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。     

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。     

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

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 ofA.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 d.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.     

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.     

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     

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.     

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。     

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。     

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.     

濒危植物裂叶沙参种群生殖力表的研究

本文分别以当年产种子数量和一年生幼苗数量为基础,编制了裂叶沙参种群(总和)和海拔2300-2400m, 2700-2900m, 2900-3100m, 3100-3400m 4个不同海拔区域种群的生殖力表,分析了不同种群两种生殖表的种群内禀增长率(r_m),周限增殖率(λ),净增长率(R₀),毛增长率(G)等动态参数与种群动态的关系;比较了两种生殖力表的特点和利弊;讨论了裂叶沙参种群生殖生态学特性和种群发展趋势及其与环境因素的关系。以种子数量为基础编制的生殖力表的各种群动态参数表明,裂叶沙参具有较强的产生种子的能力,特别是在生境条件较好的海拔2700-3100m区域,种群个体结实量可达到很高的水平。以一年生幼苗数量为基础编制的生殖力表表明,裂叶沙参种群从整体上呈衰退趋势。但不同种群间存在差异:在海拔2300-2400m,水份条件较好的特殊生境和海拔2900-3100m的种群呈稳定特征;而海拔2700-2900m,3100-3400m的种群呈缓慢衰退特征。以种子数量为基础编制的生殖力表可以反映裂叶沙参种群产生种子的能力和生殖生态学特征;而以一年生幼苗数量为基础编制的生殖力表包含了种子库的生态学过程,能较好的反映种群动态特征。在实践中,两种生殖力表应配合使用,才能全面刻划种群生殖生态学和种群动态特征。     

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。     

Gas Exchange and Water Use Efficiency of Adenophora lobophylla (Campanulaceae) at Different Altitudes on the East Boundary of the Qing- Zang Plateau

Field measurements were made of leaf net photosynthetic rate (Pn), apparent quantum yield (AQY), dark respiration (Rd), transpiration (Tr), water use efficiency (WUE), stomatal conductance (Gs), intercellular CO2 concentration (Ci), leaf temperature (TI), stomatal density, air temperature (Ta) and relative soil water content (SWC) for Adenophora lobophylla Hong in August, 1996. The species grows in the region from an altitude of 2 300 m to 3 400 m on the eastern boundary of the Qing-Zang Plateau in Sichuan Province, China. Leaf gas exchange, water use efficiency and plasticity of populations in field were compared among different altitudes to evaluate the possible interactions between adaptation of A. lobophylla and environmental factors in these habitats. Pn and AQY at low altitudes were lower than those at high altitudes. They strongly responded to SWC and Ta. On the other hand, Rd at low altitudes was higher than that at high altitudes because of the higher air temperature there. The growth rates at low altitudes were associated with the increases in Rd and a relevant less Pn. Stomata showed strong responses to leaf-to-air vapor pressure deficit at the leaf surface (Vpdl) and Tl in these habitats. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in Pn at high Tl, Vpdl and low available soil water for A. lobophylla at low ahitudes. Nonstomatal limitation to photosynthesis also happened at extreme soil water deficits and high Tl and Vpdl at an altitude of 2 300 m. Tr had a close relationship with stomatal conductance and was also affected by leaf temperature and leaf-to-air vapor pressure deficit at the leaf surface among habitats grown in different ahitudes. WUE increased with altitude. Increasing stomatal densities showed different plasticity of A. lobophylla as altitude increased. SWC and Ta appeared to be important factors to limit carbon assimilation in A. lobophylla at low altitudes, primarily through the process of stomatal closure. The overall results are in consistence with the hypothesis that strong pressure from tmfavorable environmental factors to gas exchange and wateruse of A. lobophylla may prevent their population expansion at low altitudes, which indicate that the above-mentioned restrictions might lead to the endangerment of A. lobophylla.     

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。     

裂叶沙参分布区域和生物学生态学习性的调查

濒危植物裂叶沙参(Adenophora lobophylla Hong)已发现的天然分布区在四川西北部金川县的安宁、曾大、卡撒、马尔邦、观音桥五个乡,分布面积估计不超过5000km2,根据文献记载和50年代采集的标本分析,裂叶沙参的分布区正在缩小。裂叶沙参以有性生殖为主,在外界干挠和春旱严重的条件下也营无性生殖。在海拔2700m以下地区,裂叶沙参星散分布,生长发育不良,年龄结构老化,种群呈衰退趋势。在较为适生的海拔区间(2700-2900m),裂叶沙参生长正常,花果产量高,年龄结构合理。在海拔2900m以上地区,裂叶沙参分蘖数、花果量减少,个体寿命缩短。裂叶沙参所处群落可分为山杨、山棉花林;旱生多刺灌丛或灌丛;旱生灌草丛;和旱生草丛。裂叶沙参种群在各群落中均为伴生种,受到群落不同程度的影响。     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.     

The effects of seed weight on growth and competitive ability of Rumex acetosella from two successional old-fields

Summary The effects of seed size on growth, biomass allocation and competitive ability of Rumex acetosella plants grown either individually or in competition were studied in two populations (6 months and 15 years old respectively) sampled from a postcultivation successional gradient. For plants grown individually there were highly significant effects of seed weight on growth after 43 days, with a higher relative growth rate (RGR) observed for plants raised from heavier seeds. However at the end of the experiment, seedlings developed from lighter seeds had a RGR 2 times greater than those from heavier seeds. Final biomass of the two types was not significantly different after 73 days of growth. When plants were grown individually, there were only slight differences between populations, but when grown in monocultures of 4 plants per pot, plants from the old population had higher root and leaf biomass per pot whereas those from the young population had a higher reproductive effort per pot. This suggests that a trade-off between allocation to sexual and vegetative reproduction occurs over successional time. In mixtures of light and heavy seeds, plants from light seeds were shorter, had fewer leaves and lower biomass than plants from heavy seeds, which were also taller and produced more dry matter than plants grown from heavy seeds in monoculture. The significant effects of seed weight and population on biomass parameters persisted unit the end of the experiment. Seedlings from heavy seeds were strong competitors: those from the young population grew better in the presence of neighbors than in monoculture and those from the late successional population suppressed the more the growth of their partners. Seedlings from light seeds were subordinate competitors. These results suggest that seedlings from seeds of different sizes benefit from contrasting ecological conditions and that selection acts on reproductive output along successional gradients.     

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。     

The unmanaged reproductive ecology of domesticated plants in traditional agroecosystems: An example involving cassavaand a call for data

Although cassava is a strictly vegetatively propagated crop, in many traditional Amazonian agroecosystems, Amerindian farmers recognise volunteer seedlings of cassava and allow them to grow. If their properties are deemed desirable, plants originating from seedlings are included in the harvest of tuberous roots, and their stems are used to prepare cuttings for propagation. Incorporation of these products of spontaneous sexual reproduction appears to be important in origin and maintenance of genetic diversity in this clonally propagated plant. Our observations conducted in an Amerindian village in Guyana suggest that volunteer seedlings arise from a bank of viable seeds stored in soil, and that dispersal and burial of seeds by ants may be important in its constitution. Future investigations of the dynamics of genetic diversity in this crop in traditional agroecosystems must consider the role of the ‘wild’ sexual reproduction that occurs in parallel with vegetative propagation. We suggest that unmanaged processes of sexual reproduction play important but neglected roles in the evolutionary ecology of many domesticated plants in traditional agroecosystems.     

濒危植物明党参种子散布和种子库动态研究

对伞形科濒危植物明党参（Changium smyrnioides)的种子生产、散布和种子库动态进行了研究。明党参平均每株产生完好种子132粒,86.36%来自顶生花序,不同生境中个体产生的种子千粒重有差异;其平均1.06m高的花葶在种子散布过程中易倒伏,使种子远离母株;种子散布格局为聚集型;种子库密度随时间逐渐下降。明党参种子产量和不同时期种子库密度均低于同科非濒危植物峨参（Anthriscus sylvestris),2个种的种子库均为非持续型;明党参具有低种子数、大粒种子、种子散布远离母株以减少种内竞争的K对策,在受到人类大量采挖和生境干扰后种群不容易迅速恢复;而峨参采取高种子数、小粒种子、种子近母株密集分布和产生无性系的r对策,种群受干扰后易恢复。     

川西北泡沙参种群根系生长动态及其与环境因素关系研究

对川西北地区泡沙参种群根系生物量累积和形态发育过程及其与环境因素的关系进行了研究。结果表明,泡沙参种群根系生物量累积一般规律符合Logisticc增长过程。较高的泡沙参根系生物量累积和形态发育时期可以持续到15～17年生以后,根系采收直径应在1．7cm以上。动物啃食、人为采挖、土壤、气候等环境条件与不同海拔各种群的根系生物量累积等有着密切联系,中海拔地区(2800～3300m)人为干扰少,土壤和水热条件适宜,根系生物量和形态发育达到较高水平,适合高产栽培。而低海拔和高海拔地区的不利环境条件限制了泡沙参根系生长,在进行野生资源保护利用以及人工栽培泡沙参时应充分考虑环境因素,努力减少放牧、采挖等人为破坏,有条件的地区应实行分区禁牧、禁采挖,为泡沙参种群恢复和药材品质提高创造条件。