青藏高原东缘大耳叶风毛菊繁殖特征对海拔的响应

研究了青藏高原东缘不同海拔12个居群大耳叶风毛菊(Saussurea macrota Franch.)繁殖分配对海拔的响应。结果表明:(1)随着海拔的升高,大耳叶风毛菊个体大小呈现减小的趋势,但其繁殖分配随海拔升高而增大,这是为了保证顺利完成有性繁殖而采取的生存策略;(2)大耳叶风毛菊部分花部特征发生变异,以响应随海拔升高面临的传粉困难问题,保证花期传粉的最大成功几率;(3)随着海拔的升高,大耳叶风毛菊产生的种子数目减少,种子百粒重增加;花序和管状小花的数量均减少,而它们的重量均增加,即权衡现象在大耳叶风毛菊的这些器官中普遍存在。     

青藏高原东缘常见种长毛风毛菊(Saussurea hieracioides)的繁殖分配

对分布在青藏高原东缘高寒草甸常见种长毛风毛菊的6个海拔梯度(3200～3850m)18个居群的繁殖分配进行了研究,结果表明:长毛风毛菊随海拔的升高个体和营养器官减小,繁殖的投入增加;随着海拔的升高,长毛风毛菊的个体管状小花数目和种子数目减少,管状小花重量和百粒重增大;个体管状小花的数目及重量和个体种子的数目及重量间存在权衡关系;长毛风毛菊的个体大小和繁殖分配存在负相关关系。在资源受限的环境下,长毛风毛菊以减少管状小花数目和种子的数目来增加管状小花重量和百粒重,是保证有性繁殖成功的策略之一。     

青藏高原3种风毛菊属植物的繁殖分配与海拔高度的相关性

对产于青藏高原东缘的风毛菊属植物柳叶菜风毛菊(Saussurea epilobioides)、尖苞风毛菊(S. subulisquama)和钝苞雪莲(S. nigrescens)的30个居群的繁殖分配及其与海拔间的相关性进行了初步研究。通过采样调查法和烘干重量法, 检测了不同海拔3种风毛菊属植物的繁殖分配。结果显示: 1)海拔与植株个体大小呈负相关关系; 2)海拔与繁殖分配呈正相关关系, 将个体大小作为协变量, 去除其影响, 海拔与繁殖分配仍呈正相关关系; 3)个体大小与繁殖分配呈负相关关系, 将海拔作为协变量, 去除其影响, 个体大小和繁殖分配的相关性消失。研究表明, 海拔作为外界因子对繁殖分配有显著的影响, 这种影响不依赖于植物的内部因素(如个体大小的差异)。     

青藏高原东缘不同海拔波缘风毛菊资源分配及花部特征对种子数目和质量的影响

通过采样调查法和烘干称量法检测了青藏高原东缘不同海拔种群波缘风毛菊(Saussurea undulata)花期与果期的形态特征, 研究了其花期与果期的资源分配, 并对花期花部特征对果期种子特征的影响作出分析。结果表明: 1)波缘风毛菊在花期将更多的资源投入繁殖器官, 而在果期增加了对种子的资源投资; 2)波缘风毛菊花期部分花部特征的变异导致其果期产生的种子随海拔升高数量减少、百粒重增加, 因为波缘风毛菊需要在高寒草甸的极端环境下保证每粒种子的存活能力, 因此采取了产生更少但更大种子的生存策略。     

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。     

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。     

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。     

川西风毛菊果期资源分配对海拔的响应

以分布于青藏高原东缘的川西风毛菊为试验材料,研究了其不同海拔高度16个居群的果期资源分配。结果显示:1)随着海拔的升高,个体大小、繁殖器官生物量、营养器官生物量、根系质量、茎叶质量以及每株植物种子总数量均不断减小,但种子百粒重不断增加;2)繁殖分配和根系分配与海拔呈正相关关系,营养分配和茎叶分配与海拔呈负相关关系;3)果期繁殖分配和营养分配、根系分配和茎叶分配以及种子数量与百粒重之间均存在资源分配上的权衡。研究结论:1)海拔作为外界因子对川西风毛菊果期各生物量及资源分配有显著的影响;2)随着海拔的升高,川西风毛菊通过增加繁殖分配,根系分配以及种子百粒重来适应胁迫环境,提高自身的适合度。     

小花风毛菊的性器官在青藏高原的海拔变异

 小花风毛菊(Saussurea parviflora)是菊科风毛菊属的一种多年生草本植物,是该属在青藏高原东缘的一个优势种。该文研究了小花风毛菊11个居群性器官的变异与11个海拔高度(2 100～3 500 m)之间的相关性,以及花丝、花药长和花柱、花柱分枝长与花粉数之间的相关性;探讨了小花风毛菊的性器官是如何随海拔高度发生变异,如何与胁迫环境相适应,实现交配成功,提高繁殖成功率,使其成为该属植物“现代 分布中心"的优势种。结果表明：1)花丝、花药长与海拔高度呈极显著的正相关关系(p<0.01);2)花粉数与海拔高度呈极显著的负相关关系(p<0.01);3)花柱、花柱分枝长及花柱分枝长和雌蕊长的比与海拔高度呈极显著的正相关关系(p<0.01);4)花柱、花柱分枝长和花丝长与花粉数呈显著的负相关关系(p<0.05);5)种子的成熟率与海拔高度呈显著的正相关关系(p<0.05)。说明小花风毛菊种内性器官在特定的生存环境下发生了变异,特别是在花粉数随海拔的升高明显减少以及昆虫的多样性、丰富性和活动性明显降低的情况下,它能够通过花柱分枝长度的变异来提高柱头对传粉昆虫的感受性,能通过种子成熟率的提高来保证高海拔数目显著减少的种子的成功发育,进而在一定程度上导致了该物种繁殖成功率的上升,使它成为青藏高原这一胁迫环境中的优势种。     

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。     

Geographic variation in primary sex allocation per flower within and among 12 species of Pedicularis (Orobanchaceae): Proportional male investment increases with elevation

? Premise of the study: The study of geographic variation in ecologically important traits within and among taxa is a first step toward understanding the environmental factors that contribute to population differentiation and species divergence. This study examines variation in mean sex allocation per flower (androecium mass/gynoecium mass) among 49 wild populations representing 12 Pedicularis species across an elevation gradient on the eastern Tibetan Plateau. ? Methods: We used population means to evaluate sources of variation in per-flower sex allocation within and across species. In particular, we evaluate the relative influence of intrinsic (i.e., plant size, estimated as aboveground stem biomass) vs. extrinsic factors affecting mean sex allocation among populations. ? Key results: Mean sex allocation per flower (the relative investment in male floral organs) is negatively correlated with mean plant size; populations of large plants produce relatively female-biased flowers. This relationship between mean plant size and mean sex allocation is not statistically significant, however, when the effect of elevation is controlled statistically. Among populations within and across species, mean sex allocation increases with elevation. This relationship persists even when the effect of mean plant size is controlled statistically. Factors associated with increasing elevation appear to favor genotypes and/or taxa with male-biased flowers. ? Conclusion: Extrinsic environmental conditions may be more important than intrinsic resource status in determining patterns of geographic variation in mean sex allocation among populations or species of Pedicularis. We cannot conclude whether the effect of elevation on mean sex allocation is the result of environmentally induced plasticity, genetically based adaptation, or species sorting, but it is only partly mediated by mean plant size.     

不同密度猪毛菜形态结构性状及生物量分配策略的异速关系

植物的生长特性随环境条件的变化具有可塑性,而不同的环境因素对植物可塑性的影响也不尽相同。利用异速分析的方法,通过模拟退化草地恢复过程中猪毛菜（Salsola collina）的不同种群密度（16、44、100、400株/m²）,研究其形态结构性状及生物量分配策略的异速关系在种群密度间的差异。结果表明,种群密度增大能抑制猪毛菜的生长,而且对猪毛菜的株高、根长、一级分枝数、二级分枝数、三级分枝数以及总分枝长均产生了极显著的影响,表明种群密度的变化使得植物的高生长和侧向生长发生了显著变化。种群密度的变化也引起了植物生物量的变化,其中植物根、茎、叶间的生物关系是一种表观可塑性,植物的生长策略未发生改变,只是植物个体大小发生改变引起的生物量分配的变化。植物株高、总分枝长、一级分枝数及繁殖分配的变化,是由种群密度变化引起的,植物的适应策略发生了改变,是真正的可塑性。种群密度改变了植物的繁殖分配策略,而未改变植物叶的分配策略,说明当环境发生变化时,植物调整了其繁殖策略以适应环境因素的改变,以保证种群的生存繁衍。     

SEX ALLOCATION IN THE MONOECIOUS HERB BEGONIA SEMIOVATA

Sex-allocation models predict that the evolution of self-fertilization should result in a reduced allocation to male function and pollinator attraction in plants. The evolution of sex allocation may be constrained by both functional and genetic factors, however. We studied sex allocation and genetic variation for floral sex ratio and other reproductive traits in a Costa Rica population of the monoecious, highly selfing annual Begonia semiovata. Data on biomass of floral structures, flower sex ratios, and fruit set in the source population were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and genetic correlations for floral sex ratio and for floral traits related to male and female function were estimated from the greenhouse-grown progeny of field-collected maternal families. The proportion of reproductive biomass invested in male function was low (0.34 at flowering, and 0.07 for total reproductive allocation). Significant among-family variation was detected in the size (mass) of individual male and female flowers, in the proportion of male flowers produced, and in the proportion of total flower mass invested in male flowers. Significant among-family variation was also found in flower number per inflorescence, petal length of male and female flowers, and petal number of female flowers. Except for female petal length, we found no difference in the mean value of these characters between selfed and outcrossed progeny, indicating that, with the possible exception of female petal length, the among-family variation detected was not the result of variation among families in the level of inbreeding. Significant positive phenotypic and broad-sense genetic correlations were detected between the mass of individual male and female flowers, between male and female petal length, and between number of male and number of female flowers per inflorescence. The ratio of stamen-to-pistil mass (0.33) was low compared to published data for autogamous species with hermaphroditic flowers, suggesting that highly efficient selfing mechanisms may evolve in monoecious species. Our results indicate that the study population harbors substantial genetic variation for reproductive characters. The positive genetic correlation between investment in male and female flowers may reflect selection for maximum pollination efficiency, because in this self-pollinating species, each female flower requires a neighboring male flower to provide pollen.     

Visibility vs. biomass in flowers: exploring corolla allocation in Mediterranean entomophilous plants

Background and Aims

While pollinators may in general select for large, morphologically uniform floral phenotypes, drought stress has been proposed as a destabilizing force that may favour small flowers and/or promote floral variation within species.

Methods

The general validity of this concept was checked by surveying a taxonomically diverse array of 38 insect-pollinated Mediterranean species. The interplay between fresh biomass investment, linear size and percentage corolla allocation was studied. Allometric relationships between traits were investigated by reduced major-axis regression, and qualitative correlates of floral variation explored using general linear-model MANOVA.

Key Results

Across species, flowers were perfectly isometrical with regard to corolla allocation (i.e. larger flowers were just scaled-up versions of smaller ones and vice versa). In contrast, linear size and biomass varied allometrically (i.e. there were shape variations, in addition to variations in size). Most floral variables correlated positively and significantly across species, except corolla allocation, which was largely determined by family membership and floral symmetry. On average, species with bilateral flowers allocated more to the corolla than those with radial flowers. Plant life-form was immaterial to all of the studied traits. Flower linear size variation was in general low among conspecifics (coefficients of variation around 10 %), whereas biomass was in general less uniform (e.g. 200–400 mg in Cistus salvifolius). Significant among-population differences were detected for all major quantitative floral traits.

Conclusions

Flower miniaturization can allow an improved use of reproductive resources under prevailingly stressful conditions. The hypothesis that flower size reflects a compromise between pollinator attraction, water requirements and allometric constraints among floral parts is discussed.Key words: Allometry, biomass, corolla, drought, evolution, flower, Mediterranean, sclerophyllous, size, variation, water     

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.     

青藏高原不同生境下湿生扁蕾(Gentianopsis paludosa)个体大小依赖的繁殖分配

植物对资源的投资和分配是生态学中的重要问题,它反映了植物应对环境变化时的生活史策略。选择青藏高原东缘同一海拔下的嵩草草甸(Kobresia sp.meadow)、金露梅灌丛(Potentilla fruticosa shrub meadow)以及草甸-灌丛交错带3种生境类型,并以3种生境下的湿生扁蕾(Gentianopsis paludosa)为对象,研究了其繁殖分配特征。结果发现:(1)在种群水平上,在生境从草甸经交错带到灌丛的变化中,湿生扁蕾个体大小和繁殖分配比例逐渐增加;3个种群湿生扁蕾的总花数目没有显著差异,但草甸生境湿生扁蕾的蕾期花数目显著高于灌丛生境,而果期花数目则显著低于灌丛生境;(2)在个体水平上,湿生扁蕾的繁殖绝对投入与个体大小显著正相关,且各种群植株都存在繁殖所需的个体大小阈值,而繁殖阈值在生境从草甸经交错带到灌丛的过渡中逐渐减小;湿生扁蕾的繁殖相对投入与个体大小负相关,但相关系数随着生境从草甸经交错带到灌丛的过渡中逐渐减小;各种群花数目与湿生扁蕾植株个体大小显著正相关。研究表明,湿生扁蕾的繁殖投资存在大小依赖效应,但生境差异会对其繁殖投资和生活史策略造成显著影响,而这种影响主要是由不同生境下自然条件的不同造成的。同时,资源分配也与湿生扁蕾的遗传特性和延迟自交的繁育系统特征有关。湿生扁蕾这种不同生境下个体大小依赖的繁殖投资差异是湿生扁蕾与其生境长期适应和进化(生境选择)的结果。