种子萌发对策:理论与实验

植物种子的萌发／休眠现象有复杂的原因和机制,综述了理论生态学家的研究结果。应用的理论基础是最优化理论和进化稳定对策(Ess)理论。当环境条件随机波动,种群受非密度依赖因素调节时,采用最优化理论的两头下注对策预测休眠一定会得到进化且萌发率与环境条件直接相关。环境条件稳定时采用进化稳定对策理论可得到在亲属竞争,种子扩散,基因冲突等等因素影响下的进化稳定休眠／萌发率,预测了休眠／萌发与它们之间的相互关系。以上各种环境条件影响种子萌发行为的方式可以表述为若种子立即萌发会遭遇到不良环境使适合度下降,那么就会推迟萌发,出现休眠,形成土壤种子库。萌发率应使种群适合度最优或具有进化稳定性。一些实验也部分验证了理论预测。     

An Application of the Sex Allocation Theory to Arisaema serratum

Abstract The sex allocation theory based on size-specific demography is applied to sex change phenomena exhibited by Arisaema serratum , which changes sex expression according to its size. In the evolutionary analysis where adopted strategy correlates with resource level, identifying equal resource level is very important. But directly measured size does not seem to represent the plant's real resource level perfectly. In the analysis presented here, we introduce a new variable that represents the resource level. According to the sex allocation theory, we calculated the transition matrix and other demographic parameters, the stable sex-and-size distribution, and reproductive values. The obtained results were: (1) reproductive values generally increased in accordance with size, but in a different manner for different sexes; (2) in the size class where different sex expressions were adopted, reproductive values for the equal resources were the same; and (3) plant individuals seemed to adopt evolutionarily stable strategies. The adaptive nature of the plant is further discussed.     

Offspring allocation in externally ovipositing fig wasps with varying clutch size and sex ratio

The simultaneous optimization of clutch size and sex ratio isa tricky problem. Unless parameters such as host size or fecundityexist to pin down the optimal clutch size, this problem remainselusive to analytical analysis. This is because the fitnesslandscape with respect to clutch size and sex ratio does nothave one single evolutionarily stable peak toward which thepopulation can evolve. To solve this problem, I used a computeremulation to optimize both clutch size and sex ratio using externallyovipositing fig wasps as a model taxon. The simulation approachallows the use of integer numbers of eggs rather than assumingthat females can produce any sex ratio between 0 and 1. Whenfemales have no information about the patches on which theyoviposit, they produce either large clutches with a strong femalebias or clutches of a single male egg. When females have completeknowledge of their oviposition site, a set of conditional substrategiesis evolutionarily stable. Again, these substrategies are eitherlarge clutches with a female bias or dutches consisting of asingle male egg. This dichotomous oviposition pattern resultsin unrelated males sharing a fig, a condition conducive to theevolution of fatal fighting. Selection on female ovipositionstrategies may therefore be an important driving force behindhigh levels of fighting observed between male fig wasps.     

ATTRACTIVE STRUCTURES AND THE STABILITY OF HERMAPHRODITIC SEX EXPRESSION IN FLOWERING PLANTS

Plant attributes serving both male and female sex functions are thought to favor hermaphroditic breeding systems over systems where sex expression is separated between individuals. Morphological features used by plants to attract biotic pollinators can be considered in this light, because such “attractive” structures are prerequisite for both pollen donation and receipt. A model involving allocation of a limiting resource between attractive structures benefiting both sex functions, structures benefiting male function only, and structures benefiting female function only, is constructed and analyzed using the evolutionary stable strategy (ESS) approach. Contrary to expectation, conditions resulting in large allocation to attractive structures tend to destabilize hermaphroditism, although the effect is slight. Substantial asymmetry in the benefits accruing to male and female fertility for a given investment in pollinator attraction produces more pronounced effects, reducing the parameter space in which hermaphroditism is expected. Results of this model are used to assess the importance of “accelerating gain” curves previously suggested for the evolution of dioecy in flowering plants. Accelerating gains associated with fruit production and dispersal represent female function only, and can be sufficient to initiate evolution toward dioecy. Accelerating gains associated with pollinator attraction may contribute to the evolution of separate sexes only if male function benefits from increased investment in attractive structures at a disproportionate rate compared to female function. Even in this instance, though, hermaphroditism may be evolutionarily stable.     

Landscape dynamics and evolution of colonizer syndromes: interactions between reproductive effortand dispersal in a metapopulation

The evolutionary consequences of changes in landscape dynamics for the evolution of life history syndromes are studied using a metapopulation model. We consider in turn the long-term effects of a change in the local disturbance rate, in the maximal local population persistence, in habitat productivity, and in habitat fragmentation. We examine the consequences of selective interactions between dispersal and reproductive effort by comparing the outcome of joint evolution to a situation where the species has lost the potential to evolve either its reproductive effort or its dispersal rate. We relax the classical assumption that any occupied site in the metapopulation reaches its carrying capacity immediately after recolonization. Our main conclusions are the following: (1) genetic diversity modifies the range of landscape parameters for which the metapopulation is viable, but it alters very little the qualitative evolutionary trends observed for each trait within this range. Although they are both part of a competition/colonization axis, reproductive effort and dispersal are not substitutable traits: their evolution reflects more directly the change in the landscape dynamics, than a selective interaction among them. (2) no general syndrome of covariation between reproductive effort and dispersal can be predicted: the pattern of association between the two traits depends on the type of change in landscape dynamics and on the saturation level. We review empirical evidence on colonizer syndromes and suggest lines for further empirical work. This revised version was published online in July 2006 with corrections to the Cover Date.     

韭菜迟眼蕈蚊的性别分化及其生态与进化意义

性别分化是遗传学、进化生物学和生殖行为生态学研究的重要领域之一,对于研究生物的遗传与进化关系、分析生物种群数量动态具有重要意义。对韭菜迟眼蕈蚊BradysiaodoriphagaYangetZhang的性别分化研究结果表明,韭菜迟眼蕈蚊的生殖对策主要以产单性别方式为主,其次是产优势性别方式,2种方式占92%,且产雌产雄的比例基本呈对称现象。用雌雄试虫都来自单性别的种群,连续繁殖3代,其性别分化如同上述,每一代种群的性比基本保持在1∶1。用双性别后代雌虫所产的雌雄虫进行近亲繁殖,连续繁殖3代,结果表明双性别后代的比例明显增大。鉴于以上实验结果,对韭菜迟眼蕈蚊生殖对策的生态与进化意义进行了分析。     

Biased Sex Allocation in Hermaphroditic Plants

Received 8 June 2000/ Accepted in revised form 7 July 2000     

Classic predictions about sex change do not hold under all types of size advantage

Theory predicts that the ‘size advantage’ (rate of increase in male and female fitness with age or size) determines the direction and the timing of sex change in sequential hermaphrodites. Whereas the size advantage is generated by the mating system and would be expected to vary within and between species, the shape or form of the size advantage has rarely been estimated directly. Here, we ask whether theoretical predictions about the timing of sex change hold under different types of size advantage. We model two biological scenarios representing different processes generating the size advantage and find that different types of size advantage can produce patterns that qualitatively differ from classic predictions. Our results demonstrate that a good understanding of sequentially hermaphroditic mating systems, and specifically, a direct assessment of the processes underlying the size advantage is crucial to reliably predict and explain within‐species patterns of the timing of sex change.     

Multi-species evolutionary dynamics

Dynamical attainability of an evolutionarily stable strategy (ESS) through the process of mutations and natural selection has mostly been addressed through the use of the continuously stable strategy (CSS) concept for species evolutionary games in which strategies are drawn from a continuum, and by the adaptive trait dynamics method. We address the issue of dynamical attainability of an ESS in coevolving species through the use of the concept of an ESNIS. It is shown that the definition of an ESNIS coalition for coevolving species is not in general equivalent to other definitions for CSS given in the literature. We show under some additional conditions that, in a dynamic system which involves the strategies of a dimorphic ESNIS coalition and at most two strategies that are not members of ESNIS coalition, the ESNIS coalition will emerge as the winner. In addition an ESNIS will be approached because of the invasion structure of strategies in its neighborhood. This proves that under the above conditions an ESNIS has a better chance of being attained than a strategy coalition which is a CSS. The theory developed is applied to a class of coevolutionary game models with Lotka–Volterra type interactions and we show that for such models, an ESS coalition will be dynamically attainable through mutations and natural selection if the ESS coalition is also an ESNIS coalition.Co-ordinating editor: Metz     

MODELS OF CHARACTER DISPLACEMENT AND THE THEORETICAL ROBUSTNESS OF TAXON CYCLES

The appropriateness of the techniques used in modeling character displacement has been the focus of vigorous debate. In this paper, the three competing methods (the coevolutionarily stable community (CSC), the evolutionarily stable strategy (ESS), and quantitative genetic recursion (QGR)) are compared in models using a common ecological setting. Specific predictions of the CSC model have been used to understand features of character displacement among Cnemidophorous lizards on islands off Mexico, Anolis lizards in the Lesser Antilles and Galápagos finches. Nonetheless, the validity of the approach has been repeatedly questioned. Conceptually the three formalisms vary in the degree to which within species variability is allowed in the models. The predictions of the CSC are found not to be robust to even small violation of its fundamental assumption of absolute species monomorphy. We show by simulation and analytical observations that the CSC is not valid under frequency dependent selection, and that the ESS is the limiting case of QGR as intraspecific phenotypic variation goes to zero. Thus the ESS and the QGR models agree closely when the between-phenotype component (BPC) of the niche width is small. However, as the BPC increases, quantitative discrepancies between ESS and QGR predictions increase, although model behavior remains qualitatively similar. A fourth approach, termed “Quantitative Genetic Optimization” (QGO) analysis, is suggested, combining advantages of both the ESS and QGR. Although all approaches support the possibility of taxon cycles, the cycle patterns predicted are qualitatively different and strongly model dependent.     

Evolution of learning capacities and learning levels

Humans strongly depend on individual and social learning, both of which are highly effective and accurate. I study the effects of environmental change on the evolution of the effectiveness and accuracy of individual and social learning (individual and social learning levels) and the number of pieces of information learned individually and socially (individual and social learning capacities) by analyzing a mathematical model. I show that individual learning capacity decreases and social learning capacity increases when the environment becomes more stable; both decrease when the environment becomes milder. I also show that individual learning capacity increases when individual learning level increases or social learning level decreases, while social learning capacity increases when individual or social learning level increases. The evolution of high learning levels can be triggered when the environment becomes severe, but a high social learning level can evolve only when a high individual learning level can simultaneously evolve with it.