Biosystematic studies on Adenophora potaninii Korsh. complex (Campanulaceae) V. A taxonomic treatment

Abstract Adenophora is an extremely variable genus, and its taxonomy is very controversial. Of the genus, Adenophora potaninii complex, including A. potaninii, A. bockiana, A. wawreana, A. lobophylla, A. biformifolia, A. polydentata, and A. wawreana var. lanceifolia, is a typical group with different taxonomical treatments due to high level of morphological diversity. We carried out extensive biosystematic studies based on population sampling, transplantation experiments and offspring tests, cluster analysis, and a crossing experiment. The results reveal four main findings. (i) Leaf forms of the A. potaninii complex were extremely polymorphic; the leaf form of A. potaninii and A. bockiana, and that of A. wawreana and A. biformifolia could be found, respectively, on a single population or among the offsprings of a single plant. (ii) Cluster analysis and a crossing experiment indicated that A. bockiana and A. polydentata could not be separated from A. potaninii, nor A. biformifolia from A. wawreana. (iii) Adenophora potaninii and A. wawreana were gradational in morphology and their compatibility value was slightly reduced compared to that within each entity. (iv) Adenophora lobophylla was distinct from the other members of the complex in shape and size of corolla, relative length of style, and shape of capsule. This species was incompatible reproductively with the other members of the complex, but partly compatible with A. stenanthina, a species in another section. Therefore, we recognized only one species with two subspecies in the complex, A. potaninii subsp. potaninii and subsp. wawreana, moved A. lobophylla out of the complex, and reduced all the other names as new synonyms.     

Biosystematic Studies on Adenophora potaninii Korsh. Complex (Campanulaceae) III. Genetic Variation and Taxonomic Value of Morphological Characters

Adenophora potaninii Korsh．complex is a morphologically variable group including six species．As shown in many species in the genus Adenophora,the variability of many morphological characters in the complex under investigation is amazingly great,which brings difficulties in the delimitation and indentification of taxa．In the present study,six populations representing five species in the complex were systematically sampled to investigate the pattern of morphological variation within population and to detect the genetic basis of the variation with progeny tests and controlled crosses．The results are as follows： Great morphological variation within population is found (Fig.1),including characters such as leaf shape,teeth number and size of leaf margins,teeth number of calyx lobes,and indumentum on the surface of stems and leaves,which were previously considered as diagnotic．Systematic sampling and statistical analysis show that the differences in the above characters exhibit continuous patterns of variation within population (Fig．1,2),though they are highly correlated (Table 2)．From progeny testing and crossing between two contrast types of individuals (oblong,dentate and pubescent leaves vs．narrow,entire and glabrous leaves)it is evident that those characters show continuous variability in segregating progenies (Fig.3,4)and appear to be influenced by large numbers of loci with individually slight effects．Since most wild populations were highly heterozygous for those characters,the individuals in one extreme of variation can produce their variable offsprings including individuals similar to those in the other extreme (Fig. 3). This is also true for the indumentum density on stems and teeth number of calyx lobes. In this complex, A. biformifolia Y. Z. Zhao, A. bockiana Diels and A. polydentata P.F. Tu et G. J. Xu were described exclusively based on leaf shapes, teeth number of leaf margins and calyx lobes, teeth size of leaf margins, and indumentum on the surface of stemsand leaves. It is demonstrated, however, that the variation of those characters is of a quantitative nature and show no discontinuities. As a result, genetic analysis of diagnostic characters along with their sympatric distribution and same habitats, strongly suggest that A.biformifolia as well as A. bockiana and A. polydentata are actually the extreme individuals within A. wawreana and A. potaninii respectively and should not be recognized as taxa at any taxonomic level.     

泡沙参复合体(桔梗科)的物种生物学研究——Ⅲ.性状的遗传变异及其分类价值

阐明性状变异的遗传基础是评估性状分类价值以及揭示类群关系和进化过程的前提。本文根据泡沙参复合体内6个天然居群的材料,利用居群样本分析、子代测定和杂交试验等手段,结合统计学方法对该复合体一些在以往类群划分中起鉴别作用的形态性状进行了遗传学分析。结果表明,叶形、叶缘锯齿数目和大小、茎叶被毛、花萼裂片齿数、花梗长短等性状都是遗传性比较强的性状,而且性状之间有很强的相关性。但是,这些性状在居群内呈现连续变异且变异幅度非常大,以致于变异的两个极端个体被分别作为不同的种处理。通过对野外居群样本的统计分析和对极端变异类型个体所进行的子代测定及其人工杂交,进一步证实上述性状是受多基因决定的数量性状,其变异幅度大,变异式样中并不存在任何程度的间断性,因而至少在本复合体内没有鉴别意义。因此,以往建立在上述性状变异基础上的3个类群(A.biformifolia Y.Z.Zhao;A.bockiana Diels和A.polydentata P.F.Tu et G.I.Xu)均不成立。最后,本文对形态性状变异的遗传基础及其与类群划分的关系进行了讨论。     

用核糖体DNA的ITS序列探讨裂叶沙参的系统位置——兼论ITS片段在沙参属系统学研究中的价值

本文采用PCR直接测序方法,对来自沙参属全部2组7亚组的10个种和作为外类群的风铃草属2个种的核糖体DNA　ITS片段进行了序列分析。在重点探讨裂叶沙参分类地位的同时,分析了ITS片段序列在沙参属系统发育重建中的价值。结果表明,在沙参属中,ITS片段在长度、GC含量和位点变异量上均比较一致;长度539bp～541 bp,GC含量57％～60％,信息位点只占总位点的3．9～6％。采用PAUP软件进行的系统发育分析表明,裂叶沙参A．lobophylla与大花盘亚组的A．himalayana组成一支,而不是象以往形态学和杂交试验所推断的与泡沙参A．potaninii或A．stenanthina近缘。可见,ITS序列进一步支持将裂叶沙参移出泡沙参复合体甚至移出有齿亚组的推论,但同时也表明将其作为A．stenanthina的近缘种是不适宜的。尽管本研究所测定的类群已涵盖了整个沙参属,但种间序列的两两比较表明,沙参属种间在ITS片段上的分化很小(0.0～3.9％),相比之下,沙参属类群与风铃草属类群间的分化却很高(17.8％～19.2％)。这大概和沙参属起源较晚、遗传分化较小有关。本文还就ITS片段在沙参属和桔梗科系统学研究中的价值进行了讨论。     

A Reevaluation of the Status of A. lobophylla Based on ITS Sequence,with Reference to the Utility of ITS Sequence in Adenophora

Adenophora Fisch. (Campanulaceae)is a medium-sized and diverse genus, which occurs mainly in eastern Asia, particularly in China. Although several taxonomic systems have been proposed, there is still much debate on the delimitation and systematic arrangement of the species within the genus, due mainly to high variation in morphology, habitats, phenotypic plasticity, and potential hybridization. A. lobophylla Hong is a rare perennial endemic to several counties of Sichuan, China. Since this species was recognized, much work has been done on its taxonomic status, including morphology, crossing experiment and allozymes. However, evidence available is not concordance so far, and its relationship with other species within the genus remains largely uncertain. In the present paper, the internal transcribed spacer(ITS) region of 18s～26s nuclear ribosomal DNA was sequenced in 10 representatives of Adenophora and two outgroup species of Campanula to assess its utility for phylogenetic reconstruction in Adenophora, in addition to reevaluation of the taxonomic status of A. lobophylla. The results indicate that there exist high sequence alignability and length conservation among ITS sequences, with informative sites being 3.9% and length ranging from 539 to 541 bp and G + C content varying from 57% to 60%. Phylogenetic　analyses using Fitch parsimony show that A. lobophylla is closely related to A. himalayana instead of A. potaninii or A. stenanthina as revealed by previous morphological work and crossing experiment. The ITS phylogeny is generally congruent with other evidence in that A. lobophylla should be removed from the A. potaninii complex or even subsect. Microdiscus. Its appropriate placement, however, should be further explored. In pairwise comparisons at unambiguously aligned sites, sequence divergence was relatively low among Adenophora species with the values ranging from 0.0 to 3.9% in spite of the fact that the species surveyed covered all of the sections and subsections of the genus. In contrast, high sequence divergence is found among Adenophora and Campanula species with the values from 17.8 % to 19.2 %. The low divergence among Adenophora species is justified considering its recent origin, and also suggests that ITS sequence would provide a promising source of nuclear phylogenetic markers to reconstruct the phylogeny within Campanulaceae on the tribe and family levels.     

遮光条件下裂叶沙参和泡沙参气孔行为的对比研究

通过遮光实验研究了裂叶沙参(Adenophora lobophylla)和泡沙参(A. potaninii)叶片的气孔行为,比较了不同遮光处理对裂叶沙参和泡沙参气孔的生理特性以及生态适应的影响。结果表明:裂叶沙参对不同遮光处理的适应性较差,相反,泡沙参具有较强的适应性和抗逆性;在自然光照和轻度遮光条件下,中午时泡沙参叶片的气孔开度逐渐减小和部分气孔关闭;裂叶沙参光合作用的非气孔限制占主要地位,泡沙参光合作用的气孔限制较明显。     

裂叶沙参与泡沙参种群分布格局分形特征的分析

 裂叶沙参与泡沙参种群在不同海拔区域分布格局的计盒维数与其在群落中占据空间的能力成正比,并且与环境因素密切相关。在裂叶沙参与泡沙参种群较为适生的区域,海拔2700～3100m,两者在群落中占据空间的能力最强,他们的计盒维数达到最高。在低于海拔2700m或高于海拔3100m,环境条件严酷的区域,裂叶沙参种群与泡沙参种群占据空间的能力不同程度的下降。在不同的海拔区域,广布种泡沙参种群占据空间的能力均比裂叶沙参种群高,而其受环境因素影响较弱。在海拔2700m以下地区,泡沙参种群水平分布格局的计盒维数比裂叶沙参种群高4.5倍;在两种群较为适生的区域,海拔2700～3100m,泡沙参计盒维数比裂叶沙参种群高1.5倍,差异最小;在泡沙参与裂叶沙参种群分布的上限,海拔3100m以上地区,两者计盒维数差异又增加,泡沙参计盒维数比裂叶沙参种群高1.8倍。这说明裂叶沙参种群的内在适应力、竞争力比泡沙参种群相对较弱。     

濒危物种裂叶沙参及其近缘广布种泡沙参的遗传多样性研究

根据对１２个形态性状的统计分析和１０个基因位点的等位酶检测,探讨了濒危植物裂叶沙参及其近缘广布种泡沙参的遗传多样性水平,对３个裂叶沙参和６个泡沙参天然群体的遗传分析表明,两种沙参属植物均具有很高的遗传变异水平,这种变异性既体现在形态学水平上,也体现在酶位点水平上,在２个茎叶形状以及１０个花果和种子形状上,濒危种裂叶沙参的变异性均与广布种相当,同样,根据７个酶系统１０个等位酶位点的度量,裂叶沙参群体     

裂叶沙参与泡沙参解剖生态学的比较研究

借助于光镜和扫描电镜,系统观察了濒危植物裂叶沙参(Adenophora lobo-phylla Hong.)的根、芦头、茎和叶的解剖特征及茎、叶、种子的表面形态特征,并将有关性状与广布种泡沙参(A.potaninii Korsh.)作了对比研究。结果表明,两种沙参在叶表皮细胞形状,垂周壁式样,角质层初级雕纹及次级雕纹;叶片气孔形状及外拱盖外缘纹饰及内缘形状,保卫细胞上角质层薄厚及蜡质纹饰;叶表毛状体形状及纹饰,茎中皮层占茎横切面的比例;茎中有无髓纤维等方面存在着明显差异。相对于泡沙参,裂叶沙参单株叶面积大,生物量高,表皮角质层薄,单位面积内被毛数量少,栅栏组织薄,气孔周围蜡质沉积颗粒少;茎的直径大,皮层较薄等。这些特征表明裂叶沙参对旱生生境适应能力弱,而泡沙参对旱生生境适应能力强。     

裂叶沙参和泡沙参气孔行为与蒸腾特性的关系

采用LI-6400便携式光合分析系统对裂叶沙参和泡沙参进行了不同遮光水平蒸腾速率的测定,并同步测定了叶片气孔大小、气孔密度、气孔开度和气孔导度等气孔行为.采用相关系数和通径系数分析方法分析了不同遮光水平裂叶沙参和泡沙参气孔行为与蒸腾特性的关系.结果表明:不同遮光水平测定的气孔行为对裂叶沙参和泡沙参蒸腾作用的直接影响大致相同,其中,两种沙参的气孔密度对蒸腾速率的直接影响较大.但泡沙参的气孔导度与蒸腾作用具有极显著的相关关系.     

Comparative Studies of Sexual Reproduction and Asexual Propagation Between Populations of Adenophora lobophylla and A. potaninii

To reveal the internal mechanism which lead to the endangerment of the AdenophoUa lobophylla Hong population, comparative studies of sexual reproduction and asexual propagation between the populations of A. lobophylla and A. potaninii Korsh were carried out. Young seedlings of A. lobophylla had less ability to occupy the uper space of the herbaceous layer than those of A. potaninii. The A. lobophylla population produced much more seeds and lived a long reproductive life to adapt the ambient environment. Though the number of flowers and fruits of A. lobophylla population was more than those of A.potaninii population, the number of its ripe seeds as well as the quality was less. Thus, the development rate from its seeds to one-year old seedlings was very low, and viability of its young seedlings was very poor. This could be the important factor causing the imminent population endangerment of A. lobophylla. Besides the sexual reproduction, the A. lobophylla population also had a comparatively tiny ability of asexual propagation, which was advantageously complementary to its sexual reproduction.     

Isolation and Characterization of Total DNA from Several Endangered Species and Their Allies

A low pH extraction medium with high salts, which avoids ionization and subsequent oxidation of phenolic compounds during tissue grinding and precipitation of large amounts of materials, were successfully used to obtain total DNA from Cathaya argyrophylla Chun et Kuang, Paeonia suffruticosa var. spontanea, Cimici fuga nanchuanensis Hsiao, Adenophora potaninii (Congeneric species with A. lobophylla) etc. The DNA yields, quality and purity were characterized. These isolated DNA could be used directly for RFLP and RAPD analysis which are useful as molecular genetic markers without sedimentation in cesium chloride gradient or column chromatography. A fast, inexpensive and reliable procedure has been developed for detecting the genetic diversity of endangered plants.     

Biosystematic Studies on Adenophora potaninii Korsh. Complex (Campanulaceae). Ⅰ. Phenotypic Plasticity

Phenotypic plasticity is the environmental modification of genotypic expression and an important means by which individual plants respond to environmental heterogeneity. The study of phenotypic plasticity in the genus Adenophora, which is very complicated taxo nomically because of great morphological variation, proves to be helpful in both investigating the phenotypic variation so as to evaluate potential taxonomic value of their characters and providing important sources of information on the variation, adaptation and evolution of the genus. Twenty-three populations representing all the six species in Adenophora potaninii complex were transplanted into the garden. Of them six populations were selected for study ing their performance in the field and in the garden, in addition to cultivation experiment under different treatments. The results show that there exists considerable developmental plasticity in some leaf, floral and capsule characters. In particular, the leaf shape and length of calyx lobe display significant developmental variation with the maximum being three times as great as the minimum, which is noteworthy because they were previously considered as diagnostic. The characters of root, caudex, stem and inflorescence are found to be very plastic, especially the root diameter, the number of stems, stem height and inflorescence length with great environmental plasticity. In addition, the populations from different habi tats show distinct amounts of plasticity. On the contrary, the characters of leaf, floral, cap sule and seed are less influenced by environments. It seems that the considerable variation in the characters of leaf is attributed mainly to genetic differences. Finally, the phenotypic plasticity of morphological characters of A. potaninii complex and its taxonomic significanceis discussed.     

几种濒危植物及其近缘类群总DNA的提取与鉴定

用低ｐＨ 介质,高盐沉淀蛋白质方法成功地从银杉（Ｃａｔｈａｙａ ａｒｇｙｒｏｐｈｙｌｌａ Ｃｈｕｎ ｅｔＫｕａｎｇ）、矮牡丹（Ｐａｅｏｎｉａ ｓｕｆｆｒｕｔｉｃｏｓａ ｖａｒ． ｓｐｏｎｔａｎｅａ）、南川升麻（Ｃｉｍ ｉｃｉｆｕｇａ ｎａｎｃｈｕａｎｅｎｓｉｓＨｓｉａｏ）、裂叶沙参（Ａｄｅｎｏｐｈｏｒａ ｌｏｂｏｐｈｙｌｌａ）的同属种泡沙参（Ａ． ｐｏｔａｎｉｎｉｉ）等植物中提取和部分纯化了细胞总ＤＮＡ,并对其产率、质量和纯度作了鉴定。此方法的关键是用了一个低ｐＨ提取介质,它能有效防止组织破碎及沉淀大量材料时的电离化作用及酚化合物的进一步氧化。所得ＤＮＡ 不需经氯化铯梯度离心或柱层析,直接可用于限制性片断长度多态性（ＲＦＬＰ）及随机扩增的ＤＮＡ多态性（ＲＡＰＤ）等分子水平的遗传标记。为检测濒危植物的遗传多样性提供了一套迅速、简便和可靠的技术方案     

泡沙参复合体（桔梗科）的物种生物学研究：Ⅰ表型的可塑性

表型可塑性是生物变异中由环境引起的一种变异,是植物适应的一种重要方式。对沙参属这样一个形态上复杂多变、分类上很难处理的类群,研究其表型可塑性不仅为探讨性状变异、判断其系统学意义及选择可靠的分类性状提供了有益的资料,而且有助于揭示沙参属植物变异、适应和进化的机制。本文从泡沙参复合体中选择了6个居群,利用播种和移栽试验,通过对不同个体和居群在一致条件下的表现及野外和移栽后的对比,对根、茎、叶、花和果等形态性状的表型可塑性进行了初步的观测分析。结果表明,一些叶片、花部和果实性状具有较大的发育可塑性,尤其是叶形、花萼裂片不仅发育变化大,而且随发育过程定向变化。环境可塑性较大的性状主要是根、茎、花序分枝等性状,而叶片、花部、果实和种子性状的环境饰变能力都较小。最后,对泡沙参复合体形态性状的变异从发育可塑性和环境可塑性的角度进行了讨论。     

裂叶沙参和泡沙参种群有性生殖和无性繁殖的比较

通过对裂叶沙参(Adenophora lobophylla Hong)和泡沙参(A.potaninii Korsh.)种群有性生殖和无性繁殖特性的对比,揭示濒危植物裂叶沙参的种群内部致濒机制。结果表明:裂叶沙参种群幼苗抢占草本层上层空间能力弱于泡沙参种群;裂叶沙参种群是以相对长的生殖期和高产籽量来适应环境;裂叶沙参种群开花结实量虽高于泡沙参种群,但其中成熟种子少,种子质量差,致使其种群由种子到一年生幼苗的转化率极低,并且幼苗生活力弱,是导致其种群濒危的重要内部原因。裂叶沙参种群除有性生殖外还兼有较弱的无性繁殖,是对其有性生殖的一个重要补充。     

Biosystematic Studies on Adenophora potaninii Korsh. Complex (Campanulaceae) Ⅳ. Allozyme Variation and Differentiation

Fifteen natural populations representing all the taxa in Adenophora potaninii complex were investigated using vertical slab polyacrylamide gel electrophoresis. Allozyme data for ten loci of seven enzymes demonstrate high levels of genetic variation within populations, with the values of P＝0.40～0.70, A＝1.5～2.5 and He＝O.102～0.291. At specific level, there existed the highest variation in A. wawreana (A= 2.10, P=0.63, He ＝0.237), followed by those of A. lobophylla (A＝1.85, P=0.60, He＝0.204) and A. potaninii (A＝1.83, P＝0.60, He ＝O.188). Based on Wright's F-statistics, the outcrossing rates( t ) were 0.86 ( A.wawreana ), O.74 ( A.potaninii ) and 0.63 ( A.lobophylla). The analysis on population genetic structure indicates that most variation resided within populations, which is consistent with the high rate of outcrossing rates in the complex. On the basis of genetic identity(I), cluster analysis shows that there was high genetic divergence between A. lobophylla populations and A. wawreana-A, potaninii clade, while no clear genetic difference was found between A. wawreana populations and A. potaninii populations. Consequently, the present allozyme data justify the establishment of A. lobophylla, but give no support for treating A. wawreana and A. potaninii as a sepa-rate species. Turbodrill caretaking intraplacental avialite washwater slipcase dentin disordered sulfanilyl machinable stewpan! Netherward pressbodies horror abscissa, keratosis frieze. 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Sporogenesis and Gametogenesis of Endangered Adenophora lobophylla Hong (Campanulaceae)

Reported in this paper are the microsporogenesis, megasporogenesis and the development of male and female gametophytes of the endangered species Adenophora lobophylla. The anther is four-sporangiate with its wall composed of four layers: epidermis, fibrous endothelium, middle layer and glandular tapetum of binucleate cells. The cytokinesis of microspore mother cell in meiosis is simultaneous, and tetrads are tetrahedral. Pollen grains are 2-celled at the dispersal stage. Ovule is anatropous, unitegmic and tenuinucellate with a hypodermal archesporial cell developing directly as the megaspore mother cell which undergoes meiotic division and then forms a linear tetrad. The embryo sac is of Polygonum type. The polar nuclei fuse before fertilization. An endothelium differentiates when the uninucleate embryo sac forms. Comparative studies were made with the closely related and widely distributed species A. potaninii. No differences between them were found. No obstacle to sexual reproduction in this endangered species was observed. As a conclusion, no endangering factors were found to influence the sporogenesis and gametogenesis of A. lobophylla. According to our observision and the references on embryological studies of the Campanulaceae(s. l. ), there is no evident differentiation in sporogenesis and gametogenesis in this family.     

濒危植物裂叶沙参生境条件及外界致危因素分析

裂叶沙参分布于四川西北地区金川县,受到当地地质、地貌、气候、土壤、生物和人为因素等多方面的综合影响,形成了裂叶沙参种群生长发育的外部特定环境条件。裂叶沙参种群生长发育最主要的外界致危因素是:人为采挖、森林过伐、开荒、放牧所导致的生境破坏;频繁的地震、滑坡、泥石流等地质灾害;周期性的春旱、伏旱和暴雨所引发的洪涝灾害;锈病、地下害虫和野生动物的危害。这些致危因素周期性或经常性地对裂叶沙参种群数量、个体生长产生着长期而广泛的危害。应立即停止人为对当地生境条件的一切破坏活动;在金川县建立以保护裂叶沙参为主的自然保护区,并在适生地引种栽培,实行迁地保护;对裂叶沙参种群继续跟踪研究,为揭示一般濒危植物的内部致危机理及外界致危因素提供科学参考。     

濒危植物裂叶沙参的大小孢子发生及雌雄配子体发育

本文以近缘广布种泡沙参为对照,对濒危物种裂叶沙参进行了大小孢子发生和雌雄配子体发育的研究。裂叶沙参的药壁发育为双子叶型,绒毡层为腺质型,细胞含两核。小孢子母细胞在减数分裂过程中胞质分裂为同时型。小孢子四分体为四面体型,成熟花粉粒为2-细胞型。胚珠倒生,单珠被,薄珠心,大孢子四分体为线形排列,胚囊发育为蓼型。成熟胚囊中两极核在受精之前融合为一个大的次生核。当胚囊发育至单核胚囊时,珠被的最内层细胞发育为珠被绒毡层。濒危植物裂叶沙参在大、小孢子发生及雌、雄配子体的发育过程中,未见有败育及其它异常现象;与对照种泡沙参相比,也未见有差异,这说明裂叶沙参的致濒原因不在于有性生殖过程。