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。     

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.bockinan 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。     

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。     

泡沙参复合体(桔梗科)的物种生物学研究 Ⅳ. 等位酶水平的变异和分化

利用聚丙烯酰胺凝胶电泳技术,对采自泡沙参复合体整个分布区的15个天然居群进行了等位酶 分析。7种酶系统共10个等位酶基因位点的检测表明,本复合体植物具有高水平的遗传变异性,居群 水平上的遗传多样性指标分别为:多态位点百分率P=0.40～0.70,等位基因平均数A=1.5～2.5,平 均期望杂合度He=0.102～0.291。3个类群在居群水平上的平均多样性水平由高至低依次为;A． wawreana(A=2.10,P=0.63, He=0.237),A．lobophylla(A=1.85,P=0.60,He=0.204)和A． potaninii(A=1.83,P=O.60,He=0.188)。居群遗传结构和交配系统的分析表明,本复合体植物的大 部分遗传变异存在于居群之内,Fst=0.176(A．wawreana)和0.115(A．potaninii),并以异交为主,异 交率估计值分别为t=0.86(A．wawreana),0.74(A．potaninii)和0.63(A．lobophylla),这是本复合 体植物具有丰富遗传变异的主要原因之一。根据15个居群的遗传一致度(I)值进行聚类分析发现, 15 个居群可分为明显的2支,由A．lobophylla 2个居群构成的一支与由A. wawreana 和A．potaninii 13 个居群构成的一支在遗传上的分化十分明显,而6个A．wawreana 居群和7个A．potaninii 居群间的 遗传差异不明显,是两个分化不充分的类群。因此,本文的等位酶证据说明,A．lobophylla是遗传上独立的好种,同时不支持将A．wawreana 和A. potaninii 分别作为独立种的处理。     

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     

狭叶沙参复合体(桔梗科)的物种生物学研究

通过对狭叶沙参复合体进行居群(共46个)采样、引种、栽培实验,生物学特性观察,染色体观察 (已发表),杂交、同工酶比较、形态性状的定性定量分析、茎解剖和花粉形态的观察,发现：(1)该 复合体主要进行异花授粉,实生苗第二年才抽茎生花;(2)32个居群的染色体计数表明,在辽宁东部、 辽宁西部(绝大部分地区)和山西霍县的居群都为二倍体(2n=34),而其它地方的居群为四倍体(2n 68);(3)叶的酯酶同工酶的变异大,和一些形态特征,尤其是叶的形态和大小的变异相似,而种子的酯 酶同工酶则相当稳定;(4)通过9个居群的杂交试验,在狭叶沙参和石沙参四倍体居群间得到了一 些F1种子,而辽宁西部二倍体与上述的四倍体居群间的杂交未产生任何F1种子;(5)性状分析(包括 聚类分析与主成份分析)和茎解剖的研究,揭示了它们在叶、果实、花和种子的形态、髓纤维组织的有无、 叶锯齿和花萼裂片齿的有无和多少等形状上的变异幅度,并据此分辨出8个宗。其中辽宁西部的二倍 体宗形态上相当独特,被提升为种的等级。 其余7个宗,根据不同的分化程度被处理为三个种和5个 亚种;(6)运用杂交指数法分析了石沙参和狭叶沙参在太行山一带杂交的可能性;(7)分析讨论了复合体内各类群的起源和进化关系。     

河北木蓝的叶表型可塑性研究

以中国分布最广、形态变异复杂且分类上存在争议的木蓝属植物河北木蓝(Indigofera bungeana Walp.)为研究对象,运用GIS技术从较大尺度上(17省28县29个居群)进行叶表型可塑性分析,利用表型可塑性指数和变异系数对叶表型可塑性进行评价,并对叶表型性状与环境因子的相关性进行分析。结果显示:河北木蓝叶表型性状在居群间的变异大于居群内;叶长、叶柄长、最少小叶数、最多小叶数、小叶长、小叶宽6个叶表型性状均具有可塑性,其中叶长的可塑性最大,小叶数目的可塑性最小;年均降水量是对叶表型可塑性影响最大的环境因子;6个叶表型性状与海拔均呈负相关,与年均气温呈正相关。研究结果可为河北木蓝的分类、适应性进化和开发利用奠定基础。     

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。     

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.     

泡沙参同工酶基因位点的遗传分析

利用聚丙烯酰胺凝胶电泳技术 ,对来自天然群体 (居群 )的泡沙参 (Adenophora potaninii Korsh.)及其人工杂交子代进行了 8种同工酶的电泳检测和谱带遗传分析 ,以确定编码这些酶系统的基因位点和等位基因。选用 4种不同的凝胶缓冲系统 ,对下列不同酶系统进行了酶谱的遗传分析 :天冬氨酸转氨酶 (AAT)、酯酶 (EST)、甲酸脱氢酶 (FDH)、谷氨酸脱氢酶 (GDH)、异柠檬酸脱氢酶 (IDH)、乳酸脱氢酶(LDH)、苹果酸酶 (ME)和超氧化物歧化酶 (SOD)。结果表明 ,这 8种酶系统至少由 1 8个基因位点编码 ,其中 1 2个位点为遗传稳定的等位酶位点 ,是可靠的遗传标记。酶谱的分离式样表明 ,EST为单聚体结构 ,AAT、FDH、IDH、SOD为二聚体结构 ,GDH为六聚体结构。最后对同工酶的器官和发育特异性以及同工酶基因位点的遗传分析进行了讨论     

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.     

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。     

PHENOTYPIC PLASTICITY IN DRAPARNALDIA (CHLOROPHYTA: CHAETOPHORACEAE). I. EFFECTS OF THE CHEMICAL ENVIRONMENT1

Eight axenic isolates of Draparnaldia spp. were in defined media under controlled condidom of day length, light intensity and temperature to study effects of the nutrientions on genera; pheuotype and development of main text cells. Phosphate (0–6,8 mg/l P), magnesium (0-9.6 mg/l Mg), sulfate (1.2 –14.0 mg/l S) and chloride (0.2-48.2mg/l Cl) had no effect. Small, but significant, concentration-independent, isolate-specific differences due to sodium and patassium (as the phosphate salt) occured in main axis cell length. The presence of calcium (>1.7 mg/l) was required by all isolates for main axis differentiation. In the absence of calcium, only zoospores and stunted Stigeoclonium- like plants were produced. Nitrate (>5 mg/l N) depressed main axis cell size;at higher levels (> 15 mg/l N) hair prooduction was depressed and several isolates developed a very typical Cloniophora appearance, Several of the isolates did not produce typical Draparnaldia- like phenotypes under any of the chemical formulations tried; therefore, it is suggested that chemical factors alone do not account for either the phemotypec plasticity of Draparnaldia in the field or the failure of others to grow typical plants in culture.     

PHENOTYPIC PLASTICITY IN SCENEDESMUS COMMUNIS (CHLOROPHYCEAE). I. RELATIONSHIP OF S. COMMUNIS TO S. KOMAREKII1

When scenedesmus communis Hegew. (UTEX 76) was transferred daily in dilute media and a low cell density was maintained (ca. 1000 cells · mL^?1), up to 30% unicells were produced in that population. Unlike previously described uncell-coenobium-unicell transformation with other species, these unicells never produced S. communis coenobia (large coenobium type, LCT) but rather small coenobium type (SCT) resembling S. komarekii Hegew. Growth and morphological development of the paratype strain of S. komarekii (UTEX 1236) was compared with an isolated SCT strain (SCT 76–8). SCT 76–8 never produced LCTs and grew significantly faster than UTEX 1236. Both SCT 76–8 and UTEX 1236 produced uncells at low cell densities. Coenobia formed when cell densities increased over time in batch cultures. SCT 76–8 and UTEX 1236 did not differ morphologically when viewed with the light microscope. Under scanning electron microscopy, an outer opaque layer covered an inner warty layer on unicells. The outer layer was reduced or absent in coenobia from batch cultures in stationary growth. In addition, long spikelets, not present on the walls of unicells, were prominent on coenobial walls. The spikelets of UTEX 1236 appeared smaller and more uniformly distributed than in strain 76–8. In contrast, the surface wall morphology of LCT S. communis was composed of an outer reticulate layer supported by spikelets and appeared as a pentagonal meshwork covering the cell walls. This phenotypic plasticity, as demonstrated by SEM and light microscopy, provides further evidence needed for an understanding of Scenedesmus evolution.     

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。