中国串珠藻属植物分子系统学研究
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摘要
本论文通过分子生物学、形态学、分类学、生物信息学和统计学的一些手段,对中国串珠藻属植物的形态学特征、区系分布特点以及分子系统发育等方面进行了研究。对中国串珠藻属植物30个分类单位的形态特征进行了描述,对其区系分布特点进行了总结；运用多重的分子数据,包括一个核基因组基因(SSU),一个叶绿体基因(rbcL)和两个线粒体基因(COI, cox2-3spacer),对串珠藻属植物的分子系统发生关系进行了探讨；对中国串珠藻属四个特有种进行了系统发育研究；对弧形串珠藻的进行了分子系统学研究并对其传统的分类特征提出了质疑。
通过对串珠藻样本的形态观察和测量显示,本研究所描述的串珠藻属植物的种类形态特征与国内、外描述的有较小差异。其中最主要的差异在于本研究所采集到的弧形串珠藻,有4个地点的标本为雌雄同株,与其雌雄异株的分类特征不相一致。
对中国串珠藻属植物区系研究的结果表明,目前我国的串珠藻属植物有30个分类单位,分布于20个省(市、自治区),是淡水红藻中最重要的类群。区系分析的结果包括8种分布区类型,其中以中国特有分布占绝对优势,有11种,3变种,占48.3%；温带性质明显,种类达到82.8%。种类数的分布以华东、华中和西南地区较为丰富,西北、华北和华东地区的分布相对较贫乏,特有种的分布也以华东和西南地区为多。
通过多重分子数据对中国的串珠藻属植物进行了分子系统学研究。结果显示,采自中国的6个种在各个系统发育树中所处的位置均与形态分类相一致,并且得到很好的统计学支持。基于rbcL序列的串珠藻属系统发育研究结果显示,红索藻目植物均独立于串珠藻目植物,构成一个单独的分支,支持红索藻目的建立。串珠藻属扭曲组与杂生组的差异度较小,结合其形态特点,倾向于将杂生组并入扭曲组。胶串珠藻作为串珠藻属串珠藻组的模式种,独立于其它串珠藻属植物,单独位于一个分支,这与其它分子序列得到的系统树的结果是一致的。SSU序列对多芒组、绿色组、沼生组和杂生组的分类关系没有得到很好的解决。基于COI序列的串珠藻属系统发育研究结果显示,扭曲组的种类聚集在一起,独立于其它串珠藻属植物,结合果胞枝弯曲或扭曲的形态特征,支持该组的重新定位。基于cox2-3spacer序列的串珠藻属系统发育研究结果显示,贵州所采集的扭曲串珠藻与形态研究的结果不一致,本该位于扭曲组的它,在系统树中位于串珠藻组的位置。但是由于该标本经过短时间固定,只测得了片段相对较小的cox2-3spacer序列,其系统位置有待于进一步研究。
对四种串珠藻属中国特有种进行了分子系统学研究,包括洪洞串珠藻、长柄串珠藻、弯形串珠藻和绞扭串珠藻。多重的分子数据包括一个核基因组基因(SSU),一个叶绿体基因(rbcL)和两个线粒体基因(COI, cox2-3spacer)被用来研究这四种串珠藻属中国特有种的系统发育关系。结果显示洪洞串珠藻和长柄串珠藻,从系统发育树上看,很明显属于曾经被划分为串珠藻组的种类,但与串珠藻组的模式种胶串珠藻的亲缘关系较远。因此,应将洪洞串珠藻和长柄串珠藻列入Sect. Helminthoidea。弯形串珠藻和绞扭串珠藻在所有的进化树中,都与扭曲组的种类位于同一分枝,并与串珠藻属并列,应该将其归入新属Kumanoa。
对7个种群的弧形串珠藻的形态特征及分子系统进行了研究。通过形态观察和聚类分析来评估种群间的个体差异。叶绿体rbcL序列和线粒体cox2-3spacer序列别用来评估种群间的遗传距离。形态观察的结果显示有4个种群的藻体为雌雄同株,其它3个种群的藻体为雌雄异株。聚类分析的结果显示,雌雄同株和雌雄异株的藻体没有分开,而地理距离较近生境相似的种群聚集在一起。同时,.rbcL序列和cox2-3spacer序列的结合序列所构建的系统树也证实了雌雄同株／异株不能作为一个典型的分类特征用于弧形串珠藻的鉴定。
The means of molecular biology, morphology, taxonomy, bioinformatics and statistics were used to determine the morphological character, flora, molecular systematic and phylogeny of Batrachospermum Roth in China. Morphological and floristics studies were made among30taxa of Batrachospermum in China. Multilocus molecular data from one nuclear gene (the ribosomal SSU), one chloroplast gene (rbcL), and two mitochondrial genes (COI, cox2-3spacer) were used to determine the phylogenetic placement of these taxa in Batrachospermum. Meanwhile, the studies on four endemic species and Batrachospermum arcuatum were also undertake to determine there phylogenetic relationship.
The studies on taxonomy and flora of Batrachospermum in China were made. Morphological studies show that Batrachospermum arcuatum specimens for4streams were monoecious; while for3streams were dioecious. The descriptions of other taxa are accord with the previous report. Thirty species (including varieties) of Batrachospermum were reported in China, which distributed in20provinces. The analysis of flora shows that there are8areal types of Batrachospermum, among which the endemic to China is predominant, including15species and accounting for48.3%. The temperature distribution of Batrachospermum is obvious and the number of the species takes82.8%totally. Batrachospermum is rich in East China, Central China and Southwest China; and rare in Northwest China, North China and Northeast China. Endemic species are distributed mainly in East China.
Multilocus molecular data were used to determine the phylogenetic relationship of taxa in genus Batrachospermum. The placement of the6taxa we collected in the phylogenetic trees was similar and the results accord with the morphological observation.The results based on rbcL sequence suggested that the placement of Thoreales was similar in each phylogenetic tree and formed a monophyletic group with high bootstrap support. Section Hybrida and section Contorta had a relatively small sequence divergence. The results of this work suggested combining section Hybrida as section Contorta. B. gelatinosum is the type of paraphyletic genus Batrachospermum. However, most of the species currently included in the section Batrachospermum are in a well-supported clade distant to B. gelatinosum. The same result was got with other sequence data. The results of phylogenetic relationship within Batrachospermum species based on SSU sequences showed that the taxonomic relationships of section Aristata, section Virescentia, section Turfosa and section Contorta had not been satisfactorily resolved. The results based on COI sequence showed that there was a good support for a clade of section Contorta species, which was apart from most Batrachospermum taxa. So we agreed with the proposal that taxa formerly in Batrachospermum sections Contorta and Hybrida were elevated into a new genus Kumanoa. The results based on cox2-3spacer sequence showed that Batrachospermum torsivum was sister to section Batrachospermum taxa, morphologically very distant entities. But only cox2-3spacer sequence of B. torsivum was got, and more sequence data were needed to solve this problem.
Four endemic Batrachospermum species in China, B. hongdongense, B. longipedicellatum, B. curvatum and B. intortum, were analyzed with multilocus data in order to assess their phylogenetic placement. Multilocus molecular data from one nuclear gene (the ribosomal SSU), one chloroplast gene (rbcL), and two mitochondrial genes (COI, cox2-3spacer) as well as morphological study were used to determine the phylogenetic relationships of these endemic species within the Batrachospermales. B. hongdongense and B. longipedicellatum had previously been placed in Batrachospermum section Batrachospermum, but based on the data from this study should be transferred into section Helminthoidea. B. curvatum and B. intortum clearly belong to the newly established genus Kumanoa.
Batrachospermum arcuatum specimens were analyzed from seven stream segments in North China. Morphological characteristics were observed and cluster analysis was used to evaluate the divergence among thalli from. Sequence data of the rbcL gene (chloroplast gene) and cox2-3spacer region (mitochondrial gene) were also utilized to evaluate genetic variation in specimens among stream segments. The specimens from four of the streams were monoecious, while the individuals at the other three locations were dioecious. Cluster analysis showed that the monoecious specimens were not separated from the dioecious specimens, based on morphology, but rather the specimens were grouped by geographical closeness and habitat similarity. Likewise, the combined analyses of rbcL and the cox2-3spacer data from provided more evidence that breeding system (monoecy vs. dioecy) is not a good morphological character to distinguish species.
通过对串珠藻样本的形态观察和测量显示,本研究所描述的串珠藻属植物的种类形态特征与国内、外描述的有较小差异。其中最主要的差异在于本研究所采集到的弧形串珠藻,有4个地点的标本为雌雄同株,与其雌雄异株的分类特征不相一致。
对中国串珠藻属植物区系研究的结果表明,目前我国的串珠藻属植物有30个分类单位,分布于20个省(市、自治区),是淡水红藻中最重要的类群。区系分析的结果包括8种分布区类型,其中以中国特有分布占绝对优势,有11种,3变种,占48.3%；温带性质明显,种类达到82.8%。种类数的分布以华东、华中和西南地区较为丰富,西北、华北和华东地区的分布相对较贫乏,特有种的分布也以华东和西南地区为多。
通过多重分子数据对中国的串珠藻属植物进行了分子系统学研究。结果显示,采自中国的6个种在各个系统发育树中所处的位置均与形态分类相一致,并且得到很好的统计学支持。基于rbcL序列的串珠藻属系统发育研究结果显示,红索藻目植物均独立于串珠藻目植物,构成一个单独的分支,支持红索藻目的建立。串珠藻属扭曲组与杂生组的差异度较小,结合其形态特点,倾向于将杂生组并入扭曲组。胶串珠藻作为串珠藻属串珠藻组的模式种,独立于其它串珠藻属植物,单独位于一个分支,这与其它分子序列得到的系统树的结果是一致的。SSU序列对多芒组、绿色组、沼生组和杂生组的分类关系没有得到很好的解决。基于COI序列的串珠藻属系统发育研究结果显示,扭曲组的种类聚集在一起,独立于其它串珠藻属植物,结合果胞枝弯曲或扭曲的形态特征,支持该组的重新定位。基于cox2-3spacer序列的串珠藻属系统发育研究结果显示,贵州所采集的扭曲串珠藻与形态研究的结果不一致,本该位于扭曲组的它,在系统树中位于串珠藻组的位置。但是由于该标本经过短时间固定,只测得了片段相对较小的cox2-3spacer序列,其系统位置有待于进一步研究。
对四种串珠藻属中国特有种进行了分子系统学研究,包括洪洞串珠藻、长柄串珠藻、弯形串珠藻和绞扭串珠藻。多重的分子数据包括一个核基因组基因(SSU),一个叶绿体基因(rbcL)和两个线粒体基因(COI, cox2-3spacer)被用来研究这四种串珠藻属中国特有种的系统发育关系。结果显示洪洞串珠藻和长柄串珠藻,从系统发育树上看,很明显属于曾经被划分为串珠藻组的种类,但与串珠藻组的模式种胶串珠藻的亲缘关系较远。因此,应将洪洞串珠藻和长柄串珠藻列入Sect. Helminthoidea。弯形串珠藻和绞扭串珠藻在所有的进化树中,都与扭曲组的种类位于同一分枝,并与串珠藻属并列,应该将其归入新属Kumanoa。
对7个种群的弧形串珠藻的形态特征及分子系统进行了研究。通过形态观察和聚类分析来评估种群间的个体差异。叶绿体rbcL序列和线粒体cox2-3spacer序列别用来评估种群间的遗传距离。形态观察的结果显示有4个种群的藻体为雌雄同株,其它3个种群的藻体为雌雄异株。聚类分析的结果显示,雌雄同株和雌雄异株的藻体没有分开,而地理距离较近生境相似的种群聚集在一起。同时,.rbcL序列和cox2-3spacer序列的结合序列所构建的系统树也证实了雌雄同株／异株不能作为一个典型的分类特征用于弧形串珠藻的鉴定。
The means of molecular biology, morphology, taxonomy, bioinformatics and statistics were used to determine the morphological character, flora, molecular systematic and phylogeny of Batrachospermum Roth in China. Morphological and floristics studies were made among30taxa of Batrachospermum in China. Multilocus molecular data from one nuclear gene (the ribosomal SSU), one chloroplast gene (rbcL), and two mitochondrial genes (COI, cox2-3spacer) were used to determine the phylogenetic placement of these taxa in Batrachospermum. Meanwhile, the studies on four endemic species and Batrachospermum arcuatum were also undertake to determine there phylogenetic relationship.
The studies on taxonomy and flora of Batrachospermum in China were made. Morphological studies show that Batrachospermum arcuatum specimens for4streams were monoecious; while for3streams were dioecious. The descriptions of other taxa are accord with the previous report. Thirty species (including varieties) of Batrachospermum were reported in China, which distributed in20provinces. The analysis of flora shows that there are8areal types of Batrachospermum, among which the endemic to China is predominant, including15species and accounting for48.3%. The temperature distribution of Batrachospermum is obvious and the number of the species takes82.8%totally. Batrachospermum is rich in East China, Central China and Southwest China; and rare in Northwest China, North China and Northeast China. Endemic species are distributed mainly in East China.
Multilocus molecular data were used to determine the phylogenetic relationship of taxa in genus Batrachospermum. The placement of the6taxa we collected in the phylogenetic trees was similar and the results accord with the morphological observation.The results based on rbcL sequence suggested that the placement of Thoreales was similar in each phylogenetic tree and formed a monophyletic group with high bootstrap support. Section Hybrida and section Contorta had a relatively small sequence divergence. The results of this work suggested combining section Hybrida as section Contorta. B. gelatinosum is the type of paraphyletic genus Batrachospermum. However, most of the species currently included in the section Batrachospermum are in a well-supported clade distant to B. gelatinosum. The same result was got with other sequence data. The results of phylogenetic relationship within Batrachospermum species based on SSU sequences showed that the taxonomic relationships of section Aristata, section Virescentia, section Turfosa and section Contorta had not been satisfactorily resolved. The results based on COI sequence showed that there was a good support for a clade of section Contorta species, which was apart from most Batrachospermum taxa. So we agreed with the proposal that taxa formerly in Batrachospermum sections Contorta and Hybrida were elevated into a new genus Kumanoa. The results based on cox2-3spacer sequence showed that Batrachospermum torsivum was sister to section Batrachospermum taxa, morphologically very distant entities. But only cox2-3spacer sequence of B. torsivum was got, and more sequence data were needed to solve this problem.
Four endemic Batrachospermum species in China, B. hongdongense, B. longipedicellatum, B. curvatum and B. intortum, were analyzed with multilocus data in order to assess their phylogenetic placement. Multilocus molecular data from one nuclear gene (the ribosomal SSU), one chloroplast gene (rbcL), and two mitochondrial genes (COI, cox2-3spacer) as well as morphological study were used to determine the phylogenetic relationships of these endemic species within the Batrachospermales. B. hongdongense and B. longipedicellatum had previously been placed in Batrachospermum section Batrachospermum, but based on the data from this study should be transferred into section Helminthoidea. B. curvatum and B. intortum clearly belong to the newly established genus Kumanoa.
Batrachospermum arcuatum specimens were analyzed from seven stream segments in North China. Morphological characteristics were observed and cluster analysis was used to evaluate the divergence among thalli from. Sequence data of the rbcL gene (chloroplast gene) and cox2-3spacer region (mitochondrial gene) were also utilized to evaluate genetic variation in specimens among stream segments. The specimens from four of the streams were monoecious, while the individuals at the other three locations were dioecious. Cluster analysis showed that the monoecious specimens were not separated from the dioecious specimens, based on morphology, but rather the specimens were grouped by geographical closeness and habitat similarity. Likewise, the combined analyses of rbcL and the cox2-3spacer data from provided more evidence that breeding system (monoecy vs. dioecy) is not a good morphological character to distinguish species.
引文
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