蜡梅属植物谱系地理学、系统发育及遗传多样性研究
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摘要
蜡梅属(Chimonanthus Lindl.)隶属于蜡梅科(Calycanthaceae),是我国特有的多年生灌木,具有重要的观赏价值和药用价值。主要分布在秦岭以南、云贵高原以西、南岭以北的亚热带河谷山地区域。现今种群的分布区范围很小,易受扰动。蜡梅属在上新世初分化形成,是新近纪(晚第三纪)孑遗植物,现存物种进化历史上受第四纪冰川的影响较大。本文利用叶绿体DNA (cpDNA)的IGS区.trnL-F、trnS-G、trnH-psbA和核糖体DNA (nrDNA)的ITS区对蜡梅属植物25个种群的遗传结构、亲缘地理、系统发育关系进行了研究。进一步利用古地质、古气候资料及末次冰期植被重建资料,推测蜡梅属植物在第四纪冰期的避难所。同时,通过ISSR分子标记对其中的两种蜡梅进行了遗传结构和遗传多样性分析,结合叶绿体序列数据,提出了蜡梅属不同物种不同种群相对应的保护策略。主要研究结果表述如下：
1)蜡梅属25个种群264个个体中,叶绿体cpDNA trnL-F、trnS-G、trnH-psbA联合数据分析结果共检测到56个变异位点,40种单倍型类型。cpDNA单倍型关系呈现明显的“双星状”分支关系,各种群的特有单倍型分别与内部的古老单倍型H1和H3相连。H1为柳叶蜡梅和浙江蜡梅种群所共有,主要分布于我国华东地区；H3为蜡梅种群所共有,主要分布于华中到西南地区。根据种群分子遗传变异分析,结合地质历史事件,推测现今蜡梅属各种群是在各自的避难所独立演化而来,形成了多个避难所,包括大巴山、巫山、武陵山、雪峰山、云贵高原、黄山、庐山和武夷山。蜡梅属植物在冰期后没有经历过明显的种群扩张事件。
2)采用cpDNA trnL-F和ITS两个片段,分别运用最大似然法、邻接法和最大简约法对蜡梅属下6种1变种进行了系统发育树的构建。蜡梅属中,首先分化出来的种包括蜡梅和贵州蜡梅,并且有较高的支持率(BS>81%)。合并的分子数据显示,形态上作为原变种的西南蜡梅没有与变种贵州蜡梅聚在一起,而是与其他5个类群聚为一支,表明西南蜡梅与贵州蜡梅亲缘关系较远。蜡梅属中的第二大分支中主要包括浙江蜡梅L、柳叶蜡梅P、山蜡梅K、突托蜡梅O和西南蜡梅N,除西南蜡梅分布在云南,其他物种均分布在华东和华中地区,交叉分布而且范围均较小。支持将柳叶蜡梅和浙江蜡梅归并为一种,并且支持率较高(BS>87%)。
3)叶绿体DNA3个片段的综合数据表明蜡梅属物种水平的单倍型多态性(h=0.90500士0.01200)和核酸多态性(π=0.00339士0.00013)相对较高。遗传变异主要存在于种群间,并且基因流较小(Fst=0.92736,Gst=0.66184,Nst=0.88072,Nm=0.13)。为了进一步分析两种古老单倍型所在种群的遗传多样性水平,本文还采用ISSR分子标记对蜡梅和柳叶蜡梅进行了研究。分析结果发现,两种蜡梅在种群水平上的遗传多样性不高(Ch.praecox:PPB=32.49%,h=0.0968,I=0.1488；Ch.salicifolius:PPB=34.25%,h=0.1192,I=0.1777),但物种水平上的遗传多样性相对较高(Ch.praecox:PPB=86.03%,h=0.1552,I=0.2635:Ch.salicifolius:PPB=81.25%,h=0.2401,I=0.3684)。遗传结构分析发现Ch.praecox:ΦST=0.4650,GST=0.3784,Nm=0.8213;Ch.salicifolius:ΦsT=0.5912,GsT=0.5036,Nm=0.4929。种子流的cpDNA和花粉流的ISSR对蜡梅属植物遗传多样性的丰富程度和遗传结构的分析检测结果较为一致。
4)母系遗传的cpDNA和核基因的ISSR分子标记均表明,种群间存在明显遗传分化,为了保护众多避难所地区的遗传多样性,应该主要实施就地保护策略。重点应该包括蜡梅的湖南石门SMF、湖北保康BKH、湖南吉首JSJ、浙江临安LAI和湖南新宁XNS共5个种群；柳叶蜡梅的浙江松阳SYC和江西广丰GFD共2个种群；浙江蜡梅的的福建光泽GZG种群；山蜡梅的广西资源ZYM和广西阳朔YSK共两个种群；突托蜡梅的江西会昌HCO种群；西南蜡梅的云南禄劝LQN种群。如果要进行迁地保护,应该根据不同种群的特点来取样。重点涵盖江西修水的柳叶蜡梅、贵州安龙的山蜡梅和贵州兴义的贵州蜡梅种群并尽可能采集更多的个体。
Chimonanthus Lindl. is a perennial shrub endemic to China with important ornamental and medicinal values, belongs to the family Calycanthaceae. The main distribution of wild populations is limited in the subtropical hills and valleys of the south of Qinling Mountains, the west of Yunnan-Guizhou Plateau, and the north of the Nanling Mountains regions. The present distributions of it are vulnerable to disturbance. The molecular clock estimates suggest that relict Ch. of the Neogene (late Tertiary) diverged from Calycanthus in the early Pliocene. The current genetic structure and geographical distribution were influenced significantly by the climate of Quaternary glacial.
Our present work investigated the genetic structure, phylogeography and phlogeny of Ch. including25populations from China. The analysis was carried out based on nuclear sequence data from the internal transcribed specer (ITS) region of the ribosomal DNA and chloroplast (cp) DNA data from trnL-F, trnS-G, and trnH-psbA intergenic spacer, located in the large single-copy region of the cp genome. Using of the ancient geological, paleoclimatic data and revegetation information of the last glacial period, the inference was made that the possible glacial refuge of Ch. plants in the Quaternary Ice Age. In additional, the genetic structure and genetic diversity were estimated based on inter-simple sequence repeat (ISSR) markers from Ch. praecox and Ch. salicifolius. Combined with results of the cpDNA sequence data, we proposed an appropriate protection strategy for different populations of different species. The main results were listed as follows:
1)25populations (264individuals) were analyzed using cpDNA trnL-F、trnS-G、and trnH-psbA intergenic spacer, and56variable sites were detected, which produced40haplotypes. Analysis of cpDNA haplotypes from different populations suggested that two phylogeographical groups existed a double star-like relationship in this genus. Populations from Dabashan, Wushan, Wulingshan Mountains, mainly distributed in the Central China to southwest, had a unique ancestral haplotypes (H3). Areas around Lushan, Tianmushan, Wuyishan, Xuefengshan mountains, mainly distributed in the East China, had another unique ancestral haplotypes (H3).All other private haplotypes from different populations originated from the two ancestral haplotypes. Accoding to the analysis of the molecular genetic variation of different populations, combined with the geological history events, we speculated that the present populations are evolved independently in their respective refuge, i.e. their present distribution areas. Neutrality test and mismatch distribution neither support any significant postglacial population expansion.
2) Chose trnL-F genes of chloroplast genome and ITS sequences of nuclear genome, we adopted maximum likelihood, maximum parsimony and neighbor joining method to resolve the systematic problems of6species and1variety of this genus. In the end, we combined both data of trnL-F and ITS to explain the relationships among these species. Ch. praecox and Ch. campanulatus var. guizhouensis diverged from this genus firstly grouped into one branch (BS>81%). And leftover of this genus grouped into another main branch including five species, such as Ch. salicifolius, Ch. grammatus, Ch. nitens, Ch. zhejiangensis, and Ch. campanulatus. Ch. campanulatus var guizhouensis determined previously as a variety of Ch. campanulatus by the flexible morphological and closed geographical characteristics. Inconsistent results with previous showed a far relationship in molecular evidence between them. Our results supported (BS>87%) that Ch. zhejiangensis was treated as a synonyms of Ch. salicifolius.
3)According to cpDNA data, haplotype diversity (h=0.90500±0.01200) and nucleotide diversity (π=0.00339±0.00013) of the genus is relatively high. In order to estimate the genetic diversity of populations from two ancestral haplotypes, two species from17populations were chose based on ISSR. For both species, the genetic diversity was low at the species level(Ch.praecox:PPB=32.49%, h=0.0968,I=0.1488; Ch. salicifolius:PPB=34.25%, h=0.1192,I=0.1777), but relatively higher at the species level (Ch. praecox:PPB=86.03%, h=0.1552,I=0.0.2635; Ch. salicifolius:PPB=81.25%, h=0.2401,I=0.3684). The hierarchical AMOVA revealed high levels of among-population genetic differentiation in both species, in line with the gene differentiation coefficient and the limited among-population gene flow (Ch. praecox:ΦST=0.4650, GST=0.3784, Nm=0.8213; Ch. salicifolius:ΦST=0.5912, GST=0.5036, Nm=0.4929). Both neighbor-joining (NJ) cluster analysis and Principal Coordinate Analysis (PCoA) clustered all17populations into two major groups, which corresponded to the two separate species. An isolation-by-distance pattern was revealed in Ch. salicifolius (r=0.703, P=0.048,999permutations), but not in Ch. praecox (r=0.168, P=0.185,999permutations). For the respect of description of the richness of the species genetic diversity, results from cpDNA and ISSR data are consistent.
4) Maternal inheritance of cpDNA and nuclear gene ISSR markers showed a significant differentiation among these populations. In order to protect the genetic diversity of these plants distributed in these refuge areas, we should use in situ conservation strategies. Highlights include the following populations:SMF, BKH, JSJ, LAI, XNS of Ch. praecx; SYC and GFD of Ch. salicifolius; GZG of Ch. zhejiangensis; ZYM and YSK of Ch. nitens; HCO of Ch. grammatus; LQN of Ch. campanulatus. If need an ex situ conservation strategy in emergency, sampling method should be in the light of the characteristics of above-metioned populations, also should include XSZ of Jiangxi Province, ALU and XYT of Guizhou Province, especially.
1)蜡梅属25个种群264个个体中,叶绿体cpDNA trnL-F、trnS-G、trnH-psbA联合数据分析结果共检测到56个变异位点,40种单倍型类型。cpDNA单倍型关系呈现明显的“双星状”分支关系,各种群的特有单倍型分别与内部的古老单倍型H1和H3相连。H1为柳叶蜡梅和浙江蜡梅种群所共有,主要分布于我国华东地区；H3为蜡梅种群所共有,主要分布于华中到西南地区。根据种群分子遗传变异分析,结合地质历史事件,推测现今蜡梅属各种群是在各自的避难所独立演化而来,形成了多个避难所,包括大巴山、巫山、武陵山、雪峰山、云贵高原、黄山、庐山和武夷山。蜡梅属植物在冰期后没有经历过明显的种群扩张事件。
2)采用cpDNA trnL-F和ITS两个片段,分别运用最大似然法、邻接法和最大简约法对蜡梅属下6种1变种进行了系统发育树的构建。蜡梅属中,首先分化出来的种包括蜡梅和贵州蜡梅,并且有较高的支持率(BS>81%)。合并的分子数据显示,形态上作为原变种的西南蜡梅没有与变种贵州蜡梅聚在一起,而是与其他5个类群聚为一支,表明西南蜡梅与贵州蜡梅亲缘关系较远。蜡梅属中的第二大分支中主要包括浙江蜡梅L、柳叶蜡梅P、山蜡梅K、突托蜡梅O和西南蜡梅N,除西南蜡梅分布在云南,其他物种均分布在华东和华中地区,交叉分布而且范围均较小。支持将柳叶蜡梅和浙江蜡梅归并为一种,并且支持率较高(BS>87%)。
3)叶绿体DNA3个片段的综合数据表明蜡梅属物种水平的单倍型多态性(h=0.90500士0.01200)和核酸多态性(π=0.00339士0.00013)相对较高。遗传变异主要存在于种群间,并且基因流较小(Fst=0.92736,Gst=0.66184,Nst=0.88072,Nm=0.13)。为了进一步分析两种古老单倍型所在种群的遗传多样性水平,本文还采用ISSR分子标记对蜡梅和柳叶蜡梅进行了研究。分析结果发现,两种蜡梅在种群水平上的遗传多样性不高(Ch.praecox:PPB=32.49%,h=0.0968,I=0.1488；Ch.salicifolius:PPB=34.25%,h=0.1192,I=0.1777),但物种水平上的遗传多样性相对较高(Ch.praecox:PPB=86.03%,h=0.1552,I=0.2635:Ch.salicifolius:PPB=81.25%,h=0.2401,I=0.3684)。遗传结构分析发现Ch.praecox:ΦST=0.4650,GST=0.3784,Nm=0.8213;Ch.salicifolius:ΦsT=0.5912,GsT=0.5036,Nm=0.4929。种子流的cpDNA和花粉流的ISSR对蜡梅属植物遗传多样性的丰富程度和遗传结构的分析检测结果较为一致。
4)母系遗传的cpDNA和核基因的ISSR分子标记均表明,种群间存在明显遗传分化,为了保护众多避难所地区的遗传多样性,应该主要实施就地保护策略。重点应该包括蜡梅的湖南石门SMF、湖北保康BKH、湖南吉首JSJ、浙江临安LAI和湖南新宁XNS共5个种群；柳叶蜡梅的浙江松阳SYC和江西广丰GFD共2个种群；浙江蜡梅的的福建光泽GZG种群；山蜡梅的广西资源ZYM和广西阳朔YSK共两个种群；突托蜡梅的江西会昌HCO种群；西南蜡梅的云南禄劝LQN种群。如果要进行迁地保护,应该根据不同种群的特点来取样。重点涵盖江西修水的柳叶蜡梅、贵州安龙的山蜡梅和贵州兴义的贵州蜡梅种群并尽可能采集更多的个体。
Chimonanthus Lindl. is a perennial shrub endemic to China with important ornamental and medicinal values, belongs to the family Calycanthaceae. The main distribution of wild populations is limited in the subtropical hills and valleys of the south of Qinling Mountains, the west of Yunnan-Guizhou Plateau, and the north of the Nanling Mountains regions. The present distributions of it are vulnerable to disturbance. The molecular clock estimates suggest that relict Ch. of the Neogene (late Tertiary) diverged from Calycanthus in the early Pliocene. The current genetic structure and geographical distribution were influenced significantly by the climate of Quaternary glacial.
Our present work investigated the genetic structure, phylogeography and phlogeny of Ch. including25populations from China. The analysis was carried out based on nuclear sequence data from the internal transcribed specer (ITS) region of the ribosomal DNA and chloroplast (cp) DNA data from trnL-F, trnS-G, and trnH-psbA intergenic spacer, located in the large single-copy region of the cp genome. Using of the ancient geological, paleoclimatic data and revegetation information of the last glacial period, the inference was made that the possible glacial refuge of Ch. plants in the Quaternary Ice Age. In additional, the genetic structure and genetic diversity were estimated based on inter-simple sequence repeat (ISSR) markers from Ch. praecox and Ch. salicifolius. Combined with results of the cpDNA sequence data, we proposed an appropriate protection strategy for different populations of different species. The main results were listed as follows:
1)25populations (264individuals) were analyzed using cpDNA trnL-F、trnS-G、and trnH-psbA intergenic spacer, and56variable sites were detected, which produced40haplotypes. Analysis of cpDNA haplotypes from different populations suggested that two phylogeographical groups existed a double star-like relationship in this genus. Populations from Dabashan, Wushan, Wulingshan Mountains, mainly distributed in the Central China to southwest, had a unique ancestral haplotypes (H3). Areas around Lushan, Tianmushan, Wuyishan, Xuefengshan mountains, mainly distributed in the East China, had another unique ancestral haplotypes (H3).All other private haplotypes from different populations originated from the two ancestral haplotypes. Accoding to the analysis of the molecular genetic variation of different populations, combined with the geological history events, we speculated that the present populations are evolved independently in their respective refuge, i.e. their present distribution areas. Neutrality test and mismatch distribution neither support any significant postglacial population expansion.
2) Chose trnL-F genes of chloroplast genome and ITS sequences of nuclear genome, we adopted maximum likelihood, maximum parsimony and neighbor joining method to resolve the systematic problems of6species and1variety of this genus. In the end, we combined both data of trnL-F and ITS to explain the relationships among these species. Ch. praecox and Ch. campanulatus var. guizhouensis diverged from this genus firstly grouped into one branch (BS>81%). And leftover of this genus grouped into another main branch including five species, such as Ch. salicifolius, Ch. grammatus, Ch. nitens, Ch. zhejiangensis, and Ch. campanulatus. Ch. campanulatus var guizhouensis determined previously as a variety of Ch. campanulatus by the flexible morphological and closed geographical characteristics. Inconsistent results with previous showed a far relationship in molecular evidence between them. Our results supported (BS>87%) that Ch. zhejiangensis was treated as a synonyms of Ch. salicifolius.
3)According to cpDNA data, haplotype diversity (h=0.90500±0.01200) and nucleotide diversity (π=0.00339±0.00013) of the genus is relatively high. In order to estimate the genetic diversity of populations from two ancestral haplotypes, two species from17populations were chose based on ISSR. For both species, the genetic diversity was low at the species level(Ch.praecox:PPB=32.49%, h=0.0968,I=0.1488; Ch. salicifolius:PPB=34.25%, h=0.1192,I=0.1777), but relatively higher at the species level (Ch. praecox:PPB=86.03%, h=0.1552,I=0.0.2635; Ch. salicifolius:PPB=81.25%, h=0.2401,I=0.3684). The hierarchical AMOVA revealed high levels of among-population genetic differentiation in both species, in line with the gene differentiation coefficient and the limited among-population gene flow (Ch. praecox:ΦST=0.4650, GST=0.3784, Nm=0.8213; Ch. salicifolius:ΦST=0.5912, GST=0.5036, Nm=0.4929). Both neighbor-joining (NJ) cluster analysis and Principal Coordinate Analysis (PCoA) clustered all17populations into two major groups, which corresponded to the two separate species. An isolation-by-distance pattern was revealed in Ch. salicifolius (r=0.703, P=0.048,999permutations), but not in Ch. praecox (r=0.168, P=0.185,999permutations). For the respect of description of the richness of the species genetic diversity, results from cpDNA and ISSR data are consistent.
4) Maternal inheritance of cpDNA and nuclear gene ISSR markers showed a significant differentiation among these populations. In order to protect the genetic diversity of these plants distributed in these refuge areas, we should use in situ conservation strategies. Highlights include the following populations:SMF, BKH, JSJ, LAI, XNS of Ch. praecx; SYC and GFD of Ch. salicifolius; GZG of Ch. zhejiangensis; ZYM and YSK of Ch. nitens; HCO of Ch. grammatus; LQN of Ch. campanulatus. If need an ex situ conservation strategy in emergency, sampling method should be in the light of the characteristics of above-metioned populations, also should include XSZ of Jiangxi Province, ALU and XYT of Guizhou Province, especially.
引文
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