Evolution of phenotypic plasticity: Genetic differentiation and additive genetic variation for induced plant defence in wild arugula Eruca sativa

Phenotypic plasticity is the primary mechanism of organismal resilience to abiotic and biotic stress, and genetic differentiation in plasticity can evolve if stresses differ among populations. Inducible defence is a common form of adaptive phenotypic plasticity, and long‐standing theory predicts that its evolution is shaped by costs of the defensive traits, costs of plasticity and a trade‐off in allocation to constitutive versus induced traits. We used a common garden to study the evolution of defence in two native populations of wild arugula Eruca sativa (Brassicaceae) from contrasting desert and Mediterranean habitats that differ in attack by caterpillars and aphids. We report genetic differentiation and additive genetic variance for phenology, growth and three defensive traits (toxic glucosinolates, anti‐nutritive protease inhibitors and physical trichome barriers) as well their inducibility in response to the plant hormone jasmonic acid. The two populations were strongly differentiated for plasticity in nearly all traits. There was little evidence for costs of defence or plasticity, but constitutive and induced traits showed a consistent additive genetic trade‐off within each population for the three defensive traits. We conclude that these populations have evolutionarily diverged in inducible defence and retain ample potential for the future evolution of phenotypic plasticity in defence.     

On the fate of seasonally plastic traits in a rainforest butterfly under relaxed selection

Many organisms display phenotypic plasticity as adaptation to seasonal environmental fluctuations. Often, such seasonal responses entails plasticity of a whole suite of morphological and life‐history traits that together contribute to the adaptive phenotypes in the alternative environments. While phenotypic plasticity in general is a well‐studied phenomenon, little is known about the evolutionary fate of plastic responses if natural selection on plasticity is relaxed. Here, we study whether the presumed ancestral seasonal plasticity of the rainforest butterfly Bicyclus sanaos (Fabricius, 1793) is still retained despite the fact that this species inhabits an environmentally stable habitat. Being exposed to an atypical range of temperatures in the laboratory revealed hidden reaction norms for several traits, including wing pattern. In contrast, reproductive body allocation has lost the plastic response. In the savannah butterfly, B. anynana (Butler, 1879), these traits show strong developmental plasticity as an adaptation to the contrasting environments of its seasonal habitat and they are coordinated via a common developmental hormonal system. Our results for B. sanaos indicate that such integration of plastic traits – as a result of past selection on expressing a coordinated environmental response – can be broken when the optimal reaction norms for those traits diverge in a new environment.     

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.     

Role of phenotypic plasticity and population differentiation in adaptation to novel environmental conditions

Species can adapt to new environmental conditions either through individual phenotypic plasticity, intraspecific genetic differentiation in adaptive traits, or both. Wild emmer wheat, Triticum dicoccoides, an annual grass with major distribution in Eastern Mediterranean region, is predicted to experience in the near future, as a result of global climate change, conditions more arid than in any part of the current species distribution. To understand the role of the above two means of adaptation, and the effect of population range position, we analyzed reaction norms, extent of plasticity, and phenotypic selection across two experimental environments of high and low water availability in two core and two peripheral populations of this species. We studied 12 quantitative traits, but focused primarily on the onset of reproduction and maternal investment, which are traits that are closely related to fitness and presumably involved in local adaptation in the studied species. We hypothesized that the population showing superior performance under novel environmental conditions will either be genetically differentiated in quantitative traits or exhibit higher phenotypic plasticity than the less successful populations. We found the core population K to be the most plastic in all three trait categories (phenology, reproductive traits, and fitness) and most successful among populations studied, in both experimental environments; at the same time, the core K population was clearly genetically differentiated from the two edge populations. Our results suggest that (1) two means of successful adaptation to new environmental conditions, phenotypic plasticity and adaptive genetic differentiation, are not mutually exclusive ways of achieving high adaptive ability; and (2) colonists from some core populations can be more successful in establishing beyond the current species range than colonists from the range extreme periphery with conditions seemingly closest to those in the new environment.     

Estimation of fitness from energetics and life‐history data: An example using mussels

Changing environments have the potential to alter the fitness of organisms through effects on components of fitness such as energy acquisition, metabolic cost, growth rate, survivorship, and reproductive output. Organisms, on the other hand, can alter aspects of their physiology and life histories through phenotypic plasticity as well as through genetic change in populations (selection). Researchers examining the effects of environmental variables frequently concentrate on individual components of fitness, although methods exist to combine these into a population level estimate of average fitness, as the per capita rate of population growth for a set of identical individuals with a particular set of traits. Recent advances in energetic modeling have provided excellent data on energy intake and costs leading to growth, reproduction, and other life‐history parameters; these in turn have consequences for survivorship at all life‐history stages, and thus for fitness. Components of fitness alone (performance measures) are useful in determining organism response to changing conditions, but are often not good predictors of fitness; they can differ in both form and magnitude, as demonstrated in our model. Here, we combine an energetics model for growth and allocation with a matrix model that calculates population growth rate for a group of individuals with a particular set of traits. We use intertidal mussels as an example, because data exist for some of the important energetic and life‐history parameters, and because there is a hypothesized energetic trade‐off between byssus production (affecting survivorship), and energy used for growth and reproduction. The model shows exactly how strong this trade‐off is in terms of overall fitness, and it illustrates conditions where fitness components are good predictors of actual fitness, and cases where they are not. In addition, the model is used to examine the effects of environmental change on this trade‐off and on both fitness and on individual fitness components.     

Changes in reproductive investment with altitude in an alpine plant

Aims In perennial species, the allocation of resources to reproduction results in a reduction of allocation to vegetative growth and, therefore, impacts future reproductive success. As a consequence, variation in this trade-off is among the most important driving forces in the life-history evolution of perennial plants and can lead to locally adapted genotypes. In addition to genetic variation, phenotypic plasticity might also contribute to local adaptation of plants to local conditions by mediating changes in reproductive allocation. Knowledge on the importance of genetic and environmental effects on the trade-off between reproduction and vegetative growth is therefore essential to understand how plants may respond to environmental changes.Methods We conducted a transplant experiment along an altitudinal gradient from 425 to 1?921 m in the front range of the Western Alps of Switzerland to assess the influence of both altitudinal origin of populations and altitude of growing site on growth, reproductive investment and local adaptation in Poa alpina .Important findings In our study, the investment in reproduction increased with plant size. Plant growth and the relative importance of reproductive investment decreased in populations originating from higher altitudes compared to populations originating from lower altitudes. The changes in reproductive investment were mainly explained by differences in plant size. In contrast to genetic effects, phenotypic plasticity of all traits measured was low and not related to altitude. As a result, the population from the lowest altitude of origin performed best at all sites. Our results indicate that in P. alpina genetic differences in growth and reproductive investment are related to local conditions affecting growth, i.e. interspecific competition and soil moisture content.     

To grow or to seed: ecotypic variation in reproductive allocation and cone production by young female Aleppo pine (Pinus halepensis, Pinaceae)

Age and size at the first reproduction and the reproductive allocation of plants are linked to different life history strategies. Aleppo pine only reproduces through seed, and, as such, early female reproduction confers high fitness in its infertile highly fire-prone habitats along the Mediterranean coast because life expectancy is short. We investigated the extent of ecotypic differentiation in female reproductive allocation and examined the relation between early female reproduction and vegetative growth. In a common-garden experiment, the threshold age and size at first female reproduction and female reproductive allocation at age seven differed significantly among Aleppo pine provenances of ecologically distinct origin. Significant correlations among reproductive features of the provenances and the ecological traits of origin were found using different analytical tools. In nonlinear models of cone counts vs. stem volume, medium-sized trees (not the largest trees) produced the highest cone yield, confirming that, at the individual level, early female reproduction is incompatible with fast vegetative growth. The contribution of founder effects and adaptation to contrasting fire regimes may be confounding factors. But considering all traits analyzed, the geographical patterns of resource allocation by Aleppo pine suggest ecotypic specialization for either resource-poor (favoring early reproduction) or resource-rich (favoring vegetative growth) habitats.     

Differences in developmental strategies between long‐settled and invasion‐front populations of the cane toad in Australia

Phenotypic plasticity can enhance a species’ ability to persist in a new and stressful environment, so that reaction norms are expected to evolve as organisms encounter novel environments. Biological invasions provide a robust system to investigate such changes. We measured the rates of early growth and development in tadpoles of invasive cane toads (Rhinella marina) in Australia, from a range of locations and at different larval densities. Populations in long‐colonized areas have had the opportunity to adapt to local conditions, whereas at the expanding range edge, the invader is likely to encounter challenges that are both novel and unpredictable. We thus expected invasion‐vanguard populations to exhibit less phenotypic plasticity than range‐core populations. Compared to clutches from long‐colonized areas, clutches from the invasion front were indeed less plastic (i.e. rates of larval growth and development were less sensitive to density). In contrast, those rates were highly variable in clutches from the invasion front, even among siblings from the same clutch under standard conditions. Clutches with highly variable rates of growth and development under constant conditions had lower phenotypic plasticity, suggesting a trade‐off between these two strategies. Although these results reveal a strong pattern, further investigation is needed to determine whether these different developmental strategies are adaptive (i.e. adaptive phenotypic plasticity vs. bet‐hedging) or instead are driven by geographic variation in genetic quality or parental effects.     

Are plasticity in functional traits and constancy in performance traits linked with invasiveness? An experimental test comparing invasive and naturalized plant species

The role of phenotypic plasticity in plant invasions is among the most often discussed relationships in invasion ecology. However, despite the large number of studies on this topic, there is little consistency. Reconsideration of the role of plasticity by distinguishing two substantially distinct trait-groups, performance traits (contributing directly to fitness) and functional traits (influencing fitness indirectly), could form a more operative framework for comparative studies. In the current study we expect that invasive species benefit from being plastic in functional traits, which allows them to maintain a more constant performance across different environmental conditions compared to non-invasive alien species. We compared invasive and naturalized non-invasive alien plant species by their germination (20 species), their vegetative (10 species) and their reproductive (four species) responses to three different levels of water, light and nutrient availability in a common garden experiment. Used traits were classified into performance (germination ratio, total biomass, seed number) and functional traits (time to germination, root:shoot ratio, specific leaf area, reproductive allocation). We found that invasive and non-invasive species responded similarly to environmental factors, except for example that invasive species germinated earlier with decreasing light conditions or, surprisingly, non-invasive species reacted more intensely to increased nitrogen availability by having a superior ability to achieve greater biomass. The two groups were equally plastic in all the germination and vegetative traits measured but the reproductive traits, since higher plasticity in relative reproductive allocation and higher constancy in reproductive performance showed a pronounced relation with invasiveness.     

Trade‐offs between life‐history traits at range‐edge and central locations

The allocation of resources to different life‐history traits should represent the best compromise in fitness investment for organisms in their local environment. When resources are limiting, the investment in a specific trait must carry a cost that is expressed in trade‐offs with other traits. In this study, the relative investment in the fitness‐related traits, growth, reproduction and defence were compared at central and range‐edge locations, using the seaweed Ascophyllum nodosum as a model system. Individual growth rates were similar at both sites, whereas edge populations showed a higher relative investment in reproduction (demonstrated by a higher reproductive allocation and extended reproductive periods) when compared to central populations that invested more in defence. These results show the capability of A. nodosum to differentially allocate resources for different traits under different habitat conditions, suggesting that reproduction and defence have different fitness values under the specific living conditions experienced at edge and central locations. However, ongoing climate change may threaten edge populations by increasing the selective pressure on specific traits, forcing these populations to lower the investment in other traits that are also potentially important for population fitness.     

Stress promotes changes in resource allocation to growth and reproduction in a simultaneous hermaphrodite with indeterminate growth

In iteroparous animals, investment in growth is compromised by investment in reproduction, especially in species with indeterminate growth. Life‐history theory predicts that growth should be favoured over reproduction, assuming size‐related fecundity or survival. Hence, increase body condition represents an increase in reproductive potential. Simultaneous hermaphrodites should adjust their resource allocation to each sex function in response to current conditions but, recently, it has been suggested that, in hermaphrodites, gender allocation should be considered as a three‐way trade‐off, including the investment in somatic growth. Due to the higher costs involved, the female function is affected to a greater extent by environmentally stressful conditions rather than the male function. To examine this, we induced stress in the hermaphroditic earthworm Eisenia fetida (Savigny, 1826) and looked for changes in resource allocation in nonreproductive and reproductive individuals. Experimental stress was induced by using tweezers to elicit contractile escape movements. We predicted that stressed earthworms would preferentially allocate resources to growth. In nonreproductive individuals, however, stress had a negative effect on growth, although weight recovery was rapid once manipulation ceased, indicating the importance of body condition, as well as the existence of mechanisms of compensatory growth for growth trajectories in this earthworm species. The response of reproductive individuals was consistent with our expectation: (1) stressed worms maintained their growth rate at the expense of current reproduction and (2) stressed earthworms laid 25% fewer cocoons, which were 30% lighter than cocoons laid by control earthworms. The present results suggest that E. fetida regulates its reproductive effort and that future reproduction has more impact on its fitness than current reproduction. The trade‐off between current and future reproduction should be taken into consideration in models of sex allocation in simultaneous hermaphrodites. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 593–600.     

Cross-generational environmental effects and the evolution of offspring size in the Trinidadian guppy Poecilia reticulata

Abstract The existence of adaptive phenotypic plasticity demands that we study the evolution of reaction norms, rather than just the evolution of fixed traits. This approach requires the examination of functional relationships among traits not only in a single environment but across environments and between traits and plasticity itself. In this study, I examined the interplay of plasticity and local adaptation of offspring size in the Trinidadian guppy, Poecilia reticulata. Guppies respond to food restriction by growing and reproducing less but also by producing larger offspring. This plastic difference in offspring size is of the same order of magnitude as evolved genetic differences among populations. Larger offspring sizes are thought to have evolved as an adaptation to the competitive environment faced by newborn guppies in some environments. If plastic responses to maternal food limitation can achieve the same fitness benefit, then why has guppy offspring size evolved at all? To explore this question, I examined the plastic response to food level of females from two natural populations that experience different selective environments. My goals were to examine whether the plastic responses to food level varied between populations, test the consequences of maternal manipulation of offspring size for offspring fitness, and assess whether costs of plasticity exist that could account for the evolution of mean offspring size across populations. In each population, full‐sib sisters were exposed to either a low‐ or high‐food treatment. Females from both populations produced larger, leaner offspring in response to food limitation. However, the population that was thought to have a history of selection for larger offspring was less plastic in its investment per offspring in response to maternal mass, maternal food level, and fecundity than the population under selection for small offspring size. To test the consequences of maternal manipulation of offspring size for offspring fitness, I raised the offspring of low‐ and high‐food mothers in either low‐ or high‐food environments. No maternal effects were detected at high food levels, supporting the prediction that mothers should increase fecundity rather than offspring size in noncompetitive environments. For offspring raised under low food levels, maternal effects on juvenile size and male size at maturity varied significantly between populations, reflecting their initial differences in maternal manipulation of offspring size; nevertheless, in both populations, increased investment per offspring increased offspring fitness. Several correlates of plasticity in investment per offspring that could affect the evolution of offspring size in guppies were identified. Under low‐food conditions, mothers from more plastic families invested more in future reproduction and less in their own soma. Similarly, offspring from more plastic families were smaller as juveniles and female offspring reproduced earlier. These correlations suggest that a fixed, high level of investment per offspring might be favored over a plastic response in a chronically low‐resource environment or in an environment that selects for lower reproductive effort     

Water availability and population origin affect the expression of the tradeoff between reproduction and growth in Plantago coronopus

Investment in reproduction and growth represent a classic tradeoff with implication for life history evolution. The local environment can play a major role in the magnitude and evolutionary consequences of such a tradeoff. Here, we examined the investment in reproductive and vegetative tissue in 40 maternal half‐sib families from four different populations of the herb Plantago coronopus growing in either a dry or wet greenhouse environment. Plants originated from populations with an annual or a perennial life form, with annuals prevailing in drier habitats with greater seasonal variation in both temperature and precipitation. We found that water availability affected the expression of the tradeoff (both phenotypic and genetic) between reproduction and growth, being most accentuated under dry condition. However, populations responded very differently to water treatments. Plants from annual populations showed a similar response to drought condition with little variation among maternal families, suggesting a history of selection favouring genotypes with high allocation to reproduction when water availability is low. Plants from annual populations also expressed the highest level of plasticity. For the perennial populations, one showed a large variation among maternal families in resource allocation and expressed significant negative genetic correlations between reproductive and vegetative biomass under drought. The other perennial population showed less variation in response to treatment and had trait values similar to those of the annuals, although it was significantly less plastic. We stress the importance of considering intraspecific variation in response to environmental change such as drought, as conspecific plants exhibited very different abilities and strategies to respond to high versus low water availability even among geographically close populations.     

Effects of four generations of density-dependent selection on life history traits and their plasticity in a clonally propagated plant

Life history evolution of many clonal plants takes place with long periods of exclusively clonal reproduction and under largely varying ramet densities resulting from clonal reproduction. We asked whether life history traits of the clonal herb Ranunculus reptans respond to density-dependent selection, and whether plasticity in these traits is adaptive. After four generations of exclusively clonal propagation of 16 low and 16 high ramet-density lines, we studied life history traits and their plasticities at two test ramet-densities. Plastic responses to higher test-density consisted of a shift from sexual to vegetative reproduction, and reduced flower production, plant size, branching frequency, and lengths of leaves and internodes. Plants of high-density lines tended to have longer leaves, and under high test-density branched less frequently than those of low-density lines. Directions of these selection responses indicate that the observed plastic branching response is adaptive, whereas the plastic leaf length response is not. The reverse branching frequency pattern at low test-density, where plants of high-density lines branched more frequently than those of low-density lines, indicates evolution of plasticity in branching. Moreover, when grown under less stressful low test-density, plants of high-density lines tended to grow larger than the ones of low-density lines. We conclude that ramet density affects clonal life-history evolution and that under exclusively clonal propagation clonal life-history traits and their plasticities evolve differently at different ramet densities.     

Phenotypic plasticity in vegetative and reproductive traits in an invasive weed, Lythrum salicaria (Lythraceae), in response to soil moisture

In colonizing species, high phenotypic plasticity can contribute to survival and propagation in heterogenous adventive environments, and it has been suggested as a predictor of invasiveness. Observation of natural populations of an invasive species, Lythrum salicaria salicaria, indicated extensive variation in its growth and reproductive traits. Phenotypic plasticity of different life history traits of L. salicaria was investigated using vegetative clones of each of 12 genotypes from one population in Ontario, Canada. We chose soil moisture as the treatment factor because of its importance in wetland species and raised all 12 genotypes in each of four soil moisture treatments. We examined an array of vegetative and reproductive traits, including root and shoot mass, shoot and inflorescence length, total seed set, floral mass, and morphometric variables. All observed vegetative as well as reproductive traits demonstrated significant phenotypic plasticity in response to soil moisture treatment. Even the stigma-anther separation involved significant genotype by environment interactions, suggesting that soil moisture may modify the relative positions of anthers and stigma. Compared to vegetative traits, most reproductive traits demonstrated crossing reaction norms, implying that the average differences in those traits among genotypes vary with the environment maintaining the genetic variation in a population.     

Life history consequences of developing in anthropogenic noise

When environments change rapidly, adaptive phenotypic plasticity can ameliorate negative effects of environmental change on survival and reproduction. Recent evidence suggests, however, that plastic responses to human‐induced environmental change are often maladaptive or insufficient to overcome novel selection pressures. Anthropogenic noise is a ubiquitous and expanding disturbance with demonstrated effects on fitness‐related traits of animals like stress responses, foraging, vigilance, and pairing success. Elucidating the lifetime fitness effects of noise has been challenging because longer‐lived vertebrate systems are typically studied in this context. Here, we follow noise‐stressed invertebrates throughout their lives, assessing a comprehensive suite of life history traits, and ultimately, lifetime number of surviving offspring. We reared field crickets, Teleogryllus oceanicus, in masking traffic noise, traffic noise from which we removed frequencies that spectrally overlap with the crickets’ mate location song and peak hearing (nonmasking), or silence. We found that exposure to masking noise delayed maturity and reduced adult lifespan; crickets exposed to masking noise spent 23% more time in juvenile stages and 13% less time as reproductive adults than those exposed to no traffic noise. Chronic lifetime exposure to noise, however, did not affect lifetime reproductive output (number of eggs or surviving offspring), perhaps because mating provided females a substantial longevity benefit. Nevertheless, these results are concerning as they highlight multiple ways in which traffic noise may reduce invertebrate fitness. We encourage researchers to consider effects of anthropogenic disturbance on growth, survival, and reproductive traits simultaneously because changes in these traits may amplify or nullify one another.     

Life history variation in an annual plant under two opposing environmental constraints along an aridity gradient

Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species.     

Interpreting reproductive allometry: Individual strategies of allocation explain size-dependent reproduction in plant populations

Size-dependent or allometric relationships between reproductive and vegetative size are extremely common in plant populations. Reproductive allometry where plant size differences are due to environmental variability has been interpreted both as an adaptive strategy of plant growth and allocation, and as the product of fixed developmental constraints. Patterns of development are crucial in defining reproductive allometry but development is not fixed across individuals. For example, environmental adversity (e.g. resource impoverishment) tends to favor reproduction at relatively small sizes – an adaptive response to environmental adversity. While small individuals may have lower reproductive output than large individuals, all plants should maximize their reproductive output and relative allocation to reproduction may be constant across sizes. Thus, where individual plants within a population initiate reproduction at different sizes, no significant reproductive allometry is an appropriate null expectation. Reproductive allometry occurs in plant populations where initiating reproduction at small sizes yields relatively high or low reproductive size at final development. Both of these outcomes are common in plant populations. Our interpretation of reproductive allometry combines previous adaptive and developmental constraint interpretations, and is the first to successfully explain the range of relationships in plant populations where relative allocation has been observed to increase, decrease or remain constant will increasing plant size.     

The ubiquity of phenotypic plasticity in plants: a synthesis

Adaptation to heterogeneous environments can occur via phenotypic plasticity, but how often this occurs is unknown. Reciprocal transplant studies provide a rich dataset to address this issue in plant populations because they allow for a determination of the prevalence of plastic versus canalized responses. From 31 reciprocal transplant studies, we quantified the frequency of five possible evolutionary patterns: (1) canalized response–no differentiation: no plasticity, the mean phenotypes of the populations are not different; (2) canalized response–population differentiation: no plasticity, the mean phenotypes of the populations are different; (3) perfect adaptive plasticity: plastic responses with similar reaction norms between populations; (4) adaptive plasticity: plastic responses with parallel, but not congruent reaction norms between populations; and (5) nonadaptive plasticity: plastic responses with differences in the slope of the reaction norms. The analysis included 362 records: 50.8% life‐history traits, 43.6% morphological traits, and 5.5% physiological traits. Across all traits, 52% of the trait records were not plastic, and either showed no difference in means across sites (17%) or differed among sites (83%). Among the 48% of trait records that showed some sort of plasticity, 49.4% showed perfect adaptive plasticity, 19.5% adaptive plasticity, and 31% nonadaptive plasticity. These results suggest that canalized responses are more common than adaptive plasticity as an evolutionary response to environmental heterogeneity.     

Costs of inducible defence along a resource gradient

In addition to having constitutive defence traits, many organisms also respond to predation by phenotypic plasticity. In order for plasticity to be adaptive, induced defences should incur a benefit to the organism in, for example, decreased risk of predation. However, the production of defence traits may include costs in fitness components such as growth, time to reproduction, or fecundity. To test the hypothesis that the expression of phenotypic plasticity incurs costs, we performed a common garden experiment with a freshwater snail, Radix balthica, a species known to change morphology in the presence of molluscivorous fish. We measured a number of predator-induced morphological and behavioural defence traits in snails that we reared in the presence or absence of chemical cues from fish. Further, we quantified the costs of plasticity in fitness characters related to fecundity and growth. Since plastic responses may be inhibited under limited resource conditions, we reared snails in different densities and thereby levels of competition. Snails exposed to predator cues grew rounder and thicker shells, traits confirmed to be adaptive in environments with fish. Defence traits were consistently expressed independent of density, suggesting strong selection from predatory molluscivorous fish. However, the expression of defence traits resulted in reduced growth rate and fecundity, particularly with limited resources. Our results suggest full defence in predator related traits regardless of resource availability, and costs of defence consequently paid in traits related to fitness.