中华山蓼不同海拔居群的繁殖分配研究

研究了具有克隆繁殖和雌雄异株繁育特性的中华山蓼Oxyria sinensis Hemsl.在5个不同海拔居群的繁殖分配.结果表明:(1)雄株高度和茎叶总生物量仪在海拔2780 m的居群显著高于雌株,在另外4个居群与雌株高度和茎叶总生物量均不存在显著差异:雄株地上部分总生物量在海拔较高的3个届群显著高于雌株的地上部分总生物量;雄株的花生物量和繁殖分配在海拔1978 m的居群与雌株没有显著差异,但在另外4个居群均显著高于雌株,表明在海拔较高的地区,中华山蓼增加了对雄性植株的资源配置,可能是对海拔较高地区不可预见性降雨和降低昆虫访花频率的适应.(2)雌雄株高度、地上部分总生物量和茎叶生物量以及雌株的花生物量和繁殖分配随海拔升高表现出降低的趋势,但雄花的生物量和繁殖分配随海拔升高显著增加,进一步证明中华山蓼在海拔较高地区的居群增加了对雄性资源的投资.(3)雄雌株繁殖分配受个体大小制约(个体大小依赖性),但并不支持"植物开始繁殖必须达到一定的大小(阈值)"的观点.这可能因为中华山蓼具有较强的无性繁殖能力,而同一植株上不同分株间能通过地下根状茎达到资源共享,因此中华山蓼分株的开花繁殖不需要达到一定的大小.     

Response of resource allocation of Saussurea leontodontoides during its fruiting stage to the elevation

植物资源分配是目前植物生态学研究的热点问题, 主要集中在性分配和繁殖分配两个方面。该研究以分布在青藏高原的狮牙草状风毛菊(Saussurea leontodontoides)作为研究材料, 研究了6个海拔高度上果期植株的繁殖特征及资源分配的差异, 并用异速模型分析了繁殖性状及资源分配与个体大小的关系。结果显示: 1)狮牙草状风毛菊的个体大小、繁殖器官生物量、营养器官生物量、种子数和营养分配均与海拔存在极显著负相关关系, 百粒质量和繁殖分配与海拔存在极显著正相关关系。2)在不同海拔高度下, 百粒质量、种子数、繁殖器官生物量及营养分配与植株个体大小呈正相关关系; 繁殖分配与植株个体大小呈负相关关系; 而营养器官生物量与植株个体大小呈极显著正相关关系。这表明海拔和个体大小对狮牙草状风毛菊的繁殖对策有不同程度的影响, 狮牙草状风毛菊通过增加繁殖部分的生物量和百粒质量来适应高海拔的胁迫环境。     

狮牙草状风毛菊果期资源分配对海拔的响应

植物资源分配是目前植物生态学研究的热点问题, 主要集中在性分配和繁殖分配两个方面。该研究以分布在青藏高原的狮牙草状风毛菊(Saussurea leontodontoides)作为研究材料, 研究了6个海拔高度上果期植株的繁殖特征及资源分配的差异, 并用异速模型分析了繁殖性状及资源分配与个体大小的关系。结果显示: 1)狮牙草状风毛菊的个体大小、繁殖器官生物量、营养器官生物量、种子数和营养分配均与海拔存在极显著负相关关系, 百粒质量和繁殖分配与海拔存在极显著正相关关系。2)在不同海拔高度下, 百粒质量、种子数、繁殖器官生物量及营养分配与植株个体大小呈正相关关系; 繁殖分配与植株个体大小呈负相关关系; 而营养器官生物量与植株个体大小呈极显著正相关关系。这表明海拔和个体大小对狮牙草状风毛菊的繁殖对策有不同程度的影响, 狮牙草状风毛菊通过增加繁殖部分的生物量和百粒质量来适应高海拔的胁迫环境。     

展毛翠雀花期繁殖特征和生物量分配对放牧的响应

生物量分配影响植物生长和繁殖,是植物生活史研究的重要内容。为了了解植物生活史性状对放牧的响应,该研究以青藏高原高寒草甸毒杂草展毛翠雀为对象,分析了放牧干扰对展毛翠雀的花期繁殖分配和性分配的影响。结果表明:放牧显著降低了展毛翠雀的总生物量、个体大小和繁殖投入; 放牧未改变展毛翠雀的营养部分与繁殖部分的等速生长关系,但显著增加了繁殖部分的生物量分配和总花数; 展毛翠雀的个体大小与总花数呈显著的正相关关系,但与性分配呈显著的负相关关系; 展毛翠雀的总花数与单花大小、单花的花瓣比例均表现出负相关关系,表明总花数与单花大小之间、总花数与单花的花瓣比例之间均存在权衡。因此,在放牧条件下,展毛翠雀的繁殖分配和性分配均表现出显著的可塑性。     

川西风毛菊花期资源分配随海拔的变化

以分布于青藏高原东缘的川西风毛菊(Saussurea dzeure)为试验材料, 研究了其位于不同海拔高度的16个种群的花期资源分配。结果显示: 1)花期植物个体大小、头状花序数量、繁殖器官及营养器官生物量、花瓣质量、雌蕊及雄蕊群质量均与海拔呈负相关关系, 每个头状花序质量与海拔呈正相关关系; 2)繁殖分配和雄性分配与海拔呈正相关关系, 营养分配和雌性分配与海拔呈负相关关系; 3)花期头状花序的数量和大小、繁殖分配和营养分配以及雄性分配和雌性分配之间均存在资源分配上的权衡。由此推论: 1)海拔作为外界因子对川西风毛菊花期各生物量及资源分配有显著的影响; 2)在资源有限的情况下, 川西风毛菊权衡对各结构的资源投入, 通过增加繁殖分配和雄性分配来适应胁迫环境, 提高繁殖的成功率。     

青藏高原东缘禾叶风毛菊的繁殖分配与海拔的相关性

通过采样调查法和烘干称重法,对分布在青藏高原东缘不同海拔高度的禾叶风毛菊的繁殖分配的特征进行研究。结果表明：（1）随着海拔的升高,禾叶风毛菊的个体大小、营养器官生物量、繁殖器官生物量、个体管状小花数目、雄蕊质量均与海拔呈负相关关系（P<0.01）;繁殖分配、管状小花生物量、雌蕊质量均与海拔呈正相关关系（P<0.01）;（2）繁殖分配是依赖个体大小的,个体越大,繁殖分配越小（P<0.01）;（3）禾叶风毛菊个体管状小花的数目及重量（P<0.05）、雌雄蕊重量（P<0.05）之间存在权衡关系。由此推论：（1）海拔作为外界因子对禾叶风毛菊花期各生物量及繁殖分配有显著的影响,但海拔并不是影响禾叶风毛菊繁殖分配唯一生态因子,植株个体大小也与其繁殖分配策略密切相关;（2）禾叶风毛菊的垂直分布的特征很有可能就是海拔通过影响植株个体大小变化来完成的。     

高寒草地狼毒种群繁殖分配对海拔的响应

繁殖与物种的生存和发展有着极为密切的关系,是植物生活史研究的核心问题,海拔制约下的环境因子对植物繁殖分配产生重要影响.2010年7月采用样方调查法,研究了祁连山北坡高寒草地2700～3000 m范围内4个海拔梯度下狼毒(Stellera chamaejasme)种群繁殖分配特征.结果表明:随着海拔的升高,狼毒地上生物量呈降低趋势,总花干重、株高呈先减小后增大的趋势,繁殖分配呈增大趋势;在海拔梯度下,狼毒种群繁殖分配与地上生物量均呈极显著负相关(海拔2800 m除外),与株高呈不显著负相关(海拔3000 m除外).狼毒种群繁殖分配存在大小依赖性且生物量的依赖性远大于株高,说明随海拔升高,狼毒种群通过减小个体大小来降低资源消耗,把获取的有限资源分配到繁殖部分,从而达到繁殖成功.     

岩白菜（虎耳草科）不同海拔居群的繁殖分配

资源分配策略是植物生活史研究的重要内容之一,植物用于繁殖的相对资源比例（即繁殖分配）与植株的生活史特征、个体大小及植株的生境密切相关。本文研究了藏东南色季拉山一个阴坡上海拔4200m～4640m范围内6个不同居群的虎耳草科多年生草本植物岩白菜（Bergenia purpurascens）的繁殖分配特征,结果发现：（1）繁殖器官生物量、营养器官生物量、地上部分总生物量、花数目、花序轴长度均随海拔的升高而显著降低,而叶数目随海拔变化不大,繁殖分配值则先降低后升高,转折点在林线过渡带（海拔4400m）处;（2）各居群（海拔4300m居群除外）营养器官生物量与繁殖器官生物量均显著正相关,而营养器官生物量与繁殖分配则负相关,但各居群的显著性不同;（3）各居群繁殖器官生物量与植株个体大小（营养器官生物量）呈不同程度的异速增长,而繁殖分配则与植株个体大小负相关;（4）各居群植株都存在一个繁殖所需的个体大小阈值,而且这一阈值在林线以下区域随海拔的升高而显著增大,在林线以上区域变化不显著。研究结果表明,海拔并不是影响岩白菜繁殖分配策略的唯一生态因子,不同居群的生境状况和植株个体大小都与其资源分配策略密切相关,高山地区林线的存在对植物资源的权衡方式会产生巨大影响。     

唐古特雪莲花部特征及生殖分配的海拔差异

为探究高山植物生殖分配策略以及分析唐古特雪莲花部特征对海拔梯度的响应机制,该研究利用采样调查法和烘干称重法,对分布在青藏高原东缘不同海拔14个居群的唐古特雪莲花部特征和生殖分配进行了研究。结果表明:(1)繁殖分配随个体大小(地上生物量和株高)的增大呈线性递减趋势。(2)花期植株地上生物量、株高、管状小花数目、繁殖器官及营养器官生物量均随海拔升高呈线性递减趋势,管状小花生物量随海拔升高呈线性递增趋势。(3)花期管状小花数量及大小、繁殖器官生物量与营养器官生物量、雄蕊重量和雌蕊重量以及花粉数目与花丝长度之间均存在权衡现象。由此推论:(1)海拔作为外界因子对唐古特雪莲花部特征具有显著的影响,个体大小对其繁殖分配也存在潜在调控作用。(2)在不同海拔梯度上,唐古特雪莲有效地整合了有限的资源,其适应性特征之一就是通过减小个体大小来削弱营养生长以达到促进生殖生长的目的。     

紫翅猪毛菜、钠猪毛菜不同个体大小繁殖分配差异及随海拔的变化

以新疆准噶尔盆地藜科猪毛菜属植物紫翅猪毛菜(Salsola affinis C.A.Mey)、钠猪毛菜(Salsola nitraria Pall)为研究对象,用繁殖分配比例的方法对比分析了两种猪毛菜不同海拔同一种群内不同个体大小繁殖分配的特点,并用异速生长模型分析了不同海拔繁殖生物量与营养生物量之间分配与个体大小的依赖关系。结果发现:1)不同海拔繁殖生物量(R)与营养生物量(V)呈不同程度的异速生长。紫翅猪毛菜随海拔的升高R-V的异速生长斜率显著升高,截距随海拔的升高没有显著增加;而钠猪毛菜的斜率随海拔升高显著降低,截距则显著升高。2)紫翅猪毛菜在较低海拔个体大小与繁殖分配呈负相关,在较高海拔呈正相关;钠猪毛菜在较低海拔个体大小与繁殖分配呈正相关,在较高海拔呈负相关;两种猪毛菜繁殖分配的适应对策相反。3)将同一种群个体大小分成大、中、小3种类型,多重比较发现紫翅猪毛菜在较低海拔,中小个体的繁殖分配显著高于大个体的繁殖分配;在较高海拔,大个体的繁殖分配显著高于中小个体的繁殖分配。钠猪毛菜在较低海拔,大个体的繁殖分配显著高于中、小个体的繁殖分配;在较高海拔,小个体的繁殖分配显著高于大、中个体的繁殖分配。综合分析认为:两个物种随海拔变化产生不同的繁殖分配策略,除遗传效应外,环境和个体大小对钠猪毛菜繁殖分配的变化均产生重要影响,而紫翅猪毛菜繁殖分配的变化主要由海拔差异导致。由于微生境对同一种群的个体大小产生影响,进而产生不同的繁殖分配模式,所以在干旱区更应重视个体大小对繁殖分配的影响。     

Sex allocation and mating systems in pigeonpea,Cajanus cajan (Fabaceae)

Sex allocation theory predicts that: (1) resources allocated to androecium should decrease with an increase in selfing, (2) a decrease in androecium biomass should be accompanied by an increase in the biomass of pistils, and (3) a decrease in androecium biomass should be coupled with a decrease in flower size, specifically corolla biomass. Another predicted change in reproductive traits associated with variation in selfing concerns seed to ovule ratios, but does not directly stem from sex allocation theory. It has been postulated that seed to ovule ratios should be positively correlated with the amount of selfing. These predictions were tested for six accessions of pigeonpea,Cajanus cajan L., that differed in selfing rates. The results were remarkably in accordance with the predictions. We conclude that sex allocation theory provides a powerful tool to understand the evolution of many reproductive traits in plants.     

Altitudinal Variations in Reproductive Allocation of Bergenia purpurascens (Saxifragaceae)

The strategy of resource allocation between vegetative and reproductive functions, quantitative relationship between size and reproductive output are central aspects of plant life history. To test the tactics of resource allocation and its altitudinal trend, we examined the reproductive allocation (RA) of Bergenia purpurascens (Saxifragaceae), in six populations along a shady slope in Sejila Mountain of southeast Tibet, at an altitude gradient from 4 200 m to 4 640 m. Our results showed that (1) with increasing altitude, vegetative biomass, reproductive biomass, total aboveground biomass, flower number per plant and length of flower stalk decreased significantly, but the number of leaves did not change greatly. However, the change of RA did not show a monotonic trend when altitude increased, shifting from significantly decreasing below the tree line to slightly increasing above it; (2) vegetative biomass was positively correlated with reproductive biomass, but negatively correlated with RA in all populations, but the level of significance was different among the populations; (3) RA decreased with individual size in all populations, whereas the relationship between absolute resource allocated to reproduction and individual size was allometric; (4) reproductive allometry and a size threshold for reproduction did exist in this alpine perennial, but the obvious altitudinal trend was only found along the populations below the tree line, not above it. We then concluded the altitude could not fully explain the change of resource allocation strategy of this alpine perennial, and different effects of size and habitat on RA may result from various environmental constraints along the altitudinal gradient or genetic background. Therefore, each individual within a population will follow its own developmental trajectory shaped by its genotype and the habitats. The most innovative finding was plant adaptation and resource trade off might be sharply altered at the tree line, which is a sensitive area in alpine mountains. Further investigations are needed to better understand the relationship between the reproductive allocation and changing environmental conditions.     

Variations of flower size and reproductive traits in self-incompatible <Emphasis Type="Italic">Trollius ranunculoides</Emphasis> (Ranunculaceae) among local habitats at Alpine Meadow

For plants that rely on animals for pollination, the ability to attract the animals to their flowers can be a crucial component of fitness. A large number of studies have documented pollinators to be important selective agents driving the evolution of flower size and correlated traits on a large scale. In this paper, we studied variations of reproductive traits in self-incompatible Trollius ranunculoides (Ranunculaceae) among local habitats at Alpine Meadow. The results showed significant variations of floral size, seed mass per fruit and sex allocation (male/female mass ratio) between different habitats, where floral size and seed mass was not explained fully by variation of plant size among habitats. It suggested that other factors unrelated to plant size might also influence floral variation. However, in our manipulated experiment, it showed no effects of manipulated floral size not only on visit rate of effective pollinators (bees and flies) but also on female success (seed set, seed mass per fruit), irrespective of flower density. Consequently, we could not conclude that the variation of floral size in T. ranunculoides was due to phenotypic plasticity, or natural selection. But if selection occurred, it should not be mediated by pollinators. It was likely that variation of sex allocation between habitats lead to changes of flower or corolla size, because plant invested much less to male function (female-biased sex allocation and larger single seed mass) in shade habitat (bottom of bush) than other exposed habitats, to gain higher fitness. In addition, high-floral density in T. ranunculoides had a negative effect on service of main pollinator (bees) and female success. This situation would influence the strength of selection on floral size.     

Selecting for floral character associations in wild radish,Raphanus sativus L.

ESS models of reproductive allocation have been used extensively to explain patterns of floral diversity in angiosperms. These theoretical explorations assume that proportional allocation to pollen, ovules, and seeds, as well as to secondary features such as showy petals and nectar rewards, can evolve independently within the limits set by total resource availability. In populations of California wild radish, we have shown previously that petal size, a strong determinant of visitation by honey bee pollinators, is positively correlated with both pollen and nectar production, but not with ovule or seed number per flower. These phenotypic associations may reflect selection, environmental correlation, and/or genetic constraint. By exerting selection on the petal size : pollen number ratio over two generations, we eliminated the positive correlation between petal size and pollen production, with both characters showing significant change after a single selection episode. Once these two floral traits became uncoupled, nectar sugar production was significantly correlated only with petal size. Our results suggest that natural selection could readily alter reproductive allocation in these flowers, and that the phenotypic correlations observed in nature may be maintained by selection for effective reproductive phenotypes.     

Sex allocation of females and hermaphrodites in the gynodioecious Geranium sylvaticum

Seed production and patterns of sex allocation were studied in female and hermaphroditic plants in two gynodioecious populations of Geranium sylvaticum (Geraniaceae). Females produced more flower buds and seeds than hermaphrodites in one of the two study populations. The other female traits measured (pistil biomass, seed number per fruit, individual seed mass) did not differ between the gender morphs. The relative seed fitness of hermaphrodites differed between the study populations, with hermaphrodites gaining less of their fitness through female function in the population with a high frequency of females. However, the amount and size of pollen produced by hermaphrodites did not differ between populations. The number of flower buds was positively correlated with seed production in females, whereas in hermaphrodites a positive correlation between number of buds and seed production was found in only one of the two study populations. These results suggest that fitness gain through female function is labile in hermaphrodites of this species, and is probably affected by environmental factors such as the sex ratio of the population.     

羌塘雪兔子繁殖特征及资源分配的海拔差异

植物种群的生殖特性和资源配置策略是植物生活史研究的重要内容之一,对于理解和预测植被结构和全球变化过程中的适应策略至关重要。该文以青藏高原特有风毛菊属植物羌塘雪兔子(Saussurea wellbyi)为试验材料,分析6个海拔梯度18个居群的花期繁殖特征的变化,并用异速生长模型分析资源投入及资源分配的个体大小依赖关系。结果显示:(1)个体大小、繁殖器官生物量、营养器官生物量、管状小花数目、雄蕊重量、花药长度及花粉数与海拔高度呈负相关(P<0.05),管状小花重量、雌蕊重量、花丝长度及花柱长度与海拔高度呈正相关(P<0.01);(2)营养分配与海拔高度呈负相关(P<0.01),繁殖分配与海拔高度呈正相关(P<0.01);(3)管状小花的数量与重量(P<0.05)、雌蕊重量与雄蕊重量(P<0.01)、花丝长度与花药长度(P<0.01)、花丝长度与花粉数(P<0.01)之间均存在权衡关系;(4)资源投入及资源分配在不同海拔间与个体大小都呈显著正相关(P<0.05),且截距在海拔间差异极显著(P<0.01)。综合分析认为:羌塘雪兔子在不同海拔下的繁殖特征以及大小依赖的资源投入和资源分配模式是羌塘雪兔子与其所处的高山环境长期适应和进化的结果。     

海拔对全缘叶绿绒蒿植株性状和花特征的表型选择分析

为了研究海拔差异对植株性状、花特征表型选择的影响,以青藏高原高寒草甸的全缘叶绿绒蒿（Meconopsis integrifolia）为研究材料,于盛花期内,测定不同海拔（4 452、4 081和3 681 m）种群中个体植株性状、花特征、单果结实数并进行统计分析,采用线性回归模型估计不同海拔种群间植株性状、花特征所受的表型选择（选择差与选择梯度）。结果表明：（1）随着海拔升高,全缘叶绿绒蒿植株性状、花特征及单果结实数显著降低,海拔越高的种群中株高越矮、叶面积越小、花数越少、花越小、单果结实数越低。（2）不同海拔种群中各性状的表型选择存在差异,较低海拔（3 681 m）种群中花数、花大小具有显著的选择差和选择梯度,表现为花越多、花越大的个体雌性适合度越高;海拔较高（4 081 m）的种群中株高、叶面积及花数更容易受到选择,表现为植株越高、叶面积越大、花越多的个体雌性适合度越高;海拔最高（4 452 m）的种群中叶面积与花数的选择梯度接近显著。（3）植物性状分化伴随着海拔的变化而呈现出差异,较低海拔种群中花特征容易受到选择,而较高海拔种群中可能由于传粉者稀少、资源限制等因素使得株高、叶面积更容易受到选择。     

入侵植物小花山桃草种群构件生物量结构及种子萌发特征

通过野外设置样方调查和室内萌发试验,研究小花山桃草种群各构件生物量的结构特征和它们之间的关系模型、繁殖分配以及种子萌发特点。结果表明:(1)小花山桃草根、茎、叶、花(果)序生物量与植株高度之间以及各构件生物量之间均呈正相关关系,可用幂函数模型或线性函数模型较好地表达;(2)各构件生物量在个体生物量中所占的比率表现为茎>花序>叶>根;(3)小花山桃草的繁殖投入和繁殖分配都随植株个体的增大而增加;(4)小花山桃草个体大小和繁殖投入之间为线性关系,而个体大小和繁殖分配之间为幂函数关系;(5)小花山桃草存在一个较小的繁殖阈值(0.6043g);(6)小花山桃草种子在有光照(12h)和黑暗条件下发芽率均可达到85%以上;未经贮藏的种子不萌发,低温沙藏(1～2℃)和室温干藏(14～32℃)一个半月种子萌发率分别可达92.5%和79%;低温沙藏时种子即可发芽,且发芽率可达61%。在研究地区,小花山桃草几乎整个生长季都可萌发,甚至初冬还有幼苗产生。小花山桃草构件生物量结构和繁殖分配特征、种子萌发特点等都有助于其入侵能力的提高,是其成功入侵我国的重要原因。