毛茛科植物花形态发育性状的演化研究
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摘要
花是被子植物特有的繁殖器官,没有花的进化就没有被子植物的繁盛和多样化。花的起源和演化一直是系统与进化植物学家研究的焦点和热点,相对于成熟结构,花器官在发生发育过程中表现出了更为丰富的系统演化信息。毛茛科是联系基部被子植物和核心真双子叶植物的关键类群,花器官表现出了丰富的多样性,是研究花起源和演化的理想类群。本文以毛茛科植物为研究对象,在系统发育框架重建和类群分化时间估算的基础上,对花形态发育性状的演化趋势进行了深入分析,主要内容如下：
(1)利用扫描电子显微镜(SEM)对毛茛科Ranunculaceae 6族12属12种植物花器官的形态发生和发育过程进行了全面或补充研究,进一步丰富了毛茛科花器官发生发育的基本资料。
(2)利用NCBI数据库中的叶绿体rbcL, matK, trnL-F三个基因/DNA片段联合分析,分别用MP、ML、Bayes三种方法重建了毛茛科的系统发育树,结果表明前人认为的毛茛亚科不是单系,唐松草亚科镶嵌于毛茛亚科内,和该亚科的部分成员为姊妹群关系,花发育的证据表明唐松草亚科无区别于毛茛亚科的独征,我们建议将唐松草亚科归并入毛茛亚科,因此毛茛科应包括4个亚科,Glaucidioideae, Hydrastidoideae,黄连亚科Coptidoideae和毛茛亚科Ranunculoideae。
(3)利用Beast软件估算了毛茛科的分子钟,指出毛茛科内四个亚科级分支在晚白垩世时期出现(90-68.97 Ma)；随后族级分支集中于早古新世-早始新世时期出现(62.22-51.3 Ma)；属级分支集中于晚始新世-中新世时期出现(38.55-5.17 Ma)；推测毛茛科亚科级的祖先类群是白垩纪生物大灭绝后的残余类群,而族和属在两个时期的快速辐射分化与地球的两次温度变暖相关。
(4)基于毛茛科系统发育框架,利用Mesquite软件重建了一些花发育性状的演化历史,研究表明：
花发育性状的平行演化现象普遍存在于不同的分类阶元。
黄连亚科和毛茛亚科的共同祖先存在花瓣,无花瓣出现在不同的分支中,表明是次生性丢失和平行演化的结果；在铁线莲属和白头翁属中有花瓣被认为是花瓣丢失后再重新获得的结果。
68.97 Ma分化出的黄连亚科和毛茛亚科共同祖先的花瓣发育过程中没有附属结构,蜜腺组织集中于表面的特定区域,而花瓣在发育过程中腹面凹陷下方产生两枚凸起是毛茛亚科的共同祖征,一枚凸起的产生是次生的结果；两枚凸起通常与花瓣瓣片相互愈合(或不愈合),进而形成距、囊等结构,一枚凸起通常发育为鳞片状,二者都加强了对蜜腺组织的保护作用,进而增强了对传粉者选择的专一性,这些现象在不同分支中出现是平行演化的结果。
对折心皮是毛茛科心皮的共同祖征,囊状心皮在不同分支中独立发生是次生的结果；心皮的中等数目(5-10)是祖先状态,向数目减少和增多两个方向演化；心皮数目减少和器官轮状排列相关,心皮数目增加和器官不规则排列相关。
辐射对称是毛茛科的共同祖征,约53.19 Ma前分化出的翠雀族花表现出的两侧对称是花瓣退化导致的次生性状,花瓣退化是该族的独征。
花器官轮状排列是毛茛科的祖先状态,而85.15 Ma分化出的黄连亚科和毛茛亚科的共同祖先为螺旋状排列；轮状排列出现在该科系统演化的不同分支上,是平行演化的结果；54.94 Ma分化出的毛茛族和银莲花族花器官排列不规则式样的出现是雄蕊和心皮数目次生增加的结果。
Flowers are the reproductive organs of angiosperms, and they play a crucial role in the diversity of angiosperms. The issue of floral origin and evolution has been a long-standed question for the systematic and evolutionary botanists. There is much more important evolutionary information during floral devlopment comparing with the mature structures. Ranunculaceae, which links basal angiosperms and core eudicots, has shown great floral morphological diversity. Ranunculaceae is thus ideally suited for examination of floral origin and evolution. Reconstruction of the phylogeny and bayesian dating are conducted based on three regions in plastid and evolution of floral developmental characters are analysed, and the results are as follows:
(1) The development of flowers of 12 species from 12 genera,6 tribes in Ranunculaceae were studied using scanning electron microscopy (SEM).
(2) Plastid (rbcL, matK and trnL-F) sequences were analyzed with parsimony, likelihood, and bayesian inference. Ranunculoideae is paraphyletic and Thalictroideae is nested within the Ranunculoideae clade. The Thalictroideae has not autapomorphy in flora development and should be merged with Ranunculoideae. Ranunculaceae is thus comprised of four subfamilies, i.e. Glaucidioideae, Hydrastidoideae, Coptidoideae and Ranunculoideae.
(3) Divergence times of disjunct lineages were estimated with relaxed Bayesian dating. The divergence of the four subfamilies occurred 90-68.97 Ma (late Cretaceous); the divergence of the tribes occurred 62.22-51.3 Ma (early Paleocene to early Eocene); the divergence of the genera occurred 38.55-5.17 Ma (late Eocene to Miocene). The ancestor taxa of the subfamilies maybe the relictual lineage after the mass extinction in Cretaceous. The quick radiation of the tribes and genera may be related with temperature warming.
(4) Evolution of the floral developmental characters was assessed by tracing these characters onto Bayesian trees using the Trace Character Over Trees option in the program Mesquite, and the results showed that:
Parallel evolution occurs for different characters in different taxa. The petals are present in the ancestor node of Coptidoideae and Ranunculoideae, while the petals are absent in several unrelated clades, which is the results of secondary missing and parallel evolution. The petals appear again in some species of Clematis and Pulsatilla is the result of secondary acquisition.
There are no appendages during the petal development in the ancestor of Coptidoideae and Ranunculoideae, and the nectar tissue appears on a specific region. During the petal development in the ancestor of Ranunculoideae, a shallow depression appears firstly in the centre of the ventral side, then two bulges arise in a short succession at the base of the depression. Only one bulge at the base of the depression is considered to be derived. The two bulges always become connate and then fuse with the blade and forming a tubular, spur-shaped or sac shaped structure where the nectar tissue appears. The single bulge always develops into the small scale beneath which the nectar tissue appears. We consider that the developmental result of the appendage may be a highly modified structural and functional feature for concealing the nectar and controlling the access of the visitors.
Epeltate carpel is the plesiomorphy while ascidiate carpel in unrelated clades is the apomorph in Ranunculaceae. The moderate number of carpels is the plesiomorphy and less or more numbers are derived. The less number of carpels is related with whorled phyllotaxis and the large number is related with spiral phyllotaxis.
Radial symmetry is plesiomorphy in Ranunculaceae. Bisymmetry in Delphinieae is caused by the reduction of several petal primordia. Bisymmetry is thus derived characters and the autapomorphy of the tribe Delphinieae.
Wholed phyllotaxis for floral organs is the plesiomorphy in Ranunculaceae, and the floral phyllotaxis are spiral in the ancestral taxa of Coptidoideae and Ranunculoideae. The whorled phyllotaxis are present in different clades, which is the result of parallel or convergent evolution. The irregular phyllotaxis in Ranunculeae and Anemoneae is caused by redundant stamens and carpels.
(1)利用扫描电子显微镜(SEM)对毛茛科Ranunculaceae 6族12属12种植物花器官的形态发生和发育过程进行了全面或补充研究,进一步丰富了毛茛科花器官发生发育的基本资料。
(2)利用NCBI数据库中的叶绿体rbcL, matK, trnL-F三个基因/DNA片段联合分析,分别用MP、ML、Bayes三种方法重建了毛茛科的系统发育树,结果表明前人认为的毛茛亚科不是单系,唐松草亚科镶嵌于毛茛亚科内,和该亚科的部分成员为姊妹群关系,花发育的证据表明唐松草亚科无区别于毛茛亚科的独征,我们建议将唐松草亚科归并入毛茛亚科,因此毛茛科应包括4个亚科,Glaucidioideae, Hydrastidoideae,黄连亚科Coptidoideae和毛茛亚科Ranunculoideae。
(3)利用Beast软件估算了毛茛科的分子钟,指出毛茛科内四个亚科级分支在晚白垩世时期出现(90-68.97 Ma)；随后族级分支集中于早古新世-早始新世时期出现(62.22-51.3 Ma)；属级分支集中于晚始新世-中新世时期出现(38.55-5.17 Ma)；推测毛茛科亚科级的祖先类群是白垩纪生物大灭绝后的残余类群,而族和属在两个时期的快速辐射分化与地球的两次温度变暖相关。
(4)基于毛茛科系统发育框架,利用Mesquite软件重建了一些花发育性状的演化历史,研究表明：
花发育性状的平行演化现象普遍存在于不同的分类阶元。
黄连亚科和毛茛亚科的共同祖先存在花瓣,无花瓣出现在不同的分支中,表明是次生性丢失和平行演化的结果；在铁线莲属和白头翁属中有花瓣被认为是花瓣丢失后再重新获得的结果。
68.97 Ma分化出的黄连亚科和毛茛亚科共同祖先的花瓣发育过程中没有附属结构,蜜腺组织集中于表面的特定区域,而花瓣在发育过程中腹面凹陷下方产生两枚凸起是毛茛亚科的共同祖征,一枚凸起的产生是次生的结果；两枚凸起通常与花瓣瓣片相互愈合(或不愈合),进而形成距、囊等结构,一枚凸起通常发育为鳞片状,二者都加强了对蜜腺组织的保护作用,进而增强了对传粉者选择的专一性,这些现象在不同分支中出现是平行演化的结果。
对折心皮是毛茛科心皮的共同祖征,囊状心皮在不同分支中独立发生是次生的结果；心皮的中等数目(5-10)是祖先状态,向数目减少和增多两个方向演化；心皮数目减少和器官轮状排列相关,心皮数目增加和器官不规则排列相关。
辐射对称是毛茛科的共同祖征,约53.19 Ma前分化出的翠雀族花表现出的两侧对称是花瓣退化导致的次生性状,花瓣退化是该族的独征。
花器官轮状排列是毛茛科的祖先状态,而85.15 Ma分化出的黄连亚科和毛茛亚科的共同祖先为螺旋状排列；轮状排列出现在该科系统演化的不同分支上,是平行演化的结果；54.94 Ma分化出的毛茛族和银莲花族花器官排列不规则式样的出现是雄蕊和心皮数目次生增加的结果。
Flowers are the reproductive organs of angiosperms, and they play a crucial role in the diversity of angiosperms. The issue of floral origin and evolution has been a long-standed question for the systematic and evolutionary botanists. There is much more important evolutionary information during floral devlopment comparing with the mature structures. Ranunculaceae, which links basal angiosperms and core eudicots, has shown great floral morphological diversity. Ranunculaceae is thus ideally suited for examination of floral origin and evolution. Reconstruction of the phylogeny and bayesian dating are conducted based on three regions in plastid and evolution of floral developmental characters are analysed, and the results are as follows:
(1) The development of flowers of 12 species from 12 genera,6 tribes in Ranunculaceae were studied using scanning electron microscopy (SEM).
(2) Plastid (rbcL, matK and trnL-F) sequences were analyzed with parsimony, likelihood, and bayesian inference. Ranunculoideae is paraphyletic and Thalictroideae is nested within the Ranunculoideae clade. The Thalictroideae has not autapomorphy in flora development and should be merged with Ranunculoideae. Ranunculaceae is thus comprised of four subfamilies, i.e. Glaucidioideae, Hydrastidoideae, Coptidoideae and Ranunculoideae.
(3) Divergence times of disjunct lineages were estimated with relaxed Bayesian dating. The divergence of the four subfamilies occurred 90-68.97 Ma (late Cretaceous); the divergence of the tribes occurred 62.22-51.3 Ma (early Paleocene to early Eocene); the divergence of the genera occurred 38.55-5.17 Ma (late Eocene to Miocene). The ancestor taxa of the subfamilies maybe the relictual lineage after the mass extinction in Cretaceous. The quick radiation of the tribes and genera may be related with temperature warming.
(4) Evolution of the floral developmental characters was assessed by tracing these characters onto Bayesian trees using the Trace Character Over Trees option in the program Mesquite, and the results showed that:
Parallel evolution occurs for different characters in different taxa. The petals are present in the ancestor node of Coptidoideae and Ranunculoideae, while the petals are absent in several unrelated clades, which is the results of secondary missing and parallel evolution. The petals appear again in some species of Clematis and Pulsatilla is the result of secondary acquisition.
There are no appendages during the petal development in the ancestor of Coptidoideae and Ranunculoideae, and the nectar tissue appears on a specific region. During the petal development in the ancestor of Ranunculoideae, a shallow depression appears firstly in the centre of the ventral side, then two bulges arise in a short succession at the base of the depression. Only one bulge at the base of the depression is considered to be derived. The two bulges always become connate and then fuse with the blade and forming a tubular, spur-shaped or sac shaped structure where the nectar tissue appears. The single bulge always develops into the small scale beneath which the nectar tissue appears. We consider that the developmental result of the appendage may be a highly modified structural and functional feature for concealing the nectar and controlling the access of the visitors.
Epeltate carpel is the plesiomorphy while ascidiate carpel in unrelated clades is the apomorph in Ranunculaceae. The moderate number of carpels is the plesiomorphy and less or more numbers are derived. The less number of carpels is related with whorled phyllotaxis and the large number is related with spiral phyllotaxis.
Radial symmetry is plesiomorphy in Ranunculaceae. Bisymmetry in Delphinieae is caused by the reduction of several petal primordia. Bisymmetry is thus derived characters and the autapomorphy of the tribe Delphinieae.
Wholed phyllotaxis for floral organs is the plesiomorphy in Ranunculaceae, and the floral phyllotaxis are spiral in the ancestral taxa of Coptidoideae and Ranunculoideae. The whorled phyllotaxis are present in different clades, which is the result of parallel or convergent evolution. The irregular phyllotaxis in Ranunculeae and Anemoneae is caused by redundant stamens and carpels.
引文
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