密度制约作用对箭叶淫羊藿有性繁殖时期形态特征的影响

运用回归分析和非参数检验方法,研究了箭叶淫羊藿(Epimedium sagittatum)在有性繁殖的三个时期克隆片段密度对其各个形态特征的影响,以揭示其资源和繁殖投资情况。结果表明：箭叶淫羊藿在三个有性繁殖期间克隆片段密度对其各个形态特征的影响显著。从回归分析的R2指数来看,其克隆片段密度增大对三个生长时期的制约作用是由弱到强,同时在有性繁殖期间随克隆片段密度的增加其资源投资趋向于有性繁殖;从非参数检验结果来看,有性繁殖期间箭叶淫羊藿明显增加其叶数、叶宽等来获取更多的资源用于提高其繁殖能力,以取得生存。     

箭叶淫羊藿与近缘种天平山淫羊藿花粉形态对比研究

该研究采用扫描电子显微镜(SEM,scanning electron microscope)对箭叶淫羊藿13个居群(含箭叶淫羊藿光叶变种1个居群)、天平山淫羊藿7个居群进行花粉形态和外壁纹饰的观察及对比研究。结果表明:两个类群的花粉均为近球形,具三孔沟,大小相近。箭叶淫羊藿的花粉大小为(18.21～21.66)μm×(19.55～23.63)μm,天平山淫羊藿的花粉大小为(18.32～21.43)μm×(19.73～23.48)μm。外壁纹饰是花粉形态的重要性状。不同居群间花粉外壁纹饰存在较大差异,主要分为网纹纹饰、条网纹纹饰和颗粒状纹饰三大类,继而根据网脊类型和网眼密度分为六亚类。其中,箭叶淫羊藿的花粉形态包含了全部的三大类六亚类,天平山淫羊藿包含其中的三大类四亚类。天平山淫羊藿花粉特征为箭叶淫羊藿花粉形态变异的子集。整体而言,外壁纹饰类型随纬度和海拔增加呈现出由网状纹饰向条网状纹饰过渡的趋势。花粉主要形态性状特征均与海拔存在密切相关性。在花粉形态及外壁纹饰特征上,天平山淫羊藿在箭叶淫羊藿变异范围之内,与其他宏观形态性状变异一致。花粉形态性状无法将两类群有效区分,未能提供天平山淫羊藿...     

三种药用淫羊藿的地理分布与资源调查

淫羊藿属(Epimedium L.)植物为传统的中草药,也是新优地被观赏植物,具有悠久的药用历史和巨大的开发潜力.对3种重要的药用淫羊藿(朝鲜淫羊藿E.koreanum、心叶淫羊藿E.brevicornum和箭叶淫羊藿E.sagittatum)的地理分布、形态特征和资源现状进行了调查.调查发现,朝鲜淫羊藿分布于中国东北部,主要沿长白山脉分布,长期的过度采收使野生资源遭到严重破坏,保护工作迫在眉睫;心叶淫羊藿分布于西北各省以及山西、河南等省,野生资源丰富,但目前开发利用较少;箭叶淫羊藿广泛分布于华东、华南各省区,但野生资源蕴藏量并不丰富,且不同分布区形态变异较大,物种鉴定尚存诸多疑问.为保证野生资源的可持续利用,未来应控制朝鲜淫羊藿的过度采收,加强心叶淫羊藿的合理开发利用,不同地区的箭叶淫羊藿形态差异较大,质量不稳定,建议对其进行系统的质量评估之后择优开发利用.     

川产淫羊藿属（Epimedium）药用植物的无性系构件特征和生物量配置比较

在构件水平上,对川产的3种药用淫羊藿属植物（淫羊藿（Epimedium brevicornu）、箭叶淫羊藿（E.sagittatum）、柔毛淫羊藿（E.pubescens））的无性系构件特征、克隆构型及构件生物量配置进行比较,结果表明在野生状态下：该属3个种无性系构件的形态特征差异显著,且存在较大的变异性,变异系数范围在29.29%~48.31%之间;箭叶淫羊藿更趋于“游击型”克隆构型,而淫羊藿和柔毛淫羊藿则更趋于“密集型”克隆构型;该属3个种都将高比例生物量配置到叶片或者根茎,其次为茎,再其次为根。柔毛淫羊藿具有更强的克隆繁殖能力,单位面积上各构件生物量均最大,应作为人工引种栽培的首选种。     

箭叶淫羊藿居群形态及遗传多样性比较研究

箭叶淫羊藿（Epimedium sagittatum）是淫羊藿属中分布最广、形态变异最大、分类学最难处理的一个物种,不同箭叶淫羊藿居群其形态、活性成分等差异较大,质量极其不稳定。本实验选择湖北罗田等12个不同的箭叶淫羊藿地理居群,在武汉植物园进行同园栽培,分析其主要形态数量性状及遗传多样性。结果发现箭叶淫羊藿不同居群在形态上表现出各自明显的差异。基于AFLP数据进行的遗传多样性分析显示,各居群聚类关系与地理分布密切相关,柳州居群（LZ）、西南居群（CL、HH、ZY）、华东华中居群（YT、WN、NF、HS、QZ、SG、LY、LT）依次分出。而形态分层聚类分析显示各居群形态变异复杂,只有部分与遗传多样性有一定的相关性。本研究结果对箭叶淫羊藿分类研究及资源筛选具有重要指导作用。     

淫羊藿分株种群特征及其与箭叶淫羊藿空间分布的点格局分析

对四川省南充市金城山3个不同海拔梯度(560m、653m、774m)上的淫羊藿分株种群特征以及用聚块性指标(m*/m)、聚集指数、Cassie指标、扩散系数和点格局分析法,分别对淫羊藿基株种群与海拔771m处箭叶淫羊藿基株种群的空间分布格局进行了研究。结果显示:在低海拔梯度560m处淫羊藿分株种群密度达到最大,随着海拔的升高,淫羊藿分株种群密度显著减小。不同海拔高度下,淫羊藿分钻种群根冠比在774m处最高。3种海拔梯度上,淫羊藿基株种群空间分布格局为集群分布;箭叶淫羊藿基株种群在尺度(t)0~0.01之间呈随机分布,在0.01~0.5之间呈集群分布。最后,结合保护淫羊藿和箭叶淫羊藿的药用资源以及点格局分析方法在研究草本植物种群空间分布格局中的优越性展开了讨论。     

箭叶淫羊藿同质园栽培居群非腺毛多样性及其分类学启示

淫羊藿是国际上多年来重点研究的药用植物.箭叶淫羊藿(Epimedium sagittatum)是淫羊藿属分布最广,也是形态变异最大的物种,箭叶淫羊藿复合群种分类中存在诸多疑点和争议.为了重建淫羊藿属的系统发育,我们利用数码显微摄影技术对10省16个箭叶淫羊藿同质园栽培居群的叶背非腺毛性状进行了比较研究.结果表明,箭叶淫羊藿非腺毛性状的主要特征包括:细胞数、非腺毛长度、是否存在伸长细胞、顶细胞形态及由此导致的顶细胞直径和壁厚差异.不同居群的非腺毛形态和特征存在巨大差异和丰富的多样性,其中项细胞形状是最重要的特征,并与全长、有无伸长细胞、项细胞比例、顶细胞直径和壁厚等多个性状密切关联.聚类分析将16个居群划分为两个类型,即梭形组和剑形组.变异式样的分析表明个体间和居群间整体呈现连续变异,丰富的多样性可能是多型种的表现.但形态上的分化往往来源于关键性状特征的强烈选择作用引起的居群间适应性分化,同质园栽培个体和居群间的形态差异暗示着适应性相关性状的遗传分化,居群间的形态差异应为地方适应性的结果,这种差异可能是物种形成的雏形.     

盐胁迫对两种淫羊藿属植物生长及各器官总黄酮含量的影响

采用不同浓度NaCl(0、100、200、300和500 mmol/L)的Hoagland's培养液处理两种淫羊藿属植物--淫羊藿、箭叶淫羊藿幼苗,处理30 d后分别测定其株高、叶表面积、干物质生物量以及根、茎、叶的总黄酮含量,以明确淫羊藿属两种药用植物的抗盐特性.结果显示:(1)盐胁迫对淫羊藿和箭叶淫羊藿的生长均有影响,随着NaCl处理浓度的增加,淫羊藿、箭叶淫羊藿的株高和叶表面积均逐渐显著减小;各器官干物质生物量逐渐降低,但淫羊藿的干物质下降量较小(不显著),且同一处理下淫羊藿的生物量明显小于箭叶淫羊藿.(2)随着NaCl胁迫浓度的增加,除淫羊藿茎的总黄酮含量一直呈下降趋势外,淫羊藿根、叶和箭叶淫羊藿根、茎、叶均呈现在低浓度范围内总黄酮含量增加,且箭叶淫羊藿的含量高于淫羊藿;而在高浓度盐胁迫下,其总黄酮含量均呈下降的趋势,且淫羊藿的含量高于箭叶淫羊藿.研究表明,淫羊藿的抗盐性明显高于箭叶淫羊藿,适于高盐土壤环境种植,但在低盐环境条件下,种植箭叶淫羊藿可显著提高总黄酮产量并提高经济效益.     

柔毛淫羊藿生物量与生殖配置研究

探索了柔毛淫羊藿在有性繁殖时期不同生境和不同发育阶段的生殖配置,以探讨柔毛淫羊藿的生物量及生殖配置的内在规律.结果如下:随着生长季节的变化,柔毛淫羊藿总生物量呈现出低-高-低的趋势,柔毛淫羊藿生物量积累最快的时间为柔毛淫羊藿花期和果期.其中地上部分生物量在不断增加,地下部分生物量变化较小,从而根冠比在有性繁殖时期逐渐下...     

不同生境中淫羊藿克隆构型和分株种群特征

为探求淫羊藿资源的保护策略,通过野外样地调查、室内测定和统计分析等方法,对林缘旷地、林缘和林下三种不同生境中淫羊藿(Epimedium brevicornum)克隆构型及其分株种群特征进行了研究,并初步分析环境因子对淫羊藿克隆构型和分株种群特征的影响.结果表明:分枝强度、间隔子长度、分枝高度、根状茎长度及分株种群密度在3种生境中差异显著,且与生境相对光强和土壤含水量有较强的回归关系.群落相对光强、土壤含水量在淫羊藿的克隆生长过程中起着重要作用.淫羊藿以地下根茎为克隆器官进行克隆生长,有着较强的克隆构型可塑性.结合克隆植物对资源的利用策略,讨论不同生境中淫羊藿克隆构型和分株种群特征可塑性的生态适应意义及淫羊藿保护机制.     

The effect of local resource availability and clonal integration on ramet functional morphology in Hydrocotyle bonariensis

Summary Within a physiologically integrated clone, the structure and functioning of an individual ramet is determined by: 1) the response of that ramet to its local environment and 2) its response to resource integration within the clone. In a multifactorial experiment, Hydrocotyle bonariensis ramets were grown in limiting resource environments with and without the benefit of basipetal resource movement from another branch of the clone. Ramets were analyzed for their morphological responses to variation in local light, water and nitrogen availability and to the superimposed effect of resource integration on these conditions. The expression of ramet morphology, from induction to development, was highly plastic in response to variable local resource availability. Resource integration changed a ramet's local response in a variety of ways depending on the resource(s) being translocated and the character involved. Among leaf characteristics (leaf weight, petiole height, blade area), resource translocation into the shade resulted in an enhancement of the local response. Similarly, the translocation of nitrogen and water generally increased clonal proliferation and sexual reproduction among ramets. In contrast, the translocation of water reversed the effect of local low water conditions on ramets by inhibiting root production. Some characters such as internode distance and leaf allometry were unaffected by integration. The maintenance of connections between ramets as a Hydrocotyle clone expands allows for resource sharing among widely separated ramets and can result in an integrated morpological response to a resource environment that is patchy in time and space.     

初始克隆分株数对大米草表型可塑性及生物量分配的影响

克隆植物大米草 (Spartina anglica) 目前在我国出现了严重的自然衰退 (Dieback),为了阐明大米草衰退的机理,分析影响大米草形态可塑性的因素与自然衰退之间的相关性,以期为近缘植物互花米草 (S. alterniflora) 这一爆发种群的生物控制提供借鉴,对3种不同初始克隆分株数 (单克隆、三克隆和五克隆) 大米草的克隆生长、生物量累积与分配和异速生长特征进行了野外栽培试验。研究结果表明,初始克隆分株数对间隔子长度影响较弱;初始多克隆的分支强度高于初始单克隆;初始三克隆和五克隆在总生物量 (7.921 5~10.431 7 g 和 8.903 9~10.431 7 g)、地上生物量 (3.396 1~4.255 8 g 和3.618 4~4.338 9 g)、地下生物量 (4.286 9~5.206 6 g 和 5.298 8~6.079 3 g)和根状茎生物量 (1.318 6~1.767 7 g 和 1.499 1~2.038 7 g) 积累上均显著高于初始单克隆,不同初始克隆分株数条件下根生物量差异不显著;初始多克隆倾向于将资源更多地分配给根状茎,而初始单克隆倾向于将更多的资源分配给根系。由此推断,在不同初始克隆分株数条件下,大米草的形态可塑性和生物量分配格局的差异显示出在同样资源格局下,初始多克隆的克隆生殖能力较初始单克隆强。初始多克隆生长的大米草较初始单克隆生长的大米草更能占据优势生境,选择生境“觅养”的能力与克隆繁殖能力更强。     

Effects of clonal integration on plant plasticity in Fragaria chiloensis

The ability of clonal plants to transport substances between ramets located in different microsites also allows them to modify the plastic responses of individual ramets to local environmental conditions. By equalising concentrations of substances between ramets, physiological integration might decrease responses to local conditions. However, integration has also been observed to increase plasticity and induce novel plastic responses in ramets. To ask how integration modifies plant plasticity in the clonal herb, Fragaria chiloensis, ramets were given either low light and high nitrogen or high light and low nitrogen, simulating a pattern of resource patchiness in their native habitat. Ramets in contrasting light/nitrogen treatments were either connected or single. Effects of light/nitrogen and connection were measured at three levels of morphological organisation, the organ, the ramet, and the clonal fragment. Connection between ramets reduced or had no effect on plastic responses in leaf size at the level of the plant organ. This suggested that integration dampened certain plastic responses. Connection induced a new plastic response at the level of the clonal fragment, an increase in allocation to vegetative reproduction in patches of low light and high nitrogen. It is concluded that clonal integration can have different effects on plant plasticity at different levels of plant organisation. It appears that, at least in this species, integration can increase plasticity at the level of the clonal fragment and concentrate vegetative reproduction in particular microsite types.     

Population demographics and trade-offs to reproduction of an invasive and noninvasive species of <Emphasis Type="Italic">Rubus</Emphasis>

Do trade-offs between growth and reproduction differ between an invasive and noninvasive plant species and how do such trade-offs relate to population demographics? To help address these questions, we compared demographics for an invasive plant species, Rubus discolor, with a noninvasive congener, R. ursinus, in several populations of varying density. Removal of floral buds from reproductive canes increased the size of juvenile canes that arose from clonal sprouting in R. ursinus, suggesting a trade-off between current reproduction and growth. Removal of floral buds had no effect on growth of R. discolor. R. ursinus displayed trade-offs between reproduction (sexual and vegetative) and future growth based on negative correlations between leaf area production and both clonal sprouting and seedling production during the previous year. R. discolor did not exhibit these trade-offs. Both species had high population growth rates in low-density populations, but exhibited little or no growth in high-density populations. A life table response experiment was used to determine the underlying cause for the effect of density on population growth. For R. ursinus, lack of population growth in high-density populations was due primarily to increased mortality of clonally sprouting canes, while for R. discolor, it was due to decreased clonal cane production. Elasticity analysis revealed that clonal growth was more important than sexual reproduction for population growth of both species. However, elasticity values for sexual reproduction in R. discolor were greater in high- than low-density populations. This suggests an increased reliance on sexual reproduction in populations that had reached stable sizes, which could increase the capacity of R. discolor to disperse to new sites. Elasticity analyses were also used to simulate the efficacy of various control strategies for R. discolor. Control of this species could be attained by reducing clonal production within existing populations while reducing seed production to limit establishment of new populations.     

高寒草甸退化对鹅绒委陵菜克隆生长特征的影响

为了揭示高寒草甸典型匍匐茎克隆植物对不同生境的生态适应对策,验证生境适应假说,并为高寒草地的退化演替机理研究提供依据,以调查统计和比较样地法研究了江河源区高寒草甸退化对鹅绒委陵菜克隆生长特征的影响。研究表明,重度退化草甸的植物群落结构、功能以及土壤特征发生了明显的变化,继而对鹅绒委陵菜无性系的克隆生长行为和形态特性产生了影响。高寒草甸退化后鹅绒委陵菜的匍匐茎增多,分支强度加大。退化草甸内鹅绒委陵菜的基株高度小于未退化草甸,根长大于未退化草甸,基株的叶片数目问没有明显差别。退化草甸的分株高度显著小于未退化草甸,而分株叶数大于未退化草甸,根长小于未退化革甸且差异不显著。未退化草甸内鹅绒委陵菜无性系的问隔子长度、粗度和匍匐茎长度大于退化草甸,间隔子平均数目少于退化草甸,差异都不显著。随着鹅绒委陵菜无性系匍匐茎数目的增加,不论重度退化草甸与未退化草甸,用于鹅绒委陵菜克隆繁殖的能量投资也逐渐增加。鹅绒委陵菜在未退化草甸用于克隆繁殖的能量投资比例高于退化草甸,其中未退化草甸内鹅绒委陵菜基株的干重占无性系总生物量的比例略低于退化草甸,分株和匍匐茎的干重占无性系总生物量的比例高于退化草甸。高寒草甸退化对鹅绒委陵菜克隆生长特征的这些影响,与植物群落结构和功能的变化导致生境变异密切相关,是其对高寒草甸退化导致的资源和生境差异的反应,也是对资源利用达到的最合理状态,是一种选择适应的结果,有利于克隆繁殖潜力的发挥。同时为生境适应假说提供了又一例证。     

不同生境条件下羊草种群无性系和有性生殖特征的比较研究

比较研究了东北草原不同生境条件下4个羊草种群无性系和有性生殖数量特征的差异,结果表明,由于各种群所处立地的资源差异(特别是土壤水分),导致各种群在无性系(包括营养枝和根茎芽密度)和有性生殖(生殖枝密度、生殖枝分化率、生殖贡献)等数量特征方面差异显著,尤其在低平原种群与沙丘种群之间最为显著.与沙丘种群相比,低平原种群营养枝平均密度高50%;根茎芽平均密度高30%以上;生殖枝密度平均值高200%以上.     

Sexual and vegetative reproduction in the aboveground part of a dioecious clonal plant, <Emphasis Type="Italic">Dioscorea japonica</Emphasis> (Dioscoreaceae)

In dioecious clonal plants, the reproductive effort required to set seeds will be responsible for the larger investment in sexual reproduction by females. If there will be a trade-off in resource allocation between sexual and clonal reproduction, this differential sexual reproduction will lead to sexual differentiation in the relative amount of clonal reproduction. To test this prediction, we studied differences between the sexes in their phenologies and investments in sexual and vegetative reproduction (clonal reproduction by means of bulbils) with respect to ramet size in a dioecious clonal plant, Dioscorea japonica Thunb. The period of bulbil production overlapped the period during which infructescences developed. Females flowered later, produced heavier inflorescences, and fewer flowers per inflorescence than did males. Regression analysis using the size of the individual plants demonstrated that large females made smaller investments in inflorescences and larger investments in sexual reproduction than did large males. In contrast, females invested fewer resources in vegetative reproduction than did males. However, the total investments in sexual and vegetative reproduction did not differ between the sexes. These results supported our hypothesis on the sexual differentiation in sexual and clonal reproduction.     

Demographic consequences of greater clonal than sexual reproduction in Dicentra canadensis

Clonality is a widespread life history trait in flowering plants that may be essential for population persistence, especially in environments where sexual reproduction is unpredictable. Frequent clonal reproduction, however, could hinder sexual reproduction by spatially aggregating ramets that compete with seedlings and reduce inter‐genet pollination. Nevertheless, the role of clonality in relation to variable sexual reproduction in population dynamics is often overlooked. We combined population matrix models and pollination experiments to compare the demographic contributions of clonal and sexual reproduction in three Dicentra canadensis populations, one in a well‐forested landscape and two in isolated forest remnants. We constructed stage‐based transition matrices from 3 years of census data to evaluate annual population growth rates, λ. We used loop analysis to evaluate the relative contribution of different reproductive pathways to λ. Despite strong temporal and spatial variation in seed set, populations generally showed stable growth rates. Although we detected some pollen limitation of seed set, manipulative pollination treatments did not affect population growth rates. Clonal reproduction contributed significantly more than sexual reproduction to population growth in the forest remnants. Only at the well‐forested site did sexual reproduction contribute as much as clonal reproduction to population growth. Flowering plants were more likely to transition to a smaller size class with reduced reproductive potential in the following year than similarly sized nonflowering plants, suggesting energy trade‐offs between sexual and clonal reproduction at the individual level. Seed production had negligible effects on growth and tuber production of individual plants. Our results demonstrate that clonal reproduction is vital for population persistence in a system where sexual reproduction is unpredictable. The bias toward clonality may be driven by low fitness returns for resource investment in sexual reproduction at the individual level. However, chronic failure in sexual reproduction may exacerbate the imbalance between sexual and clonal reproduction and eventually lead to irreversible loss of sex in the population.     

Effects of intraspecific competition on size variation and reproductive allocation in a clonal plant

Clonal plants grow in diameter rather than height, and therefore competition among genets is likely to be symmetric and to result in smaller variation in size of genets than in non-clonal plants. Moreover, clonal plants can reproduce both sexually and vegetatively. We studied the effects of density on the size of rosettes and of clones, variation in the size of rosettes and of clones, and allocation to sexual and vegetative reproduction in the clonal herb Ranunculus reptans . We grew plants from an artificial population of R. reptans in 32 trays at two densities. After four months, differences in density were still apparent, although clones in the low-density treatment had on average 155% more rosettes and 227% more rooted rosettes than clones in the high-density treatment. The coefficient of variation of these measures of clone size was 15% and 83% higher, respectively, in the low-density treatment. This indicates that intraspecific competition among clones of R. reptans is symmetric and increases the effective population size. Rooted rosettes were larger and varied more in size in the low-density treatment. The relative allocation of the populations to sexual and to vegetative reproduction was 19% and 13% higher, respectively, in the high-density treatment. Moreover, seeds produced in the high-density treatment had a 24% higher mass and a 7% higher germination percentage. This suggests that with increasing density, allocation to sexual reproduction increases more than allocation to vegetative reproduction in R. reptans , which corresponds to the response of some other species with a spreading growth form but not of species with a compact growth form. We conclude that intraspecific competition is an important factor in the life-history evolution of R. reptans because intraspecific competition affects its clonal life-history traits and may affect evolutionary processes such as genetic drift and selection through its effect on the effective population size.