The evolution of genetic architecture under frequency-dependent disruptive selection

We propose a model to analyze a quantitative trait under frequency-dependent disruptive selection. Selection on the trait is a combination of stabilizing selection and intraspecific competition, where competition is maximal between individuals with equal phenotypes. In addition, there is a density-dependent component induced by population regulation. The trait is determined additively by a number of biallelic loci, which can have different effects on the trait value. In contrast to most previous models, we assume that the allelic effects at the loci can evolve due to epistatic interactions with the genetic background. Using a modifier approach, we derive analytical results under the assumption of weak selection and constant population size, and we investigate the full model by numerical simulations. We find that frequency-dependent disruptive selection favors the evolution of a highly asymmetric genetic architecture, where most of the genetic variation is concentrated on a small number of loci. We show that the evolution of genetic architecture can be understood in terms of the ecological niches created by competition. The phenotypic distribution of a population with an adapted genetic architecture closely matches this niche structure. Thus, evolution of the genetic architecture seems to be a plausible way for populations to adapt to regimes of frequency-dependent disruptive selection. As such, it should be seen as a potential evolutionary pathway to discrete polymorphisms and as a potential alternative to other evolutionary responses, such as the evolution of sexual dimorphism or assortative mating.     

Assessing evolutionary consequences of size-selective recreational fishing on multiple life-history traits,with an application to northern pike (<Emphasis Type="Italic">Esox lucius</Emphasis>)

Despite mounting recognition of the importance of fishing-induced evolution, methods for quantifying selection pressures on multiple adaptive traits affected by size-selective harvesting are still scarce. We study selection differentials on three life-history traits—reproductive investment, size at maturation, and growth capacity—under size-selective exploitation of northern pike (Esox lucius L.) with recreational-fishing gear. An age-structured population model is presented that accounts for the eco-evolutionary feedback arising from density-dependent and frequency-dependent selection. By introducing minimum-length limits, maximum-length limits, and combinations of such limits (resulting in harvestable-slot length limits) into the model, we examine the potential of simple management tools for mitigating selection pressures induced by recreational fishing. With regard to annual reproductive investment, we find that size-selective fishing mortality exerts relatively small positive selection differentials. By contrast, selection differentials on size at maturation are large and consistently negative. Selection differentials on growth capacity are often large and positive, but become negative when a certain range of minimum-length limits are applied. In general, the strength of selection is reduced by implementing more stringent management policies, but each life-history trait responds differently to the introduction of specific harvest regulations. Based on a simple genetic inheritance model, we examine mid- and long-term evolutionary changes of the three life-history traits and their impacts on the size spectrum and yield of pike. Fishing-induced evolution often reduces sizes and yields, but details depend on a variety of factors such as the specific regulation in place. We find no regulation that is successful in reducing to zero all selection pressures on life-history traits induced by recreational fishing. Accordingly, we must expect that inducing some degree of evolution through recreational fishing is inevitable.     

Sexual dimorphism and adaptive speciation: two sides of the same ecological coin

Models of adaptive speciation are typically concerned with demonstrating that it is possible for ecologically driven disruptive selection to lead to the evolution of assortative mating and hence speciation. However, disruptive selection could also lead to other forms of evolutionary diversification, including ecological sexual dimorphisms. Using a model of frequency-dependent intraspecific competition, we show analytically that adaptive speciation and dimorphism require identical ecological conditions. Numerical simulations of individual-based models show that a single ecological model can produce either evolutionary outcome, depending on the genetic independence of male and female traits and the potential strength of assortative mating. Speciation is inhibited when the genetic basis of male and female ecological traits allows the sexes to diverge substantially. This is because sexual dimorphism, which can evolve quickly, can eliminate the frequency-dependent disruptive selection that would have provided the impetus for speciation. Conversely, populations with strong assortative mating based on ecological traits are less likely to evolve a sexual dimorphism because females cannot simultaneously prefer males more similar to themselves while still allowing the males to diverge. This conflict between speciation and dimorphism can be circumvented in two ways. First, we find a novel form of speciation via negative assortative mating, leading to two dimorphic daughter species. Second, if assortative mating is based on a neutral marker trait, trophic dimorphism and speciation by positive assortative mating can occur simultaneously. We conclude that while adaptive speciation and ecological sexual dimorphism may occur simultaneously, allowing for sexual dimorphism restricts the likelihood of adaptive speciation. Thus, it is important to recognize that disruptive selection due to frequency-dependent interactions can lead to more than one form of adaptive splitting.     

浅水湖泊四种渔具捕捞选择性及对鱼类群落影响比较研究

了解不同种类和大小的鱼类对各种捕捞方式的脆弱性, 对于制定最佳的捕捞策略和合理开发鱼类资源至关重要。研究从渔获物组成、捕捞效率和个体大小等方面, 对长江中游典型湖泊牛山湖的四种渔具(网簖、刺网、电捕和鸬鹚)的捕捞选择性进行了比较分析。使用相对重要性指数(IRI)评价渔具捕捞对鱼类物种的选择性, 同时通过计算渔获物的平均营养水平(MTL)和饵料鱼与凶猛性鱼类的重量比, 分析捕捞对鱼类群落营养结构的潜在影响。结果表明, 网簖和电捕捕获的鱼类种数多、体长范围广, 对鱼类资源的破坏性最大, 应当加以限制或废除; 鸬鹚是捕获鳜(尤其是高龄鳜)的一种有效方式, 但对幼鳜和其他幼鱼也有较大损害, 应控制其捕捞强度; 刺网主要捕获几种经济鱼类, 并可平衡捕食者和饵料鱼之间数量关系, 是利用鱼类资源的一种较好方式, 关键在于对刺网网目大小和捕捞强度的限制。建议湖泊捕捞应从单一物种转向多物种综合管理, 并考虑物种之间的营养生态关系, 以更好地保护和可持续利用渔业资源。     

Rapid selection effects in a short-lived semelparous squid species exposed to exploitation: inferences from the optimisation of life-history functions

The short life span of cephalopods suggests a potential for high sensitivity to the artificial selective effects of human exploitation. To explore the effects of such selection life-history optimisation was applied using data for a semelparous squid, Illex argentinus with a life span of one year. Survival and fecundity functions were combined to generate a life time reproductive potential function. The maximum reproductive potential identified the optimum age for the squid to mature. In a situation of a constant mortality rate the maximum reproductive potential was achieved at an earlier age of maturation as mortality rate increased. The exact age when the optimum maturation occurs is sensitive to the rate of mortality and the form of the assumed growth curve but covers the age range when maturation is known to occur in this species. A more realistic seasonal fishing mortality function produced a more complex fitness curve with a temporally more restricted optimal age of maturity. The selection effects will be stronger in a seasonal fishery suggesting potentially very rapid evolutionary rates. Developing analyses specifically considering frequency-dependence and environmental-feedbacks will be valuable to clarify the potential of squid to show rapid evolutionary responses to selection. Strong selection for early age of maturation could affect the yield from the fishery but more importantly, could also make the migratory strategy, on which the fishery is based, an unviable option, resulting in collapse of the fishery that exploits the migratory component of the species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Spatial Distribution and Population Structure of the Pike <Emphasis Type="Italic">Esox lucius</Emphasis> L. in the Rybinsk Reservoir during the Climate Warming Period

Data on the distribution, size and age structure, and growth of the pike Esox lucius collected in the Rybinsk Reservoir in 1953–2015 were analyzed. The main factors that determine the population size of the pike and dynamics of its structural indices are the intensity of the fishery and changes in the temperature regime of the reservoir. Since the mid-1990s, under the influence of high harvesting load, the population of the pike decreased and its age structure changed toward the predominance of individuals of younger age groups. As a result of climate warming, the water column warming of the Rybinsk Reservoir in the 2000s affected the growth character of large individuals. The ratio of phenotypes with small and high number of vertebrae in the first year depends on the water temperature during embryonic development. Among the large individuals preferring deep-water areas with a lower temperature, approximately 50% have 61–62 vertebrae in the axial skeleton. It is suggested that, with climate warming and the development of pike spawn at a higher temperature, the portion of individuals with a high number of vertebrae may decrease, which will lead to a decrease in the number of recruits of the deep-water part of the population.     

Weak disruptive selection and incomplete phenotypic divergence in two classic examples of sympatric speciation: cameroon crater lake cichlids

Abstract Recent documentation of a few compelling examples of sympatric speciation led to a proliferation of theoretical models. Unfortunately, plausible examples from nature have rarely been used to test model predictions, such as the initial presence of strong disruptive selection. Here I estimated the form and strength of selection in two classic examples of sympatric speciation: radiations of Cameroon cichlids restricted to Lakes Barombi Mbo and Ejagham. I measured five functional traits and relative growth rates in over 500 individuals within incipient species complexes from each lake. Disruptive selection was prevalent in both groups on single and multivariate trait axes but weak relative to stabilizing selection on other traits and most published estimates of disruptive selection. Furthermore, despite genetic structure, assortative mating, and bimodal species-diagnostic coloration, trait distributions were unimodal in both species complexes, indicating the earliest stages of speciation. Long waiting times or incomplete sympatric speciation may result when disruptive selection is initially weak. Alternatively, I present evidence of additional constraints in both species complexes, including weak linkage between coloration and morphology, reduced morphological variance aligned with nonlinear selection surfaces, and minimal ecological divergence. While other species within these radiations show complete phenotypic separation, morphological and ecological divergence in these species complexes may be slow or incomplete outside optimal parameter ranges, in contrast to rapid divergence of their sexual coloration.     

Consumption of crucian carp (Carassius carassius L., 1758) by restocked pike (Esox lucius L., 1758) in a lake with frequent winter hypoxia

To assess the effects of stocked pike (Esox lucius L.) on crucian carp (Carassius carassius L.) biomasses, the annual consumption of pike was estimated and compared with removal fishery catches. The studied lake, Lake Savijärvi, is a small (40 ha), shallow and eutrophicated lake in southern Finland with frequent algal blooms during summers and fish kills during winters. Until the 1980s, the fish fauna consisted of pike, perch (Perca fluviatilis L.), roach [Rutilus rutilus (L.)], crucian carp, and tench (Tinca tinca L.). Since 2003, when crucian carp were abundant in the extreme and with only a few roach and tench individuals, the lake has been biomanipulated by removal seining. To enhance the effects of seining, piscivorous pike were restocked in the spring of 2008. During the 3‐year study, the catch of crucian carp decreased from 243 to 136 kg while the catch of pike increased from 0.1 to 5.7 kg per seining hectare due to their spawning in the lake as of 2009. The total crucian carp consumption by captured pike was 588 kg during the 2008–2010 study period. When applying the number of pike estimated with the mark‐recapture method, consumption estimate increased to 917 kg, or 22.9 kg per lake hectare in 2010. Thus, pike consumption of crucian carp was about 17% of the removal catch in 2010.     

Competition and the Effect of Spatial Resource Heterogeneity on Evolutionary Diversification

A model is presented to explore how the form of selection arising from competition for resources is affected by spatial resource heterogeneity. The model consists of a single species occupying two patches connected by migration, where the two patches can differ in the type of resources that they contain. The main goal is to determine the conditions under which competition for resources results in disruptive selection (i.e., selection favoring a polymorphism) since it is this form of selection that will give rise to the evolutionary diversification of resource exploitation strategies. In particular, comparing the conditions giving rise to disruptive selection when the two patches are identical to the conditions when they contain different resources reveals the effect of spatial resource heterogeneity. Results show that when the patches are identical, the conditions giving rise to disruptive selection are identical to those that give rise to character displacement in previous models. When the patches are different, the conditions giving rise to disruptive selection can be either more or less stringent depending upon demographic parameters such as the intrinsic rate of increase and the migration rate. Surprisingly, spatial resource heterogeneity can actually make forms of evolutionary diversification such as character displacement less likely. It is also found that results are dependent on how the resource exploitation strategies and the spatial resource heterogeneity affect the population dynamics. One robust conclusion however, is that spatial resource heterogeneity always has a disruptive effect when the migration rate between patches is low.     

Evolution of dispersal in spatially and temporally variable environments: The importance of life cycles

Spatiotemporal variability of the environment is bound to affect the evolution of dispersal, and yet model predictions strongly differ on this particular effect. Recent studies on the evolution of local adaptation have shown that the life cycle chosen to model the selective effects of spatiotemporal variability of the environment is a critical factor determining evolutionary outcomes. Here, we investigate the effect of the order of events in the life cycle on the evolution of unconditional dispersal in a spatially heterogeneous, temporally varying landscape. Our results show that the occurrence of intermediate singular strategies and disruptive selection are conditioned by the temporal autocorrelation of the environment and by the life cycle. Life cycles with dispersal of adults versus dispersal of juveniles, local versus global density regulation, give radically different evolutionary outcomes that include selection for total philopatry, evolutionary bistability, selection for intermediate stable states, and evolutionary branching points. Our results highlight the importance of accounting for life‐cycle specifics when predicting the effects of the environment on evolutionarily selected trait values, such as dispersal, as well as the need to check the robustness of model conclusions against modifications of the life cycle.     

ASYMMETRIC PATCH SIZE DISTRIBUTION LEADS TO DISRUPTIVE SELECTION ON DISPERSAL

Numerous models have been designed to understand how dispersal ability evolves when organisms live in a fragmented landscape. Most of them predict a single dispersal rate at evolutionary equilibrium, and when diversification of dispersal rates has been predicted, it occurs as a response to perturbation or environmental fluctuation regimes. Yet abundant variation in dispersal ability is observed in natural populations and communities, even in relatively stable environments. We show that this diversification can operate in a simple island model without temporal variability: disruptive selection on dispersal occurs when the environment consists of many small and few large patches, a common feature in natural spatial systems. This heterogeneity in patch size results in a high variability in the number of related patch mates by individual, which, in turn, triggers disruptive selection through a high per capita variance of inclusive fitness. Our study provides a likely, parsimonious and testable explanation for the diversity of dispersal rates encountered in nature. It also suggests that biological conservation policies aiming at preserving ecological communities should strive to keep the distribution of patch size sufficiently asymmetric and variable.     

Bioeconomic dynamics of a fishery modeled as an S-system

A bioeconomic model of a single species fishery is developed by using a realistic catch-rate function and assuming that a regulatory agency controls exploitation of the fishery by imposing a suitable tax on the catch. The existence of a nontrivial steady state and its stability are examined. The optimal harvesting policy is discussed from the viewpoints of variational calculus and control theory. The fishery is then modeled as an S-system by following the recasting techniques of Savageau and Voit. Numerical examples of the optimal control curves and the yield-effort curves are obtained by executing the ESSYNS algorithm.     

Evolutionary rebound from selective harvesting

Size-selective harvesting favours reduced investment in growth and earlier age at reproduction. A recent study by Edeline and colleagues has shown that trait dynamics of Lake Windermere pike reflect the interplay of 50 years of harvest and natural selection. Growth and investment in early reproduction decreased under harvesting, and then recovered when fishing pressures declined. This is the first study to use contrasting temporal patterns of natural and harvest selection pressures to account for observed trait changes.     

Sexual selection's impacts on ecological specialization: an experimental test

In many species, individuals specialize on different resources, thereby reducing competition. Such ecological specialization can promote the evolution of alternative ecomorphs—distinct phenotypes adapted for particular resources. Elucidating whether and how this process is influenced by sexual selection is crucial for understanding how ecological specialization promotes the evolution of novel traits and, potentially, speciation between ecomorphs. We evaluated the population-level effects of sexual selection (as mediated by mate choice) on ecological specialization in spadefoot toad tadpoles that express alternative ecomorphs. We manipulated whether sexual selection was present or reversed by mating females to their preferred versus non-preferred males, respectively. We then exposed their tadpoles to resource competition in experimental mesocosms. The resulting distribution of ecomorphs was similar between treatments, but sexual selection generated poorer trait integration in, and lower fitness of, the more specialized carnivore morph. Moreover, disruptive and directional natural selection were weaker in the sexual selection present treatment. Nevertheless, this effect on disruptive selection was smaller than previously documented effects of ecological opportunity and competitor density. Thus, sexual selection can inhibit adaptation to resource competition and thereby hinder ecological specialization, particularly when females obtain fitness benefits from mate choice that offset the cost of producing competitively inferior offspring.     

Four decades of opposing natural and human-induced artificial selection acting on Windermere pike (Esox lucius)

The ability of natural selection to drive local adaptation has been appreciated ever since Darwin. Whether human impacts can impede the adaptive process has received less attention. We tested this hypothesis by quantifying natural selection and harvest selection acting on a freshwater fish (pike) over four decades. Across the time series, directional natural selection tended to favour large individuals whereas the fishery targeted large individuals. Moreover, non-linear natural selection tended to favour intermediate sized fish whereas the fishery targeted intermediate sized fish because the smallest and largest individuals were often not captured. Thus, our results unequivocally demonstrate that natural selection and fishery selection often acted in opposite directions within this natural system. Moreover, the two selective factors combined to produce reduced fitness overall and stronger stabilizing selection relative to natural selection acting alone. The long-term ramifications of such human-induced modifications to adaptive landscapes are currently unknown and certainly warrant further investigation.     

Ecosystem services of Lake Võrtsjärv under multiple stress: a case study

This study is the first attempt at the European scale to make an inventory of ecosystem services (ESS) of a large lake. We analysed a set of ESS indicators against the annual mean values of environmental parameters for 2006–2013. According to principal component analysis, the trophic state- and hydrology-related factors explained about 70% of the environmental variability of the lake and showed strong relationships with some ESS. Among the provisioning ESS, the annual eel catch and the total fish catch were positively related to different eutrophication indicators while the catches of pike, bream, and burbot depended rather on hydrological factors. Reed harvesting efficiency was related to the lake’s water level. The indicators of regulating, maintenance, and cultural ESS showed very high variability in different years, the latter depending on socio-economic conditions rather than environmental factors. We discovered numerous trade-offs between ESS benefitting from higher trophic state or regulated water level of the lake and the goals of good ecological status of the lake. Our analysis showed a clear need for rules prioritizing life supporting regulatory services against other ESS.     

Molecular ecological approaches to studying the evolutionary impact of selective harvesting in wildlife

Harvesting of wildlife populations by humans is usually targeted by sex, age or phenotypic criteria, and is therefore selective. Selective harvesting has the potential to elicit a genetic response from the target populations in several ways. First, selective harvesting may affect population demographic structure (age structure, sex ratio), which in turn may have consequences for effective population size and hence genetic diversity. Second, wildlife-harvesting regimes that use selective criteria based on phenotypic characteristics (e.g. minimum body size, horn length or antler size) have the potential to impose artificial selection on harvested populations. If there is heritable genetic variation for the target characteristic and harvesting occurs before the age of maturity, then an evolutionary response over time may ensue. Molecular ecological techniques offer ways to predict and detect genetic change in harvested populations, and therefore have great utility for effective wildlife management. Molecular markers can be used to assess the genetic structure of wildlife populations, and thereby assist in the prediction of genetic impacts by delineating evolutionarily meaningful management units. Genetic markers can be used for monitoring genetic diversity and changes in effective population size and breeding systems. Tracking evolutionary change at the phenotypic level in the wild through quantitative genetic analysis can be made possible by genetically determined pedigrees. Finally, advances in genome sequencing and bioinformatics offer the opportunity to study the molecular basis of phenotypic variation through trait mapping and candidate gene approaches. With this understanding, it could be possible to monitor the selective impacts of harvesting at a molecular level in the future. Effective wildlife management practice needs to consider more than the direct impact of harvesting on population dynamics. Programs that utilize molecular genetic tools will be better positioned to assess the long-term evolutionary impact of artificial selection on the evolutionary trajectory and viability of harvested populations.     

On the Optimal Size of Marine Reserves

The excessive and unsustainable exploitation of our marine resources has led to the promotion of marine reserves as a fisheries management tool. Marine reserves, areas in which fishing is restricted or prohibited, can offer opportunities for the recovery of exploited stock and fishery enhancement. This study examines the impact of the creation of marine protected areas, from both economic and biological perspectives. The consequences of reserve establishment on the long-run equilibrium fish biomass and fishery catch levels are evaluated. We include reserve size as control variable to maximize catch at equilibrium. A continuous time model is used to simulate the effects of reserve size on fishing catch. Fish movements between the sites is assumed to take place at a faster time scale than the variation of the stock and the change of the fleet size. We take advantage of these two time scales to derive a reduced model governing the dynamics of the total fish stock and the fishing effort. Simulation results suggest that the establishment of a protected marine reserve will always lead to an increase in total fish biomass, an optimal size of a marine reserve can achieve to maximize the catch at equilibrium.     

Pathogen-induced rapid evolution in a vertebrate life-history trait

Anthropogenic factors, including climate warming, are increasing the incidence and prevalence of infectious diseases worldwide. Infectious diseases caused by pathogenic parasites can have severe impacts on host survival, thereby altering the selection regime and inducing evolutionary responses in their hosts. Knowledge about such evolutionary consequences in natural populations is critical to mitigate potential ecological and economic effects. However, studies on pathogen-induced trait changes are scarce and the pace of evolutionary change is largely unknown, particularly in vertebrates. Here, we use a time series from long-term monitoring of perch to estimate temporal trends in the maturation schedule before and after a severe pathogen outbreak. We show that the disease induced a phenotypic change from a previously increasing to a decreasing size at maturation, the most important life-history transition in animals. Evolutionary rates imposed by the pathogen were high and comparable to those reported for populations exposed to intense human harvesting. Pathogens thus represent highly potent drivers of adaptive phenotypic evolution in vertebrates.     

A framework to compare theoretical predictions on trait evolution in temporally varying environments under different life cycles

Predicting the evolution of traits such as dispersal or local adaptation, in a variable environment is an important issue in theoretical evolutionary ecology. With concepts such as hard selection vs. soft selection or fine-grained vs. coarse-grained environmental variability, this issue has attracted much attention, and yet different models seldom agree on qualitative predictions about, e.g. the evolution of generalist or specialist strategies, or the occurrence of stabilizing or disruptive selection on studied traits.Here, I investigate the effect of the order of events in the life cycle on trait evolution in a spatially heterogeneous, temporally varying landscape using a Wright–Fisher island model. I first develop a methodological framework allowing for different life cycles. Then I illustrate the importance of life cycles on selection regimes by looking more closely at the evolution of local adaptation.Model results show that the occurrence of disruptive selection and bi- or tristability mainly depends on the life cycle, the convexity of the trade-off behind local adaptation, the immigration rate, and the autocorrelation in patch state. With the same forces driving the evolution of local adaptation, different life cycles induce different evolutionary outcomes. Model results highlight the importance of accounting for life cycle specificities when attempting to predict the effects of the environment on evolutionarily selected trait values, as well as the need to check the robustness of evolutionary model conclusions against modifications of the life cycle.