The combined effects of temporal autocorrelation and the costs of plasticity on the evolution of plasticity

Adaptive phenotypic plasticity is an important source of intraspecific variation, and for many plastic traits, the costs or factors limiting plasticity seem cryptic. However, there are several different factors that may constrain the evolution of plasticity, but few models have considered costs and limiting factors simultaneously. Here we use a simulation model to investigate how the optimal level of plasticity in a population depends on a fixed maintenance fitness cost for plasticity or an incremental fitness cost for producing a plastic response in combination with environmental unpredictability (environmental fluctuation speed) limiting plasticity. Our model identifies two mechanisms that act, almost separately, to constrain the evolution of plasticity: (i) the fitness cost of plasticity scaled by the nonplastic environmental tolerance, and (ii) the environmental fluctuation speed scaled by the rate of phenotypic change. That is, the evolution of plasticity is constrained by the high cost of plasticity in combination with high tolerance for environmental variation, or fast environmental changes in combination with slow plastic response. Qualitatively similar results are found when maintenance and incremental fitness costs of plasticity are incorporated, although a larger degree of plasticity is selected for with an incremental cost. Our model highlights that it is important to consider direct fitness costs and phenotypic limitations in relation to nonplastic environmental tolerance and environmental fluctuations, respectively, to understand what constrains the evolution of phenotypic plasticity.     

How to balance the offspring quality–quantity tradeoff when environmental cues are unreliable

Maternal effects on offspring size can have a strong effect on fitness, as larger offspring often survive better under harsh environmental conditions. Selection should hence favour mothers that find an optimal solution to the offspring size versus number tradeoff. If environmental conditions are variable, there will not be a single optimal offspring size, as predicted in a constant environment, but plastic responses can be favoured. To be able to adjust offspring size in an adaptive manner, mothers have to use environmental cues to predict offspring environmental conditions. Cues can be unreliable, however, particularly in species where individuals occupy different niches at different life stages. Here we model the evolution of plasticity of offspring size when the environmental cues mothers use to predict the conditions experienced by their offspring are not perfectly reliable. Our results show that plastic strategies are likely to be superior to fixed strategies in a stochastically varying environment when the environmental cues are at least moderately reliable, with the threshold depending on plasticity costs and the difference of resources available to mothers. Plasticity is more likely to occur if resource availability is not too different between environments. For any given scenario, plasticity in offspring size is favoured if offspring survival varies greatly between environmental states. Whenever plastic strategies are optimal, the occurring switches performed by mothers between small and large offspring are predicted to be substantial, as small adjustments are unlikely to reap fitness benefits great enough to overcome the costs of plasticity.     

A heuristic model on the role of plasticity in adaptive evolution: plasticity increases adaptation,population viability and genetic variation

An ongoing new synthesis in evolutionary theory is expanding our view of the sources of heritable variation beyond point mutations of fixed phenotypic effects to include environmentally sensitive changes in gene regulation. This expansion of the paradigm is necessary given ample evidence for a heritable ability to alter gene expression in response to environmental cues. In consequence, single genotypes are often capable of adaptively expressing different phenotypes in different environments, i.e. are adaptively plastic. We present an individual-based heuristic model to compare the adaptive dynamics of populations composed of plastic or non-plastic genotypes under a wide range of scenarios where we modify environmental variation, mutation rate and costs of plasticity. The model shows that adaptive plasticity contributes to the maintenance of genetic variation within populations, reduces bottlenecks when facing rapid environmental changes and confers an overall faster rate of adaptation. In fluctuating environments, plasticity is favoured by selection and maintained in the population. However, if the environment stabilizes and costs of plasticity are high, plasticity is reduced by selection, leading to genetic assimilation, which could result in species diversification. More broadly, our model shows that adaptive plasticity is a common consequence of selection under environmental heterogeneity, and hence a potentially common phenomenon in nature. Thus, taking adaptive plasticity into account substantially extends our view of adaptive evolution.     

Plastic inducible morphologies are not always adaptive: The importance of time delays in a stochastic environment

Summary We present a mathematical model for predicting the expected fitness of phenotypically plastic organisms experiencing a variable environment. We assume that individuals experience two discrete environments probabilistically in time (as a Markov process) and that there are two different phenotypic states, each yielding the highest fitness in one of the two environments. We compare the expected fitness of a phenotypically fixed individual to that of an individual whose phenotype is induced to produce the better phenotype in each environment with a time lag between experiencing a new environment and realization of the new phenotype. Such time lags are common in organisms where phenotypically plastic, inducible traits have been documented. We find that although plasticity is generally adaptive when time lags are short (relative to the time scale of environmental variability), plasticity can be disadvantageous for longer lag times. Asymmetries in environmental change probabilities and/or the relative fitnesses of each phenotype strongly influence whether plasticity is favoured. In contrast to other models, our model does not require costs for plasticity to be disadvantageous; costs affect the results quantitatively, not qualitatively.     

Comparing the effects of rapid evolution and phenotypic plasticity on predator-prey dynamics

Ecologists have increasingly focused on how rapid adaptive trait changes can affect population dynamics. Rapid adaptation can result from either rapid evolution or phenotypic plasticity, but their effects on population dynamics are seldom compared directly. Here we examine theoretically the effects of rapid evolution and phenotypic plasticity of antipredatory defense on predator-prey dynamics. Our analyses reveal that phenotypic plasticity tends to stabilize population dynamics more strongly than rapid evolution. It is therefore important to know the mechanism by which phenotypic variation is generated for predicting the dynamics of rapidly adapting populations. We next examine an advantage of a phenotypically plastic prey genotype over the polymorphism of specialist prey genotypes. Numerical analyses reveal that the plastic genotype, if there is a small cost for maintaining it, cannot coexist with the pairs of specialist counterparts unless the system has a limit cycle. Furthermore, for the plastic genotype to replace specialist genotypes, a forced environmental fluctuation is critical in a broad parameter range. When these results are combined, the plastic genotype enjoys an advantage with population oscillations, but plasticity tends to lose its advantage by stabilizing the oscillations. This dilemma leads to an interesting intermittent limit cycle with the changing frequency of phenotypic plasticity.     

The genetics of phenotypic plasticity. XV. Genetic assimilation,the Baldwin effect,and evolutionary rescue

We used an individual‐based simulation model to examine the role of phenotypic plasticity on persistence and adaptation to two patterns of environmental variation, a single, abrupt step change and continual, linear change. Our model tested the assumptions and predictions of the theory of genetic assimilation, explored the evolutionary dynamics of the Baldwin effect, and provided expectations for the evolutionary response to climate change. We found that genetic assimilation as originally postulated is not likely to occur because the replacement of plasticity by fixed genetic effects takes much longer than the environment is likely to remain stable. On the other hand, trait plasticity as an enhancement to continual evolutionary change may be an important evolutionary mechanism as long as plasticity has little or no costs. Whether or not plasticity helps or hinders evolutionary rescue following a step change in the environment depends on whether plasticity is costly. For linear environmental change, noncostly plasticity always decreases extinction rates, while costly plasticity can create a fitness drag and increase the chance of extinction. Thus, with changing climates plasticity can enhance adaptation and prevent extinction under some conditions, but not others.     

Plasticity versus canalization: population differences in the timing of shade-avoidance responses

The reliability of environmental cues and costs of a fixed phenotype are two factors determining whether selection favors phenotypic plasticity or environmental specialization. This study examines the relationship between these two factors and the evolution of plant competitive strategies (plastic vs. fixed morphologies). In natural plant populations, shifts in light quality associated with foliar shade reliably indicate the presence of neighbors. These cues mediate plastic stem-elongation responses that often increase competitive ability and access to light. Using experimental light treatments (full sun, neutral shade, and foliar shade), genetic differences among populations of Abutilon theophrasti (velvetleaf) in average elongation and plasticity to foliar-shade cues were examined. Six populations, two from each of three site types (fields in continuous corn cultivation, fields undergoing corn-soy rotation, and weedy sites), were exposed to the light treatments at two stages in their life history. At the seedling stage, populations derived from cornfield sites exhibited higher, average elongation than populations from either rotating corn-soy fields or weedy areas. Because seedling elongation may delay shading of velvetleaf by corn, population differences may reflect adaptive responses to directional selection imposed by competitive conditions. However, the effects of simulated foliar shade on elongation were three times as great as the average population differences, and these comparatively higher levels of elongation were associated with an allocation cost. These results are consistent with the hypothesis that phenotypic plasticity may limit the evolution of specialists; reliable environmental cues enable individuals to facultatively adopt highly elongated, costly phenotypes in crowded patches while avoiding the costs of that phenotype in less crowded microsites. At later life-history stages, populations experiencing competition with corn exhibited lower plasticity to light quality than populations derived from weedy areas. Elongation at later nodes is maladaptive in cornfields because velvetleaf is ultimately incapable of overtopping corn; individuals that elongate therefore experience the cost of allocating to stems but fail to improve leaf exposure. The decreased responsiveness of cornfield populations to light quality is consistent with theoretical predictions in which reduced plasticity is favored when environmental cues fail to mediate an adaptive response.     

Evolution of environmental cues for phenotypic plasticity

Phenotypically plastic characters may respond to multiple variables in their environment, but the evolutionary consequences of this phenomenon have rarely been addressed theoretically. We model the evolution of linear reaction norms in response to several correlated environmental variables, in a population undergoing stationary environmental fluctuations. At evolutionary equilibrium, the linear combination of environmental variables that acts as a developmental cue for the plastic trait is the multivariate best linear predictor of changes in the optimum. However, the reaction norm with respect to any single environmental variable may exhibit nonintuitive patterns. Apparently maladaptive, or hyperadaptive plasticity can evolve with respect to single environmental variables, and costs of plasticity may increase, rather than reduce, plasticity in response to some variables. We also find conditions for the evolution of an indirect environmental indicator that affects expression of a plastic phenotype, despite not influencing natural selection on it.     

Phenotypic integration and plastic correlations in Phlox drummondii (Polemoniaceae)

Clones from two populations of Phlox drummondii were grown in three different nutrient environments to determine the extent to which the overall level and pattern of correlation among traits within an environment changes across environments. With one exception, the level of phenotypic correlation in both populations was the same across environments. Plants from Lexington, Texas exhibited a significantly lower level of phenotypic correlation when grown at a high nutrient concentration. The two populations did not differ from one another in their levels of phenotypic correlation when compared within environments. The pattern of correlation was homogenous both within populations across environments and among populations within environments. Tests of a priori hypotheses regarding the associations among functionally or developmentally related traits suggest that the correlations among traits are higher in traits that share a common function or developmental origin. We also compared the level and pattern of plasticity correlations among populations for three different components of the plastic response. We found that the level and pattern of plastic correlation for the average, linear, and nonlinear components of the plastic response did not differ among the two populations. With only one exception, the relationships among the plastic responses of different traits fit our model of functional and developmental integration. The results from our analyses of phenotypic and plastic correlations support the hypothesis that plastic correlations determine the extent to which phenotypic correlations are environment-dependent.     

Biogeochemical Change During Climate-Driven Afforestation: A Paleoecological Perspective from the Rocky Mountains

Shifts in an ecosystem’s state can alter biogeochemical cycling and the extent of nutrient conservation within a terrestrial landscape on multiple time scales. Transient biogeochemical changes may follow disturbance and succession, although persistent long-term differences may exist under different climates and vegetation types. We evaluate the potential for such biogeochemical changes in the context of long-term ecological history by measuring the nitrogen isotope composition of organic matter in a lake sediment core. We targeted Little Windy Hill Pond (LWH) in the Medicine Bow Mountains, Wyoming because reconstructions of the lake level, fire, and vegetation histories from the lacustrine sediments indicated a century-scale transformation from an arid, shrub-dominated landscape to a sub-alpine, tree-dominated ecosystem with extensive woody cover and large, live biomass pools. We demonstrate that the afforestation at the beginning of the Holocene transformed the Artemisia-dominated ecosystem, which had persisted for millennia during the Pleistocene. The changes affected nitrogen cycling dynamics, especially through intensified nutrient conservation when live biomass pools increased with greater woody cover. The LWH sediments record a baseline δ¹⁵N shift from 2.2–3.0 to 0.3–2.0‰ as less ¹⁵N-enriched organic matter accumulated in the lake. We also observed a transient pattern of maximum nutrient conservation and minimum δ¹⁵N values as terrestrial biomass increased during the aggradation (~175 years) and transition phases of ecological succession. Our nitrogen isotope results support theoretical expectations of long-term biogeochemical dynamics as nutrient conservation increases during afforestation.     

Deepened snow increases late thaw biogeochemical pulses in mesic low arctic tundra

Pulses of plant-available nutrients to the soil solution are expected to occur during the dynamic winter–spring transition in arctic tundra. Our aims were to quantify the magnitude of these potential nutrient pulses, to understand the sensitivity of these pulses to winter conditions, and to characterize and integrate the environmental and biogeochemical dynamics of this period. To test the hypotheses that snow depth, temperature and soil water—and not snow nutrient content—are important controls on winter and spring biogeochemistry, we sampled soil from under ambient and deepened snow every 3 days from late winter to spring, in addition to the snowpack at the start of thaw. Soil and microbial biogeochemical dynamics were divided into distinct phases that correlated with steps in soil temperature and soil water. Soil solution and microbial pools of C, N and P fluctuated with strong peaks and declines throughout the thaw, especially under deepened snow. Snowpack nutrient accumulation was negligible relative to these biogeochemical peaks. All nutrient and microbial peaks declined simultaneously at the end of snowmelt and so this decline was delayed by 15 days under deepened snow. The timing of these nutrient pulses is critical for plant species nutrient availability and landscape nutrient budgets. This detailed and statistically-based characterisation of the winter–spring transition in terms of environmental and biogeochemical variables should provide a useful foundation for future biogeochemical process-based studies of thaw, and indicate that spring thaw and possibly growing season biogeochemical dynamics are sensitive to present and future variability in winter snow depth.     

How humans alter dissolved organic matter composition in freshwater: relevance for the Earth’s biogeochemistry

Dissolved organic matter (DOM) is recognized for its importance in freshwater ecosystems, but historical reliance on DOM quantity rather than indicators of DOM composition has led to an incomplete understanding of DOM and an underestimation of its role and importance in biogeochemical processes. A single sample of DOM can be composed of tens of thousands of distinct molecules. Each of these unique DOM molecules has their own chemical properties and reactivity or role in the environment. Human activities can modify DOM composition and recent research has uncovered distinct DOM pools laced with human markers and footprints. Here we review how land use change, climate change, nutrient pollution, browning, wildfires, and dams can change DOM composition which in turn will affect internal processing of freshwater DOM. We then describe how human-modified DOM can affect biogeochemical processes. Drought, wildfires, cultivated land use, eutrophication, climate change driven permafrost thaw, and other human stressors can shift the composition of DOM in freshwater ecosystems increasing the relative contribution of microbial-like and aliphatic components. In contrast, increases in precipitation may shift DOM towards more relatively humic-rich, allochthonous forms of DOM. These shifts in DOM pools will likely have highly contrasting effects on carbon outgassing and burial, nutrient cycles, ecosystem metabolism, metal toxicity, and the treatments needed to produce clean drinking water. A deeper understanding of the links between the chemical properties of DOM and biogeochemical dynamics can help to address important future environmental issues, such as the transfer of organic contaminants through food webs, alterations to nitrogen cycling, impacts on drinking water quality, and biogeochemical effects of global climate change.

     

Unraveling the ecosystem functions in the Amazonia–Cerrado transition: evidence of hyperdynamic nutrient cycling

The contact between savanna and forest in the Amazonia–Cerrado transition zone is characterized by the hyperdynamics of the vegetation (recruitment vs. mortality). However, the related nutrient dynamics under these conditions are not well understood. We determined for the first time the biogeochemical cycles of the vegetation in the zone of transition estimating the litterfall, nutrient input, decomposition rates, and nutrient release in cerradão and cerrado plots. We examine the hypothesis that nutrient cycling is strongly associated with the vegetation dynamics. The litterfall was sampled in 30 traps placed within 1-ha vegetation plots for 2 years. The release of nutrients from the litterfall back to the soil was also estimated using decomposition bags in the two areas. The decomposition rates did not vary between areas, although in the cerradão the input of total biomass (9.27 Mg ha^?1 year^?1) and total nutrients (219.17 kg ha^?1 year^?1), the decomposition of the total biomass, and the cycling of most nutrients through litterfall and decomposition were at least twice higher than in the cerrado. These results confirmed the hypothesis concerning the differences between vegetation types in nutrient cycling, suggesting for the first time that the hyperdynamics observed in both vegetations were also reflected in the biogeochemical cycle, particularly in the cerradão. Thus, it is likely that the rapid and effective cycling of nutrients observed in the cerradão might be a key condition guaranteeing the ability of the cerradão to colonize new areas previously occupied by the typical cerrado.     

The timescale of environmental fluctuations determines the competitive advantages of phenotypic plasticity and rapid evolution

Organisms can respond to fluctuating environments by phenotypic plasticity and rapid evolution, both occurring on similar timescales to the environmental fluctuations. Because each adaptation mechanism has been independently studied, the effects of different adaptation mechanisms on ecological dynamics are not well understood. Here, using mathematical modeling, we compared the advantages of phenotypic plasticity and rapid evolution under conditions where the environment fluctuated between two states on various timescales. The results indicate that the advantages of phenotypic plasticity under environmental fluctuations on different timescales depend on the cost and the speed of plasticity. Both the speed of plastic adaptation and the cost of plasticity affect competition results, while the quantitative effects of them vary depending on the timescales. When the environment fluctuates on short timescales, the two populations with evolution and plasticity coexist, although the population with evolution is dominant. On moderate timescales, the two populations also coexist; however, the population with plasticity becomes dominant. On long timescales, whether the population with phenotypic plasticity or evolution is more advantageous depended on the cost of plasticity. Moreover, our results indicate that the mechanisms resulting in the dominance of the plastic population over the population with evolution are different depending on the timescales of environmental fluctuations. Therefore, the timescales of environmental fluctuations deserve more attention if we are to better understand the detailed competition results underlying phenotypic variation.     

Multivariate selection shapes environment-dependent variation in the clonal morphology of a red seaweed

Within-individual strategies of variation (e.g., phenotypic plasticity) are particularly relevant to modular organisms, in which ramets of the same genetic individual may encounter diverse environments imposing diverse patterns of selection. Hence, measuring selection in heterogeneous environments is essential to understanding whether environment-dependent phenotypic change enhances the fitness of modular individuals. In sublittoral marine habitats, competition for light and space among modular taxa generates extreme patchiness in resource availability. Little is known, however, of the potential for plasticity within individuals to arise from spatially-variable selection in such systems. We tested whether plasticity enhances genet-level fitness in Asparagopsis armata, a clonal seaweed in which correlated traits mediate morphological responses to variation in light. Using the capacity for rapid, clonal growth to measure fitness, we identified aspects of ramet morphology targeted by selection in two contrasting light environments and compared patterns of selection across environments. We found that directional selection on single traits, coupled with linear and nonlinear selection on multi-trait interactions, shape ramet morphology within environments and favor different phenotypes in each. Evidence of environment-dependent, multivariate selection on correlated traits is novel for any marine modular organism and demonstrates that seaweeds, such as A. armata, may potentially adapt to environmental heterogeneity via plasticity in clonal morphology.     

Differentiation in populations of Hordeum spontaneum Koch along a gradient of environmental productivity and predictability: plasticity in response to water and nutrient stress

Plants from four populations of Hordeum spontaneum originating in distinct environments of Israel were compared for stress induced phenotypic plasticity. The environments ranged along a gradient of increasing rainfall amount and predictability from low (desert) to moderate (semisteppe batha) to high (Mediterranean grassland and mountain, the latter also experiencing frost stress). The plants were exposed to a set of four treatments: no stress (optimum water and nutrients), water, nutrient and both water and nutrient stress. Plants from the four populations (or ecotypes) exhibited different patterns of plasticity in response to the different stresses (water and nutrients) and in different trait categories (reproductive, fitness and resource allocation). The importance of plasticity in response to water stress appears to decrease, and to nutrient stress appears to increase along the increasing rainfall gradient. The mountain ecotype, growing in an area with high potential productivity (amount of rainfall) but experiencing periodic frosts, was the most plastic among ecotypes in resource allocation under both water and nutrient stress, but exhibited low plasticity in other trait categories. In contrast, the desert ecotype had low plasticity in resource allocation under water stress and the lowest plasticity among the four ecotypes in all trait categories in response to nutrient stress. The ecotype originating in Mediterranean grassland, a predictable and most favourable environment, was highly plastic in fitness and allocation traits in response to low nutrient levels which is likely to occur due to competition in productive environment. We discuss the observed differences in ecotype plasticity as part of their environmentally induced adaptive ‘strategies’. We found no support for the hypothesis that plants originating in environments with greater variation and unpredictability are more plastic. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society 2002, 75 , 301–312.     

季节性雪被变化对森林凋落物分解及土壤氮动态的影响

全球气候变化引发的雪被格局变化将深刻影响植被的凋落物分解、陆地生态系统的土壤养分循环等过程.森林是陆地生态系统的主体,在全球生物地球化学循环中起着不可替代的作用.本研究综述了季节性雪被变化对森林凋落物分解及土壤氮动态的影响.全球气候变化情景下季节性雪被表现出因地域而异的增加或减少的变化格局,一方面通过改变环境温湿度、凋落物质量、分解者动态等直接影响分解过程,另一方面通过改变森林群落结构、植被物候、土壤养分等间接地作用于凋落物分解.同时,季节性雪被通过影响氮富集作用、雪被下土壤温湿度、冻融循环、森林群落、雪下动物和微生物等相关因子而改变森林土壤氮循环.本领域未来应开展的研究是: 1) 全面考虑全球气候变化情景下季节性雪被格局的变异性,开展不同季节性雪被格局变化的模拟研究;2) 开展季节性雪被融雪水淋溶作用对森林凋落物分解和土壤氮动态的影响研究;3) 阐明不同生态系统和气候带中季节性雪被格局变化对森林凋落物分解过程和土壤氮动态的驱动机制研究;4) 量化季节性雪被变化对森林凋落物分解和土壤氮动态在雪被覆盖期的瞬时影响和无雪期的延续影响,为阐明和模型预测陆地生态系统生物地球化学循环对全球气候变化的响应提供理论基础和数据支持.     

The Plastic Growth Responses of Three Agamospecies of Dandelion to Two Levels of Nutrient

The ability of a genotype to produce different phenotypes inresponse to variable environments is a crucial aspect of lifehistory strategies as it determines the shape of the fitnessset for the population. Apomictic dandelions generate littlegenetic variation between parent and offspring and plasticityis the main strategy in the face of environmental variability.The plastic response of three coexisiting dandelions has beenmeasured over two nutrient regimes. Cyclical growth patternsare species specific and in some cases independent of nutrientlevels. Differences between the agamospecies are greater athigh nutrient levels and the agamospecies appear to produceonly one phenotype at low nutrient levels. Taraxacum, plasticity, phenotypes, nutrient level     

Ecological limits to plant phenotypic plasticity

Phenotypic plasticity is considered the major means by which plants cope with environmental heterogeneity. Although ubiquitous in nature, actual phenotypic plasticity is far from being maximal. This has been explained by the existence of internal limits to its expression. However, phenotypic plasticity takes place within an ecological context and plants are generally exposed to multifactor environments and to simultaneous interactions with many species. These external, ecological factors may limit phenotypic plasticity or curtail its adaptive value, but seldom have they been considered because limits to plasticity have typically addressed factors internal to the plant. We show that plastic responses to abiotic factors are reduced under situations of conservative resource use in stressful and unpredictable habitats, and that extreme levels in a given abiotic factor can negatively influence plastic responses to another factor. We illustrate how herbivory may limit plant phenotypic plasticity because damaged plants can only rarely attain the optimal phenotype in the challenging environment. Finally, it is examined how phenotypic changes involved in trait-mediated interactions can entail costs for the plant in further interactions with other species in the community. Ecological limits to plasticity must be included in any realistic approach to understand the evolution of plasticity in complex environments and to predict plant responses to global change.     

Metapopulation structure favors plasticity over local adaptation

We describe a model for the evolutionary consequences of plasticity in an environmentally heterogeneous metapopulation in which specialists for each of two alternative environments and one plastic type are initially present. The model is similar to that proposed by Moran (1992) but extends her work to two sites. We show that with migration between sites the plastic type is favored over local specialists across a broad range of parameter space. The plastic type may dominate or be fixed even in an environmentally uniform site, and even if the plasticity has imperfect accuracy or bears some cost such that a local specialist has higher fitness in that site, as long as there is some migration between sites with different distributions of environmental states. These results suggest that differences among taxa in dispersal and hence realized migration rates may play a heretofore unrecognized role in their patterns of adaptive population differentiation. Migration relaxes the thresholds for both environmental heterogeneity and accuracy of plastic response above which plasticity is favored. Furthermore, small changes in response accuracy can dramatically and abruptly alter the evolutionary outcome in the metapopulation. A fitness cost to plasticity will substantially reduce the range of conditions in which the plastic type will prevail only if the cost is both large and global rather than environment specific.