5种毛茛科植物个体大小依赖的繁殖分配和性分配

 植物繁殖分配和性分配是生活史理论的核心问题,一直受到生态学家、进化生物学家们的关注。通过对青藏高原东部高寒草甸(3 500 m)及亚高山草甸(2 900 m)毛茛科5种虫媒两性花植物花期的繁殖分配和性分配的研究发现：1）个体越大,繁殖投入越高,繁殖分配越低,与以往研究结果一致;2）性分配是个体大小依赖的,大个体更偏向雌性器官的资源投入,花粉胚珠比与个体大小的关系较复杂,因种而异;3）花期雌雄功能之间存在资源分配上的权衡（Trade-off）,并且种群之间有差异,表明其受环境条件影响。     

蒙古沙冬青花序内性分配的变化、传粉者运动与繁殖成功

性分配理论主要研究繁殖资源在雌雄功能间的最优分配,从雌雄功能的角度考虑其个体适合度.对花序内不同部位花的雌性与雄性资源分配变化的研究,对于我们理解植物采取哪种繁殖对策保障繁殖成功具有重要意义.本文对生长在中国科学院吐鲁番沙漠植物园内的蒙古沙冬青(Ammopiptanthus mongolicus)连续开花花序内不同部位...     

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.     

雌全同株植物三脉紫菀花期偏雄的个体大小依赖的性别分配

经典的虫媒传粉植物个体大小依赖的性别分配模型通常预期:分配给雌性功能的资源比例将随着个体大小的增大而增加;但一些研究表明,花期个体大小依赖的性别分配模式表现出随个体大小增大而偏雄的趋势.我们以植株高度衡量个体大小,从花和花序两个水平上研究了雌花、两性花同株植物三脉紫菀(Aster ageratoides)花期个体大小依赖的性别分配策略.随着植株高度的增大,植株产生的头状花序数量增加,表明三脉紫菀投入到繁殖的资源不是固定不变的,而是随个体大小增大而增加的.在花和花序水平上,繁殖资源在雌雄性别功能之间的分配均表现为随个体大小的增大而更偏雄的模式,即花粉/胚珠比增加,产生花粉的两性花占两性花和雌花总花数的比例升高.这些结果与花期个体越大、性别分配越偏雄的预期一致.花期更偏雄的性别分配可能有助于植物在花期通过输出花粉提高雄性适合度,从而实现个体适合度的最大化.     

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。     

三种同域分布喉毛花的繁殖分配

以青藏高原高寒草甸中三种同域分布的喉毛花为研究对象,通过比较三个种的植株性状和繁殖分配,探讨繁殖分配的种间差异及其与植株个体大小的关系。结果表明:(1)三个种的植株高度、顶花大小和单株花数目、繁殖分配均存在种间差异,这可能与其各自的交配系统和具体的生境以及相应的生活史对策有关;(2)在三种喉毛花中,投入到营养器官和繁殖器官的绝对资源量均呈显著正相关,未检测到植株生长和繁殖间的权衡关系;(3)三个种的个体大小与繁殖器官生物量均呈显著正相关,而与繁殖分配均呈显著负相关,这表明个体越大,繁殖投入越高,而繁殖分配越低,与以往研究结果一致,这可能是由于繁殖分配与个体大小之间存在异速关系。     

放牧对青藏高原东缘高寒草甸群落27种植物地上生物量分配的影响

通过对比研究青藏高原高寒草甸27种植物群落组分种在放牧和长期排除放牧生境中的生物量分配差异,试图揭示长期放牧干扰对植物生活史对策的影响.结果表明:(1)放牧对群落物种个体生物量大小和生物量分配有着显著的影响;(2)总体来看:多数物种(24)放牧生境中的平均个体生物量明显小于禁牧地中的平均个体生物量;而多数物种在放牧后(23种)繁殖分配明显增加;茎分配有增有减(15减小12种增加);叶分配呈减小趋势(20种减小7增加).(3)放牧的影响在不同物种间和功能群间都存在着明显的差异.放牧使毒草茎分配减小叶分配增加,繁殖分配几乎不受影响;豆科和杂草繁殖分配增加,茎分配和叶分配减小,其中豆科两个种的生物量分配变化都不显著;禾草叶分配减小,繁殖分配和茎分配增加; (4)在群落水平上,放牧使繁殖分配和叶分配增加,茎分配减少.     

高山垫状植物团状福禄草开花面积与方位随海拔的变化及其适应性

作为高山生态系统中的奠基种(foundation species), 垫状植物自身种群的繁殖与扩张, 对高山生态系统功能稳定性起着关键作用。但是, 垫状植物如何在极端环境条件下实现资源的有效利用与分配, 达到繁殖最优化, 至今鲜为人知。该研究在滇西北白马雪山沿海拔梯度选择具有不同坡度及坡向的5个团状福禄草(Arenaria polytrichoides)种群, 调查、比较种群内、种群间以及具有不同性系统的植株个体之间的开花面积比、开花方位, 并分析不同生态因子对其开花特性的影响。结果表明: 随着海拔的升高, 团状福禄草个体变小, 其分配到开花的资源比例总体上随海拔上升呈现下降的趋势, 说明团状福禄草的繁殖分配受到由海拔所引起的生态因子的调控。但是, 部分低海拔种群内植物个体的繁殖分配显著低于部分高海拔种群, 说明海拔并非控制植物繁殖分配的唯一因素。此外, 植株开花总面积随植株个体增大而增加, 但开花面积比却随个体增大而变小, 说明植株分配到开花的资源增长速率可能低于植株个体的增长速率。在性别差异方面, 两性植株对开花的资源分配比例要显著高于雌性植株, 但是, 其差异程度受到海拔因素的影响。最后, 在同一种群内, 团状福禄草在冠层表面不同方位上的开花面积比存在显著差异性, 这种差异性在不同种群之间又具有不同的表现形式。     

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。     

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。     

Size‐dependent resource allocation and sex allocation in herbaceous perennial plants

Individuals within a population often differ considerably in size or resource status as a result of environmental variation. In these circumstances natural selection would favour organisms not with a single, genetically determined allocation, but with a genetically determined allocation rule specifying allocation in relation to size or environment. Based on a graphical analysis of a simple evolutionarily stable strategy (ESS) model for herbaceous perennial plants, we aim to determine how cosexual plants within a population should simultaneously adjust their reproductive allocation and sex allocation to their size. We find that if female fitness gain is a linear function of resource investment, then a fixed amount of resources should be allocated to male function, and to post‐breeding survival as well, for individuals above a certain size threshold. The ESS resource allocation to male function, female function, and post‐breeding survival positively correlate if both male and female fitness gains are a saturating function of resource investment. Plants smaller than the size threshold are expected to be either nonreproductive or functionally male only.     

INTRASPECIFIC VARIATION IN SEX ALLOCATION IN A SIMULTANEOUS HERMAPHRODITE: THE EFFECT OF INDIVIDUAL SIZE

Within a population of simultaneous hermaphrodites, individuals may vary in both their current reproductive investment (biomass invested in gonads) and in how they allocate that investment between male and female function. In the chalk bass, Serranus tortugarum, estimates of both reproductive allocation and reproductive success as a male and a female can be made for individuals of different sizes. As individuals increase in size, their investment in gamete production increases, and there is a shift in allocation to a stronger female bias. Spawning frequency as a female in pair spawnings and as a male in both pair spawning and streaking (an alternative mating tactic) does not vary with individual size. As a result, larger individuals should release more sperm or eggs per spawn. Size-assortative pair spawning in this species leads to larger individuals having higher potential returns in total male reproductive success than smaller individuals, which should lead to increases in absolute levels of sperm production in larger individuals when individuals compete for fertilizations through sperm competition. However, smaller individuals contribute a smaller proportion of the sperm released in spawns with multiple spawners and thus are under more intense sperm competition than larger individuals, which should select for increases in male allocation in smaller individuals, all else equal. A local-mate-competition (LMC) model predicts that these factors select for increasing absolute male and female investment with individual size but a relative shift to more female-biased allocation as individual size increases. These predictions are supported by gonadal data. The predictions of average male allocation from the quantitative LMC model were 21.6% and 25.7%, whereas the collections averaged 21.3%. This close agreement of both the mean male allocation and its relative shift with individual size between model and data support the hypothesis that size-specific shifts in sex allocation in this species represent an adaptive response to patterns of mating success and sperm competition.     

Intraspecific variation in sex allocation in hermaphroditic Plantago coronopus (L.)

Models for sex allocation assume that increased expenditure of resources on male function decreases the resources available for female function. Under some circumstances, a negative genetic correlation between investment in stamens and investment in ovules or seeds is expected. Moreover, if fitness returns for investment in male and female function are different with respect to size, sex allocation theory predicts size‐specific gender changes. We studied sex allocation and genetic variation for investment in stamens, ovules and seeds at both the flower and the plant level in a Dutch population of the wind‐pollinated and predominantly outcrossing Plantago coronopus. Data on biomass of floral structures, stamens, ovules, seedset and seedweight were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and (genetic) correlations were estimated from the greenhouse‐grown progeny of maternal families, raised at two nutrient levels. The proportion of reproductive biomass invested in male function was high at flowering (0.86 at both nutrient levels) and much lower at fruiting (0.30 and 0.40 for the high and low nutrient treatment, respectively). Androecium and gynoecium mass exhibited moderately high levels of genetic variance, with broad‐sense heritabilities varying from 0.35 to 0.56. For seedweight no genetic variation was detected. Significant among‐family variation was also detected for the proportion of resources invested in male function at flowering, but not at fruiting. Phenotypic and broad‐sense genetic correlations between androecium and gynoecium mass were positive. Even after adjusting for plant size, as a measure of resource acquisition, maternal families that invested more biomass in the androecium also invested more in the gynoecium. This is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the overall lack of a negative correlation between male and female function. Larger plants had a more female‐biased allocation pattern, brought about by an increase in seedset and seedweight, whereas stamen biomass did not differ between small and large plants. These results are discussed in relation to size‐dependent sex allocation theory (SDS). Our results indicate that the studied population harboured substantial genetic variation for reproductive characters.     

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.     

Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae)

Abstract Patterns of reproductive and vegetative biomass allocation were compared in male and female plants of the alpine herb Aciphylla simplicifolia. Male and female plants had similar vegetative biomass but differed in the pattern of resource allocation. Inflorescences of males and females were similar in weight at the time of flowering, but differed in biomass allocation to some structures within the inflorescences, particularly those associated with ovule production and pollinator attraction (number and size of flowers). At the time of fruit production, female inflorescences were 2.6 times heavier than at flowering with developing fruit six times heavier than flowers. In addition to the increase in biomass allocated to structures associated with the provisioning and dissemination of seed, support structures (main and side stalks) were also heavier. As a result of this additional investment of resources at the time of fruit production, the reproductive effort (RE) of female plants was much higher than that of males: 37% of above ground biomass compared with 21% for males. Differences in RE did not change with plant size; however, allocation to reproduction appeared to be a constant proportion of biomass over nearly all plant sizes sampled. These results show that sex‐specific resource allocation can be a complex of temporal and morphological patterns.     

Sex allocation in a simultaneously hermaphroditic marine shrimp

Two fundamental questions dealing with simultaneous hermaphrodites are how resources are optimally allocated to the male and female function and what conditions determine shifts in optimal sex allocation with age or size. In this study, I explored multiple factors that theoretically affect fitness gain curves (that depict the relationship between sex-specific investment and fitness gains) to predict and test the overall and size-dependent sex allocation in a simultaneously hermaphroditic brooding shrimp with an early male phase. In Lysmata wurdemanni, sperm competition is absent as hermaphrodites reproducing in the female role invariably mated only once with a single other shrimp. Shrimps acting as females preferred small over large shrimps as male mating partners, male mating ability was greater for small compared to large hermaphrodites, and adolescent males were predominant in the population during the breeding season. In addition, brooding constraints were not severe and varied linearly with body size whereas the ability to acquire resources increased markedly with body size. Using sex allocation theory as a framework, the findings above permitted to infer the shape of the male and female fitness gain curves for the hermaphrodites. The absence of sperm competition and the almost unconstrained brooding capacity imply that both curves saturate, however the male curve levels off much more quickly than the female curve with increasing level of investment. In turn, the predominance of adolescent males in the population implies that the absolute gain of the female curve is greater than that of the male curve. Last, the size-dependent female preference and male mating ability of hermaphrodites determines that the absolute gain of the male curve is greater for small than for large hermaphrodites. Taking into consideration the inferred shape of the fitness gain curves, two predictions with respect to the optimal sex allocation were formulated. First, overall sex allocation should be female biased; it permits hermaphrodites to profit from the female function that provides a greater fitness return than the male function. Second, sex allocation should be size-dependent with smaller hermaphrodites allocating more than proportionally resources to male reproduction than larger ones. This size-dependent sex allocation permits hermaphrodites to profit from male mating opportunities that are the greatest at small body sizes. Size-dependent sex allocation is also expected because the male fitness gain curve decelerates more quickly than the female gain curve and experiments indicated that resources are greater for large than small hermaphrodites. These two predictions were tested when determining the sex allocation of hermaphrodites by dissecting their gonad and quantifying ovaries versus testes mass. Supporting the predictions above, hermaphrodites allocated, on average, 118 times more to the female than to the male gonad and the proportion of resources devoted to male function was higher in small than in large hermaphrodites. A trade-off between male and female allocation is assumed by theory but no negative correlation between male and female reproductive investment was observed. In L. wurdemanni, the relationship between sex-specific investment and fitness changes during ontogeny in a way that is consistent with an adjustment of sex allocation to improve size-specific reproductive success.     

Size-dependent sex allocation within flowers of the annual herb Clarkia unguiculata (Onagraceae): ontogenetic and among-plant variation

The relative allocation of resources to male and female functions may vary among flowers within and among individual plants for many reasons. Several theoretical models of sex allocation in plants predict a positive correlation between the resource status of a flower or individual and the proportion of reproductive resources allocated to female function. These models assume that, independent of resource status, a negative correlation exists between male and female investment. Focusing on the allocation of resources within flowers, we tested these theoretical predictions and this assumption using the annual Clarkia unguiculata (Onagraceae). We also sought preliminary evidence for a genetic component to these relationships. From 116 greenhouse-cultivated plants representing 30 field-collected maternal families, multiple flowers and fruits per plant were sampled for gamete production, pollen?:?ovule ratio, seed number, ovule abortion, seed biomass/fruit, mean individual seed mass, and petal area. If sex allocation changes as predicted, then (1) assuming that flowers produced early have access to more resources than those produced later, basal flowers should exhibit a higher absolute and proportional investment in female function than distal flowers and (2) plants of high resource status (large plants) should produce flowers with a higher proportional investment in female function than those of low resource status. Within plants, variation in floral traits conformed to the first prediction. Among plants and families, no significant effects of plant size (dry stem biomass) on intrafloral proportional sex allocation were observed. We detected no evidence for a negative genetic correlation between male and female investment per flower, even when controlling for plant size.     

Sex allocation and reproductive success in the andromonoecious perennial Solanum carolinense (Solanaceae). I. Female

Relative allocation of resources to growth vs. reproduction has long been known to be an important determinant of reproductive success. The importance of variation in allocation to different structures within reproductive allocation is somewhat less clear. This study was designed to elucidate the importance of allocation to vegetative vs. reproductive functions, and allocation within reproductive functions (sex allocation), to realized female success in an andromonoecious plant, Solanum carolinense. Allocation measurements were taken on plants in experimental arrays exposed to natural pollination conditions. These measurements included total flower number, the proportion of flowers that were male, flower size, and vegetative size. Flower number explained the majority of the variation among individuals in their success-that is, there was strong selection for increased flower production. There was also selection to decrease the proportion of flowers that were male, but neither flower size nor vegetative size (a measure of overall resource availability) were direct determinants of female success. After Bonferroni corrections for multiple comparisons, most phenotypic correlations among the traits measured were nonsignificant. Thus, in this andromonoecious species there is not a strong relationship between resource availability (vegetative size) and female success, and female success is instead determined by the relative production of the two different flower types.