During the past decade, a substantial and rapidly expanding literature has documented that a dive... more During the past decade, a substantial and rapidly expanding literature has documented that a diversity of hymenopterans have the ability to recognize their conspecific nestmates (22). As recognition research enters its second generation, investigators are turning their attention to elucidating the mechanisms of nestmate recognition and how the features of these mechanisms, as well as recognition ability itself, relate to an animal's sociobiology. Such studies are important not only because of their relevance to an understanding of the evolution and ecology of hymenopteran sociality, but also because of their potential application for control of economically important social insects. In our review, we summarize the evidence for nestmate recognition ability in social wasps, and examine in detail the mechanism of female-female nestmate recognition using primitively eusocial wasps (Polistes) as a model. We also compare the Polistes mechanism with mechanisms proposed for other social Hymenoptera and ,with the major theoretical models of kin recognition developed for animals in general. This is followed by a discussion of the possible adaptiveness of both female-female nestmate recognition ability and the specific mechanisms underlying this ability. Finally, we explore the preadaptations
Biological Journal of The Linnean Society, Sep 9, 2020
An individual's early-life environment and phenotype often influence its traits and performance a... more An individual's early-life environment and phenotype often influence its traits and performance as an adult. We investigated whether such 'carryover effects' are associated with alternative, environmentally-induced phenotypes ('polyphenism'), and, if so, whether they influence the evolution of polyphenism. To do so, we studied Mexican spadefoot toads, Spea multiplicata, which have evolved a polyphenism consisting of two, dramatically different forms: a carnivore morph and an omnivore morph. We sampled both morphs from a fast-drying and a slow-drying pond and reared them to sexual maturity. Larval environment (pond) strongly influenced survival as well as age and size at metamorphosis and sexual maturity; i.e. environment-dependent carryover effects were present. By contrast, larval phenotype (morph) did not affect life-history traits at sexual maturity; i.e. phenotype-dependent carryover effects were absent. These results are consistent with theory, which suggests that by amplifying selective trade-offs in heterogenous environments, environment-dependent carryover effects might foster the evolution of polyphenism. At the same time, by freeing selection to refine a novel phenotype without altering the existing form, the absence of phenotype-dependent carryover effects might enable polyphenism to evolve in the first place. Generally, carryover effects might play an underappreciated role in the evolution of polyphenism.
Biological Journal of The Linnean Society, Aug 1, 2020
Intraspecific competition has long been considered a key driver of evolutionary diversification, ... more Intraspecific competition has long been considered a key driver of evolutionary diversification, but whether it can also promote evolutionary innovation is less clear. Here we examined the interplay between competition and phenotypic plasticity in fuelling the origins of a novel, complex phenotype-a distinctive carnivore morph found in spadefoot toad tadpoles (genus Spea) that specializes on fairy shrimp. We specifically sought to explore the possible origins of this phenotype by providing shrimp to Scaphiopus holbrookii tadpoles (the sister genus to Spea that does not produce carnivores) while subjecting them to competition for their standard diet of detritus. Previous research had shown that this species will eat shrimp when detritus is limited, and that these shrimp-fed individuals produce features that are redolent of a rudimentary Spea carnivore. In this study, we found that: (1) behavioural and morphological plasticity enabled some individuals to expand their diet to include shrimp; (2) there was heritable variation in this plasticity; and (3) individuals received a growth and development benefit by eating shrimp. Thus, novel resource use can arise via plasticity as an adaptive response to intraspecific competition. More generally, our results show how competition and plasticity may interact to pave the way for the evolution of complex, novel phenotypes, such as the distinctive carnivore morph in present-day Spea.
Identifying the causes of diversification is central to evolutionary biology. The ecological theo... more Identifying the causes of diversification is central to evolutionary biology. The ecological theory of adaptive diversification holds that the evolution of phenotypic differences between populations and species-and the formation of new species-stems from divergent natural selection, often arising from competitive interactions. Although increasing evidence suggests that phenotypic plasticity can facilitate this process, it is not generally appreciated that competitively mediated selection often also provides ideal conditions for phenotypic plasticity to evolve in the first place. Here, we discuss how competition plays at least two key roles in adaptive diversification depending on its pattern. First, heterogenous competition initially generates heterogeneity in resource use that favors adaptive plasticity in the form of "inducible competitors". Second, once such competitively induced plasticity evolves, its capacity to rapidly generate phenotypic variation and expose phenotypes to alternate selective regimes allows populations to respond readily to selection favoring diversification, as may occur when competition generates steady diversifying selection that permanently drives the evolutionary divergence of populations that use different resources. Thus, competition plays two important roles in adaptive diversification-one well-known and the other only now emerging-mediated through its effect on the evolution of phenotypic plasticity [Current Zoology 59 (4): 537-552, 2013].
Intraspecific variation in resource-use traits can have profound ecological and evolutionary impl... more Intraspecific variation in resource-use traits can have profound ecological and evolutionary implications. Among the most striking examples are resource polymorphisms, where alternative morphs that utilize different resources evolve within a population. An underappreciated aspect of their evolution is that the same conditions that favor resource polymorphism-competition and ecological opportunity-might foster additional rounds of diversification within already existing morphs. We examined these issues in spadefoot toad tadpoles that develop into either a generalist "omnivore" or a specialist "carnivore" morph. Specifically, we assessed the morphological diversity of tadpoles from natural ponds and experimentally induced carnivores reared on alternative diets. We also surveyed natural ponds to determine if the strength of intramorph competition and the diversity and abundance of dietary resources (measures of ecological opportunity) influenced the diversity of within-morph variation. We found that five omnivore and four carnivore types were present in natural ponds; alternative diets led to shape differences, some of which mirrored variation in the wild; and both competition and ecological opportunity were associated with enhanced morphological diversity in natural ponds. Such fine-scale intraspecific variation might represent an underappreciated form of biodiversity and might constitute a crucible of evolutionary innovation and diversification.
... DAVID W. PFENNIG2, ALAN MABRY3, AND DAVID ORANGE Department of Zoology, University of Texas, ... more ... DAVID W. PFENNIG2, ALAN MABRY3, AND DAVID ORANGE Department of Zoology, University of Texas, Austin, Texas 78712-1064 USA ... with complex life cycles, a widely held view is that a trade-off exists between these two factors (Wilbur and Collins 1973, Wassersug 1975 ...
When a population experiences severe stress from a changing environment, evolution by natural sel... more When a population experiences severe stress from a changing environment, evolution by natural selection can prevent its extinction, a process dubbed “evolutionary rescue.” However, evolution may be unable to track the sort of rapid environmental change being experienced by many modern‐day populations. A potential solution is for organisms to respond to environmental change through phenotypic plasticity, which can buffer populations against change and thereby buy time for evolutionary rescue. In this review, we examine whether this process extends to situations in which the environmentally induced response is passed to offspring. As we describe, theoretical and empirical studies suggest that such “transgenerational plasticity” can increase population persistence. We discuss the implications of this process for conservation biology, outline potential limitations, and describe some applications. Generally, transgenerational plasticity may be effective at buying time for evolutionary re...
Abstract A growing number of biologists have begun asking whether environmentally induced phenoty... more Abstract A growing number of biologists have begun asking whether environmentally induced phenotypic change—phenotypic plasticity—precedes and facilitates the origin and subsequent fixation of novel, complex phenotypes. However, such plasticity-led evolution (PLE) remains controversial. Here, we begin with a summary of the PLE hypothesis and describe how its key criteria can be evaluated in natural (or experimental) systems. We then present a case study using this approach in spadefoot toad tadpoles. We close by describing phenotypic plasticity’s relevance to macroevolutionary change. As we describe, environmentally induced traits might be a potent force during the evolution of novelty, diversity, and adaptation.
Artificial prey techniques—wherein synthetic replicas of real organisms are placed in natural hab... more Artificial prey techniques—wherein synthetic replicas of real organisms are placed in natural habitats—are widely used to study predation in the field. We investigated the extent to which videography could provide additional information to such studies. As a part of studies on aposematism and mimicry of coral snakes (Micrurus) and their mimics, observational data from 109 artificial snake prey were collected from video-recording camera traps in three locations in the Americas (terra firmeforest, Tiputini Biodiversity Station, Ecuador; premontane wet forest, Nahá Reserve, Mexico; longleaf pine forest, Southeastern Coastal Plain, North Carolina, USA). During 1,536 camera days, a total of 268 observations of 20 putative snake predator species were recorded in the vicinity of artificial prey. Predators were observed to detect artificial prey 52 times, but only 21 attacks were recorded. Mammals were the most commonly recorded group of predators near replicas (243) and were responsible fo...
Seminars in Cell & Developmental Biology, 2019
A growing number of biologists have begun asking whether environmentally induced phenotypic chang... more A growing number of biologists have begun asking whether environmentally induced phenotypic change-'phenotypic plasticity'-precedes and facilitates the origin and canalization of novel, complex phenotypes. However, such 'plasticity-first evolution' (PFE) remains controversial. Here, we summarize the PFE hypothesis and describe how it can be evaluated in natural systems. We then review the evidence for PFE from amphibians (a group in which phenotypic plasticity is especially widespread) and describe how phenotypic plasticity might have facilitated macroevolutionary change. Finally, we discuss what is known about the proximate mechanisms of PFE in amphibians. We close with suggestions for future research. As we describe, amphibians offer some of the best support for plasticity's role in the origin of evolutionary novelties.
Proceedings of the Royal Society B: Biological Sciences, 2019
Plasticity-led evolution occurs when a change in the environment triggers a change in phenotype v... more Plasticity-led evolution occurs when a change in the environment triggers a change in phenotype via phenotypic plasticity, and this pre-existing plasticity is subsequently refined by selection into an adaptive phenotype. A critical, but largely untested prediction of plasticity-led evolution (and evolution by natural selection generally) is that the rate and magnitude of evolutionary change should be positively associated with a phenotype's frequency of expression in a population. Essentially, the more often a phenotype is expressed and exposed to selection, the greater its opportunity for adaptive refinement. We tested this prediction by competing against each other spadefoot toad tadpoles from different natural populations that vary in how frequently they express a novel, environmentally induced carnivore ecomorph. As expected, laboratory-reared tadpoles whose parents were derived from populations that express the carnivore ecomorph more frequently were superior competitors fo...
The relationship between inbreeding and both conception and foaling rates in Standardbred horses ... more The relationship between inbreeding and both conception and foaling rates in Standardbred horses (trotters and pacers) was examined for 1194 breeding years. There was a statistically significant (P less than 0.05) trend for conception and foaling rates to decrease with increased inbreeding; however, this relationship accounted for less than 2 percent of the variation. Additionally, the relationship between reproductive performance and inbreeding was not consistent between trotters and pacers. For trotters (F = 0.103) there was a trend for an increase in conception and foaling rates with increased inbreeding, while for pacers (F = 0.074), reproductive performance decreased with increased inbreeding. Overall, inbreeding does not appear to be a significant factor influencing reproductive performance of Standardbred horses.
Phenotypic plasticity is ubiquitous and generally regarded as a key mechanism for enabling organi... more Phenotypic plasticity is ubiquitous and generally regarded as a key mechanism for enabling organisms to survive in the face of environmental change. Because no organism is infinitely or ideally plastic, theory suggests that there must be limits (for example, the lack of ability to produce an optimal trait) to the evolution of phenotypic plasticity, or that plasticity may have inherent significant costs. Yet numerous experimental studies have not detected widespread costs. Explicitly differentiating plasticity costs from phenotype costs, we re-evaluate fundamental questions of the limits to the evolution of plasticity and of generalists vs specialists. We advocate for the view that relaxed selection and variable selection intensities are likely more important constraints to the evolution of plasticity than the costs of plasticity. Some forms of plasticity, such as learning, may be inherently costly. In addition, we examine opportunities to offset costs of phenotypes through ontogeny, amelioration of phenotypic costs across environments, and the condition-dependent hypothesis. We propose avenues of further inquiry in the limits of plasticity using new and classic methods of ecological parameterization, phylogenetics and omics in the context of answering questions on the constraints of plasticity. Given plasticity's key role in coping with environmental change, approaches spanning the spectrum from applied to basic will greatly enrich our understanding of the evolution of plasticity and resolve our understanding of limits.
Ecological character displacement is considered crucial in promoting diversification, yet relativ... more Ecological character displacement is considered crucial in promoting diversification, yet relatively little is known of its underlying mechanisms. We examined whether evolutionary shifts in gene expression plasticity ('genetic accommodation') mediate character displacement in spadefoot toads. Where Spea bombifrons and S. multiplicata occur separately in allopatry (the ancestral condition), each produces alternative, diet-induced, larval ecomorphs: omnivores, which eat detritus, and carnivores, which specialize on shrimp. By contrast, where these two species occur together in sympatry (the derived condition), selection to minimize competition for detritus has caused S. bombifrons to become nearly fixed for producing only carnivores, suggesting that character displacement might have arisen through an extreme form of genetic accommodation ('genetic assimilation') in which plasticity is lost. Here, we asked whether we could infer a signature of this process in regulatory changes of specific genes. In particular, we investigated whether genes that are normally expressed more highly in one morph ('biased' genes) have evolved reduced plasticity in expression levels among S. bombifrons from sympatry compared to S. bombifrons from allopatry. We reared individuals from sympatry vs. allopatry on detritus or shrimp and measured the reaction norms of nine biased genes. Although different genes displayed different patterns of gene regulatory evolution, the combined gene expression profiles revealed that sympatric individuals had indeed lost the diet-induced gene expression plasticity present in allopatric individuals. Our data therefore provide one of the few examples from natural populations in which genetic accommodation/assimilation can be traced to regulatory changes of specific genes. Such genetic accommodation might mediate character displacement in many systems.
Parental effects are often considered an evolved response, in which parents transmit information ... more Parental effects are often considered an evolved response, in which parents transmit information about the environment to enhance offspring fitness. However, these effects need not be adaptive. Here, we provide a striking example by presenting evidence that overfeeding of adult Mexican spadefoot toads, Spea multiplicata, is associated with decreased offspring survival. After a temporary change to their standard feeding regimen, S. multiplicata in our captive colony developed a much higher body condition (i.e. body mass for a given body length) than those in the wild. We analysed data from three subsequent experiments and found that although the body condition of a father was positively correlated with tadpole survival, mothers with a higher condition had lower tadpole survival. Our study highlights how obesity can negatively impact future generations via maladaptive maternal effects. Such effects could be especially likely for animals living in variable environments (such as spadefo...
During the past decade, a substantial and rapidly expanding literature has documented that a dive... more During the past decade, a substantial and rapidly expanding literature has documented that a diversity of hymenopterans have the ability to recognize their conspecific nestmates (22). As recognition research enters its second generation, investigators are turning their attention to elucidating the mechanisms of nestmate recognition and how the features of these mechanisms, as well as recognition ability itself, relate to an animal's sociobiology. Such studies are important not only because of their relevance to an understanding of the evolution and ecology of hymenopteran sociality, but also because of their potential application for control of economically important social insects. In our review, we summarize the evidence for nestmate recognition ability in social wasps, and examine in detail the mechanism of female-female nestmate recognition using primitively eusocial wasps (Polistes) as a model. We also compare the Polistes mechanism with mechanisms proposed for other social Hymenoptera and ,with the major theoretical models of kin recognition developed for animals in general. This is followed by a discussion of the possible adaptiveness of both female-female nestmate recognition ability and the specific mechanisms underlying this ability. Finally, we explore the preadaptations
Biological Journal of The Linnean Society, Sep 9, 2020
An individual's early-life environment and phenotype often influence its traits and performance a... more An individual's early-life environment and phenotype often influence its traits and performance as an adult. We investigated whether such 'carryover effects' are associated with alternative, environmentally-induced phenotypes ('polyphenism'), and, if so, whether they influence the evolution of polyphenism. To do so, we studied Mexican spadefoot toads, Spea multiplicata, which have evolved a polyphenism consisting of two, dramatically different forms: a carnivore morph and an omnivore morph. We sampled both morphs from a fast-drying and a slow-drying pond and reared them to sexual maturity. Larval environment (pond) strongly influenced survival as well as age and size at metamorphosis and sexual maturity; i.e. environment-dependent carryover effects were present. By contrast, larval phenotype (morph) did not affect life-history traits at sexual maturity; i.e. phenotype-dependent carryover effects were absent. These results are consistent with theory, which suggests that by amplifying selective trade-offs in heterogenous environments, environment-dependent carryover effects might foster the evolution of polyphenism. At the same time, by freeing selection to refine a novel phenotype without altering the existing form, the absence of phenotype-dependent carryover effects might enable polyphenism to evolve in the first place. Generally, carryover effects might play an underappreciated role in the evolution of polyphenism.
Biological Journal of The Linnean Society, Aug 1, 2020
Intraspecific competition has long been considered a key driver of evolutionary diversification, ... more Intraspecific competition has long been considered a key driver of evolutionary diversification, but whether it can also promote evolutionary innovation is less clear. Here we examined the interplay between competition and phenotypic plasticity in fuelling the origins of a novel, complex phenotype-a distinctive carnivore morph found in spadefoot toad tadpoles (genus Spea) that specializes on fairy shrimp. We specifically sought to explore the possible origins of this phenotype by providing shrimp to Scaphiopus holbrookii tadpoles (the sister genus to Spea that does not produce carnivores) while subjecting them to competition for their standard diet of detritus. Previous research had shown that this species will eat shrimp when detritus is limited, and that these shrimp-fed individuals produce features that are redolent of a rudimentary Spea carnivore. In this study, we found that: (1) behavioural and morphological plasticity enabled some individuals to expand their diet to include shrimp; (2) there was heritable variation in this plasticity; and (3) individuals received a growth and development benefit by eating shrimp. Thus, novel resource use can arise via plasticity as an adaptive response to intraspecific competition. More generally, our results show how competition and plasticity may interact to pave the way for the evolution of complex, novel phenotypes, such as the distinctive carnivore morph in present-day Spea.
Identifying the causes of diversification is central to evolutionary biology. The ecological theo... more Identifying the causes of diversification is central to evolutionary biology. The ecological theory of adaptive diversification holds that the evolution of phenotypic differences between populations and species-and the formation of new species-stems from divergent natural selection, often arising from competitive interactions. Although increasing evidence suggests that phenotypic plasticity can facilitate this process, it is not generally appreciated that competitively mediated selection often also provides ideal conditions for phenotypic plasticity to evolve in the first place. Here, we discuss how competition plays at least two key roles in adaptive diversification depending on its pattern. First, heterogenous competition initially generates heterogeneity in resource use that favors adaptive plasticity in the form of "inducible competitors". Second, once such competitively induced plasticity evolves, its capacity to rapidly generate phenotypic variation and expose phenotypes to alternate selective regimes allows populations to respond readily to selection favoring diversification, as may occur when competition generates steady diversifying selection that permanently drives the evolutionary divergence of populations that use different resources. Thus, competition plays two important roles in adaptive diversification-one well-known and the other only now emerging-mediated through its effect on the evolution of phenotypic plasticity [Current Zoology 59 (4): 537-552, 2013].
Intraspecific variation in resource-use traits can have profound ecological and evolutionary impl... more Intraspecific variation in resource-use traits can have profound ecological and evolutionary implications. Among the most striking examples are resource polymorphisms, where alternative morphs that utilize different resources evolve within a population. An underappreciated aspect of their evolution is that the same conditions that favor resource polymorphism-competition and ecological opportunity-might foster additional rounds of diversification within already existing morphs. We examined these issues in spadefoot toad tadpoles that develop into either a generalist "omnivore" or a specialist "carnivore" morph. Specifically, we assessed the morphological diversity of tadpoles from natural ponds and experimentally induced carnivores reared on alternative diets. We also surveyed natural ponds to determine if the strength of intramorph competition and the diversity and abundance of dietary resources (measures of ecological opportunity) influenced the diversity of within-morph variation. We found that five omnivore and four carnivore types were present in natural ponds; alternative diets led to shape differences, some of which mirrored variation in the wild; and both competition and ecological opportunity were associated with enhanced morphological diversity in natural ponds. Such fine-scale intraspecific variation might represent an underappreciated form of biodiversity and might constitute a crucible of evolutionary innovation and diversification.
... DAVID W. PFENNIG2, ALAN MABRY3, AND DAVID ORANGE Department of Zoology, University of Texas, ... more ... DAVID W. PFENNIG2, ALAN MABRY3, AND DAVID ORANGE Department of Zoology, University of Texas, Austin, Texas 78712-1064 USA ... with complex life cycles, a widely held view is that a trade-off exists between these two factors (Wilbur and Collins 1973, Wassersug 1975 ...
When a population experiences severe stress from a changing environment, evolution by natural sel... more When a population experiences severe stress from a changing environment, evolution by natural selection can prevent its extinction, a process dubbed “evolutionary rescue.” However, evolution may be unable to track the sort of rapid environmental change being experienced by many modern‐day populations. A potential solution is for organisms to respond to environmental change through phenotypic plasticity, which can buffer populations against change and thereby buy time for evolutionary rescue. In this review, we examine whether this process extends to situations in which the environmentally induced response is passed to offspring. As we describe, theoretical and empirical studies suggest that such “transgenerational plasticity” can increase population persistence. We discuss the implications of this process for conservation biology, outline potential limitations, and describe some applications. Generally, transgenerational plasticity may be effective at buying time for evolutionary re...
Abstract A growing number of biologists have begun asking whether environmentally induced phenoty... more Abstract A growing number of biologists have begun asking whether environmentally induced phenotypic change—phenotypic plasticity—precedes and facilitates the origin and subsequent fixation of novel, complex phenotypes. However, such plasticity-led evolution (PLE) remains controversial. Here, we begin with a summary of the PLE hypothesis and describe how its key criteria can be evaluated in natural (or experimental) systems. We then present a case study using this approach in spadefoot toad tadpoles. We close by describing phenotypic plasticity’s relevance to macroevolutionary change. As we describe, environmentally induced traits might be a potent force during the evolution of novelty, diversity, and adaptation.
Artificial prey techniques—wherein synthetic replicas of real organisms are placed in natural hab... more Artificial prey techniques—wherein synthetic replicas of real organisms are placed in natural habitats—are widely used to study predation in the field. We investigated the extent to which videography could provide additional information to such studies. As a part of studies on aposematism and mimicry of coral snakes (Micrurus) and their mimics, observational data from 109 artificial snake prey were collected from video-recording camera traps in three locations in the Americas (terra firmeforest, Tiputini Biodiversity Station, Ecuador; premontane wet forest, Nahá Reserve, Mexico; longleaf pine forest, Southeastern Coastal Plain, North Carolina, USA). During 1,536 camera days, a total of 268 observations of 20 putative snake predator species were recorded in the vicinity of artificial prey. Predators were observed to detect artificial prey 52 times, but only 21 attacks were recorded. Mammals were the most commonly recorded group of predators near replicas (243) and were responsible fo...
Seminars in Cell & Developmental Biology, 2019
A growing number of biologists have begun asking whether environmentally induced phenotypic chang... more A growing number of biologists have begun asking whether environmentally induced phenotypic change-'phenotypic plasticity'-precedes and facilitates the origin and canalization of novel, complex phenotypes. However, such 'plasticity-first evolution' (PFE) remains controversial. Here, we summarize the PFE hypothesis and describe how it can be evaluated in natural systems. We then review the evidence for PFE from amphibians (a group in which phenotypic plasticity is especially widespread) and describe how phenotypic plasticity might have facilitated macroevolutionary change. Finally, we discuss what is known about the proximate mechanisms of PFE in amphibians. We close with suggestions for future research. As we describe, amphibians offer some of the best support for plasticity's role in the origin of evolutionary novelties.
Proceedings of the Royal Society B: Biological Sciences, 2019
Plasticity-led evolution occurs when a change in the environment triggers a change in phenotype v... more Plasticity-led evolution occurs when a change in the environment triggers a change in phenotype via phenotypic plasticity, and this pre-existing plasticity is subsequently refined by selection into an adaptive phenotype. A critical, but largely untested prediction of plasticity-led evolution (and evolution by natural selection generally) is that the rate and magnitude of evolutionary change should be positively associated with a phenotype's frequency of expression in a population. Essentially, the more often a phenotype is expressed and exposed to selection, the greater its opportunity for adaptive refinement. We tested this prediction by competing against each other spadefoot toad tadpoles from different natural populations that vary in how frequently they express a novel, environmentally induced carnivore ecomorph. As expected, laboratory-reared tadpoles whose parents were derived from populations that express the carnivore ecomorph more frequently were superior competitors fo...
The relationship between inbreeding and both conception and foaling rates in Standardbred horses ... more The relationship between inbreeding and both conception and foaling rates in Standardbred horses (trotters and pacers) was examined for 1194 breeding years. There was a statistically significant (P less than 0.05) trend for conception and foaling rates to decrease with increased inbreeding; however, this relationship accounted for less than 2 percent of the variation. Additionally, the relationship between reproductive performance and inbreeding was not consistent between trotters and pacers. For trotters (F = 0.103) there was a trend for an increase in conception and foaling rates with increased inbreeding, while for pacers (F = 0.074), reproductive performance decreased with increased inbreeding. Overall, inbreeding does not appear to be a significant factor influencing reproductive performance of Standardbred horses.
Phenotypic plasticity is ubiquitous and generally regarded as a key mechanism for enabling organi... more Phenotypic plasticity is ubiquitous and generally regarded as a key mechanism for enabling organisms to survive in the face of environmental change. Because no organism is infinitely or ideally plastic, theory suggests that there must be limits (for example, the lack of ability to produce an optimal trait) to the evolution of phenotypic plasticity, or that plasticity may have inherent significant costs. Yet numerous experimental studies have not detected widespread costs. Explicitly differentiating plasticity costs from phenotype costs, we re-evaluate fundamental questions of the limits to the evolution of plasticity and of generalists vs specialists. We advocate for the view that relaxed selection and variable selection intensities are likely more important constraints to the evolution of plasticity than the costs of plasticity. Some forms of plasticity, such as learning, may be inherently costly. In addition, we examine opportunities to offset costs of phenotypes through ontogeny, amelioration of phenotypic costs across environments, and the condition-dependent hypothesis. We propose avenues of further inquiry in the limits of plasticity using new and classic methods of ecological parameterization, phylogenetics and omics in the context of answering questions on the constraints of plasticity. Given plasticity's key role in coping with environmental change, approaches spanning the spectrum from applied to basic will greatly enrich our understanding of the evolution of plasticity and resolve our understanding of limits.
Ecological character displacement is considered crucial in promoting diversification, yet relativ... more Ecological character displacement is considered crucial in promoting diversification, yet relatively little is known of its underlying mechanisms. We examined whether evolutionary shifts in gene expression plasticity ('genetic accommodation') mediate character displacement in spadefoot toads. Where Spea bombifrons and S. multiplicata occur separately in allopatry (the ancestral condition), each produces alternative, diet-induced, larval ecomorphs: omnivores, which eat detritus, and carnivores, which specialize on shrimp. By contrast, where these two species occur together in sympatry (the derived condition), selection to minimize competition for detritus has caused S. bombifrons to become nearly fixed for producing only carnivores, suggesting that character displacement might have arisen through an extreme form of genetic accommodation ('genetic assimilation') in which plasticity is lost. Here, we asked whether we could infer a signature of this process in regulatory changes of specific genes. In particular, we investigated whether genes that are normally expressed more highly in one morph ('biased' genes) have evolved reduced plasticity in expression levels among S. bombifrons from sympatry compared to S. bombifrons from allopatry. We reared individuals from sympatry vs. allopatry on detritus or shrimp and measured the reaction norms of nine biased genes. Although different genes displayed different patterns of gene regulatory evolution, the combined gene expression profiles revealed that sympatric individuals had indeed lost the diet-induced gene expression plasticity present in allopatric individuals. Our data therefore provide one of the few examples from natural populations in which genetic accommodation/assimilation can be traced to regulatory changes of specific genes. Such genetic accommodation might mediate character displacement in many systems.
Parental effects are often considered an evolved response, in which parents transmit information ... more Parental effects are often considered an evolved response, in which parents transmit information about the environment to enhance offspring fitness. However, these effects need not be adaptive. Here, we provide a striking example by presenting evidence that overfeeding of adult Mexican spadefoot toads, Spea multiplicata, is associated with decreased offspring survival. After a temporary change to their standard feeding regimen, S. multiplicata in our captive colony developed a much higher body condition (i.e. body mass for a given body length) than those in the wild. We analysed data from three subsequent experiments and found that although the body condition of a father was positively correlated with tadpole survival, mothers with a higher condition had lower tadpole survival. Our study highlights how obesity can negatively impact future generations via maladaptive maternal effects. Such effects could be especially likely for animals living in variable environments (such as spadefo...
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