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。     

多年生龙胆属植物个体大小与花期资源分配研究

于各物种花中前期对青藏高原东部高寒草甸6种多年生龙胆属植物花期的繁殖分配和性分配进行分析,结果表明:(1)多年生龙胆属植物的植株个体越大,繁殖投入越高,繁殖分配越低;(2)随着植物个体的增大,对雌性、雄性和吸引结构的投入都在增加,这可保证资源的充分利用,不会因为单一部分的增加而造成资源的浪费;(3)6种龙胆属植物中,有4种其性分配结果与性别分配(SDS)的理论预测一致,即大个体更偏向雌性器官的资源投入,但麻花艽(Gentiana atraminea)和达乌里秦艽(Gentiana dahurica)的性分配与个体大小则没有表现出负相关,可能与其本身具有的雌雄异熟———雄性先熟特点有关;(4)资源在雌雄功能间的分配没有表现出权衡关系,可能是由于植物必须在许多不同生活史性状之间进行资源分配,而不是两两之间非此即彼.     

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。     

不同生境自交植物黄花大苞姜生殖分配的研究

为了探讨自交植物黄花大苞姜(Caulokaempferia coenobialis)对石壁附生这一特殊生境的生态适应,对其不同物候期和不同生境的生殖分配进行了对比研究。结果表明,在生殖生长过程中,黄花大苞姜种群用于营养生长的生物量分配占有绝对优势,而用于生殖的生物量分配比例较小(<13%)。在黄花大苞姜各构件的生物量分配中,根茎和叶的比重较大(24.22%~43.25%)。在光线较弱生境中的种群,为了提高资源获取能力,黄花大苞姜分配到叶的比重明显高于光线较强的种群,而分配给根茎的比例却明显低于光线较强的种群。随着物候期的推移,黄花大苞姜生殖分配的比例不断增加,到果期达到最大值。不同种群间和年度间黄花大苞姜分配给生殖构件的比例没有显著差异,推测其生殖分配可能受遗传因素控制。个体大小与根茎生物量呈极显著线性函数同速生长,而与生殖分配在云天海种群没有表现出相关性,在上坪和天堂顶种群表现为同速生长关系,但决定系数小于40%。因此,黄花大苞姜能有效调节其在不同生境的生物量分配以适应石壁附生的特殊生境,在光线较弱的种群提高叶的生物量分配并降低根茎的生物量分配以提高资源的获取能力。整体上投资到营养构件的生物量占比高达87%以上,生殖构件在居群间和年度间均保持稳定。这种繁殖策略,一方面较高的营养构件投资可以获得更多的资源,另一方面稳定的生殖投资可以保证种群的延续,各构件相互协调以更好适应石壁这一资源匮乏的生境。     

菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系

三叶鬼针草(Bidens pilosa)是一种危害严重的菊科入侵植物.通过实验观察和人工控制套袋等方法,对其花序开花动态、花粉胚珠比(P/O)、自交亲和性、花粉活力、访花昆虫和种子(瘦果)的萌发率等与繁殖相关的特征进行了研究,探讨了这些繁殖特征与入侵性的关系.三叶鬼针草在10-11月开花,单个花序的花期约为5-6天.每小花的花柱基部有圆筒状的蜜腺环绕.单个花序内可自交亲和,自交结实率和花粉活力均较高,其P/O值为1754.12士29.87.主要访花者为灰蝶科(Lycaenidae)、粉蝶科(Pjerjdae)和茧蜂科(Braconidae)昆虫.三叶鬼针草所具有的灵活交配机制是其入侵成功的重要因素.此外,三叶鬼针草结实量大、种子产生迅速且适于传播,以及种子萌发范围广和短期快速萌发等特性也增强了其入侵性.     

濒危植物焕镛木的兼性无融合生殖

2001—2002年连续两年在广西环江木论和广西罗城大黄泥的2个焕镛木(Woonyoungia septentrionalis (Dandy) Law)自然种群中,对单性异株的濒危植物焕镛木进行繁育系统测定,对即将开花的雌花花蕾分别进行套袋、套网、人工授粉处理,并用自然授粉雌花作对照,其座果率和结实率统计结果表明：自然授粉、人工授粉、套袋和套网处理的花均能结实,但它们的座果率和结实率存在较大的差异。在两个种群中,人工授粉和自然授粉的总结实率(PERS)均比套袋和套网处理的高,其中人工授粉的最高,套网处理的最低。由此可见,焕镛木既能通过有性生殖方式结实,又能通过无融合生殖方式结实,而且这两种生殖方式获得的种子均能萌发成幼苗,由此断定,焕镛木的繁育系统为兼性无融合生殖。这是首次报道木兰科植物存在无融合生殖现象。     

金沙江干热河谷车桑子生殖枝生物量分配的性别差异

以车桑子生殖枝为材料,通过野外调查、取样以及各器官生物量的测定,研究金沙江干热河谷地区车桑子不同性别植株生殖枝生物量分配特征。结果表明：（1）金沙江干热河谷地区车桑子种群雄∶雌性别比为0.11,极显著的偏离1∶1（P<0.001）,种群偏雌性。（2）车桑子生殖枝的形态特征和生物量分配特征均具有显著的性别差异;雌性植株生殖枝花朵生物量和总生物量显著高于雄性和两性植株（P <0.05）,而后两者之间无显著差异;生殖枝生殖分配具有显著的性别差异（P <0.05）,但生殖枝叶生物量无性别差异。（3）生殖枝生物量大小与花朵生物量、营养生物量均呈极显著正相关关系（P <0.001）。营养生物量与花朵生物量之间为显著正相关关系(P <0.01),生殖分配与生殖枝大小无相关关系。研究认为,车桑子生殖枝营养生长与生殖生长不具有权衡关系,且生殖分配不具有个体大小依赖性,特定生物量分配模式可能是对金沙江干热河谷区资源利用、环境适应的一种特殊形态模式。     

濒危植物焕镛木的兼性无融合生殖

2001～2002年连续两年在广西环江木论和广西罗城大黄泥的2个焕镛木(Woonyoungia septentrionalis(Dandy)Law)自然种群中,对单性异株的濒危植物焕镛木进行繁育系统测定,对即将开花的雌花花蕾分别进行套袋、套网、人工授粉处理,并用自然授粉雌花作对照,其座果率和结实率统计结果表明:自然授粉、人工授粉、套袋和套网处理的花均能结实,但它们的座果率和结实率存在较大的差异.在两个种群中,人工授粉和自然授粉的总结实率(PERS)均比套袋和套网处理的高,其中人工授粉的最高,套网处理的最低.由此可见,焕镛木既能通过有性生殖方式结实,又能通过无融合生殖方式结实,而且这两种生殖方式获得的种子均能萌发成幼苗,由此断定,焕镛木的繁育系统为兼性无融合生殖.这是首次报道木兰科植物存在无融合生殖现象.     

菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系

三叶鬼针草(Bidens pilosa)是一种危害严重的菊科入侵植物。通过实验观察和人工控制套袋等方法, 对其花序开花动态、花粉胚珠比 (P/O)、自交亲和性、花粉活力、访花昆虫和种子 (瘦果)的萌发率等与繁殖相关的特征进行了研究, 探讨了这些繁殖特征与入侵性的关系。三叶鬼针草在10-11月开花, 单个花序的花期约为5-6天。每小花的花柱基部有圆筒状的蜜腺环绕。单个花序内可自交亲和, 自交结实率和花粉活力均较高, 其P/O值为1 7 54.1 2±29 .8 7。主要访花者为灰蝶科(Lycaenidae)、粉蝶科(Pieridae)和茧蜂科(Braconidae)昆虫。三叶鬼针草所具有的灵活交配机制是其入侵成功的重要因素。此外, 三叶鬼针草结实量大、种子产生迅速且适于传播, 以及种子萌发范围广和短期快速萌发等特性也增强了其入侵性。     

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。     

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.     

Sex allocation in hermaphroditic plants

Hermaphroditic plants allocate their reproductive resources to different functions: male, female and pollinator attraction. While earlier sex-allocation models considered only male and female functions, more recent ones can divide reproductive resources into multiple functions. The basic predictions derived from these models are similar. While most models predict sex allocation at the fruit stage (pollen and seeds), some have examined allocation at the flower stage (pollen and ovules). Selfing rate, mode of pollination and competition among offspring of the same parent are some of the factors that can influence sex allocation among populations. Although the empirical evidence lags behind the theoretical development, sex-allocation theory has been quite successful at predicting trends among populations.     

Sex allocation and sexual conflict in simultaneously hermaphroditic animals

Links between sex allocation (SA) and sexual conflict in simultaneous hermaphrodites have been evident since Charnov''s landmark paper published 30 years ago. We discuss two links, namely the potential for sexual conflict over SA between sperm donor and recipient, and the importance of post-copulatory sexual selection and the resulting sexual conflict for the evolution of SA. We cover the little empirical and theoretical work exploring these links, and present an experimental test of one theoretical prediction. The link between SA and sexual conflict is an interesting field for future empirical and theoretical research.     

Sex allocation in an hermaphroditic plant: the case of gynodioecy in Thymus vulgaris L.

Resource allocation to male and female functions was investigated in Thymus vulgaris L. (thyme), a gynodioecious species, in which females produce twice as many seeds as hermaphrodites. Negative correlations were found between male and female fertility of hermaphrodites, providing evidence of a trade-off. There was a high variability in sexual investment, some of the hermaphrodites functioning almost as males, and others almost as females. Estimation of the relative cost of male and female gametes showed that the female advantage in seed production was mainly due to reallocation of the resources not allocated to male function into female function. The determination of sex allocation was shown to have a genetic component, and there were some evidence that an interaction between nuclear and cytoplasmic genomes was involved.     

Sex allocation in functionally hermaphroditic plants: A review and critique

A number of theoretical and methodological problems plague studies of sex allocation in hermaphroditic plants. Most models assume that consumable resources limit reproduction, yet one or both sexes may be mate-limited. Most models also assume that resource limitation causes trade-offs between the allocations to male and female functions, but the sexes may be limited by different critical resources, or their resource needs may not overlap in time. Problems arise in deciding which plant parts are reproductive, when to harvest them, how to apportion them to male and female roles, and what are the appropriate measures of their cost. When energy directly limits reproduction, or other needs can be translated into energetic costs, the great variation in composition of reproductive parts makes construction costs or respiration a more potentially accurate measure of investment than energy content or especially biomass. Some simple predictions from theoretical models include a 1:1 allocation ratio to male and female function in outcrossing hermaphrodites, a female-biassed ratio in proportion to the level of selfing, and lower average allocations to male function in animal-pollinated than in windpollinated plants. These predictions have not received much support from existing studies, thus emphasizing the need for better measures of allocation as well as better accounting of the many other possible factors that may determine individual (and population) allocation ratios.     

Sexual allocation in single-flowered hermaphroditic individuals in relation to plant and flower size

Gender expression in hermaphroditic plant species usually departs from strict equisexuality. Study of those departures can aid understanding of non hermaphroditic breeding systems and prevalence of hermaphroditism within angiosperms. Plant size is one of the most studied factors in relation to gender modification. We studied variation in gender expression in the hermaphroditic, mostly single-flowered Paeonia cambessedesii. We separately studied gender modification with increasing plant and flower size using a variety of currencies: number of ovules and stamens, dry mass, N and P. Flower size and number of floral structures (petals, stamens, carpels, and ovules) increased with plant size. Number of ovules increased more rapidly with increasing plant size than number of stamens, indicating a bias towards femaleness with increasing plant size. A similar pattern was found when regressing number of stamens and number of seeds against plant size. Number of floral structures increased with increasing flower mass, but no significant difference was found between stamens and ovules in their rate of increase. Thus, gender modification at plant level was not consistent with patterns at flower level. No differential allocation to stamens vs gynoecium, or sexual structures vs petals was found when using dry mass, N or P as currencies. However, a disproportionate increase in female allocation was found when number of structures was utilised as currency. Study of size-dependent gender expression will benefit from contrast of results obtained using several analysis levels and allocation currencies.     

Sex allocation variation in Daphnia pulex

Daphnia (Crustacea: Cladocera) reproduce by cyclical parthenogenesis in which the sex of offspring is environmentally determined. Although numerous studies have demonstrated that factors such as crowding and short-day photoperiod stimulate male production, there is limited information on variation in allocation to male and female offspring for any species of Daphnia . The present study assessed the presence or absence of male production in 96 isofemale lines (clones) from each of eight populations of Daphnia pulex . An average of 37% (range 18–51%) of clones failed to produce males under crowded conditions in the laboratory. A subset of 14 of these non-male-producing clones also failed to produce males under short-day photoperiod (8L:16D). Three male-producing clones were within-clone mated as well as crossed to three non-male-producing clones to study the inheritance of the failure to produce males. The average frequency of non-male-producing F ₁ progeny was significantly higher (58%, N = 486) among the outcrossed progeny than the inbred progeny (5%, N = 86). In addition, when sixteen of the male-producing outcrossed progeny were within-clone mated, only 7% ( N = 106) of the resulting F ₂ progeny failed to produce males. These results are consistent with a genetic basis for the absence of male production. Average survival of the progeny from the nine outcrossed matings was more than twice (67%) that of the inbred progeny from the three within-clone matings (30%), suggesting that within-clone mating would result in significant inbreeding depression. We present a model that suggests that even low levels of inbreeding could allow non-male-producing females to be maintained in a population. The co-occurrence of non-male-producing females and females that produce both males and females in Daphnia pulex bears a similarity to the gynodioecious breeding system found in some plant species.     

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.     

How geitonogamous selfing affects sex allocation in hermaphrodite plants

Does the mode of self-pollination affect the evolutionarily stable allocation to male vs. female function? We distinguish the following scenarios. (1) An ‘autogamous’ species, in which selfing occurs within the flower prior to opening. The pollen used in selfing is a constant fraction of all pollen grains produced. (2) A species with ‘abiotic pollination’, in which selfing occurs when pollen dispersed in one flower lands on the stigma of a nearby flower on the same plant (geitonogamy). The selfing rate increases with male allocation but a higher selfing rate does not mean a reduced export of pollen. (3) An ‘animal-pollinated’ species with geitonogamous selfing. Here the selfing rate also increases with male allocation, but pollen export to other plants in the population is a decelerating function of the number of simultaneously open flowers. In all three models selfing selects for increased female allocation. For model 3 this contradicts the general opinion that geitonogamous selfing does not affect evolutionarily stable allocations. In all models, the parent benefits more from a female-biased allocation than any other individual in the population. In addition, in models 2 and 3, greater male allocation results in more local mate competition. In model 3 and in model 2 with low levels of inbreeding depression, hermaphroditism is evolutionarily stable. In model 2 with high inbreeding depression, the population converges to a fitness minimum for the relative allocation to male function. In this case the fitness set is bowed inwards, corresponding with accelerating fitness gain curves. If the selfing rate increases with plant size, this is a sufficient condition for size-dependent sex allocation (more allocation towards seeds in large plants) to evolve. We discuss our results in relation to size-dependent sex allocation in plants and in relation to the evolution of dioecy.