30成为20世纪90年代的骨干亲本。20世纪90年代上半时期, 分别以早熟3号及诱变30为主要亲本育成了科新8号、中黄6号、中黄25、科丰35等品种, 这些品种主要包含在A群和B群; 以国外引进种质Clack 63为亲本育成的品系遗-4和遗-2作为主要亲本育成了中黄12、中黄34、中黄36、中黄38等, 这些品种包含在C群; 20世纪90年代后半时期, 以科新3号为主要亲本育成了中黄41、中黄44、中黄48、中黄49等, 这些品种包含在D群内。21世纪的第一个10年, 国外种质L81-4590和中品661 (L83-544)的应用, 育成了中黄30、中黄39、中黄16、中黄46等多个品种, 分别属于F群和H群。

2.4 各年代品种间的亲缘系数和遗传关系

由表2可知第1时期育成的10个品种中, 所有祖先亲本均含有滨海大白花、铜山天鹅蛋和Mamotan, 亲缘关系比率为100%。第2时期和第3时期育成品种存在遗传关系的比率分别为48.37%和49.68%, 因2个时期所用亲本的差异, 使品种间存在遗传关系的比率降低。第4时期因大量使用了中品661及前期育成品种或品系为亲本(表1), 使品种的遗传关系比率升高, 为80.92%。

第1时期与后3个时期的品种亲缘关系比率呈上升趋势, 分别为66.67%、65.83%、83.76%。第2时期与后2个时期存在遗传关系的比率分别为57.78%、70.09%, 比率值也在增加, 说明随着育成品种的增多和育种规模的扩大, 利用前期育成品种或品系作为亲本的趋势在增大, 这与王江春等^[10]以小麦为材料得出的结果类似。

亲缘系数值的大小反映了品种间遗传相似的程度。随着时间的推移, 不同年代内品种的亲缘系数值呈降低的趋势(表3), 如在第1时期、第2时期和第3时期, COP值分别为0.192、0.040和0.026, 但在第4时期COP值又增大为0.044。这说明, 在2005年之前, 随着时间推移, 北京大豆品种的遗传多样性逐渐升高, 但在2005年之后, 遗传多样性开始降低, 其原因可从不同时期内及不同时期间品种间存在遗传关系的比率得到解释。

从不同年代间品种的亲缘系数分析结果可以看出, 同一年代内的品种, 对后代的遗传贡献率随着年代的推移逐渐减小, 如第1时期品种与第2时期、第3时期和第4时期品种的亲缘系数值依次为0.087、0.026和0.014, 这与Sneller ^[15]和王江春等^[10]研究的祖先品种对育成品种的贡献率随时间的推移而下降的结果一致。

2.5 北京市育成品种骨干亲本分析

从品种聚类分析及北京市品种演化过程可见, 诱变30、早熟3号、中品661、鲁豆４号、诱变31、科新3号和早熟18及从美国引进种质Clack 63、L81-4590等9个品种对北京市育成品种贡献较大。分析表明, 上述品种衍生品种数范围为5%~23%, 参与育成品种数占育成品种总数67.78% (61/90), 总核贡献率占32.50% (29.25/90); 与其他品种的COP值总和变化范围为2.000~6.229, 除L81-4590外, 其余均大于所有单一品种与其他品种COP值总和的平均值2.730 (表4)。诱变30作为亲本参与选育品种23个, 占所有育成品种25.56%, 诱变30与其他89个品种平均COP值为0.070。以上结果显示, 诱变30等9个品种在北京市育种中利用效果显著, 发挥了骨干亲本的作用。

表2 不同年代内和年代间亲缘关系比率Table 2 Ratio of genetic relationship within and between different periods (%)

表3 不同年代内和年代间品种的平均COP值Table 3 Average variety COP within and between different periods

表4 北京市大豆育成品种骨干亲本及衍生的品种数Table 4 Backbone parents and their offsprings of soybean cultivars released in Beijing

3 讨论

3.1 北京市大豆品种的遗传多样性

Cui等^[17]于2000年对1923—1995年中国651个大豆育成品种的COP分析表明, 22%的品种组合间有亲缘关系, 平均COP值较低, 仅为0.020, 说明此阶段大豆育成品种具有较高的遗传多样性及改良潜力。其中, 330个来源于东北的品种平均COP值为0.058, 210个黄淮海品种平均COP值为0.027, 111个南方品种平均COP值为0.016, 这三大区品种间COP值较低, 说明其品种间几乎没有亲缘关系。这与不同生态区之间很少有近交品种以及国外种质的引进和利用有关。

Zhou等^[18] 于2002年对日本1950—1988年86个大豆育成品种的COP分析表明, 约20%的品种组合间有亲缘关系, 平均COP值也较低, 为0.04, 可能有几个原因, 一是较多的利用来自本土的地方品种, 二是引进和利用国外种质, 三是限制种质在国内的交流。

本研究分析的90个供试品种, 组成了4005对组合, 63.62%组合存在亲缘关系, 表明大部分品种间存在亲缘关系, 此比率均大于Cui等^[17]和Zhou等^[18]的研究结果, 这与品种的育成单位较少, 亲本来源相对比较集中有关。北京市大豆品种平均COP值为0.031, 大于Cui等^[17]的研究结果而小于Zhou等^[18]的研究结果, 说明北京市大豆品种的遗传多样相对较高, 其原因之一是引进和利用了中品661、Clark 63, Williams、Mamotan、Amsoy、Century-2.3等国外种质; 另外, 北京市90个大豆品中, 63个品种父母本间的COP值为0, 说明育种家在育种实践中, 有意识地选择亲缘关系较远的亲本进行杂交, 从而提高了大豆品种的遗传多样性。

大豆^[17]、小麦^[9,10,11]品种根据亲缘系数进行聚类, 其结果能很好地反映种质资源的利用情况及遗传多样性程度, 来自同一祖先种或亲缘关系较近的品种大多聚为一类。本研究结果也反映了上述特点, 除12个品种单独自成一类外, 其他品种根据其主要亲本来源分别聚成10类, 同时, 大部分的主要亲本也构成了北京育成品种的骨干亲本。与亲缘关系较近的亲本组配相比, 亲缘关系较远两亲杂交产生的后代, 遗传变异更大^[16], 因此, 可利用亲缘关系聚类图选择合适的亲本, 一般而言, 不同类别的品种杂交更易选育出优良品种^[17,18]。建议根据北京大豆品种聚类特点, 选择分别聚在不同类的亲本进行杂交, 并评价后代产量性状表现, 以检验利用COP值选配亲本在育种实践中的成效。

从本文的研究方法可看出, COP亲缘系数分析受品种系谱的清晰程度、原始亲本的亲缘关系等影响(如原始亲本间可能有亲缘关系, 但目前无法查证), 利用分子标记进行遗传分析则可不受上述因素的干扰, 已有许多研究者利用SSR等标记对大豆育成品种的遗传多样性进行了比较研究^[20,21,22]。目前, 尚无利用分子标记研究北京市育成品种遗传关系的报道, 因此, 将COP分析与分子标记相结合, 可能会更好的揭示北京育成品种的遗传关系。

3.2 骨干亲本的利用与研究

骨干亲本的利用加速了新品种培育。一般骨干亲本农艺性状优异, 具有较好配合力, 合理地利用可提高育种效率。如中品661具有植株高大健壮、透光性好、丰产、抗倒、高抗大豆花叶病毒病、适应性广等特点。从2001—2010年, 中国农业科学院作物科学研究所利用该品种作亲本共选育品种12个, 其中10个以中品661为亲本直接杂交选育, 3个品种通过国家审定, 2个品种同时通过国家和天津审定, 4个品种通过北京审定, 1个品种通过辽宁审定, 2个品种通过河北审定, 其亲本中品661同时通过北京和云南审定, 中品661衍生的品种中黄30现正被西北地区大面积示范推广, 尤其适合于间作套种。黄淮海大面积推广品种中黄13, 其系谱中含有2个骨干亲本诱变30和Clack 63, 其直接亲本豫豆8号来源于河南省的优异种质, 中黄13则集中了3个优异种质的优点, 所以高产而适应性广。

组合配制中高频率使用骨干亲本会降低品种的遗传多样性, 所以应挖掘地方品种潜力, 加强引进和利用国外优良种质。研究表明, 国外种质的利用可以扩大育成品种的遗传基础^[17,18]。从北京市大豆品种的演化中可以看出, 国外种质如L81-4590、Clack 63等在北京市大豆育种中发挥了骨干亲本的作用。另一方面, 骨干亲本的广泛利用, 也会降低大豆品种的遗传多样性^[23], 因此, 今后大豆育种工作不仅要关注现有的育成品种和创新种质, 还要广泛挖掘地方品种潜力, 积极引进和利用国外种质, 拓宽现有大豆育成品种的遗传基础, 以加快优良新品种的培育。

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1998

0.0

... 1923—1995年, 中国利用348个祖先亲本培育出 651个大豆品种, 其祖先亲本仅占大豆品种资源总数的1%^[1] ...

... 系谱资料来源于崔章林等^[1]编著的《中国大豆育成品种系谱分析(1923—1995)》和李卫东主编的《黄淮海大豆种质改良》书稿(尚未正式出版), 2005年以后的系谱资料由育种单位专家提供 ...

1998

0.0

... 目前广泛存在利用骨干亲本及其衍生系杂交选育新品种的倾向^[2], 另外, 长期对产量、抗性、品质等性状的连续定向选择, 有可能使品种的遗传基础日益狭窄, 遗传多样性下降^[3] ...

1972

0.0

1989

0.0

... COP的计算依赖于个体的系谱信息, 不受环境因素影响, 既可以研究单个品种的遗传信息, 还可以预测品种间的遗传背景丰富程度, 因此, 对于已知系谱信息的自花授粉植物, COP分析是一种简便的研究遗传多样性的方法^[4,5], 并已应用于玉米^[6]、小麦^{[7,8,9,10,11]}、燕麦^[4]、甘蔗^[12,13]、水稻^[14]和大豆^{[15,16,17,18]}等作物 ...

1996

0.0

1990

0.0

2000

3.297

0.0

THEOR APPL GENET. 2000, 100(2):242 - 248 DOI:10.1007/s001220050032

Application of microsatellites in wheat (Triticum aestivum L.) for studying genetic differentiation caused by selection for adaptation and use

M. Stachel(1) T. Lelley(1) H. Grausgruber(2) J. Vollmann(2)

1.Institute for Agrobiotechnology, Department of Plant Biotechnology, Konrad Lorenz Strasse 20, A-3430 Tulln, Austria e-mail: lelley@ifa-tulln.ac.at AT
2.University of Agricultural Sciences Vienna, Department of Plant Breeding, Gregor Mendel Strasse 33, A-1180 Vienna, Austria AT

For studying genetic differentiation caused by selection for adaptation and end-use, the allele frequencies of 42 microsatellites (MS), representative of the three wheat genomes, were analysed in a total of 60 wheat cultivars. The cultivars originate from three agroecological areas (AEAs) – Germany, Austria and Hungary – and represent equal numbers of ’quality wheats’ and ’feed wheats’ for each country. For the 42 loci, 202 alleles were detected using PAGE and silver staining. The average number of alleles per locus was 4.8, including four monomorphic loci. For 16 loci, null alleles were detected. Cluster analysis clearly differentiated the varieties according to the three AEAs and, within each AEA, into quality wheats from feed wheats. Analysis of variance revealed highly significant differences of distance data between AEAs as well as between quality groups. The correlation between genetic distance (GD) and pedigree data (coefficient of diversity, COD) wasr_s=0.45. The results have proven the excellent resolving power of MS in varietal differentiation, which arises through breeding under specific environmental conditions, and for different end-use.

2002

1.554

0.0

EUPHYTICA. 2002, 123(2):273 - 285 DOI:10.1023/A:1014946018765

Genetic diversity in bread wheat, as revealed by coefficient of parentage and molecular markers, and its relationship to hybrid performance

1.Istituto Sperimentale per la Cerealicoltura, Sezione di S. Angelo Lodigiano

Variable results have been obtained in different crop species using geneticsimilarity (GS) estimates based on molecular markers and coefficient ofparentage (COP) to predict heterosis.This study was designed: i) to assess the level of GS among 40 breadwheat (Triticum aestivum L.) cultivars selected in Central and SouthEurope; ii) to compare GS with COP; iii) to correlate the estimates ofparental diversity with the heterotic effects detected on 149 F₁hybrids obtained by crossing the 40 cultivars according to three matingdesigns.The parental cultivars and the F₁ hybrids were grown in severallocations at normal seed density. Significant heterotic effects were detectedfor grain yield and other traits including quality attributes. The parentalcultivars were assayed for DNA polymorphisms using two classes ofmarkers: 338 RFLP and 200 AFLP^® bands were scored. GS estimates werecomputed considering each molecular marker set separately (GS_RFLP,GS_AFLP) and together (GS_TOT). Ample differentiation amongthe parental cultivars was detected with the two marker sets. Although theaverage GS_TOT (0.43) was higher than COP (0.10), the twomeasurements were significantly correlated (r = 0.36,p< 0.01).correlations="" between="" the="" different="" estimates="" of="" genetic="" diversity="">₁ performance or mid parent heterosis for grain yield and otherrelated traits were in general low although statistically significant.A more detailed analysis was conducted on 28 F₁ hybridsproduced in a half diallel cross of eight parental cultivars characterized byhigh heterotic effects for grain yield. The GS estimates based on RFLP,AFLP^® markers and also on RAPD were partitioned into general andspecific components. Correlations with general and specific combiningability effects for the measured traits were in several cases statisticallysignificant but too low to be predictive and therefore exploitable in practicalbreeding.

2002

0.0

... 大豆^[17]、小麦^[9,10,11]品种根据亲缘系数进行聚类, 其结果能很好地反映种质资源的利用情况及遗传多样性程度, 来自同一祖先种或亲缘关系较近的品种大多聚为一类 ...

2006

0.0

1.4522

Scientia Agricultura Sinica. 2006, 39(4):664 - 672

Relationship Coefficient Analysis Among Winter Wheat Varieties and Their Parents in Shandong Province after Liberation

建国以来山东省小麦品种及其亲本的亲缘系数分析

,,,,,

王江春,胡延吉,余松烈,王振林,刘爱峰,王洪刚

山东农业大学农学院

【Objective】 This paper is to probe the relationship and genetic diversity of the winter wheat varieties that selected from liberation to now in Shandong Province. 【Method】 The coefficient parentage is used to analyze the 2145 combinations of 66 winter wheat varieties in nine varieties exchanging. 【Result】 It showed that the relationship exists among 67.44% combinations, the range of COP value of all varieties is 0.00-0.8203 with the average value 0.06249, 21.53% of pedigrees Relationship Coefficients (RC) are higher than the average. The RC of varieties of every year became bigger along with the time, fell in the 90's, and then increased again. The varieties RCs in each year were bigger than that between years, less that that between early and present varieties, while became larger between medium-term and latter varieties. The variety genetic contributions of the same age decreases with the time. The result of clustering analysis on varieties COP values on the basis of the WAP method showed that 63 varieties can be united into 6 big clusters based on different years, and the left three varieties, Xuzhou438, Qida195, Yannong 78 were made into a cluster at the value of 0. The RCs of the 17 varieties which have been extended many area for a long time in Shandong Province were little, with bigger genetic difference, and all those varieties were selected from overseas parents' combinations. The breeding genetic resources of Shandong Province can be traced back to four big core parents system of Biyu, Villa Glori (Zhongnong 28), Luofulin and Youzi wheat on the ground of pedigree analysis. 【Conclusion】 The genetic differences and diversity between varieties were little by little after 1950's.

【目的】探讨山东省小麦品种的遗传多样性。【方法】对山东省建国以来历经9次更换的66个小麦品种的2145个组合进行了亲缘系数(coefficient of parentage, COP)分析。【结果】67.44％品种组合间存在亲缘关系，所有品种COP值变异范围为0.00～0.8203，平均为 0.06249，系谱亲缘系数高于平均值的占21.53％。随着时间的推移，各个年代之内品种间亲缘系数有逐渐增大的趋势，到90年代有所降低，然后又再增大。各年代内品种的亲缘系数普遍高于年代间品种的亲缘系数。早期的品种与以后年代的品种间的亲缘系数较小，中期和近期品种与以后年代的品种间的亲缘系数较大。同一个年代的品种，对后代的遗传贡献率，则随着年代的推移而逐渐减小。对品种间COP值按WAP法进行了聚类分析，除徐州438、齐大195、烟农78在COP为0处单独聚在一起外，其余63个品种可以聚为6个大类群，大多是同一年代的品种聚在一起，体现了不同年代的特点。山东省历史上年种植面积较大或推广时间较长的17个小麦品种，大都是较早利用引进亲本进行组配选育而成，亲本的亲缘系数较小，遗传差异较大。从系谱上看，山东省小麦的种质资源可基本上追溯到碧玉麦、Villa Glori(中农28)、洛夫林系、蚰子麦等四大骨干亲本体系。【结论】自50年代以来，品种间的遗传差异日趋狭窄，遗传多样性逐渐降低。

... 参考Cox等^{[10,17,18,19]}的方法计算90个品种父母本间及成对组合的COP ...

... 09%, 比率值也在增加, 说明随着育成品种的增多和育种规模的扩大, 利用前期育成品种或品系作为亲本的趋势在增大, 这与王江春等^[10]以小麦为材料得出的结果类似 ...

... 014, 这与Sneller^[15]和王江春等^[10]研究的祖先品种对育成品种的贡献率随时间的推移而下降的结果一致 ...

2011

0.0

2002

3.297

0.0

THEOR APPL GENET. 2002, 104(1):30 - 38 DOI:10.1007/s001220200003

Analysis of genetic similarity detected by AFLP and coefficient of parentage among genotypes of sugar cane (Saccharum spp.)

M. L. A. Lima(1) A. A. F. Garcia(3) K. M. Oliveira(1) S. Matsuoka(4) H. Arizono(3) C. L. de Souza Jr.(5) A. P. de Souza(1)

1.Centro de Biologia Molecular e Engenharia Genética (CBMEG), Universidade Estadual de Campinas (UNICAMP), Cidade Universitária ”Zeferino Vaz”, CP 6010, CEP 13083-970, Campinas, SP, Brasil e-mail: anete@unicamp.br Tel.: (55-19) 3788-1132 or 3788-1130, Fax: (55-19) 3788-1089, Present address: A.P. de Souza, Centro de Biologia Molecular e Engenharia Genética (CBMEG), Universidade Estadual de Campinas (UNICAMP), Cidade Universitária ”Zeferino Vaz”, CP 6010, CEP 13083-970, Campinas, SP, Brasil BR
2.Departamento Genética e Evolu??o – Instituto de Biologia – Universidade Estadual de Campinas (UNICAMP), Cidade Universitária ”Zeferino Vaz”, CP 6109, CEP 13083-970, Campinas, SP, Brasil BR
3.Departamento de Ciências Exatas, Escola Superior de Agricultura ”Luiz de Queiroz” (ESALQ), Universidade de S?o Paulo (USP), CP 9, CEP 13400-970, Piracicaba, SP, Brasil BR
4.Departamento de Biotecnologia Vegetal (DBV), Centro de Ciências Agrárias (CCA), Universidade Federal de S?o Carlos (UFSCar), CEP 13600-970, Araras, SP, Brasil BR
5.Departamento de Genética, Escola Superior de Agricultura ”Luiz de Queiroz” (ESALQ), Universidade de S?o Paulo (USP), CP 83, CEP 13400-970, Piracicaba, SP, Brasil BR

Despite the economical importance of sugar cane, until the present-date no studies have been carried out to determine the correlation of the molecular-based genetic similarity (GS) and the coefficient of parentage (f)-estimates generated for cultivars. A comprehensive knowledge of the amount of genetic diversity in parental cultivars, could improve the effectiveness of breeding programmes. In this study, amplified fragment length polymorphism (AFLP) and pedigree data were used to investigate the genetic relationship in a group of 79 cultivars (interspecific hybrids), used as parents in one of the Brazilian breeding programmes, and four species ofSaccharum (Saccharum sinense,Saccharum barberi and two ofSaccharum officinarum) . The objectives of this study were to assess the level of genetic similarity among the sugar-cane cultivars and to investigate the correlation between the AFLP-based GSandf, based on pedigree information. Twenty one primer combinations were used to obtain the AFLP molecular markers, generating a total of 2,331 bands, of which 1,121 were polymorphic, with a polymorphism rate, on average, of 50% per primer combination. GSs were determined using Jaccard’s similarity coefficient, and a final dendrogram was constructed using an unweighted pair-group method using arithmetic average (UPGMA). AFLP-based GS ranged from 0.28 to 0.89, with a mean of 0.47, whereasf ranged from 0 to 0.503, with a mean of 0.057. Cluster analysis using GS divided the genotypes into related subgroups suggesting that there is important genetic relationship among the cultivars. AFLP-based GS andf were significantly correlated (r= 0.42,P< 0.001), thus the significance of thisr value suggests that the AFLP data may help to more-accurately quantify the degree of relationship among sugar-cane cultivars.

2010

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Sugar Tech. , :

2006

0.0

1994

0.0

... 在大豆方面, Sneller^[15]和Helms等^[16]对美国大豆育成品种、Cui等^[17]对中国1923—1995大豆育成品种、Zhou等^[18]对日本1950—1988年86个大豆育成品种进行了COP分析, Helms等^[16]及Zhou等^[18]指出, 亲缘关系较近的亲本杂交后代的遗传变异要小于亲缘关系较远的亲本杂交后代, 在亲本选配时要选择COP值较小的亲本组合 ...

... 014, 这与Sneller^[15]和王江春等^[10]研究的祖先品种对育成品种的贡献率随时间的推移而下降的结果一致 ...

1997

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... 与亲缘关系较近的亲本组配相比, 亲缘关系较远两亲杂交产生的后代, 遗传变异更大^[16], 因此, 可利用亲缘关系聚类图选择合适的亲本, 一般而言, 不同类别的品种杂交更易选育出优良品种^[17,18] ...

2000

0.0

... 参考Cox等^{[10,17,18,19]}的方法计算90个品种父母本间及成对组合的COP ...

... 1 北京市大豆品种的遗传多样性Cui等^[17]于2000年对1923—1995年中国651个大豆育成品种的COP分析表明, 22%的品种组合间有亲缘关系, 平均COP值较低, 仅为0 ...

... 62%组合存在亲缘关系, 表明大部分品种间存在亲缘关系, 此比率均大于Cui等^[17]和Zhou等^[18]的研究结果, 这与品种的育成单位较少, 亲本来源相对比较集中有关 ...

... 031, 大于Cui等^[17]的研究结果而小于Zhou等^[18]的研究结果, 说明北京市大豆品种的遗传多样相对较高, 其原因之一是引进和利用了中品661、Clark 63, Williams、Mamotan、Amsoy、Century-2 ...

... 研究表明, 国外种质的利用可以扩大育成品种的遗传基础^[17,18] ...

2002

0.0

... 参考Cox等^{[10,17,18,19]}的方法计算90个品种父母本间及成对组合的COP ...

... Zhou等^[18] 于2002年对日本1950—1988年86个大豆育成品种的COP分析表明, 约20%的品种组合间有亲缘关系, 平均COP值也较低, 为0 ...

... 研究表明, 国外种质的利用可以扩大育成品种的遗传基础^[17,18] ...

1985

0.0

... 参考Cox等^{[10,17,18,19]}的方法计算90个品种父母本间及成对组合的COP ...

2000

0.0

... 从本文的研究方法可看出, COP亲缘系数分析受品种系谱的清晰程度、原始亲本的亲缘关系等影响(如原始亲本间可能有亲缘关系, 但目前无法查证), 利用分子标记进行遗传分析则可不受上述因素的干扰, 已有许多研究者利用SSR等标记对大豆育成品种的遗传多样性进行了比较研究^[20,21,22] ...

2008

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2007

0.0

2000

0.0

... 另一方面, 骨干亲本的广泛利用, 也会降低大豆品种的遗传多样性^[23], 因此, 今后大豆育种工作不仅要关注现有的育成品种和创新种质, 还要广泛挖掘地方品种潜力, 积极引进和利用国外种质, 拓宽现有大豆育成品种的遗传基础, 以加快优良新品种的培育 ...