基于trnL-F序列推测冰岛蓼属的系统发育

冰岛蓼属的界定一直存在争议,本实验以塔黄为外类群,采用最大简约法对冰岛蓼属及其近缘类群的trnL-F序列进行了系统发育分析,结果表明:(1) 冰岛蓼(Koenigia islandica)和大铜钱叶蓼(Polygonum forrestii)为姊妹群,自展分析的支持率为100%,因此,大铜钱叶蓼应归于冰岛蓼属;(2) 冰岛蓼和分叉蓼组植物聚在一起,自展支持率为99%,说明它们之间有很近的亲缘关系.     

中国广义冰岛蓼属果实形态特征

应用扫描电镜对冰岛蓼属（Koenigia L.）8个物种的果实形态与果皮微形态进行了观察。结果表明,这些果实可分为3类：类型Ⅰ：瘦果长卵形至宽卵形,中部或中上部最宽,双凸镜状或3棱状,棱脊有或无;表面粗糙或光滑,具纵向断棱状纹饰,或具波状纹饰,这种类型有冰岛蓼、铜钱叶蓼、青藏蓼、大铜钱叶蓼4种;类型Ⅱ：瘦果卵形,中部膨大,具有3棱脊,但棱脊不显著。这种类型仅小叶蓼1种;类型Ⅲ：瘦果卵形,中间最宽但不膨大,具有3棱脊,且棱脊显著;表面光滑,具多皱纹饰,或凹凸不平纹饰,或具疣状纹饰,这种类型包括细茎蓼,蓝药蓼和柔毛蓼3种。研究结果支持将柔毛蓼与蓝药蓼作独立分种处理、而冰岛蓼与青藏蓼关系很近。     

中国冰岛蓼属植物花粉形态的研究

采用光学显微镜和扫描电镜对国产冰岛蓼属Koenigia L. 9种植物的花粉形态进行了观察。结果表明：花粉均为球形;直径为17.0-32.5 μm（包括刺长）;萌发孔有7(-8)赤道环沟、散沟、散孔;外壁纹饰均为刺状。根据萌发孔类型和外壁纹饰,将该属花粉划分为3种类型,即细冰岛蓼型（Delicatulum-type）、大连线冰岛蓼型（Forrestii-type）及冰岛蓼型（Koenigia-type）。细冰岛蓼型花粉的主要特征是具7(-8)赤道环沟,覆盖层无穿孔,外壁纹饰为显著的长刺状,此种类型的植物有细冰岛蓼K. delicatula （Meisn.） Hara。大连线冰岛蓼型花粉的主要特征是具12散沟,覆盖层无穿孔,外壁纹饰为显著的长刺状,此种类型的植物有大连线冰岛蓼K. forrestii (Diels) Měsíek & Soják 及连线冰岛蓼K. nummularifolia (Meisn.) Měsíek & Soják。冰岛蓼型花粉的主要特征是具15,20(-30)散孔,覆盖层无穿孔,外壁纹饰为显著的长刺状,此种类型的植物有冰岛蓼K. islandica L.、蓝蕊冰岛蓼K. cyanandra (Diels) Měsíek & Soják、线茎冰岛蓼K. nepalensis D. Don、柔毛蓼K. pilosa Maxim.、陕甘蓼K. hubertii (Lingelsh.) Měsíek & Soják及青藏蓼K. fertilis Maxim.。结果表明冰岛蓼属的花粉形态具有重要的分类学意义,研究结果支持Koenigia的属的地位,大铜钱叶蓼Polygonum forrestii Diels和铜钱叶蓼P. nummularifolium Meisn.应从蓼属分叉蓼组Polygonum L. sect. Aconogonon Meisn.中移入冰岛蓼属中,细冰岛蓼、蓝蕊冰岛蓼、青藏蓼、陕甘蓼、线茎冰岛蓼和柔毛蓼应从蓼属头状蓼组Polygonum L. sect. Cephalophilon Meisn.中移入冰岛蓼属中。     

中国蓼属分叉蓼组和冰岛蓼属植物叶下表皮微形态研究

采用光学显微镜对中国蓼属分叉蓼组(Polygonum section Aconogonon)和冰岛蓼属(Koenigia)的21种植物的叶下表皮微形态进行了观察研究。结果表明,其叶下表皮微形态特征可分为三种类型：（1）气孔器类型为无规则型,表皮细胞多边形,垂周壁为波状;（2）气孔器类型为无规则型兼有非典型不等型,表皮细胞为多边形或不规则形,垂周壁为平直弓形或波状;（3）气孔器类型为不等细胞型,表皮细胞多边形,垂周壁为平直弓形。分叉蓼组（西伯利亚蓼除外）植物的叶表皮气孔器类型具有高度一致性,说明该组是一个自然类群。本文研究结果不支持将大铜钱叶蓼(Polygonum forrestii Diels)、铜钱叶蓼(P. nummularifolium Meissner)划归于冰岛蓼属的处理意见;支持将西伯利亚蓼(P. sibiricum Laxm.)独立成属,即西伯利亚蓼属(Knorringia Tzvel.)的观点;同时也支持仍将多穗蓼(P. polystachyum Wall.ex Meissner)和松林蓼(P. pinetorum Hemsl.)归在分叉蓼组的处理意见。     

基于叶绿体trnL-F,rbcL序列和核核糖体ITS序列探讨蓼属（蓼科）头状蓼组的系统发育

选用叶绿体基因trnL-F,rbc-L序列和核核糖体ITS序列对蓼属头状蓼组的分类和系统发育进行了分析。以药用大黄为外类群,用最大简约法对3个片段的单独和联合矩阵分别构建系统发育树。结果表明,头状蓼组是一个较自然的类群,与春蓼组、刺蓼组和金线草属的亲缘关系较近;支持头状蓼组作为一组放在春蓼属,细茎蓼和蓝药蓼从头状蓼组转移至冰岛蓼属;不支持将刺蓼组并入头状蓼组;对于小叶蓼的系统学位置有待于进一步研究。     

基于核糖体DNA内转录间隔区ITS序列的广义蓼属及近缘属分子系统学研究

通过对广义蓼属及近缘属共32个代表种内转录间隔区ITS序列的分子系统学分析,尝试研究备受争议的广义蓼属及近缘属的物种族、属、组级的划分问题,结果显示,广义蓼属在系统发育树上并不能形成一个单系类群,这些物种共聚为3大支,分别对应春蓼族、蓼族及荞麦族,其中荞麦属与翅果蓼属形成了一支独立于春蓼族及蓼族之外的类群。在春蓼族中,冰岛蓼属与分叉蓼组形成一个单系类群。     

基于核糖体DNA内转录间隔区ITS序列的广义蓼属及近缘属分子系统学研究(英文)

通过对广义蓼属及近缘属共32个代表种内转录间隔区ITS序列的分子系统学分析,尝试研究备受争议的广义蓼属及近缘属的物种族、属、组级的划分问题,结果显示,广义蓼属在系统发育树上并不能形成一个单系类群,这些物种共聚为3大支,分别对应春蓼族、蓼族及荞麦族,其中荞麦属与翅果蓼属形成了一支独立于春蓼族及蓼族之外的类群。在春蓼族中,冰岛蓼属与分叉蓼组形成一个单系类群。     

对翼蓼(蓼科)属级位置的重新探讨

通过对翼蓼Pteroxygonum giraldii Damm. &; Diels及相关属(蓼属Polygonum、何首乌属Fallopia、虎杖属Reynoutria、荞麦属Fagopyrum和金线草属Antenoron)的形态观察、果实解剖学观察、花被片脉序观察、花粉形态、核型分析, 以及ITS序列的分析确定了翼蓼和荞麦F. esculentum Moench较远的亲缘关系。其中我们发现翼蓼果实基部有三个角状物明显不同于其他属果实的形态特征。翼蓼外果皮明显加厚, 并有零星散布的波状内腔, 而荞麦的外果皮很薄, 细胞不等径, 中果皮极厚。以上证据证明了翼蓼与荞麦属亲缘关系较远。在观察荞麦属和翼蓼的花被片脉络时发现了两种不同的脉序类型, 符合将荞麦属分为两个组的划分。翼蓼花被片脉序为三出状, 支持将翼蓼归为Persicarieae族。对翼蓼及荞麦属植物的花粉进行比较后, 发现荞麦属植物的花粉网孔有明显的内凹穿孔而翼蓼却没有, 结果表明二者亲缘关系较远。通过对nrDNA ITS区域序列分析得出翼蓼及相关属为一个单系类群, 含有两个稳定的分支: 第一个分支由蓼属(萹蓄组sect. Avicularia)、何首乌属、虎杖属的植物组成, 第二个分支由蓼属(刺蓼组sect. Echinocaulon、蓼组sect. Polygonum、分叉蓼组sect. Aconogonon、拳参组sect. Bistorta、翼蓼和荞麦属植物组成。同时第二个分支又分成了两个亚分支, 蓼属(刺蓼组、蓼组、分叉蓼组、拳参组)和翼蓼属Pteroxygonum植物属于第一个亚支而荞麦属植物属于第二个亚支。结果支持翼蓼不属于荞麦属的范畴。实验结果显示翼蓼是个单型属, 属于Persicarieae族。     

两栖蓼的分子系统学研究

两栖蓼是一种水陆两栖植物,植株在不同生态环境下外部形态差异较大,同时两栖蓼的系统位置存在争议,被归入春蓼组(sect.Persicaria)或提升为两栖蓼组(sect.Amphibium)。该文选取两栖蓼及春蓼组植物12种,以及刺蓼组、头状蓼组、神血宁组、拳参组、萹蓄组和外类群掌叶大黄共23种植物进行研究。植物总DNA的提取采用改进的CTAB法,所测序列以及从Genbank数据库下载的序列,以掌叶大黄为外类群,采用最大简约法和贝叶斯法对核糖体ITS序列和叶绿体trn L-F序列进行了系统发育分析。ITS序列对位排列的长度为735 bp,包括489个可变位点,272个位点是信息位点。简约法得到9个简约树,步长为1 084,CI指数为0.680,RI指数为0.614。trn L-F序列对位排列的长度为1 121 bp,包括427个可变位点,239个位点是信息位点。简约法寻找到9个简约树,步长为551(CI=0.911,RI=0.910)。贝叶斯法和简约法得到的树基本一致。分子序列分析结果显示,trn L-F序列树类似于ITS序列树。ITS序列构建的发育树上,两栖蓼与刺蓼组植物、春蓼组其他植物形成3个并列的分支;在trn L-F序列树上,两栖蓼则与其他春蓼组植物形成两个并列的分支。由此可见,两栖蓼与春蓼组其他植物的亲缘关系较远,成一独立的分支。两个分子证据支持将两栖蓼提升为两栖蓼组的处理意见。此外,两栖蓼的花粉具散沟,与典型的春蓼组的具散孔花粉不一致。再加上两栖蓼水陆两栖的特性,因此支持把两栖蓼提升为两栖蓼组的观点。两栖蓼组的界定为多年生草本,水陆两栖,根状茎横生,生于水中茎漂浮,叶长圆形或椭圆形,生于陆地茎直立,叶披针形或长圆状披针形,托叶鞘为筒状、薄膜质,总状花序穗状,瘦果近圆形,花粉具散沟。     

中国蓼属叉分蓼组植物花粉形态的研究

采用光学显微镜和扫描电镜对中国蓼属叉分蓼组20种3变种的花粉形态进行了观察和研究。结果表明其花粉形态大多数为近球形至近长球形,少数为扁球形或长球形;花粉大小为20.4～44.0µm×17.0～34.0µm：从萌发孔看,有3沟、3 孔沟、多沟、散沟;外壁纹饰为微刺—穴状、刺状、粗网状、皱块状。据此,该组花粉可划分为5种类型,即叉分蓼型(Aconogonon-type)、钟花蓼型(Campanulatum-type)、大连线冰岛蓼型(Forrestii-type)、西伯利亚蓼(Sibiricum-type)及多穗蓼型(Polystachyum-type),编制了这些花粉类型检索表。叉分蓼型花粉的主要特征是具3沟,外壁纹饰为微刺-穴状,此种类型的植物有14种2变种。钟花蓼型花粉的主要特征是具6散沟,外壁纹饰为微刺-穴状,此种类型的植物有钟花蓼和绒毛钟花蓼。西伯利亚蓼型花粉的主要特征是具3孔沟,外壁纹饰为皱块状,此种类型的植物有西伯利亚蓼。多穗蓼型花粉的主要特征是具6(~8)多沟,外壁纹饰为粗网状,此种类型的植物有松林蓼及多穗蓼。大连线冰岛蓼型花粉的主要特征是具散沟,外壁纹饰为显著的长刺状,此种类型的植物有大铜钱叶蓼及铜钱叶蓼,结果表明叉分蓼组的花粉形态具有重要的分类学意义,研究结果支持将叉分蓼组上升为属的等级,也支持Knorringia的属的地位,大铜钱叶蓼和铜钱叶蓼应移入Koenigia属中,而松林蓼和多穗蓼仍保留在蓼属中。     

The genus Koenigia L. emend. Hedberg (Polygonaceae)

Pollen morphological studies revealed the occurrence of the characteristic spinulose pollen type of Koenigia not only in the three species earlier recognized in the genus (A" islandka, K. nepalensis and K. pilosd) but also in three additional species earlier treated under Polygonum , viz. K delicatula (Meisn.) Hara, K. forrestii (Diels) Mesicek & Soják, and A" nummularifolia (Meisn.) Mesicek & Soják. Further studies of flower morphology, fruit and petiole anatomy, basic chromosome number, etc., revealed additional similarities between those species, which led to a taxonomic revision of the genus Koenigia. This genus seems to be most closely related to Persicaria Mill, sections Cephalophilon (Meisn.) Gross and Echinocaulon (Meisn.) Gross, with Koenigia delicatula as a connecting link. There arc also interesting similarities with the genus Aconogonon (Meisn.) Rchb. Koenigia exemplifies the derivation from montane ancestors of a high mountain-dwelling genus displaying adaptive radiation to fit diverse alpine niches. Five species out of six are confined to high mountain areas in southeastern Asia, primarily in the Himalayas, whereas the sixth has spread to Arctic and alpine areas in the northern hemisphere and even penetrated to southern South America. The latter species shows progressive reduction in size in combination with adaptation to a very short summer under severe climatic conditions.     

基于叶绿体DNA atpB-rbcL区序列探讨石蒜属种间系统发育关系

摘要：为了探讨石蒜属（Lycoris Herb.）的种间系统发育关系,对石蒜属95个材料包括15种、4变种及2个人工杂种的叶绿体 DNA atpB-rbcL间隔区进行了测序,结合花部形态和核型特征,探讨了石蒜属种间系统关系及其可能的杂交起源,结果表明：在系统发育树上亲缘关系近的材料聚在一起,其中矮小石蒜（L. radiata var. pumila）和换锦花（L. sprengeri）与2个人工杂交种（Hybrid 1、Hybrid 2）、麦秆石蒜（L. straminea）、江苏石蒜（L. houdyshelii）、短蕊石蒜（L. caldwellii）和乳白石蒜（L. albiflora）具有密切的亲缘关系。atpB-rbcL序列揭示的石蒜属种间关系与染色体核型的分类结果部分一致,主要表现在具有近端部着丝粒(A)染色体的种与具有中部(M)和端部(T)着丝粒染色体的种各成一支,与形态和染色体分类结果一致;不同之处在于具有中部、端部和近端部着丝粒染色体的种分散在两个主要分支内,进一步验证了具有中部、端部和近端部3种着丝粒类型染色体组的石蒜如麦秆石蒜、江苏石蒜、短蕊石蒜和乳白石蒜等是杂交起源的假设,结合2个人工杂交种分析,揭示了短蕊石蒜和乳白石蒜的近端部着丝粒染色体来源于换锦花;麦秆石蒜和江苏石蒜近端部着丝粒染色体来源于矮小石蒜。     

Phylogeny and diversification of Chinese Araliaceae based on nuclear and plastid DNA sequence data

Chinese Araliaceae consist of 20 genera and ca. 175 species. To assess the evolutionary relationships of Araliaceae and their biogeographic diversification in China, the phylogeny of Chinese Araliaceae was constructed by sampling 96 accessions representing 20 genera and 50 species of Chinese Araliaceae and 45 closely related taxa using sequences of the nuclear ribosomal internal transcribed spacer (ITS) region and six plastid regions (the ndhF gene, the trnL-trnF region, the rps16intron, the atpB-rbcL intergenic spacer, the rpl16 intron, and the psbA-trnH intergenic spacer). Phylogenetic analyses of the combined plastid and ITS data supported the results of the previously studies that the Chinese members of Araliaceae were scattered within the Asian Palmate group and the Aralia-Panax group withOsmoxylon at the base of core Araliaceae. The generic status of Pentapanax and Tupidanthus is not supported. Our analysis clearly places them in Aralia and AsianSchefflera, respectively. In a broader phylogenetic framework of Araliaceae, based on the fossil-calibrated Bayesian dating, Chinese Araliaceae was inferred to have originated in Asia and underwent a rapid radiation in its evolutionary history. Its diversification is hypothesized to have been driven largely by the orogenies in Asia during the Cenozoic. In China, the distribution pattern of the phylogenetic diversity of Araliaceae corresponds with its taxonomic diversity across the entire region.     

Development of PCR primer sets for intron 1 of the low-copy gene LEAFY in Davalliaceae

? Premise of the study: Primers were designed for amplifying intron 1 of the single-copy nuclear LEAFY gene for species of Davalliaceae. ? Methods and Results: New primer sets were designed and successfully amplified for intron 1 of the LEAFY gene in 13 species representing the five genera of Davalliaceae. The orthology of these sequences was further confirmed by phylogenetic analyses. Site variation in LEAFY intron 1 sequences across genera of the Davalliaceae and among accessions of the Humata repens complex were 18% and 8%, respectively. Such variation was greater than that for the cpDNA atpB-rbcL intergenic spacer region across the same taxa and accessions. ? Conclusions: Using our newly designed primers, intron 1 of the LEAFY gene could be amplified for all species tested. In addition, this single-copy, biparentally inherited, and quickly evolving region showed considerable potential for addressing infraspecific-level questions.     

Reappraisal of the generic status of Pteroxygonum (Polygonaceae) on the basis of morphology,anatomy and nrDNA ITS sequence analysi

Gross morphology, fruit anatomy, tepal venation, pollen morphology, chromosome number and ITS sequence of Pteroxygonum Damm. & Diels as well as other related genera (Polygonum, Fallopia, Reynoutria, Fagopyrum, and Antenoron) have been investigated to evaluate the generic status of Pteroxygonum. Pt. giraldii Damm. & Diels has three sharp horns at the base of fruit, which is distinctive among all the genera investigated. Upon observation of fruits under a light microscope (LM), the exocarp of Pt. giraldii is usually thickened and delimited by the rectangular cells with some sporadic undulating lumen, while that of Fagopyrum is thin-walled and isodiametric to rectangular in the cell shape. Analysis of tepal venation was performed under a stereomicroscope, and two types of tepal venation were found in Fagopyrum and Pteroxygonum. The type I is trifid, observed in Pt. giraldii, F. esculentum Moench, F. dibotrys (D. Don) Hara and F. tataricum (L.) Gaertn. The type II, found in F. caudatum (Sam.) A. J. Li, F. urophyllum (Bur. & Franch.) H. Gross and F. gracilipes (Hemsl.) Damm. ex Diels, has the main vein extending from tepal base with some secondary veins. Evidence from tepal venation supports the previous classification in which Fagopyrum can be divided into a large-achene group and a small-achene group. Pollen morphology was investigated under a scanning electron microscope (SEM). The exine ornamentation of Pt. giraldii was finely reticulate with lumina diameter wider than muri width. The exine ornamentation in all the examined Fagopyrum species is, however, prominently sunken punctuate. The phylogenetic analysis of nuclear ribosomal DNA (nrDNA) ITS sequences in Pteroxygonum and related genera indicated that all the species form a well-supported monophyletic group with two clades. One includes Polygonum sect. Avicularia Meisn., genus Fallopia and genus Reynoutria, and the other consists of other sections of Polygonum, genus Fagopyrum and Pteroxygonum. The latter clade can be divided into two subclades. Fagopyrum species compose the first one, while Pteroxygonum giraldii, species of Polygonum (except sect. Avicularia) and Antenoron form the second one. In consideration of the above evidence, we conclude that Pteroxygonum is an independent genus in tribe Persicarieae, and should not be merged into the genus Fagopyrum.