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。     

Evolutionarily unstable fitness maxima and stable fitness minima of continuous traits

Summary We present models of adaptive change in continuous traits for the following situations: (1) adaptation of a single trait within a single population in which the fitness of a given individual depends on the population's mean trait value as well as its own trait value; (2) adaptation of two (or more) traits within a single population; (3) adaptation in two or more interacting species. We analyse a dynamic model of these adaptive scenarios in which the rate of change of the mean trait value is an increasing function of the fitness gradient (i.e. the rate of increase of individual fitness with the individual's trait value). Such models have been employed in evolutionary game theory and are often appropriate both for the evolution of quantitative genetic traits and for the behavioural adjustment of phenotypically plastic traits. The dynamics of the adaptation of several different ecologically important traits can result in characters that minimize individual fitness and can preclude evolution towards characters that maximize individual fitness. We discuss biological circumstances that are likely to produce such adaptive failures for situations involving foraging, predator avoidance, competition and coevolution. The results argue for greater attention to dynamical stability in models of the evolution of continuous traits.     

The genetic mating system of spotted sunfish (Lepomis punctatus): mate numbers and the influence of male reproductive parasites

In nest-building fish species, mature males often exhibit one of two alternative reproductive behaviours. Bourgeois males build nests, court females, and guard their eggs. Parasitic cuckolders attempt to steal fertilizations from bourgeois males and do not invest in parental care. Previous evidence from the bluegill sunfish (Lepomis macrochirus) suggests that adult males are morphologically specialized for these two tactics. Here, we used microsatellite markers to determine genetic parentage in a natural population of the spotted sunfish (L. punctatus) that also displayed both bourgeois and parasitic male morphs. As gauged by relative investments in gonadal vs. somatic tissues, between 5 and 15% of the mature adult males were parasites. Multi-locus genotypes were generated for more than 1400 embryos in 30 nests, their nest-guardian males, and for other adults in the population. Progeny in approximately 57% of the nests were sired exclusively by the guardian male, but the remaining nests contained embryos resulting from cuckoldry as well. Overall, the frequency of offspring resulting from stolen fertilizations was only 1.3%, indicating that the great majority of paternity is by bourgeois nesting males. With regard to maternity, 87% of the nests had at least three dams, and computer simulations estimate that about 7.2 dams spawned per nest.     

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.     

Why Darwin would have loved evolutionary game theory

Humans have marvelled at the fit of form and function, the way organisms'' traits seem remarkably suited to their lifestyles and ecologies. While natural selection provides the scientific basis for the fit of form and function, Darwin found certain adaptations vexing or particularly intriguing: sex ratios, sexual selection and altruism. The logic behind these adaptations resides in frequency-dependent selection where the value of a given heritable phenotype (i.e. strategy) to an individual depends upon the strategies of others. Game theory is a branch of mathematics that is uniquely suited to solving such puzzles. While game theoretic thinking enters into Darwin''s arguments and those of evolutionists through much of the twentieth century, the tools of evolutionary game theory were not available to Darwin or most evolutionists until the 1970s, and its full scope has only unfolded in the last three decades. As a consequence, game theory is applied and appreciated rather spottily. Game theory not only applies to matrix games and social games, it also applies to speciation, macroevolution and perhaps even to cancer. I assert that life and natural selection are a game, and that game theory is the appropriate logic for framing and understanding adaptations. Its scope can include behaviours within species, state-dependent strategies (such as male, female and so much more), speciation and coevolution, and expands beyond microevolution to macroevolution. Game theory clarifies aspects of ecological and evolutionary stability in ways useful to understanding eco-evolutionary dynamics, niche construction and ecosystem engineering. In short, I would like to think that Darwin would have found game theory uniquely useful for his theory of natural selection. Let us see why this is so.     

The evolution of traits that determine ability in competitive contests

Summary We analyse mathematical models of the evolution of a trait that determines ability in contest competition. We assume that the value of the competitive trait affects two different components of fitness, one measuring the benefit of winning contests and the other measuring the cost of developing the competitive trait. Unlike previous analyses, we include the population dynamical consequences of larger competitive trait values. Exaggeration of the competitive trait reduces the mean probability of survival during the non-competitive stage of the life cycle. The resulting lower population density reduces competition and, therefore, reduces the advantages of greater competitive ability. Models without population dynamics often predict dimorphism in the competitive trait when resource possession is decided by interactions with many other individuals. If the competition involves a contest with a single other individual, models without population dynamics often predict cycles of increase and collapse in the trait or a continual increase, possibly resulting in extinction. When population dynamics are included, both of these results become less likely and a single stable trait value becomes more likely. Population dynamics also make it possible to have dimorphism when individuals have a single pairwise contest and alternative stable trait values when an individual has many contests. Increases in the value of the resource being contested may increase or decrease the evolutionarily stable size of the trait. Competition between very differently sized species will often decrease size in the larger species (character convergence).     

THE EVOLUTIONARILY STABLE PHENOTYPE DISTRIBUTION IN A RANDOM ENVIRONMENT

In an unpredictably changing environment, phenotypic variability may evolve as a “bet-hedging” strategy. We examine here two models for evolutionarily stable phenotype distributions resulting from stabilizing selection with a randomly fluctuating optimum. Both models include overlapping generations, either survival of adults or a dormant propagule pool. In the first model (mixed-strategies model) we assume that individuals can produce offspring with a distribution of phenotypes, in which case, the evolutionarily stable population always consists of a single genotype. We show that there is a unique evolutionarily stable strategy (ESS) distribution that does not depend on the amount of generational overlap, and that the ESS distribution generically is discrete rather than continuous; that is, there are distinct classes of offspring rather than a continuous distribution of offspring phenotypes. If the probability of extreme fluctuations in the optimum is sufficiently small, then the ESS distribution is monomorphic: a single type fitted to the mean environment. At higher levels of variability, the ESS distribution is polymorphic, and we find stability conditions for dimorphic distributions. For an exponential or similarly broad-tailed distribution of the optimum phenotype, the ESS consists of an infinite number of distinct phenotypes. In the second model we assume that an individual produces offspring with a single, genetically determined phenotype (pure-strategies model). The ESS population then contains multiple genotypes when the environmental variance is sufficiently high. However the phenotype distributions are similar to those in the mixed-strategies model: discrete, with an increasing number of distinct phenotypes as the environmental variance increases.     

The evolution of conformist transmission in social learning when the environment changes periodically

Conformity is often observed in human social learning. Social learners preferentially imitate the majority or most common behavior in many situations, though the strength of conformity varies with the situation. Why has such a psychological tendency evolved? I investigate this problem by extending a standard model of social learning evolution with infinite environmental states (Feldman, M.W., Aoki, K., Kumm, J., 1996. Individual versus social learning: evolutionary analysis in a fluctuating environment. Anthropol. Sci. 104, 209-231) to include conformity bias. I mainly focus on the relationship between the strength of conformity bias that evolves and environmental stability, which is one of the most important factors in the evolution of social learning. Using the evolutionarily stable strategy (ESS) approach, I show that conformity always evolves when environmental stability and the cost of adopting a wrong behavior are small, though environmental stability and the cost of individual learning both negatively affect the strength of conformity.     

The evolution of dispersal in a two-patch system: some consequences of differences between migrants and residents

We investigate how age-structure and differences in certain demographic traits between residents and immigrants of a single species act to determine the evolutionarily stable dispersal strategy in a two-patch environment that is heterogeneous in space but constant in time. These two factors have been neglected in previous models of the evolution of dispersal, which generally consider organisms with very simple life-cycles and assume that, whatever their origin, individuals in a given habitat have the same bio-demographic characteristics. However, there is increasing empirical evidence that dispersing individuals have different demographic properties from phylopatric ones. We develop a matrix model in which recruitment depends on local population densities. We assume that dispersal entails a proportional cost to immigrant fecundity, which can be compensated by differences in survival rates between immigrants and residents. The evolutionarily stable strategies (ESS) for dispersal are identified using a combination of analytical expressions and numerical simulations. Our results show that philopatry is selected (1) when dispersal rates do not vary in space, (2) when the metapopulation is a source-sink system and (3) when dispersal rates vary in space (asymmetric dispersal) and immigrants do not compensate for their reduced fecundity. We observe that non-zero asymmetric dispersal rates may be evolutionarily stable when (1) immigrants and residents are demographically alike and (2) immigrants compensate totally for their reduced fecundity through an increase in adult survival. Under these conditions, we find that the ESS occurs when the fitnesses at equilibrium in the two habitats, measured in our model by the realized reproductive rates, are each equal to unity. A comparison with previous studies suggests a unifying rule for the evolution of dispersal: the dispersal rates which permit the spatial homogenization of fitnesses are ESSs. This condition provides new insight into the evolutionary stability of source-sink systems. It also supports the hypothesis that immigrants have adapted demographic strategies, rather than the hypothesis that dispersal is costly and immigrants are at a disavantage compared with residents.     

POLYPHENISM IN SPADEFOOT TOAD TADPOLES AS A LOCALLY ADJUSTED EVOLUTIONARILY STABLE STRATEGY

I examined the evolutionary factors maintaining two environmentally induced morphs in ponds of variable duration. Larvae of New Mexico spadefoot toads (Scaphiopus multiplicatus) often occur in the same pond as a large, rapidly developing carnivorous morph and as a smaller, more slowly developing omnivorous morph. Previous studies revealed that carnivores can be induced by feeding tadpoles live fairy shrimp and that morph determination is reversible. Field and laboratory experiments indicated that the ability of an individual to become a carnivore or an omnivore is maintained evolutionarily as a response to variability in pond longevity and food abundance. Carnivores survived better in highly ephemeral artificial ponds, because they developed faster. Omnivores survived better in longer-duration artificial ponds, because their larger fat reserves enhanced postmetamorphic survival. The two morphs also occupy different trophic niches. Experimental manipulations of morph frequency in ponds of intermediate duration revealed that increased competition for food among individuals of the more common morph made the rarer form more successful. Morph frequency within each pond was stabilized at an equilibrium by frequency-dependent morph reversal, which reflected frequency-dependent natural selection on size at metamorphosis: larger metamorphs had higher survival, and individuals reared at a frequency above the pond's equilibrium frequency were smaller at metamorphosis than were individuals of that morph reared at a frequency below the pond's equilibrium. Because neighboring ponds often differed in pond longevity and food abundance, each pond possessed a unique equilibrium morph frequency. This implies that morph determination in Scaphiopus is a locally adjusted evolutionarily stable strategy (ESS).     

Asymmetric interaction will facilitate the evolution of cooperation

Explaining the evolution of cooperation remains one of the greatest problems for both biology and social science. The classical theories of cooperation suggest that cooperation equilibrium or evolutionary stable strategy between partners can be maintained through genetic similarity or reciprocity relatedness. These classical theories are based on an assumption that partners interact symmetrically with equal payoffs in a game of cooperation interaction. However, the payoff between partners is usually not equal and therefore they often interact asymmetrically in real cooperative systems. With the Hawk-Dove model, we find that the probability of cooperation between cooperative partners will depend closely on the payoff ratio. The higher the payoff ratio between recipients and cooperative actors, the greater will be the probability of cooperation interaction between involved partners. The greatest probability of conflict between cooperative partners will occur when the payoff between partners is equal. The results show that this asymmetric relationship is one of the key dynamics of the evolution of cooperation, and that pure cooperation strategy (i.e., Nash equilibrium) does not exist in asymmetrical cooperation systems, which well explains the direct conflict observed in almost all of the well documented cooperation systems. The model developed here shows that the cost-to-benefit ratio of cooperation is also negatively correlated with the probability of cooperation interaction. A smaller cost-to-benefit ratio of cooperation might be created by the limited dispersal ability or exit cost of the partners involved, and it will make the punishment of the non-cooperative individuals by the recipient more credible, and therefore make it more possible to maintain stable cooperation interaction.     

Evolution of unstable and stable biparental care

Evolutionarily stable strategy models suggest that biparentalcare will be stable when parents partially compensate for changesin care by the other parent. Previous work has emphasized therelationship between parental expenditure and the current componentof fitness (e. g., offspring survival and fecundity) in causingpartial compensation. This study shows that partial compensationdepends critically on the effect of current parental expenditureon a parent's future fitness (e. g., survival to and fecundityin subsequent breeding seasons). Partial compensation is favoredand biparental care is stable when future fitness is a concave-downfunction of expenditure (i. e., each increment of expenditureis more costly than the previous). However, when future fitnessis a convex-down function of expenditure (i. e., each incrementof expenditure is less costly) biparental care is unstable.(BehavEcol 7: 490–493(1996)]     

ALLOCATION TO ATTRACTIVE STRUCTURES IN ANIMAL-POLLINATED FLOWERS

An optimal allocation model was developed for the evolutionarily stable size of attractive structures of flowers (ESA) in animal-pollinated plants. It was assumed that a plant can change the sizes of attractive and sexual structures of a flower and the size and the number of flowers. In the absence of constraints on flower size, the ESA should not depend on the frequency of self-fertilization or the sexuality of plants. However, with constraints on flower size, the ESA decreases with increasing self-fertilization, except in special cases, and it is possible that males have a larger or a smaller ESA than females. Thus, differences in self-fertilization and sexuality alone cannot explain the differences in allocation among nondomesticated plants. In addition, attractive structures can contribute more to male or female function depending on the cost of gamete production, pollination efficiency for pollen and ovules, and pollinator availability.     

Maximization of fitness through limiting the number of offspring in the evolution ofHomo sapiens

It is proposed that the concealment of ovulation and the extension of sexual receptivity of the hominid female is a fitness maximizing adaptation resulting in limiting the number of offspring. To support this hypothesis a probability model is introduced. For a specific set of parameter values (including maternal mortality during delivery) through a formal deduction, it is shown that keeping the number of offspring below the physiological limit is a fitness maximizing strategy.     

On measuring selection in experimental evolution

Distributions of mutation fitness effects from evolution experiments are available in an increasing number of species, opening the way for a vast array of applications in evolutionary biology. However, comparison of estimated distributions among studies is hampered by inconsistencies in the definitions of fitness effects and selection coefficients. In particular, the use of ratios of Malthusian growth rates as ‘relative fitnesses’ leads to wrong inference of the strength of selection. Scaling Malthusian fitness by the generation time may help overcome this shortcoming, and allow accurate comparison of selection coefficients across species. For species reproducing by binary fission (neglecting cellular death), ln2 can be used as a correction factor, but in general, the growth rate and generation time of the wild-type should be provided in studies reporting distribution of mutation fitness effects. I also discuss how density and frequency dependence of population growth affect selection and its measurement in evolution experiments.     

RUNAWAY EVOLUTION TO SELF-EXTINCTION UNDER ASYMMETRICAL COMPETITION

We analyze a popular model of the evolution of traits related to performance in exploitative competition. This model has previously been used to explain a mechanism by which interspecific competition can cause taxon cycles. We show that purely intraspecific competition can cause evolution of extreme competitive abilities that ultimately result in extinction, without any influence from other species. The only change in the model required for this outcome is the assumption of a nonnormal distribution of resources of different sizes measured on a logarithmic scale. This suggests that taxon cycles, if they exist, may be driven by within- rather than between-species competition. Self-extinction does not occur when the advantage conferred by a large value of the competitive trait (e.g., size) is relatively small, or when the carrying capacity decreases at a comparatively rapid rate with increases in trait value. The evidence regarding these assumptions is discussed. The results suggest a need for more data on resource distributions and size-advantage in order to understand the evolution of competitive traits such as body size.     

On several conjectures from evolution of dispersal

We address several conjectures raised in Cantrell et al. [Evolution of dispersal and ideal free distribution, Math. Biosci. Eng. 7 (2010), pp. 17–36 [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]]] concerning the dynamics of a diffusion–advection–competition model for two competing species. A conditional dispersal strategy, which results in the ideal free distribution of a single population at equilibrium, was found in Cantrell et al. [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]]. It was shown in [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]] that this special dispersal strategy is a local evolutionarily stable strategy (ESS) when the random diffusion rates of the two species are equal, and here we show that it is a global ESS for arbitrary random diffusion rates. The conditions in [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]] for the coexistence of two species are substantially improved. Finally, we show that this special dispersal strategy is not globally convergent stable for certain resource functions, in contrast with the result from [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]], which roughly says that this dispersal strategy is globally convergent stable for any monotone resource function.