毛莨科金莲花族和升麻族细胞学的比较研究

对毛茛科金莲花族Trollieae和升麻族Cimicifugeae的细胞学进行了比较研究.发现驴蹄草Caltha palustris L. 在云南西北部形成一个多倍体系列(2n=32, 48, 64),四倍体细胞型 (2n=4x=32) 较为常见,其核型有明显的居群间变异.驴蹄草属Caltha、鸡爪草属Calathodes、Megaleranthis以及金莲花属Trollius的染色体在大小上基本相似,都属于中等大小的R-型染色体.细胞学和花粉学证据都支持鸡爪草属与Megaleranthis和金莲花属有较近的亲缘关系.铁破锣属Beesia、Anemonopsis、黄三七属Souliea、升麻属Cimicifuga以及类叶升麻属Actaea的核型彼此基本相似,在染色体大小和形态上都与驴蹄草属、鸡爪草属、Megaleranthis以及金莲花属的核型明显有别.细胞学证据表明铁破锣属应是升麻族中的成员.     

角叶铁破锣的核型及其系统学意义

本文首次报道了角叶铁破锣的核形态。其静止核和有丝分裂前期染色体分别属于复杂中央染色微粒型和中间型;中期染色体数目为2n=16;核型公式为2n＝10m+4st十2t(2sat)。根据上述结果并结合有关资料,本文讨论了铁破锣和角叶铁破锣之间的核型差异以及铁破锣属的系统位置,指出铁破锣属可能与升麻属等类群关系较近而与金莲花属等类群关系较远,因此将该属置于升麻族中比置于金莲花族中合理。     

鸡爪草属的染色体研究及其系统位置

本文研究了毛茛科鸡爪草Calathodes oxycarpa的核形态。其静止核和有丝分裂前期染色体分别属于复杂中央染色微粒型和中间型,中期染色体属于R型,核型公式为2n=16=8sm+8st(2sat)。据此并结合有关资料,讨论了鸡爪草属与毛茛科其它类群的亲缘关系,认为鸡爪草属与金莲花属和特产于南朝鲜的Megaleranthis属是极为近缘的类群,不宜将它们分开置于不同的族或亚科中。     

鸡爪草属的染色体及其系统位置

本文研究了毛茛科鸡爪草Calathodes oxycarpa的核形态。其静止核和有丝分裂前期染色体分别属于复杂中央染色微粒型和中间型,中期染色体属于R型,核型公式为2n=16=8sm 8st(2sat)。据此并结合有关资料,讨论了鸡爪草属与毛茛科其它类群的亲缘关系,认为鸡爪草属与金莲花属和特产于南朝鲜的Megaleranthis属是极为近缘的类群,不宜将它们分开置于不同的族或亚科中。     

国产毛茛科驴蹄草属两种植物的细胞学研究

为探讨国产毛茛科(Ranunculaceae)驴蹄草属(Caltha L.)植物的细胞学特征,对驴蹄草(C.palustris L.)3个居群和花葶驴蹄草(C.scaposa Hook.f.&Thoms.)5个居群进行了细胞学研究。驴蹄草贵州纳雍居群的染色体数目为2n=32(四倍体),两个云南中甸居群的染色体数目均为2n=64(八倍体)。花葶驴蹄草四川红原、康定、石渠居群的染色体数目均为2n=32(四倍体),该数目为首次报道;西藏林芝和云南德钦居群的染色体数目均为2n=64(八倍体)。驴蹄草的染色体比花葶驴蹄草大。这两种植物的32或64条染色体分别以4条或8条为单位大致能够排列为8组同源染色体,但同一组内的染色体经常具有明显的异形性(heteromorphy),不同居群的核型组成多少具有差异。同时,还分析了驴蹄草和花葶驴蹄草的不同倍性细胞型在我国的地理分布式样。     

毛茛科升麻族植物药用亲缘学初探

本文总结和归纳了在北美、欧洲以及亚洲广泛应用的毛茛科Ranunculaceae升麻族Cimicifugeae药用植物的系统分类、传统药用价值、现代药理以及化学成分研究结果,并在此基础上初步探讨了该族植物的药用亲缘学关系.升麻族药用植物主要含有9,19-阿尔廷烷三萜皂苷以及肉桂酸衍生物两大类化合物.升麻族植物在传统药用中被用来镇痛、解毒、抗炎,在北美很早就被印第安人用来治疗妇科疾病.现代药理研究发现升麻族植物还具有抗骨质疏松、抗病毒、抗肿瘤、抗过敏、类雌激素样作用等.通过对升麻族植物药用亲缘学的初步研究,我们发现由于类叶升麻属Actaea和升麻属Cimicifuga的疗效和化学成分相近,因此两属的亲缘关系也近.考虑它们果实的形态差异,以及细胞学特征不同,认为这两属为升麻族植物的一个分支,且以类叶升麻属较升麻属更为进化.从化学分类学的角度来看,铁破锣属Beesia含有特殊铁破锣皂苷可以成为一个独立的分支,而黄三七属Souliea既和铁破锣属一样含有五环三萜和铁破锣型环阿尔廷烷三萜类化合物,又和升麻属、类叶升麻属一样含有吲哚生物碱,因此可以认为它是铁破锣属和升麻属、类叶升麻属之间的一个过渡类型.升麻族植物特有的清热解毒功效与其特殊的阿尔廷烷三萜皂苷化学成分密切相关,因此我们认为阿尔廷烷三萜皂苷可以作为抗肿瘤、抗病毒的新药源进一步深入研究.     

毛茛科金莲花亚科植物的地理分布

本文对毛茛科金莲花亚科各属的地理分布作了分析,该亚科植物除了少数属的一些种分布到南半球的温带地区,一些种分布或延伸到亚热带山地、非洲东部和北部的干旱、半干旱的地区外,绝大部分的属、种均分布于泛北极区域。根据其17个属的地理分布式样,把它们划分为8个分布区类型：(1)北温带分布类型4属;(2)北温带和非洲分布类型1属;(3)北半球温带和南半球间断分布类型l属;(4) 欧洲和东亚间断分布类型l属;(5)西亚分布类型l属;(6)地中海分布类型3属;(7)欧亚和温带亚洲分布类型l属;(8)东亚分布类型5属。本文以形态特征为主,结合花粉和染色体的性状分析,认为东亚特有的鸡爪草属、Megaleranthis和铁破锣属可能分别是联系驴蹄草属和金莲花属,鸡爪草属和金莲花属以及金莲花族和升麻族的中间类型。另外,文中详细地统计了该亚科的不同等级分类群及特有种在各个植物区的分布,并从系统发育的观点讨论了各个植物区所具有的原始类群和进化类群,提出了如下论点,即东亚植物区(特别是中国西南部)不但是金莲花亚科植物分布的多度和多样性中心以及特有类群的分布中心,而且还是原始类群的保存中心,伊朗-土兰区及地中海周围是第二分布中心。     

毛茛科金莲花亚科植物的地理分布

本文对毛茛科金莲花亚科各属的地理分布作了分析,该亚科植物除了少数属的一些种分布到南半球的温带地区,一些种分布或延伸到亚热带山地、非洲东部和北部的干旱、半干旱的地区外,绝大部分的属、种均分布于泛北极区域。根据其17个属的地理分布式样,把它们划分为8个分布区类型:(1)北温带分布类型4属;(2)北温带和非洲分布类型1属;(3)北半球温带和南半球间断分布类型1属;(4)欧洲和东亚间断分布类型1属;(5)西亚分布类型1属;(6)地中海分布类型3属;(7)欧亚和温带亚洲分布类型1属;(8)东亚分布类型5属。本文以形态特征为主,结合花粉和染色体的性状分析,认为东亚特有的鸡爪草属、Megaleranthis和铁破锣属可能分别是联系驴蹄草属和金莲花属,鸡爪草属和金莲花属以及金莲花族和升麻族的中间类型。另外,文中详细地统计了该亚科的不同等级分类群及特有种在各个植物区的分布,并从系统发育的观点讨论了各个植物区所具有的原始类群和进化类群,提出了如下论点,即东亚植物区(特别是中国西南部)不但是金莲花亚科植物分布的多度和多样性中心以及特有类群的分布中心,而且还是原始类群的保存中心,伊朗-土兰区及地中海周围是第二分布中心。     

两种羽叶菊属植物(菊科:千里光族)的细胞学研究及其系统学意义

该研究首次报道了滇羽叶菊和台湾刘寄奴的染色体数据。结果表明:两者的染色体数量都为48,核型公式均为2n=2x=36m+10sm+2st,与前人报道的刻裂羽叶菊的核型稍有不同。两者的染色体形态均由大到小逐渐变化,核型二型性不明显,但前者染色体明显比后者大。这说明羽叶菊属染色体基数确实应为x=24,且染色体大小在种间有较大差异。细胞学证据表明,该属与蒲儿根属中染色体基数为x=24的类群以及狗舌草属确实近缘。     

国产驴蹄草的细胞地理学研究（英文）

为探讨国产毛茛科(Ranunculaceae)驴蹄草属(Caltha)两种植物的演化,该文利用传统染色体压片技术和流式细胞术,并结合前人染色体研究结果,对我国驴蹄草23个居群和花葶驴蹄草10个居群进行了细胞学研究。结果表明:驴蹄草是由四倍体(2n=4x=32)、六倍体(2n=6x=48)和八倍体(2n=8x=64)构成的多倍体复合群,花葶驴蹄草具有四倍体(2n=4x=32)和八倍体(2n=8x=64)两种倍性水平。驴蹄草和花葶驴蹄草均是四倍体较为常见,目前尚未见有二倍体报道。由于驴蹄草和花葶驴蹄草大部分居群采自中国青藏高原地区,可能在冰期时存在古二倍体,其适应性较弱,逐渐被其他的倍性取代,这是由于不同细胞型对环境适应性的结果。驴蹄草可能存在两条进化路线:一条是从甘肃到达云南;另一条是从西藏到达云南。前期分子系统学研究显示花葶驴蹄草与驴蹄草的亲缘关系较近,该研究结果中花葶驴蹄草染色体比驴蹄草要小,花葶驴蹄草可能比驴蹄草相对进化。目前花葶驴蹄草只有10个居群,还需进一步增加居群量来解析其演化路线。     

Tribal relationships of Beesia, Eranthis and seven other genera of Ranunculaceae: evidence from cytological characters

The karyotypes of 16 species and one variety representing nine genera in the Ranunculaceae were analysed in order to obtain information on the placement of the genera Beesia and Eranthis at tribal rank in the family. Those of Beesia , Anemonopsis , Souliea , Cimicifuga , Actaea and Eranthis were found to be very similar to each other, but remarkably different from those of Caltha , Calathodes , Megaleranthis and Trollius with respect to chromosome size and morphology. From cytological data it is clearly evident that Beesia and Eranthis have a much closer affinity to Anemonopsis , Souliea , Cimicifuga and Actaea in the tribe Cimicifugeae than to Caltha , Calathodes , Megaleranthis and Trollius , which generally have been placed together in a single tribe, albeit with different tribal names such as Caltheae, Trollieae and Helleboreae. Therefore, Beesia and Eranthis may find better placement in the tribe Cimicifugeae. The long-presumed, mainly morphology-based close affinity between Eranthis and Helleborus is not supported by both cytological and palynological data. Cytologically, both Actaea and Eranthis appear to be the specialized genera in the Cimicifugeae, through having the most asymmetrical karyotypes in this tribe. Our results strongly support the recent re-definition of the tribe Actaeeae (as Cimicifugeae) to include Beesia and Eranthis , based on maximum parsimony analysis performed separately on molecular data.  © 2006 The Linnean Society of London. Botanical Journal of the Linnean Society , 2006, 150 , 267–289.     

类叶升麻具有毛茛科中最对称和最原始的核型吗?

根据以前的报道,类叶升麻 Actaea asiatica Hara具有10条大型的中部着丝点染色体和6条较大 的近中部着丝点染色体,其核型在毛茛科中显得最为对称和原始,而类叶升麻属的其他种类具有10条 大型的中部着丝点染色体、4条较大的近中部着丝点染色体和两条没有短臂的染色体。在毛茛科中,同 一属的染色体形态通常十分相似,因此上述类叶升麻的核型分析结果十分可疑。本文重新检查子该种 的染色体。结果表明其核型与该属其他种类的核型没有明显区别。与升麻属其他4属,即 Beesia, Anemonopsis,Souliea,Cimicifuga相比,类叶升麻及该属其他种类都具有两条没有短臂的T染色体,因 此类叶升麻属 Actaea L．的核型不对称性程度在升麻族中显得最高,其核型在该族中也可能最为进化, 这两条T染色体可以作为类叶升麻属的细胞学标志,据此可以将该属与升麻族其他4属区别开来。     

Does Actaea asiatica Have the Most Symmetric and Primitive Karyotype in the Ranunculaceae?

Actaea asiatica was previously reported to have the most symmetric and primitive karyotype, consisting of 10 m- and 6 sm-chromosomes, which is quite different from those of the remaining species in the genus Actaea, consisting of 10 m-, 4 sm- and two T-chromo somes. In this paper, the chromosomes of this species were re-examined. The results show that Actaea asiatica has the same karyotype as the other species in the genus. Compared with the species in other genera in the tribe Cimicifugeae, i.e. Beesia, Anemonopsis, Souliea and Cimicifuga, Actaea asiatica, together with the remaining species of the genus, has the most asymmetric and thus probably the most advanced karyotype in this tribe because of the presence of two T- chromosomes in their chromosome complements. The two T- chromosomes may serve as one of the most important cytological markers, by which the species in Actaea are clearly distinguishable cytologically from those in Beesia, Anemonop-sis, Souliea and Cimicifuga.     

The Systematic Position of Beesia: Evidence from ITS (nrDNA) Sequence Analysis

Discussed in the present paper is the systematic position of Beesia, a small ranunculaceous genus mainly distributed in SW China. Sequences of the internal transcribed spacers (ITS) and 3’ end of 5.8 S rRNA gene of Beesia calthifolia, Trollius chinensis, Cimicifuga acerina, C. brachycarpa and Actaea asiatica were determined by direct PCR sequencing method. The sequences of ITS-1 in the five species range from 225 bp to 232 bp in size and those of ITS-2 from 201 bp to 217 bp. Beesia is very similar to Cimicifuga and Actaea not only in size, sequence and G+C content of ITS, but also in sequence of 3’ end of 5.8 S rRNA gene. In PAUP analysis, Ranunculus enysii was used as outgroup, and the most parsimonious tree was obtained through exhaustive search. The gaps were treated respectively as missing characters and the fifth base in two analyses. The two analyses show that a monophyletic group comprising Beesia calthifolia, Cimicifuga acerina, C. brachycarpa and Actaea asiatica is strongly supported by the bootstrap value. In the monophyletic group, Beesia calthifolia is basal to the other three species. The present DNA sequence analysis demonstrates that the genus Beesia should be placed in the tribe Cimicifugeae, which is consistent with the results from phytochemistry, palynology and cytology. In addition, Beesia may be the most original genus in the tribe Cimicifugeae in view of simple leaf,apetalous flower and molecular evidence.     

Comparative Anatomical Studies on Vessel Elements of Chinese Ranunculaceae

The vessel elements in 77 species of 36 genera of Chinese Ranunculaceae were studied. The vessel elements of Ranunculaceae could be divided into primitive and more specialized groups on the basis of their morphological characters through clustering analysis. The most primitive type of the vessel elements was seen in Asteropyrum, and some vessel elements were considered more primitive, as in Paeonia, Trollius, Actaea, Helleborus, Souliea, Calathodes, Beesia, Caltha, Coptis, Kingdonia, Circaeaster, Adonis, Hepatica, than others. There was a marked difference between Paeonia and other genera of Ranunculaceae. The vessel elements of Paeonioideae and Helleboroideae with follicles were more primitive than Ranunculoideae with achenes. In Thalictroideae, .the vessel elements of Thalictrum with achenes were more specialized than other genera with follicles. All these findings coincided with the evolution of fruits. The character of evolution of vessel elements ran parallel with that of other parts of plant (flower, carpel, petiole, venation). The features of vessel elements in the same genus were at similar evolutional level, however, some difference between species do exist.     

Chromosome observations of three ranunculaceous genera in relation to their systematic positions

1. The present paper describes the observations of chromosome numbers and karyomorphology of 2 species of 2 endemic genera and I endemic species of Chinese Ranunculaceae: Asteropyrum peltatum (Franch.) Drumm et Hutch. 2n=16, x=8; Kingdonia unifolia Balf. f. et W. W. Sm. 2n=18, x=9 and Calathodes oxycarpa Sprague 2n=16, x=8. The chromosome counts of three ranunculaceous genera are reported for the first time. 2. The morphylogical, palynological and cytological date in relation to the systematic postition of Asteropyrum, Kingdonia and Calathodes within the family Ranunculaceae are diseussed and resulted in following conclusions: (1). On the basis of the basic number x=8 in Asteropyrum, it is further confirmed that this genus is distinct from the r elated genera such as Isopyrum, Dichocarpum and other allied taxa. The comparison of Asteropyrum with Coptis shows that they are identical in short chromosomes, with magnoflorina and benzylisaquinodine type of alkaloides, but different from coptis in the chromosome numbers (T-type), pantocolpate pollens, united carpels and the dorsi-ventral type of petioles. In view of these fundamental morphological and cytological differences, Asterop yrum is better raised to the level of Tribe. However Asteropyrum and Coptis may represent two divaricate evolutional lines of Thalictroideae. (2). The systematic position of the genus Kingdonia has been much disputed in the past. We support the view of Sinnote (1914), namely, the trilacunar in leaf traces “the ancient type”, appeared in the angiosperm line very early, while the unilacunar of Kingdonia may be derived from the trilacunar. On the basis of the chromosome numbers and morphylogical observation, the present writer accept Tamura’s and Wang’s treatment by keeping Kingdonia in Ranunculaceae instead of raising it to a family rank as has been been done by Forster (1961). Kingdonia and Coptis are similar in having short chromosome with x=9, but with one-seeded fruits; therefore it is suggested that placed into Thalictroideae as an independent tribe, indicating its close relationship with Coptideae. (3). Comparing with its allies, Calathodes being with out petals, seems to be more primitive than Trollius. But Calathodes differs from Trollius with R-type chromosomes in having T-type chromosome with x=8 and subterminal centromere. Those characteristics show that it is very similar to the related genera of Thalictroideae. But as Kurita already pointed out that most speci es of Ranunculus have usually large long chromosomes but some species have compar ativelly short chromosomes, therefore we regard T-type and R-type chromosomes appear independently in different subfamilies of Ranunculaceae. According to Tamura, G alathodes seems to be closely related to Megaleranthis, because of the resemblance in follicles. But due to lack of cytological data of the latter genus, the relationship between the two genera still is not clear pending further studies. From the fact that the morphology and chromosomes of the Calathodes differs from that of all other genera of the Helleboroideae, we consider Calathodes may form an independent tribe of its own with a closer relationship withTrollieae.9841     

Notulae de Ranunculaceis sinensibus (ⅩⅩⅢ)

(1) In this paper, differences among the five genera constituting the tribe Cimi cifugeae of the family Ranunculaceae are discussed. Beesia, the first genus, with compound cymes and flowers bearing neither petals nor staminodes, is different from the other four genera with simple or compound racemes and flowers bearing either petals or staminodes, and may occupy a primitive position within the tribe. As to the other four genera, Souliea is characterized by the stem without basal leaf but with 2～5 sheath-like cataphylls, the sepals being deciduous but not caducous, moderate in size and petaloid, the petals being much smaller than sepals, but pink in color and more or less petaloid, the pollen grains being pan tocolpate or pantoporate, the carpels being 1～3 per flower, when mature forming dry linear follicles conspicuously reticulate on the surface and dehiscent along the ventral suture, and the seeds being reticulate-foveolate on the surface. These diagnostic characters indicate clear ly that Souliea might have deviated from the lineage formed by the next three genera, i. e. Anemopsis, Cimicifuga, and Actaea, which have their own well-recognizable diagnostic characters. Anemopsis is characterized by the normally developed basal leaf, the racemose inflorescence with sparse and few long pedicellate flowers, the sepals 7～10 in number, mod erate in size, and petaloid, the petals slightly smaller than sepals, the tricolpate pollen grains, the carpels 2～4 per flower, stalked, when mature forming dry oblong follicles with transverse veins on the surface, and the seeds with scaly membranous wings. Cimicifuga is distinguished by the normally developed basal leaf, the caducous, small, often sepaloid sepa ls, the organs of the second floral whorl sometimes with empty sterile anthers being stamin odes not petals, the tricolpate pollen grains, the carpels 1～8 per flower, when mature form ing dry oblong or ovoid follicles with transverse veins on the surface, and the seeds usually with scaly membranous wings. The last genus Actaea is different by the basal leaf trans formed into a small scale, the caducous, small, often sepaloid sepals, the organs of the sec ond floral whorl being clawed petals, the pollen grains with 3(4～6) colpi, carpel 1 per flow er, when mature forming a fleshy indehiscent berry smooth on the surface and without any veins, the seeds roughish or slightly rugose, neither foveolate nor winged on the surface, and the advanced most asymmetric karyotype. According to the diagnostic characters given above, we believe that Beesia, Souliea, Anemopsis, Cimicifuga, and Actaea do represent five independent genera, and the treatment of the tribe Cimicifugeae including these five genera in it by Hutchinson (1923), Janchen (1949) and some other authors, has precisely shown the taxonomic diversity within the tribe. We are therefore unable to accept the treatment published by Compton et al. (1998) to lump the two genera, Souliea and Cimicifuga, into the genus Actaea. (2) Compton et al. (1998, 1997) found out that the Chinese plants previously identified by various authors as Cimicifuga foetida L., in which the terminal and lateral racemes of the compound raceme flower more or less simultaneously, differ from the true C. foetida L. in northern Asia, in which the terminal raceme of the compound raceme flowers before the lateral ones, and thus restored the species name Cimicifuga mairei Lévl. , which was formerly reduced to the synonymy of C. foetida L. , for the Chinese plants. After examining the specimens collected from Siberia and from Southwest China we failed to find out any other differences in both vegetative and reproductive organs between the plants of the two regions, and we consider that it is better to treat the populations in Southwest and Central China as a geographical variety of Cimicifuga foetida L. A new combination, Cimicifuga foetida L. var. mairei (Lévl.) W. T. Wang & Zh. Wang, is thus made. (3) 3 species of Delphinium, 1 species and 1 variety of Clematis are described as new.