基于不同基因组序列的十字花科分子系统发育研究
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摘要
十字花科是一个公认的自然分类大科,尽管进行了很多的研究,十字花科族和属的划分仍然存在许多问题。许多分子生物学证据表明,具有相似形态学特征的物种有的系统学关系较远,而不同形态的物种却有着较近的系统学关系,伴随着来自于渐渗和谱系筛选所产生的问题,经常会导致物种树和基因树不一致的现象,因此需要更多的分子生物学证据来对该科内物种的系统学关系进行进一步的分析。出于这个目的,本研究使用了5个非编码序列(trnS-trnG, trnD-trnT, trnS-rpS4-trnT-trnL, nad7second intron, PI first intron)和1个编码序列(matK)来对十字花科的部分属种的系统学关系进行进一步的确认和划分。
研究表明,白花菜作为十字花科的外类群位于十字花科系统树的基部,而Aethionemeae则是十字花科内部离其它族系统学关系最远的族,位于十字花科内部的基部。该科中大多数的族可以划分到3个主要的世系Lineage Ⅰ-Ⅲ中,在三大世系外的族多为单系族。
Lineage Ⅰ包括了Cardamineae(碎米荠族),Camelineae(亚麻荠族),Descurainieae(播娘蒿族),Erysimeae(糖芥族),Lepideae(独行菜族),Smelowskieae(芹叶荠族)。糖界属和亚麻荠族的其它属在分析结果中是相互分离的,增加了将其划分为一个新族的可信度。豆瓣菜属的物种和碎米荠族的物种的系统学关系比和蔊菜属物种的关系近,不支持将豆瓣菜属作为蔊菜属合并的建议。Turritis最早被划分入鼠耳芥族中,但本研究支持将其划入亚麻荠族的结论。大多数的分子生物学证据支持将臭荠属和群心菜属放入独行菜族中。鼠耳芥族包含两个分支,一个分支位于Lineage Ⅰ中,并且和Halimolobeae, Boechereae系统学关系较近,另外一个分支在三大世系之外,由Arabis, Aubrieta, Draba和Baimashania组成。
Lineage Ⅱ包括Brassiceae(芸苔族),Isatideae(菘蓝族),Sisymbrieae(大蒜芥族)。本研究中,萝卜(Raphanus sativus)位于芸苔属的两个世系B. rapa/B. oleracea和B. nigra.之间,支持萝卜属起源于两个世系的假说。线果芥属应该从芸苔族中移出,但并不支持其作为一个独立的族存在,而应该划入菘蓝族中。中国特有的诸葛菜复合体包括太白诸葛菜,铺散诸葛菜,湖北诸葛菜和诸葛菜,本研究认为诸葛菜属应从芸苔族中移出,作为单独的一个分支存在。
Lineage Ⅲ包括Anchonieae, Chorisporeae(离子芥族),Dontostemoneae(花旗杆族),Hesperideae(香花芥族),Euclidieae(鸟头荠族)。爪花芥应该放入棒果芥当中。涩荠的系统学位置不稳定,取决于片段的选择。香花芥族是一个单属的族,并和花旗杆族系统学关系较近。Desideria和Phaeonychium都应该属于Solms-laubachia, Desideria是多系的而Solms-laubachia自身的物种则是单系存在的。
同时发现,四棱荠属被划分到了一个新界定的族Calepineae中,并且和菥莫族和山嵛菜族都不相连。沙芥属物种形成了一个单系分支,但其系统学位置还不能确定。庭荠族总是多系的,但是其系统学位置取决于片段的选择。
The Brassicaceae or mustard family is a large family. Despite much research, the tribal and genus classification of Brassicaceae has remained problematic and not well understood phylogenetically. Many molecular phylogenetic data have shown that species with similar morphology may be unrelated, whereas species with different morphology may be closely related. This situation, accompanied by questions regarding introgression and lineage sorting, often results in incongruent gene and species trees. Thus, more molecular data are needed to infer phylogenetic relationships in this family. For this purpose, five non-coding DNA sequences (trnS-trnG, trnD-trnT, trnS-rpS4-trnT-trnL, nad7second intron, PI first intron) and one coding sequence (matK) were used to study the Brassicaceae taxa to help verify the phylogenetic relationships of taxa within this family, and some results were obtained.
In current analysis, Cleome gynandra, used as the outgroup, was sister to the members of the Brassicaceae family, while Aethionemeae was the "basal" tribe and supported the sister relationship to all other tribes and taxa of Brassicaceae. Most of the tribes analyzed could be grouped into Lineages Ⅰ-Ⅲ (Beilstein et al.,2006) as well as some small monophyletic groups.
Lineage Ⅰ included Cardamineae, Camelineae, Descurainieae, Erysimeae, Lepideae, Smelowskieae. Erysimum was separated from other Camelineae genera, which increased the confidence that it is a new tribe. Nasturtium is more closely related to Cardamineae than to Rorippa, suggested that Nasturtium should be treated as an alias of Rorippa was probably unreasonable. Turritis was previously placed in Arabideae, but our analyses determined that it should be included in Camelineae. Most molecular data to date suggest place Coronopus and Cardaria within Lepideae. Arabideae included two subclades, one of which was nested within Lineage I and was closely related to Halimolobeae and Boechereae, while the other did not fit within Lineages Ⅰ-Ⅱ and comprised Arabis, Aubrieta, Draba, and Baimashania.
Lineage Ⅱ contained Brassiceae, Isatideae, Sisymbrieae. B. rapa/B. oleracea and Raphanus were combined with B. nigra, supporting that Raphanus was hybridized from B. rapa/B. oleracea and B. nigra. Our analysis supported the exclusion of Conringia from Brassiceae and Conringia should not be recognized as a new tribe and should be placed within the Isatiseae tribe. Pachypterygium and Tauscheria were considered to be included in Isatideae. O. violaceus complex included O. taibaiensis, O. diffusus, O. hupehensis, O.violaceus and should be excluded from Brassiceae.
Lineage Ⅲ comprised tribes of Anchonieae, Chorisporeae, Dontostemoneae, Hesperideae, Euclidieae. Oreoloma should be a member of Sterigmostemum. The position of Malcolmieae was unstable according to the DNA makers we used. Hesperideae was defined as a unigeneric tribe and was sister to Dontostemoneae. We found Desideria and Phaeonychium were grouped into Solms-laubach, Desideria was found to be polyphyletic, while all species of Solms-laubachia formed a monophyletic clade.
Goldbachia laevigata was assigned to a newly defined tribe, Calepineae, and it was neither related to Thlaspideae nor to Eutremeae. Pugionium formed a monophyletic clade, but its phylogenetic position remains unresolved. Alysseae is polyphyletic and its position was mainly dependent on the maker used.
研究表明,白花菜作为十字花科的外类群位于十字花科系统树的基部,而Aethionemeae则是十字花科内部离其它族系统学关系最远的族,位于十字花科内部的基部。该科中大多数的族可以划分到3个主要的世系Lineage Ⅰ-Ⅲ中,在三大世系外的族多为单系族。
Lineage Ⅰ包括了Cardamineae(碎米荠族),Camelineae(亚麻荠族),Descurainieae(播娘蒿族),Erysimeae(糖芥族),Lepideae(独行菜族),Smelowskieae(芹叶荠族)。糖界属和亚麻荠族的其它属在分析结果中是相互分离的,增加了将其划分为一个新族的可信度。豆瓣菜属的物种和碎米荠族的物种的系统学关系比和蔊菜属物种的关系近,不支持将豆瓣菜属作为蔊菜属合并的建议。Turritis最早被划分入鼠耳芥族中,但本研究支持将其划入亚麻荠族的结论。大多数的分子生物学证据支持将臭荠属和群心菜属放入独行菜族中。鼠耳芥族包含两个分支,一个分支位于Lineage Ⅰ中,并且和Halimolobeae, Boechereae系统学关系较近,另外一个分支在三大世系之外,由Arabis, Aubrieta, Draba和Baimashania组成。
Lineage Ⅱ包括Brassiceae(芸苔族),Isatideae(菘蓝族),Sisymbrieae(大蒜芥族)。本研究中,萝卜(Raphanus sativus)位于芸苔属的两个世系B. rapa/B. oleracea和B. nigra.之间,支持萝卜属起源于两个世系的假说。线果芥属应该从芸苔族中移出,但并不支持其作为一个独立的族存在,而应该划入菘蓝族中。中国特有的诸葛菜复合体包括太白诸葛菜,铺散诸葛菜,湖北诸葛菜和诸葛菜,本研究认为诸葛菜属应从芸苔族中移出,作为单独的一个分支存在。
Lineage Ⅲ包括Anchonieae, Chorisporeae(离子芥族),Dontostemoneae(花旗杆族),Hesperideae(香花芥族),Euclidieae(鸟头荠族)。爪花芥应该放入棒果芥当中。涩荠的系统学位置不稳定,取决于片段的选择。香花芥族是一个单属的族,并和花旗杆族系统学关系较近。Desideria和Phaeonychium都应该属于Solms-laubachia, Desideria是多系的而Solms-laubachia自身的物种则是单系存在的。
同时发现,四棱荠属被划分到了一个新界定的族Calepineae中,并且和菥莫族和山嵛菜族都不相连。沙芥属物种形成了一个单系分支,但其系统学位置还不能确定。庭荠族总是多系的,但是其系统学位置取决于片段的选择。
The Brassicaceae or mustard family is a large family. Despite much research, the tribal and genus classification of Brassicaceae has remained problematic and not well understood phylogenetically. Many molecular phylogenetic data have shown that species with similar morphology may be unrelated, whereas species with different morphology may be closely related. This situation, accompanied by questions regarding introgression and lineage sorting, often results in incongruent gene and species trees. Thus, more molecular data are needed to infer phylogenetic relationships in this family. For this purpose, five non-coding DNA sequences (trnS-trnG, trnD-trnT, trnS-rpS4-trnT-trnL, nad7second intron, PI first intron) and one coding sequence (matK) were used to study the Brassicaceae taxa to help verify the phylogenetic relationships of taxa within this family, and some results were obtained.
In current analysis, Cleome gynandra, used as the outgroup, was sister to the members of the Brassicaceae family, while Aethionemeae was the "basal" tribe and supported the sister relationship to all other tribes and taxa of Brassicaceae. Most of the tribes analyzed could be grouped into Lineages Ⅰ-Ⅲ (Beilstein et al.,2006) as well as some small monophyletic groups.
Lineage Ⅰ included Cardamineae, Camelineae, Descurainieae, Erysimeae, Lepideae, Smelowskieae. Erysimum was separated from other Camelineae genera, which increased the confidence that it is a new tribe. Nasturtium is more closely related to Cardamineae than to Rorippa, suggested that Nasturtium should be treated as an alias of Rorippa was probably unreasonable. Turritis was previously placed in Arabideae, but our analyses determined that it should be included in Camelineae. Most molecular data to date suggest place Coronopus and Cardaria within Lepideae. Arabideae included two subclades, one of which was nested within Lineage I and was closely related to Halimolobeae and Boechereae, while the other did not fit within Lineages Ⅰ-Ⅱ and comprised Arabis, Aubrieta, Draba, and Baimashania.
Lineage Ⅱ contained Brassiceae, Isatideae, Sisymbrieae. B. rapa/B. oleracea and Raphanus were combined with B. nigra, supporting that Raphanus was hybridized from B. rapa/B. oleracea and B. nigra. Our analysis supported the exclusion of Conringia from Brassiceae and Conringia should not be recognized as a new tribe and should be placed within the Isatiseae tribe. Pachypterygium and Tauscheria were considered to be included in Isatideae. O. violaceus complex included O. taibaiensis, O. diffusus, O. hupehensis, O.violaceus and should be excluded from Brassiceae.
Lineage Ⅲ comprised tribes of Anchonieae, Chorisporeae, Dontostemoneae, Hesperideae, Euclidieae. Oreoloma should be a member of Sterigmostemum. The position of Malcolmieae was unstable according to the DNA makers we used. Hesperideae was defined as a unigeneric tribe and was sister to Dontostemoneae. We found Desideria and Phaeonychium were grouped into Solms-laubach, Desideria was found to be polyphyletic, while all species of Solms-laubachia formed a monophyletic clade.
Goldbachia laevigata was assigned to a newly defined tribe, Calepineae, and it was neither related to Thlaspideae nor to Eutremeae. Pugionium formed a monophyletic clade, but its phylogenetic position remains unresolved. Alysseae is polyphyletic and its position was mainly dependent on the maker used.
引文
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