Genetic structure of populations and introgression of three Quercus species in mountainous area of Beijing
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摘要:目的 栎属植物种间经常发生种间杂交和基因渐渗现象，特别是同域分布的同组内栎树之间，这种情况会更加频繁。本文通过对北京山区3种栎属植物居群遗传结构遗传变异与遗传结构进行研究，为了解北京地区自然分布的栎属植物种间基因渐渗情况、种质资源现状以及经营管理提供有效数据。方法 本文使用6对SSR引物对云蒙山、上方山和北农林场同域分布的304个蒙古栎、槲树、槲栎的居群遗传多样性、遗传结构和种间基因渐渗进行了研究。结果 共检测到等位标记105个，每个位点的平均等位标记数（Na）为17.5个，期望杂合度（He）为0.660 ~ 0.911，平均为0.838，多态性信息含量指数（PIC）为0.632 ~ 0.903，平均为0.822，3种栎树在总体水平上具有较高的遗传多样性。在种的水平上，3种栎树的平均等位标记数（Na）为12.667 ~ 14.167，期望杂合度（He）为0.743 ~ 0.849，多态性信息含量指数（PIC）为0.725 ~ 0.826，3种栎属植物的遗传多样性水平为蒙古栎 > 槲树 > 槲栎。对7个栎树居群的遗传结构分析表明，遗传变异大部分发生在居群内。通过Structure软件对3种栎树种间基因渐渗进行分析，发现槲树−槲栎、槲树−蒙古栎和槲栎−蒙古栎这3个种对间均有基因渐渗发生。结论 在北京山区分布的蒙古栎、槲树和槲栎这3种栎属植物之间存在普遍的、复杂的渐渗杂交现象。Abstract:Objective Interspecific hybridization and introgression often occur among species of Quercus, especially among sympatric Quercus more frequently. In this paper, the genetic structure and genetic variation of three species of Quercus in the mountainous area of Beijing were studied. It will provide effective data for gene introgression, germplasm resources and management of Quercus species naturally distributed in Beijing.Method 304 natural oak samples of Quercus mongolica, Quercus dentate and Quercus aliena from Yunmeng Mountain, Shangfang Mountain and BUA (Beijing University of Agriculture) Forest Farm were selected as the material, and six pairs of SSR primers were used to study the genetic diversity, genetic structure and introgressive hybridization among populations.Result A total of 105 alleles were detected, the average number of alleles (Na) of each locus was 17.5, the expected heterozygosity (He) was 0.660−0.911, the polymorphism information content index (PIC) was 0.632−0.903. The three oaks had high genetic diversity at the overall level. At the species level, the average allele number (Na) of the three oaks was 12.667−14.167, the expected heterozygosity (He) was 0.743−0.849, the polymorphism information content index (PIC) was 0.725−0.826, the genetic diversity level of the three species of oaks was Q. mongolica > Q. dentate > Q. aliena. The genetic structure analysis of 7 populations of Quercus showed that most of the genetic variation was in local populations. The genetic introgression analysis by structure software found that the gene introgression occurred among three pairs of Q. mongolica, Q. dentate and Q. aliena.Conclusion There is a common and complex phenomenon of introgression hybridization among Q. mongolica, Q. dentate and Q. aliena in Beijing mountainous area.
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Keywords:
- Quercus /
- genetic diversity /
- genetic structure /
- gene introgression
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表 1 6对SSR引物多态性
Table 1 Ploymorphism of 6 SSR primers
位点
Locus引物（5′—3′）
Primer（5′−3′）重复片段
Repeat motif等位标记数
Number of alleles（Na）观测杂合度
Observation heterozygosity
（Ho）期望杂合度
Expected heterozygosity
（He）多态信息量
Polymorphism information content（PIC）Qden 03011 AACCCAACCTTCCCTTCATC （AG）8 15.0 0.823 0.836 0.817 GCAGTGGTGCCTAATGTAGAC Qden 05011 CCCACTCCCTGTCCATTGT （CT）8 19.0 0.881 0.893 0.882 CACTGTGTGCTGCGACTTG Qden 03032 AGTTGTGGTCCTGCTCGC （CT）12 16.0 0.618 0.857 0.841 GAAAAGTGCGATGACGGTTG Qden 05031 CCCCGATTCGCCATCATTGT （GT）12 19.0 0.846 0.873 0.859 GTAACGCCGTTTTTCTCCACC ssrQpZAG112 TTCTTGCTTTGGTGCGCG （GA）32 11.0 0.485 0.660 0.632 GTGGTCAGAGACTCGGTAAGTATTC ssrQpZAG96 CCCAGTCACATCCACTACTGTCC （TC）20 25.0 0.881 0.911 0.903 GGTTGGGAAAAGGAGATCAGA 平均值 Mean 17.5 0.755 0.838 0.822 
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表 2 7个栎树居群的遗传多样性
Table 2 Genetic diversity of 7 populations of oak
物种
Speices居群
Population样本数
Sample numberNa Ho He 近交系数
Inbreeding coefficient（F）PIC 槲树
Quercus dentateYM-D 45 9.833 0.819 0.762 − 0.086 0.718 SF-D 33 10.833 0.758 0.797 0.036 0.760 平均值 Mean 78 12.667 0.793 0.787 − 0.014 0.754 槲栎
Quercus alienaYM-A 27 7.833 0.685 0.671 − 0.027 0.629 LC-A 53 8.000 0.657 0.683 0.063 0.650 SF-A 71 12.833 0.753 0.771 0.007 0.750 平均值 Mean 151 14.167 0.706 0.743 0.046 0.725 蒙古栎
Quercus mongolicaYM-M 43 11.667 0.797 0.831 0.022 0.800 LC-M 32 9.833 0.854 0.839 − 0.031 0.806 平均值 Mean 75 13.167 0.826 0.849 0.016 0.826 注：YM. 云蒙山样地，SF. 上方山样地，LC.北农林场样地，D.槲树，A.槲栎，M.蒙古栎。Notes: YM refers to Yunmeng Mountain sample plot; SF refers to Shangfang Mountain sample plot; LC refers to BUA Forest Farm sample plot；D refers to Quercus dentate; A refers to Quercus aliena; M refers to Quercus mongolica.
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表 3 同种栎树居群间遗传变异和基因流
Table 3 Genetic variation and Nm values between populations within same oak species
居群1
Population 1居群2
Population 2基因分化系数
Genetic differentiation coefficient（Fst）基因流
Gene flow（Nm）YM-D SF-D 0.019 12.710 YM-A LC-A 0.044 5.472 YM-A SF-A 0.032 7.649 LC-A SF-A 0.020 12.027 YM-M LC-M 0.025 9.924
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表 4 异种栎树居群间遗传变异和基因流
Table 4 Genetic variation and Nm values between populations among varied oak species
居群1 Population 1 居群2 Population 2 Fst Nm YM-D YM-A 0.101 2.237 YM-D YM-M 0.070 3.342 YM-A YM-M 0.093 2.445 LC-A LC-M 0.108 2.068 SF-D SF-A 0.045 5.271
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表 5 居群间遗传距离
Table 5 Genetic distance between populations
居群 Population LC-A LC-M SF-A SF-D YM-A YM-D YM-M LC-A — LC-M 0.427 — SF-A 0.160 0.285 — SF-D 0.374 0.343 0.196 — YM-A 0.218 0.368 0.164 0.286 — YM-D 0.448 0.394 0.264 0.131 0.334 — YM-M 0.400 0.178 0.231 0.268 0.305 0.340 —
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