中国板栗自然居群微卫星(SSR)遗传多样性

doi:10.3724/SP.J.1003.2009.09043

摘要/Abstract

摘要：

采用8对微卫星分子标记对中国板栗(Castanea mollissima)的28个自然居群进行了遗传多样性与遗传结构分析。在849个个体上扩增得到128个等位基因, 每位点平均等位基因数(A)为16。中国板栗居群的平均预期杂合度(H_E)为0.678, 平均观察杂合度(H_O)为0.590。华中地区的中国板栗居群遗传多样性最高(A= 8.112, H_E= 0.705, H_O= 0.618), 其次为西北地区和华东地区, 而西南地区遗传多样性最低(A= 6.611, H_E= 0.640, H_O= 0.559)。基于无限等位基因模型(IAM)和基于逐步突变模型(SMM)的遗传分化系数分别为F_ST= 0.120和R_ST= 0.208。分子方差分析(AMOVA)结果表明中国板栗野生居群的遗传变异主要存在于居群内(87.16%)。Mantel检测揭示遗传距离与地理距离之间无显著相关性, 表明基因流不是主导中国板栗居群遗传结构的关键因素。华中地区(尤其是神农架及其周边地区)是中国板栗遗传多样性的现代分布中心, 因而应该得到优先保护, 同时该区域的野生板栗居群可优先作为栽培板栗遗传育种的材料和基因库。

关键词: Castanea mollissima, 种质资源, 微卫星, 遗传多样性, 居群遗传结构

Abstract

Genetic diversity and population structure of 28 natural populations of Castanea mollissima were investigated by using microsatellite markers. A total of 128 alleles were identified in 849 individuals across the eight microsatellites analysed, with a mean value of 16 alleles per locus. The mean expected heterozygosity (H_E) and observed heterozygosity (H_O) across all populations were 0.678 and 0.590, respectively, and a higher level of diversity was found in populations from central China (A= 8.112, H_E= 0.705, H_O= 0.618) than those from other three regions (eastern, southwestern and northwestern China). Indices of genetic differentiation based on stepwise mutation model (SMM) and infinite allele model (IAM) were R_ST= 0.208 and F_ST= 0.120, respectively. No significant association between genetic distance and geographic distance was detected by Mantel test, suggesting gene flow is not a dominant factor shaping genetic structure of the species. The central China, particularly the area around the Shengnongjia Mountains, can be recognized as a modern center of genetic diversity of C. mollissima. Thus, natural populations of the species in this region deserve prior conservation and utilization for breeding programmes.

Key words: Castanea mollissima, germplasm, microsatellite, genetic diversity, population structure

田华, 康明, 李丽, 姚小洪, 黄宏文 (2009) 中国板栗自然居群微卫星(SSR)遗传多样性. 生物多样性, 17, 296-302. DOI: 10.3724/SP.J.1003.2009.09043.

Hua Tian, Ming Kang, Li Li, Xiaohong Yao, Hongwen Huang (2009) Genetic diversity in natural populations of Castanea mollissima inferred from nuclear SSR markers. Biodiversity Science, 17, 296-302. DOI: 10.3724/SP.J.1003.2009.09043.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2009.09043

https://www.biodiversity-science.net/CN/Y2009/V17/I3/296

图/表 3

参考文献 31

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居群 Population	采样数 Sample size	经纬度 Locality	平均等位基因数(A)	平均有效等位基因数(A_E)	观察杂合度(H_O)	预期杂合度(H_E)
华东地区 Eastern China	142		7.500	3.802	0.615	0.685
1. 江苏宜兴 Yixing, Jiangsu	36	31.218°N, 119.693°E	10.000	4.567	0.623	0.759
2. 江西婺源 Wuyuan, Jiangxi	16	29.248°N, 117.866°E	5.500	2.938	0.594	0.619
3. 浙江临安 Lin’an, Zhejiang	29	30.368°N, 119.423°E	8.375	4.472	0.596	0.706
4. 浙江兰溪 Lanxi, Zhejiang	21	29.390°N, 119.406°E	6.250	3.391	0.655	0.668
5. 安徽岩寺 Yansi, Anhui	23	29.418°N, 118.293°E	7.000	3.427	0.684	0.680
6. 安徽绩溪 Jixi, Anhui	17	30.110°N, 118.835°E	7.875	4.017	0.536	0.677
华中地区 Central China	341		8.112	4.291	0.618	0.705
7. 湖南吉首 Jishou, Hunan	42	28.406°N, 109.811°E	7.750	3.769	0.591	0.703
8. 湖南绥宁 Suining, Hunan	26	26.492°N, 110.139°E	7.375	3.992	0.595	0.679
9. 湖南永顺 Yongshun, Hunan	10	28.991°N, 109.869°E	5.000	3.490	0.589	0.658
10. 湖北神农架 Shengnongjia, Hubei	37	31.448°N, 110.390°E	8.750	4.517	0.641	0.713
11. 湖北武当山 Wudangshan, Hubei	27	32.491°N, 110.309°E	8.125	4.744	0.644	0.707
12. 湖北房县 Fangxian County, Hubei	40	31.857°N, 110.740°E	7.750	4.328	0.575	0.683
13. 湖北房县 Fangxian County, Hubei	49	31.859°N, 110.643°E	9.875	5.061	0.631	0.735
14. 湖北秭归 Zigui, Hubei	43	30.747°N, 110.793°E	10.375	4.406	0.598	0.721
15. 湖北宜昌 Yichang, Hubei	57	30.769°N, 111.331°E	10.375	4.627	0.630	0.737
16. 湖北宜昌 Yichang, Hubei	10	30.751°N, 111.311°E	5.750	3.979	0.688	0.709
西南地区 Southwestern China	255		6.611	3.354	0.559	0.640
17. 广西南丹 Nandan, Guangxi	32	24.992°N, 107.615°E	6.875	3.178	0.605	0.672
18. 贵州荔波 Libo, Guizhou	20	25.411°N, 107.964°E	6.000	2.980	0.519	0.631
19. 贵州榕江 Rongjiang, Guizhou	31	25.922°N, 108.587°E	8.125	3.799	0.598	0.685
20. 贵州紫云 Ziyun, Guizhou	24	25.956°N, 106.115°E	6.625	3.292	0.609	0.600
21. 贵州遵义 Zunyi, Guizhou	30	27.756°N, 106.792°E	6.125	3.279	0.583	0.600
22. 云南镇雄 Zhenxiong, Yunnan	35	27.438°N, 104.851°E	6.375	3.035	0.557	0.606
23. 云南麻栗坡 Malipo, Yunnan	27	23.178°N, 104.858°E	6.375	3.683	0.497	0.673
24. 四川邛崃 Qionglai, Sichuan	21	30.440°N, 103.258°E	5.875	3.299	0.548	0.635
25. 重庆城口 Chengkou, Chongqing	35	32.030°N, 108.628°E	7.125	3.638	0.518	0.661
西北地区 Northwestern China	111		7.208	3.630	0.539	0.686
26. 甘肃徽县 Huixian, Gansu	42	33.970°N, 106.006°E	7.750	4.050	0.564	0.707
27. 甘肃徽县 Huixian, Gansu	33	34.358°N, 106.649°E	7.000	3.221	0.500	0.659
28. 山西夏县 Xiaxian, Shanxi	36	35.147°N, 111.442°E	6.875	3.620	0.553	0.693
平均 Mean	30.321		7.402	3.814	0.590	0.678
合计 Total	849		7.333	3.796	0.593	0.678

居群 Population	采样数 Sample size	经纬度 Locality	平均等位基因数(A)	平均有效等位基因数(A_E)	观察杂合度(H_O)	预期杂合度(H_E)
华东地区 Eastern China	142		7.500	3.802	0.615	0.685
1. 江苏宜兴 Yixing, Jiangsu	36	31.218°N, 119.693°E	10.000	4.567	0.623	0.759
2. 江西婺源 Wuyuan, Jiangxi	16	29.248°N, 117.866°E	5.500	2.938	0.594	0.619
3. 浙江临安 Lin’an, Zhejiang	29	30.368°N, 119.423°E	8.375	4.472	0.596	0.706
4. 浙江兰溪 Lanxi, Zhejiang	21	29.390°N, 119.406°E	6.250	3.391	0.655	0.668
5. 安徽岩寺 Yansi, Anhui	23	29.418°N, 118.293°E	7.000	3.427	0.684	0.680
6. 安徽绩溪 Jixi, Anhui	17	30.110°N, 118.835°E	7.875	4.017	0.536	0.677
华中地区 Central China	341		8.112	4.291	0.618	0.705
7. 湖南吉首 Jishou, Hunan	42	28.406°N, 109.811°E	7.750	3.769	0.591	0.703
8. 湖南绥宁 Suining, Hunan	26	26.492°N, 110.139°E	7.375	3.992	0.595	0.679
9. 湖南永顺 Yongshun, Hunan	10	28.991°N, 109.869°E	5.000	3.490	0.589	0.658
10. 湖北神农架 Shengnongjia, Hubei	37	31.448°N, 110.390°E	8.750	4.517	0.641	0.713
11. 湖北武当山 Wudangshan, Hubei	27	32.491°N, 110.309°E	8.125	4.744	0.644	0.707
12. 湖北房县 Fangxian County, Hubei	40	31.857°N, 110.740°E	7.750	4.328	0.575	0.683
13. 湖北房县 Fangxian County, Hubei	49	31.859°N, 110.643°E	9.875	5.061	0.631	0.735
14. 湖北秭归 Zigui, Hubei	43	30.747°N, 110.793°E	10.375	4.406	0.598	0.721
15. 湖北宜昌 Yichang, Hubei	57	30.769°N, 111.331°E	10.375	4.627	0.630	0.737
16. 湖北宜昌 Yichang, Hubei	10	30.751°N, 111.311°E	5.750	3.979	0.688	0.709
西南地区 Southwestern China	255		6.611	3.354	0.559	0.640
17. 广西南丹 Nandan, Guangxi	32	24.992°N, 107.615°E	6.875	3.178	0.605	0.672
18. 贵州荔波 Libo, Guizhou	20	25.411°N, 107.964°E	6.000	2.980	0.519	0.631
19. 贵州榕江 Rongjiang, Guizhou	31	25.922°N, 108.587°E	8.125	3.799	0.598	0.685
20. 贵州紫云 Ziyun, Guizhou	24	25.956°N, 106.115°E	6.625	3.292	0.609	0.600
21. 贵州遵义 Zunyi, Guizhou	30	27.756°N, 106.792°E	6.125	3.279	0.583	0.600
22. 云南镇雄 Zhenxiong, Yunnan	35	27.438°N, 104.851°E	6.375	3.035	0.557	0.606
23. 云南麻栗坡 Malipo, Yunnan	27	23.178°N, 104.858°E	6.375	3.683	0.497	0.673
24. 四川邛崃 Qionglai, Sichuan	21	30.440°N, 103.258°E	5.875	3.299	0.548	0.635
25. 重庆城口 Chengkou, Chongqing	35	32.030°N, 108.628°E	7.125	3.638	0.518	0.661
西北地区 Northwestern China	111		7.208	3.630	0.539	0.686
26. 甘肃徽县 Huixian, Gansu	42	33.970°N, 106.006°E	7.750	4.050	0.564	0.707
27. 甘肃徽县 Huixian, Gansu	33	34.358°N, 106.649°E	7.000	3.221	0.500	0.659
28. 山西夏县 Xiaxian, Shanxi	36	35.147°N, 111.442°E	6.875	3.620	0.553	0.693
平均 Mean	30.321		7.402	3.814	0.590	0.678
合计 Total	849		7.333	3.796	0.593	0.678

	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28
1		0.327^**	0.201	0.201^***	0.096	0.136	0.092	0.008	0.081	0.166^*	0.144	0.115	0.112	0.117	0.092	0.114	0.168^*	0.257^*	0.240^**	0.149	0.262	0.338^**	0.093	0.161	0.222	0.221^*	0.263^*	0.173
2	0.103^***		0.208^*	0.568^***	0.463^**	0.257	0.272^**	0.326^**	0.342^*	0.185	0.203	0.154	0.135	0.191	0.190^*	0.107	0.164^*	0.111	0.314^**	0.380^*	0.501^*	0.395^**	0.242^*	0.193	0.107	0.263	0.367^*	0.306
3	0.117^***	0.103^***		0.553^***	0.394^**	0.047	0.189^*	0.254^**	0.317^**	0.047	0.261^**	0.145	0.101	0.174	0.071	0.128	0.121	0.095	0.260	0.120	0.242	0.156	0.141	0.168	0.196	0.360^***	0.456^***	0.361^**
4	0.052^*	0.130^***	0.140^***		0.088	0.492^***	0.239^***	0.165	0.174	0.453^***	0.233	0.320^**	0.302^**	0.241^**	0.310^***	0.374^**	0.452^***	0.549^***	0.401^**	0.502^***	0.604^***	0.684^***	0.329^**	0.452^**	0.473^***	0.300^**	0.175	0.093
5	0.081^***	0.120^***	0.165^***	0.094^***		0.289^*	0.079	0.097	0.050	0.273^**	0.160	0.195^*	0.181	0.123	0.165	0.219	0.280^*	0.381^*	0.215	0.311	0.427	0.571^***	0.140	0.293^*	0.344^*	0.269^*	0.222^*	0.125
6	0.089^***	0.088^***	0.079^***	0.114^***	0.110^***		0.113	0.194	0.226^*	0.007	0.201^**	0.100^*	0.081^*	0.121^*	0.050	0.084	0.081	0.083	0.150	0.059	0.182	0.265	0.054	0.111	0.184	0.310^***	0.396^***	0.310^**
7	0.102^***	0.121^***	0.114^***	0.109^***	0.102^***	0.094^***		0.088	0.007	0.098	0.114	0.097	0.058	0.014	0.033	0.047	0.095	0.145	0.130	0.162	0.303	0.376^**	0.022	0.153	0.194	0.242^*	0.252^*	0.163
8	0.077^***	0.118^***	0.132^***	0.140^***	0.127^***	0.107^***	0.086^***		0.061	0.205^**	0.111	0.109	0.105	0.099	0.106	0.105	0.174^*	0.280^*	0.266	0.228	0.365^*	0.412^***	0.102	0.172	0.218	0.181	0.216	0.122
9	0.122^***	0.153^**	0.140^***	0.153^***	0.123^***	0.109^**	0.016^**	0.094^***		0.176^**	0.085	0.122	0.072	0.006	0.061	0.070	0.185^*	0.261^**	0.201	0.265	0.424	0.535^**	0.074	0.229^*	0.253^*	0.211^*	0.163	0.080
10	0.082^***	0.085^***	0.099^***	0.125^***	0.085^***	0.064^***	0.069^***	0.090^***	0.049^***		0.188	0.099	0.068	0.090	0.035	0.049	0.043	0.039	0.156	0.080	0.207	0.255^*	0.053	0.120^*	0.173	0.292^***	0.370^**	0.293^*
11	0.137^***	0.108^***	0.122^***	0.157^***	0.114^***	0.083^***	0.101^***	0.138^***	0.117^***	0.101^***		0.022	0.042	0.066	0.122	0.051	0.123	0.212	0.135	0.330	0.442^*	0.433^**	0.095	0.080	0.079	0.028	0.100	0.051
12	0.108^***	0.081^***	0.098^***	0.137^***	0.102^***	0.032^***	0.088^***	0.097^***	0.091^**	0.068^***	0.030^***		0.011	0.071	0.069	0.015	0.036	0.122	0.097	0.241	0.351^*	0.329^*	0.034	0.003	0.025	0.100	0.226^**	0.143
13	0.103^***	0.075^***	0.086^***	0.118^***	0.103^***	0.029^**	0.062^***	0.101^***	0.067^***	0.056^***	0.032^***	0.017^***		0.024	0.020	0.004	0.039	0.087	0.118	0.195	0.322^*	0.260	0.033	0.054	0.066	0.139^*	0.219^**	0.144
14	0.071^***	0.080^***	0.102^***	0.073^***	0.121^***	0.067^***	0.091^***	0.104^***	0.099^***	0.070^***	0.103^***	0.078^***	0.065^***		0.027	0.011	0.080	0.117	0.146	0.198	0.341^*	0.341^***	0.047	0.136^**	0.158	0.172^**	0.179	0.118
15	0.073^***	0.071^***	0.077^***	0.077^***	0.111^***	0.039^***	0.079^***	0.094^***	0.094^***	0.061^***	0.074^***	0.046^***	0.031^***	0.029^***		0.028	0.051	0.085	0.148	0.108	0.249	0.236^*	0.027	0.119	0.159	0.236^**	0.285^**	0.199^*
16	0.082^***	0.099^***	0.113^***	0.144^***	0.112^***	0.102^**	0.029	0.060^**	0.021	0.056^***	0.117^***	0.094^**	0.077^***	0.097^***	0.095^***		0.012	0.045	0.146	0.205	0.356	0.372^**	0.022	0.040	0.053	0.153	0.236^*	0.155
17	0.095^***	0.068^***	0.102^***	0.121^***	0.097^***	0.075^***	0.083^***	0.116^***	0.076^***	0.060^***	0.091^***	0.067^***	0.057^***	0.100^***	0.071^***	0.082^***		0.063	0.157	0.211	0.357	0.387^**	0.024	0.056	0.096	0.225^*	0.339^*	0.243
18	0.138^***	0.110^***	0.141^***	0.165^***	0.128^***	0.102^***	0.080^***	0.123^***	0.084^***	0.037^*	0.117^***	0.094^***	0.076^***	0.088^***	0.089^***	0.082^***	0.084^***		0.193	0.210	0.360	0.340^*	0.106	0.144	0.144	0.326^***	0.406^**	0.326^*
19	0.070^***	0.152^***	0.179^***	0.105^***	0.108^***	0.158^***	0.131^***	0.174^***	0.149^***	0.117^***	0.162^***	0.156^***	0.137^***	0.101^***	0.113^***	0.143^***	0.120^***	0.165^***		0.288	0.380	0.457^**	0.079	0.120	0.185	0.267	0.329^*	0.258^*
20	0.165^***	0.172^***	0.140^***	0.195^***	0.202^***	0.108^***	0.169^***	0.151^***	0.181^***	0.151^***	0.199^***	0.162^***	0.138^***	0.148^***	0.112^***	0.178^***	0.191^***	0.187^***	0.241^***		0.071	0.290^*	0.155	0.275	0.346^*	0.430^**	0.478^*	0.389^*
21	0.172^***	0.212^***	0.174^***	0.226^***	0.235^***	0.156^***	0.205^***	0.181^***	0.243^***	0.180^***	0.234^***	0.196^***	0.176^***	0.183^***	0.161^***	0.190^***	0.223^***	0.230^***	0.246^***	0.135^***		0.409^*	0.299	0.381	0.454^**	0.520^***	0.572^**	0.489^**
22	0.178^***	0.193^***	0.152^***	0.219^***	0.238^***	0.165^***	0.172^***	0.155^***	0.217^***	0.161^***	0.224^***	0.200^***	0.170^***	0.154^***	0.144^***	0.175^***	0.201^***	0.186^***	0.255^***	0.161^***	0.217^***		0.379^***	0.392^***	0.374^*	0.497^***	0.576^**	0.510^**
23	0.099^***	0.097^***	0.128^***	0.114^***	0.079^***	0.072^***	0.074^***	0.106^***	0.065^***	0.058^***	0.071^***	0.052^***	0.050^***	0.089^***	0.059^***	0.073^***	0.039^***	0.093^***	0.139^***	0.184^***	0.225^***	0.188^***		0.063	0.128	0.222^*	0.295^*	0.195
24	0.113^***	0.098^***	0.117^***	0.146^***	0.104^***	0.080^**	0.103^***	0.110^***	0.105^**	0.055^*	0.098^***	0.071^**	0.081^***	0.073^***	0.083^***	0.091^**	0.061^***	0.050^**	0.158^***	0.202^***	0.237^***	0.192^***	0.071^***		0.009	0.166	0.325	0.234
25	0.134^**	0.130^***	0.167^***	0.158^***	0.082^***	0.136^***	0.138^***	0.158^**	0.170^**	0.118^***	0.128^***	0.118^***	0.120^***	0.129^***	0.133^***	0.133^**	0.149^***	0.137^***	0.157^***	0.215^***	0.236^***	0.242^***	0.141^***	0.106^**		0.145	0.301^*	0.235^*
26	0.114^***	0.088^***	0.134^***	0.140^***	0.103^***	0.091^***	0.128^***	0.120^***	0.120^***	0.082^***	0.052^***	0.058^***	0.065^***	0.087^***	0.084^***	0.100^***	0.088^***	0.095^***	0.151^***	0.187^***	0.214^***	0.212^***	0.082^***	0.063^***	0.117^***		0.090	0.076
27	0.137^***	0.137^***	0.167^***	0.157^***	0.090^***	0.123^***	0.128^***	0.170^***	0.131^***	0.113^***	0.076^***	0.093^***	0.086^***	0.130^***	0.120^***	0.134^***	0.111^***	0.147^***	0.150^***	0.238^***	0.256^***	0.278^***	0.123^***	0.143^***	0.146^***	0.071^***		0.023
28	0.089^***	0.129^***	0.167^***	0.126^***	0.081^***	0.114^***	0.138^***	0.132^***	0.168^***	0.110^***	0.109^***	0.108^***	0.104^***	0.118^***	0.113^***	0.132^***	0.133^***	0.157^***	0.114^***	0.195^***	0.207^***	0.248^***	0.135^***	0.150^***	0.100^***	0.105^***	0.090^***

	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28
1		0.327^**	0.201	0.201^***	0.096	0.136	0.092	0.008	0.081	0.166^*	0.144	0.115	0.112	0.117	0.092	0.114	0.168^*	0.257^*	0.240^**	0.149	0.262	0.338^**	0.093	0.161	0.222	0.221^*	0.263^*	0.173
2	0.103^***		0.208^*	0.568^***	0.463^**	0.257	0.272^**	0.326^**	0.342^*	0.185	0.203	0.154	0.135	0.191	0.190^*	0.107	0.164^*	0.111	0.314^**	0.380^*	0.501^*	0.395^**	0.242^*	0.193	0.107	0.263	0.367^*	0.306
3	0.117^***	0.103^***		0.553^***	0.394^**	0.047	0.189^*	0.254^**	0.317^**	0.047	0.261^**	0.145	0.101	0.174	0.071	0.128	0.121	0.095	0.260	0.120	0.242	0.156	0.141	0.168	0.196	0.360^***	0.456^***	0.361^**
4	0.052^*	0.130^***	0.140^***		0.088	0.492^***	0.239^***	0.165	0.174	0.453^***	0.233	0.320^**	0.302^**	0.241^**	0.310^***	0.374^**	0.452^***	0.549^***	0.401^**	0.502^***	0.604^***	0.684^***	0.329^**	0.452^**	0.473^***	0.300^**	0.175	0.093
5	0.081^***	0.120^***	0.165^***	0.094^***		0.289^*	0.079	0.097	0.050	0.273^**	0.160	0.195^*	0.181	0.123	0.165	0.219	0.280^*	0.381^*	0.215	0.311	0.427	0.571^***	0.140	0.293^*	0.344^*	0.269^*	0.222^*	0.125
6	0.089^***	0.088^***	0.079^***	0.114^***	0.110^***		0.113	0.194	0.226^*	0.007	0.201^**	0.100^*	0.081^*	0.121^*	0.050	0.084	0.081	0.083	0.150	0.059	0.182	0.265	0.054	0.111	0.184	0.310^***	0.396^***	0.310^**
7	0.102^***	0.121^***	0.114^***	0.109^***	0.102^***	0.094^***		0.088	0.007	0.098	0.114	0.097	0.058	0.014	0.033	0.047	0.095	0.145	0.130	0.162	0.303	0.376^**	0.022	0.153	0.194	0.242^*	0.252^*	0.163
8	0.077^***	0.118^***	0.132^***	0.140^***	0.127^***	0.107^***	0.086^***		0.061	0.205^**	0.111	0.109	0.105	0.099	0.106	0.105	0.174^*	0.280^*	0.266	0.228	0.365^*	0.412^***	0.102	0.172	0.218	0.181	0.216	0.122
9	0.122^***	0.153^**	0.140^***	0.153^***	0.123^***	0.109^**	0.016^**	0.094^***		0.176^**	0.085	0.122	0.072	0.006	0.061	0.070	0.185^*	0.261^**	0.201	0.265	0.424	0.535^**	0.074	0.229^*	0.253^*	0.211^*	0.163	0.080
10	0.082^***	0.085^***	0.099^***	0.125^***	0.085^***	0.064^***	0.069^***	0.090^***	0.049^***		0.188	0.099	0.068	0.090	0.035	0.049	0.043	0.039	0.156	0.080	0.207	0.255^*	0.053	0.120^*	0.173	0.292^***	0.370^**	0.293^*
11	0.137^***	0.108^***	0.122^***	0.157^***	0.114^***	0.083^***	0.101^***	0.138^***	0.117^***	0.101^***		0.022	0.042	0.066	0.122	0.051	0.123	0.212	0.135	0.330	0.442^*	0.433^**	0.095	0.080	0.079	0.028	0.100	0.051
12	0.108^***	0.081^***	0.098^***	0.137^***	0.102^***	0.032^***	0.088^***	0.097^***	0.091^**	0.068^***	0.030^***		0.011	0.071	0.069	0.015	0.036	0.122	0.097	0.241	0.351^*	0.329^*	0.034	0.003	0.025	0.100	0.226^**	0.143
13	0.103^***	0.075^***	0.086^***	0.118^***	0.103^***	0.029^**	0.062^***	0.101^***	0.067^***	0.056^***	0.032^***	0.017^***		0.024	0.020	0.004	0.039	0.087	0.118	0.195	0.322^*	0.260	0.033	0.054	0.066	0.139^*	0.219^**	0.144
14	0.071^***	0.080^***	0.102^***	0.073^***	0.121^***	0.067^***	0.091^***	0.104^***	0.099^***	0.070^***	0.103^***	0.078^***	0.065^***		0.027	0.011	0.080	0.117	0.146	0.198	0.341^*	0.341^***	0.047	0.136^**	0.158	0.172^**	0.179	0.118
15	0.073^***	0.071^***	0.077^***	0.077^***	0.111^***	0.039^***	0.079^***	0.094^***	0.094^***	0.061^***	0.074^***	0.046^***	0.031^***	0.029^***		0.028	0.051	0.085	0.148	0.108	0.249	0.236^*	0.027	0.119	0.159	0.236^**	0.285^**	0.199^*
16	0.082^***	0.099^***	0.113^***	0.144^***	0.112^***	0.102^**	0.029	0.060^**	0.021	0.056^***	0.117^***	0.094^**	0.077^***	0.097^***	0.095^***		0.012	0.045	0.146	0.205	0.356	0.372^**	0.022	0.040	0.053	0.153	0.236^*	0.155
17	0.095^***	0.068^***	0.102^***	0.121^***	0.097^***	0.075^***	0.083^***	0.116^***	0.076^***	0.060^***	0.091^***	0.067^***	0.057^***	0.100^***	0.071^***	0.082^***		0.063	0.157	0.211	0.357	0.387^**	0.024	0.056	0.096	0.225^*	0.339^*	0.243
18	0.138^***	0.110^***	0.141^***	0.165^***	0.128^***	0.102^***	0.080^***	0.123^***	0.084^***	0.037^*	0.117^***	0.094^***	0.076^***	0.088^***	0.089^***	0.082^***	0.084^***		0.193	0.210	0.360	0.340^*	0.106	0.144	0.144	0.326^***	0.406^**	0.326^*
19	0.070^***	0.152^***	0.179^***	0.105^***	0.108^***	0.158^***	0.131^***	0.174^***	0.149^***	0.117^***	0.162^***	0.156^***	0.137^***	0.101^***	0.113^***	0.143^***	0.120^***	0.165^***		0.288	0.380	0.457^**	0.079	0.120	0.185	0.267	0.329^*	0.258^*
20	0.165^***	0.172^***	0.140^***	0.195^***	0.202^***	0.108^***	0.169^***	0.151^***	0.181^***	0.151^***	0.199^***	0.162^***	0.138^***	0.148^***	0.112^***	0.178^***	0.191^***	0.187^***	0.241^***		0.071	0.290^*	0.155	0.275	0.346^*	0.430^**	0.478^*	0.389^*
21	0.172^***	0.212^***	0.174^***	0.226^***	0.235^***	0.156^***	0.205^***	0.181^***	0.243^***	0.180^***	0.234^***	0.196^***	0.176^***	0.183^***	0.161^***	0.190^***	0.223^***	0.230^***	0.246^***	0.135^***		0.409^*	0.299	0.381	0.454^**	0.520^***	0.572^**	0.489^**
22	0.178^***	0.193^***	0.152^***	0.219^***	0.238^***	0.165^***	0.172^***	0.155^***	0.217^***	0.161^***	0.224^***	0.200^***	0.170^***	0.154^***	0.144^***	0.175^***	0.201^***	0.186^***	0.255^***	0.161^***	0.217^***		0.379^***	0.392^***	0.374^*	0.497^***	0.576^**	0.510^**
23	0.099^***	0.097^***	0.128^***	0.114^***	0.079^***	0.072^***	0.074^***	0.106^***	0.065^***	0.058^***	0.071^***	0.052^***	0.050^***	0.089^***	0.059^***	0.073^***	0.039^***	0.093^***	0.139^***	0.184^***	0.225^***	0.188^***		0.063	0.128	0.222^*	0.295^*	0.195
24	0.113^***	0.098^***	0.117^***	0.146^***	0.104^***	0.080^**	0.103^***	0.110^***	0.105^**	0.055^*	0.098^***	0.071^**	0.081^***	0.073^***	0.083^***	0.091^**	0.061^***	0.050^**	0.158^***	0.202^***	0.237^***	0.192^***	0.071^***		0.009	0.166	0.325	0.234
25	0.134^**	0.130^***	0.167^***	0.158^***	0.082^***	0.136^***	0.138^***	0.158^**	0.170^**	0.118^***	0.128^***	0.118^***	0.120^***	0.129^***	0.133^***	0.133^**	0.149^***	0.137^***	0.157^***	0.215^***	0.236^***	0.242^***	0.141^***	0.106^**		0.145	0.301^*	0.235^*
26	0.114^***	0.088^***	0.134^***	0.140^***	0.103^***	0.091^***	0.128^***	0.120^***	0.120^***	0.082^***	0.052^***	0.058^***	0.065^***	0.087^***	0.084^***	0.100^***	0.088^***	0.095^***	0.151^***	0.187^***	0.214^***	0.212^***	0.082^***	0.063^***	0.117^***		0.090	0.076
27	0.137^***	0.137^***	0.167^***	0.157^***	0.090^***	0.123^***	0.128^***	0.170^***	0.131^***	0.113^***	0.076^***	0.093^***	0.086^***	0.130^***	0.120^***	0.134^***	0.111^***	0.147^***	0.150^***	0.238^***	0.256^***	0.278^***	0.123^***	0.143^***	0.146^***	0.071^***		0.023
28	0.089^***	0.129^***	0.167^***	0.126^***	0.081^***	0.114^***	0.138^***	0.132^***	0.168^***	0.110^***	0.109^***	0.108^***	0.104^***	0.118^***	0.113^***	0.132^***	0.133^***	0.157^***	0.114^***	0.195^***	0.207^***	0.248^***	0.135^***	0.150^***	0.100^***	0.105^***	0.090^***

变异来源 Source of variation	自由度 d.f.	离差平方和(SSD) Sum of squares	方差分量 Variance components	方差分量比例 Percentage of variation(%)	P-value
地区间 Among regions	3	100.241	0.03734	1.68	P = 0.00196
地区内居群间 Among populations within regions	24	408.499	0.24851	11.16	P<0.001
居群内个体间 Within populations	1,670	3,233.717	1.93636	87.16	P<0.001