生物多样性 ›› 2008, Vol. 16 ›› Issue (1): 53-62. DOI: 10.3724/SP.J.1003.2008.07050
王建民1,2, 岳文斌1,*
收稿日期:
2007-02-15
接受日期:
2007-07-30
出版日期:
2008-01-27
发布日期:
2008-01-27
通讯作者:
岳文斌
作者简介:
*E-mail:wb-yue@126.com基金资助:
Jianmin Wang1,2, Wenbin Yue1,*
Received:
2007-02-15
Accepted:
2007-07-30
Online:
2008-01-27
Published:
2008-01-27
Contact:
Wenbin Yue
摘要:
为了探讨家养绵羊与山羊的属间遗传关系, 我们利用13个定位于绵羊染色体上的微卫星基因座, 分析了黄河下游4个地方绵羊品种、4个地方山羊品种和1个杂交绵羊类群的遗传结构及其系统发生关系。经Hardy-Weinberg平衡检验和中性测试, 发现地方绵羊和山羊种群均处于不平衡状态(P< 0.01), 61.53%的基因座属于中性位点, 说明所研究种群属于非随机交配, 可能受到选择、迁移等进化因素的影响。对有效等位基因数、多态信息含量、Shannon信息指数、观察杂合度和期望杂合度等遗传多样性参数进行比较, 发现绵羊种群的遗传变异程度明显(P<0.01或P<0.05)高于山羊种群, 但不同基因座上的差异效应不一致; 结合F统计量和亲缘关系等参数, 可推测绵羊和山羊虽然均存在不同程度的近交现象(He>Ho, FIS>0),但分别属于杂交和近交繁殖。通过群体遗传分化和系统发育拓扑结构分析, 证明绵羊属由共同祖先分化而来的时间晚于山羊属, 两属间的遗传距离为1.0708-1.5927, 遗传分化时间约为19,807-28,955年; 绵羊属内品种间的遗传分化程度(FST<0.05)均低于山羊属内品种间的分化(FST>0.15)。本研究揭示了人工选择对同域家养绵羊与山羊交配系统的形成及群体遗传分化具有深刻的影响。
王建民, 岳文斌 (2008) 黄河下游家绵羊与家山羊遗传关系的微卫星分析. 生物多样性, 16, 53-62. DOI: 10.3724/SP.J.1003.2008.07050.
Jianmin Wang, Wenbin Yue (2008) Genetic relationships of domestic sheep and goats in the lower reaches of the Yellow River based on microsatellite analysis. Biodiversity Science, 16, 53-62. DOI: 10.3724/SP.J.1003.2008.07050.
品种名称 Breeds | 采样地点 Sampling location | 采样只数(羊群数) Sample size (flock number) |
---|---|---|
绵羊品种 Native or crossbred sheep breeds | ||
小尾寒羊 Small-tailed Han (STH) | 山东嘉祥县 Jiaxiang County | 42 (14) |
大尾寒羊 Large-tailed Han (LTH) | 山东临清市 Linqing City | 42 (22) |
山地绵羊 Shandi (SD) | 山东莱芜市 Laiwu City | 29 (17) |
洼地绵羊 Wadi (WD) | 山东滨州市 Binzhou City | 40 (18) |
杂交绵羊 Crossbred (CB) | 山东莱芜市 Laiwu City | 20 (1) |
山羊品种 Native goat breeds | ||
济宁青山羊 Jining grey (JNG) | 山东嘉祥县 Jiaxiang County | 40 (5) |
鲁北白山羊 Lubei white (LBW) | 山东无棣县 Wudi County | 40 (23) |
莱芜黑山羊 Laiwu black (LWB) | 山东莱芜市 Laiwu City | 40 (26) |
崂山奶山羊 Laoshan milk (LSM) | 山东临朐县 Linqu County | 40 (21) |
表1 所研究山东省羊品种的名称、采样地点和样本数量
Table 1 Names, sampling localities and sample sizes of studied breeds from Shandong Province
品种名称 Breeds | 采样地点 Sampling location | 采样只数(羊群数) Sample size (flock number) |
---|---|---|
绵羊品种 Native or crossbred sheep breeds | ||
小尾寒羊 Small-tailed Han (STH) | 山东嘉祥县 Jiaxiang County | 42 (14) |
大尾寒羊 Large-tailed Han (LTH) | 山东临清市 Linqing City | 42 (22) |
山地绵羊 Shandi (SD) | 山东莱芜市 Laiwu City | 29 (17) |
洼地绵羊 Wadi (WD) | 山东滨州市 Binzhou City | 40 (18) |
杂交绵羊 Crossbred (CB) | 山东莱芜市 Laiwu City | 20 (1) |
山羊品种 Native goat breeds | ||
济宁青山羊 Jining grey (JNG) | 山东嘉祥县 Jiaxiang County | 40 (5) |
鲁北白山羊 Lubei white (LBW) | 山东无棣县 Wudi County | 40 (23) |
莱芜黑山羊 Laiwu black (LWB) | 山东莱芜市 Laiwu City | 40 (26) |
崂山奶山羊 Laoshan milk (LSM) | 山东临朐县 Linqu County | 40 (21) |
基因座 Loci | Na | Ne | Dk | PIC | I | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | |||||
MAF70 | 12.00a | 6.75b | 6.10 | 4.52 | 0.83 | 0.79 | 0.81 | 0.74 | 2.06a | 1.60b | ||||
BMS1248 | 9.80 | 6.25 | 5.28 | 3.91 | 0.83 | 0.74 | 0.75 | 0.69 | 1.85 | 1.52 | ||||
BM6404 | 11.60 | 6.50 | 5.28 | 3.81 | 0.84 | 0.73 | 0.77 | 0.68 | 1.88 | 1.49 | ||||
BMS1714 | 10.40a | 4.67b | 5.97 | 2.94 | 0.83 | 0.72 | 0.79 | 0.65 | 1.94 | 1.35 | ||||
BMS1724 | 12.60A | 2.67B | 8.55A | 1.98B | 0.91A | 0.62B | 0.80a | 0.52b | 2.27A | 0.96B | ||||
BMS875 | 9.00a | 3.00b | 4.54a | 1.98b | 0.80a | 0.60b | 0.74a | 0.50b | 1.72A | 0.96B | ||||
BMS1678 | 10.80a | 6.00b | 7.29a | 3.51b | 0.89a | 0.69b | 0.83 | 0.62 | 2.10a | 1.37b | ||||
BM203 | 13.40A | 1.75B | 8.44A | 1.62B | 0.91A | 0.46B | 0.85A | 0.26B | 2.26A | 0.48B | ||||
MB023 | 11.80A | 2.67B | 7.66A | 1.70B | 0.88A | 0.56B | 0.85A | 0.45B | 2.20A | 0.84B | ||||
BM3033 | 12.80A | 6.25B | 9.41A | 4.79B | 0.92A | 0.80B | 0.88A | 0.76B | 2.33A | 1.65B | ||||
BMC1206 | 8.20A | 3.50B | 5.56A | 2.44B | 0.83a | 0.57b | 0.79A | 0.50B | 1.84A | 0.98B | ||||
BM6526 | 11.60a | 4.25b | 6.79 | 2.86 | 0.88a | 0.60b | 0.82a | 0.54b | 2.09A | 1.11B | ||||
BM6444 | 4.20b | 7.00a | 2.51 | 4.60 | 0.61 | 0.70 | 0.54 | 0.66 | 1.07 | 1.54 | ||||
平均 Average | 10.63A | 4.77B | 6.41A | 3.36B | 0.84A | 0.66B | 0.79A | 0.58B | 1.97A | 1.23B |
表2 绵羊和山羊种群在13个基因座上的等位基因数(Na)、有效等位基因数(Ne)、基因多样度(Dk)、多态信息含量(PIC)和Shannon指数(I)
Table 2 Number of alleles (Na), number of effective alleles (Ne), gene diversity (Dk), polymorphism information content (PIC) and Shannon's information index (I) of 13 loci in sheep and goat populations
基因座 Loci | Na | Ne | Dk | PIC | I | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | |||||
MAF70 | 12.00a | 6.75b | 6.10 | 4.52 | 0.83 | 0.79 | 0.81 | 0.74 | 2.06a | 1.60b | ||||
BMS1248 | 9.80 | 6.25 | 5.28 | 3.91 | 0.83 | 0.74 | 0.75 | 0.69 | 1.85 | 1.52 | ||||
BM6404 | 11.60 | 6.50 | 5.28 | 3.81 | 0.84 | 0.73 | 0.77 | 0.68 | 1.88 | 1.49 | ||||
BMS1714 | 10.40a | 4.67b | 5.97 | 2.94 | 0.83 | 0.72 | 0.79 | 0.65 | 1.94 | 1.35 | ||||
BMS1724 | 12.60A | 2.67B | 8.55A | 1.98B | 0.91A | 0.62B | 0.80a | 0.52b | 2.27A | 0.96B | ||||
BMS875 | 9.00a | 3.00b | 4.54a | 1.98b | 0.80a | 0.60b | 0.74a | 0.50b | 1.72A | 0.96B | ||||
BMS1678 | 10.80a | 6.00b | 7.29a | 3.51b | 0.89a | 0.69b | 0.83 | 0.62 | 2.10a | 1.37b | ||||
BM203 | 13.40A | 1.75B | 8.44A | 1.62B | 0.91A | 0.46B | 0.85A | 0.26B | 2.26A | 0.48B | ||||
MB023 | 11.80A | 2.67B | 7.66A | 1.70B | 0.88A | 0.56B | 0.85A | 0.45B | 2.20A | 0.84B | ||||
BM3033 | 12.80A | 6.25B | 9.41A | 4.79B | 0.92A | 0.80B | 0.88A | 0.76B | 2.33A | 1.65B | ||||
BMC1206 | 8.20A | 3.50B | 5.56A | 2.44B | 0.83a | 0.57b | 0.79A | 0.50B | 1.84A | 0.98B | ||||
BM6526 | 11.60a | 4.25b | 6.79 | 2.86 | 0.88a | 0.60b | 0.82a | 0.54b | 2.09A | 1.11B | ||||
BM6444 | 4.20b | 7.00a | 2.51 | 4.60 | 0.61 | 0.70 | 0.54 | 0.66 | 1.07 | 1.54 | ||||
平均 Average | 10.63A | 4.77B | 6.41A | 3.36B | 0.84A | 0.66B | 0.79A | 0.58B | 1.97A | 1.23B |
基因座 Loci | Ho | He | FST | FIS | Relat | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | |||||
MAF70 | 0.599A | 0.225B | 0.850 | 0.783 | 0.017 | 0.084 | 0.330 | 0.715 | 0.025 | 0.096 | ||||
BMS1248 | 0.401 | 0.400 | 0.796 | 0.737 | 0.061 | 0.066 | 0.543 | 0.460 | 0.078 | 0.088 | ||||
BM6404 | 0.489B | 0.850A | 0.811 | 0.727 | 0.006 | 0.158 | 0.487 | -0.172 | 0.008 | 0.311 | ||||
BMS1714 | 0.395 | 0.731 | 0.839 | 0.538 | 0.046 | 0.091 | 0.622 | -0.364 | 0.057 | 0.239 | ||||
BMS1724 | 0.419 | 0.281 | 0.896a | 0.460b | 0.024 | 0.392 | 0.538 | 0.121 | 0.032 | -1.288 | ||||
BMS875 | 0.361 | 0.256 | 0.787a | 0.447b | 0.052 | 0.070 | 0.474 | 0.430 | 0.069 | 0.096 | ||||
BMS1678 | 0.525 | 0.563 | 0.875A | 0.689B | 0.022 | 0.204 | 0.377 | 0.185 | 0.031 | 0.301 | ||||
BM203 | 0.631A | 0.219B | 0.884A | 0.340B | 0.013 | 0.359 | 0.375 | 0.360 | 0.019 | 0.451 | ||||
MB023 | 0.699A | 0.031B | 0.889A | 0.414B | 0.034 | 0.371 | 0.256 | 0.925 | 0.054 | 0.379 | ||||
BM3033 | 0.804 | 0.856 | 0.913A | 0.799B | 0.002 | 0.065 | 0.132 | -0.073 | 0.003 | 0.130 | ||||
BMC1206 | 0.583A | 0.000B | 0.837A | 0.566B | 0.021 | 0.122 | 0.338 | 1.000 | 0.031 | 0.122 | ||||
BM6526 | 0.540 | 0.488 | 0.869a | 0.595b | 0.012 | 0.294 | 0.370 | 0.183 | 0.017 | 0.413 | ||||
BM6444 | 0.260B | 0.750A | 0.610 | 0.699 | 0.171 | 0.165 | 0.594 | -0.075 | 0.205 | 0.300 | ||||
平均 Average | 0.516 | 0.464 | 0.835A | 0.646B | 0.035 | 0.164 | 0.412 | 0.280 | 0.049b | 0.234a |
表3 绵羊和山羊种群在13个基因座上的观测杂合度(Ho)、期望杂合度(He)、固定指数(FST)、近交系数((FIS)和亲缘关系指数(Relat)
Table 3 Observed and expected heterozygosity (Ho, He), fixation index (FST), inbreeding coefficient (FIS) and relatedness (Relat) of 13 loci in sheep and goat populations
基因座 Loci | Ho | He | FST | FIS | Relat | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | 绵羊 Sheep | 山羊 Goat | |||||
MAF70 | 0.599A | 0.225B | 0.850 | 0.783 | 0.017 | 0.084 | 0.330 | 0.715 | 0.025 | 0.096 | ||||
BMS1248 | 0.401 | 0.400 | 0.796 | 0.737 | 0.061 | 0.066 | 0.543 | 0.460 | 0.078 | 0.088 | ||||
BM6404 | 0.489B | 0.850A | 0.811 | 0.727 | 0.006 | 0.158 | 0.487 | -0.172 | 0.008 | 0.311 | ||||
BMS1714 | 0.395 | 0.731 | 0.839 | 0.538 | 0.046 | 0.091 | 0.622 | -0.364 | 0.057 | 0.239 | ||||
BMS1724 | 0.419 | 0.281 | 0.896a | 0.460b | 0.024 | 0.392 | 0.538 | 0.121 | 0.032 | -1.288 | ||||
BMS875 | 0.361 | 0.256 | 0.787a | 0.447b | 0.052 | 0.070 | 0.474 | 0.430 | 0.069 | 0.096 | ||||
BMS1678 | 0.525 | 0.563 | 0.875A | 0.689B | 0.022 | 0.204 | 0.377 | 0.185 | 0.031 | 0.301 | ||||
BM203 | 0.631A | 0.219B | 0.884A | 0.340B | 0.013 | 0.359 | 0.375 | 0.360 | 0.019 | 0.451 | ||||
MB023 | 0.699A | 0.031B | 0.889A | 0.414B | 0.034 | 0.371 | 0.256 | 0.925 | 0.054 | 0.379 | ||||
BM3033 | 0.804 | 0.856 | 0.913A | 0.799B | 0.002 | 0.065 | 0.132 | -0.073 | 0.003 | 0.130 | ||||
BMC1206 | 0.583A | 0.000B | 0.837A | 0.566B | 0.021 | 0.122 | 0.338 | 1.000 | 0.031 | 0.122 | ||||
BM6526 | 0.540 | 0.488 | 0.869a | 0.595b | 0.012 | 0.294 | 0.370 | 0.183 | 0.017 | 0.413 | ||||
BM6444 | 0.260B | 0.750A | 0.610 | 0.699 | 0.171 | 0.165 | 0.594 | -0.075 | 0.205 | 0.300 | ||||
平均 Average | 0.516 | 0.464 | 0.835A | 0.646B | 0.035 | 0.164 | 0.412 | 0.280 | 0.049b | 0.234a |
品种 Breeds | 品种 Breeds | ||||||||
---|---|---|---|---|---|---|---|---|---|
STH | LTH | WD | SD | CB | JNG | LBW | LSM | LWB | |
STH | ** | ** | ** | NS | ** | ** | ** | ** | |
LTH | 0.0196 | ** | ** | NS | ** | ** | ** | ** | |
WD | 0.0267 | 0.0312 | ** | NS | ** | ** | ** | ** | |
SD | 0.0405 | 0.0408 | 0.0440 | ** | ** | ** | ** | ** | |
CB | 0.0212 | 0.0472 | 0.0599 | 0.0595 | ** | ** | ** | ** | |
JNG | 0.2243 | 0.2126 | 0.2472 | 0.2334 | 0.2791 | ** | ** | ** | |
LBW | 0.1597 | 0.1549 | 0.1784 | 0.1677 | 0.1790 | 0.1283 | ** | ** | |
LSM | 0.2131 | 0.2138 | 0.2418 | 0.2292 | 0.2697 | 0.2645 | 0.1579 | ** | |
LWB | 0.1446 | 0.1400 | 0.1726 | 0.1503 | 0.1612 | 0.1669 | 0.0954 | 0.1535 |
表4 基于9个基因座的9个品种间成对固定指数(FST, 对角线下方)和遗传分化配对测试(对角线上方)
Table 4 Fixation index (FST, below the diagonal) and pairwise testing of genetic differentiation (above the diagonal) among nine breeds based on nine loci
品种 Breeds | 品种 Breeds | ||||||||
---|---|---|---|---|---|---|---|---|---|
STH | LTH | WD | SD | CB | JNG | LBW | LSM | LWB | |
STH | ** | ** | ** | NS | ** | ** | ** | ** | |
LTH | 0.0196 | ** | ** | NS | ** | ** | ** | ** | |
WD | 0.0267 | 0.0312 | ** | NS | ** | ** | ** | ** | |
SD | 0.0405 | 0.0408 | 0.0440 | ** | ** | ** | ** | ** | |
CB | 0.0212 | 0.0472 | 0.0599 | 0.0595 | ** | ** | ** | ** | |
JNG | 0.2243 | 0.2126 | 0.2472 | 0.2334 | 0.2791 | ** | ** | ** | |
LBW | 0.1597 | 0.1549 | 0.1784 | 0.1677 | 0.1790 | 0.1283 | ** | ** | |
LSM | 0.2131 | 0.2138 | 0.2418 | 0.2292 | 0.2697 | 0.2645 | 0.1579 | ** | |
LWB | 0.1446 | 0.1400 | 0.1726 | 0.1503 | 0.1612 | 0.1669 | 0.0954 | 0.1535 |
品种 Breeds | 品种 Breeds | ||||||||
---|---|---|---|---|---|---|---|---|---|
STH | LTH | WD | SD | CB | JNG | LBW | LSM | LWB | |
STH | 4,569 | 3,972 | 5,725 | 5,396 | 27,154 | 23,441 | 23,105 | 19,807 | |
LTH | 0.2513 | 5,229 | 6,174 | 8,405 | 25,063 | 24,943 | 25,639 | 19,467 | |
WD | 0.2185 | 0.2876 | 5,092 | 8,894 | 28,713 | 22,550 | 24,521 | 24,490 | |
SD | 0.3149 | 0.3396 | 0.2801 | 9,286 | 28,955 | 24,108 | 25,823 | 21,002 | |
CB | 0.2968 | 0.4623 | 0.4892 | 0.5108 | 34,493 | 26,606 | 30,001 | 21,022 | |
JNG | 1.4936 | 1.3786 | 1.5794 | 1.5927 | 1.8973 | 10,626 | 17,940 | 12,901 | |
LBW | 1.2894 | 1.3720 | 1.2404 | 1.3261 | 1.4635 | 0.5845 | 6,743 | 7,561 | |
LSM | 1.2709 | 1.4103 | 1.3488 | 1.4204 | 1.6502 | 0.9868 | 0.3709 | 8,826 | |
LWB | 1.0895 | 1.0708 | 1.3471 | 1.1552 | 1.1563 | 0.7096 | 0.4159 | 0.4855 |
表5 基于13个基因座的9个品种间Nei氏无偏遗传距离(对角线下方)和遗传分化时间(年, 对角线上方)
Table 5 Nei′s unbiased genetic distance (below the diagonal) and genetic divergence time (years, above the diagonal) among nine breeds based on 13 loci
品种 Breeds | 品种 Breeds | ||||||||
---|---|---|---|---|---|---|---|---|---|
STH | LTH | WD | SD | CB | JNG | LBW | LSM | LWB | |
STH | 4,569 | 3,972 | 5,725 | 5,396 | 27,154 | 23,441 | 23,105 | 19,807 | |
LTH | 0.2513 | 5,229 | 6,174 | 8,405 | 25,063 | 24,943 | 25,639 | 19,467 | |
WD | 0.2185 | 0.2876 | 5,092 | 8,894 | 28,713 | 22,550 | 24,521 | 24,490 | |
SD | 0.3149 | 0.3396 | 0.2801 | 9,286 | 28,955 | 24,108 | 25,823 | 21,002 | |
CB | 0.2968 | 0.4623 | 0.4892 | 0.5108 | 34,493 | 26,606 | 30,001 | 21,022 | |
JNG | 1.4936 | 1.3786 | 1.5794 | 1.5927 | 1.8973 | 10,626 | 17,940 | 12,901 | |
LBW | 1.2894 | 1.3720 | 1.2404 | 1.3261 | 1.4635 | 0.5845 | 6,743 | 7,561 | |
LSM | 1.2709 | 1.4103 | 1.3488 | 1.4204 | 1.6502 | 0.9868 | 0.3709 | 8,826 | |
LWB | 1.0895 | 1.0708 | 1.3471 | 1.1552 | 1.1563 | 0.7096 | 0.4159 | 0.4855 |
图1 基于Nei’s无偏遗传距离的9个品种的UPGMA(A)和ME(B)系统发生树。树图系由MEGA3.1软件绘制。品种代号同表1。
Fig. 1 UPGMA (A) and ME (B) phylogenetic dendrograms based on Nei's unbiased genetic distance of nine breeds. Dendrograms are drawn using MEGA 3.1 software. Breed codes are given in Table 1.
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