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Responding to environmental change: plastic responses vary little in a synchronous breeder.

The impact of environmental change on animal populations is strongly influenced by the ability of individuals to plastically adjust key life-history events. There is therefore considerable interest in establishing the degree of plasticity in traits and how selection acts on plasticity in natural populations. Breeding time is a key life-history trait that affects fitness and recent studies have found that females vary significantly in their breeding time-environment relationships, with selection often favouring individuals exhibiting stronger plastic responses. In contrast, here, we show that although breeding time in the common guillemot, Uria aalge, is highly plastic at the population level in response to a large-scale environmental cue (the North Atlantic Oscillation, NAO), there is very little between-individual variation-most individuals respond to this climate cue very similarly. We demonstrate strong stabilizing selection against individuals who deviate from the average population-level response to NAO. This species differs significantly from those previously studied in being a colonial breeder, in which reproductive synchrony has a substantial impact on fitness; we suggest that counter selection imposed by a need for synchrony could limit individuals in their response and potential for directional selection to act. This demonstrates the importance of considering the relative costs and benefits of highly plastic responses in assessing the likely response of a population to the environmental change.

Adaptation, Physiological↗

Plasticity in life-history traits.

We describe the impact of recent life-history plasticity theory on insect studies, particularly on the interface between genetics and plasticity. We focus on the three-dimensional relationship between three key life-history traits: adult size (or mass), development time and growth rate, and the connections to life cycle regulation, host plant choice, and sexual selection in seasonal environments. The review covers fitness consequences of variation in size, development time and growth rate, and effects of sex, photoperiod, temperature, diet, and perceived mortality risk on these traits. We give special attention to evidence for adaptive plasticity in growth rates because of the important effects of such plasticity on the expected relationships between development time and adult size and, hence, on the use of life-history, fitness, and optimality approaches in ecology, as well as in genetics.

Animals↗

Frequency and microenvironmental pattern of selection on plastic shade-avoidance traits in a natural population of Impatiens capensis.

The frequency and predictability of different selective environments are important parameters in models for the evolution of plasticity but have rarely been measured empirically in natural populations. We used an experimental phytometer approach to examine the frequency, predictability, and environmental determinants of heterogeneous selection on phytochrome-mediated shade-avoidance responses in a natural population of the annual plant Impatiens capensis. The strength and direction of selection on shade-avoidance traits varied substantially on a fine spatial scale. The shade-avoidance phenotype had high relative fecundity in some microsites but was disadvantageous in other microsites. Local seedling density proved to be a surprisingly poor predictor of microenvironmental variation in the strength and direction of selection on stem elongation in this study population. At least some of this unpredictability resulted from microenvironmental variation in water availability; the shade-avoidance phenotype was more costly in dry microsites. Thus, environmental heterogeneity in resource availability can affect the relative costs and benefits of expressing shade-avoidance traits independent of local seedling density, the inductive environmental cue. Theory predicts that these conditions may promote local genetic differentiation in reaction norms in structured populations, as observed in I. capensis.

Adaptation, Physiological↗

Phenotypic plasticity and interpopulation differences in life history traits of Armadillidium vulgare (Isopoda:Oniscidae).

The hypothesis that the balance of trade-offs between survivorship, growth and reproductive allocation in the terrestrial isopod Armadillidium vulgare will change when resource input is increased has been investigated experimentally. When the quality of food available was increased, by adding a mixture of litter from herbaceous dicotyledonous plants to a background low-quality food of dead grasses, survivorship was found to be the most phenotypically plastic trait, increasing by 168%. Growth rates increased by 99% but reproductive allocation by only 21%. In the field, members of a population from a site with more high-quality food grew more than twice as fast as those from a site where less high-quality food was available. The population from the site with higher food availability, contrary to predictions from the laboratory study, did not survive as well as that from the site with less available high-quality food. This may be because the site that is more favourable for growth has a more stressful physical environment due to much bigger temperature fluctuations, which are known to be an important cause of mortality in this species. When individuals from both populations were reared under controlled laboratory conditions, both the parental and F1 generations from the poor growth environment survived better than those from the good growth habitat. However, even when given an excess of high-quality food those from the poor growth environment continued to grow more slowly and had a lower reproductive allocation than those from the site with higher food availability. We conclude that microevolutionary changes may have occurred in the balance of resource allocation between survivorship, growth and reproductive allocation, to favour higher survivorship during the longer prereproductive period at the site where growth to the threshold size for reproduction takes longer.

Animals↗

Ontogenetic reaction norm for binary traits: the timing of phallus development in the snail Bulinus truncatus.

The ontogenetic trajectory of plastic binary traits may provide valuable insights into their evolutionary rate of change. In this paper, the timing of the plastic response of a temperature-dependent sexual polymorphism, aphally, is investigated in the freshwater snail Bulinus truncatus. Aphally is defined as the loss of the male copulatory organ in otherwise hermaphroditic animals. Individuals from two inbred lines were switched at various times during their early development between 25 and 30 degrees C, and their phally status ascertained, in order to evaluate the parameters characterising the ontogenetic reaction norm of aphally to temperature. A series of nested models including parameters for the onset, offset, and the intensity of the response to temperature were fitted to the data, allowing for a wide range of reaction norms. One genotype did not show any variation in aphally ratio with switching temperature, while a switch-point model (onset and offset corresponding to the same developmental point in time) best fitted the second genotype. The results suggest that the plasticity of aphally is expressed before eggs hatch. Their consequences on the evolution of aphally are discussed. More generally, the methodology proposed here can be used to analyse variation in ontogenetic parameters of discrete traits.

Animals↗

Mapping phenotypic plasticity and genotype-environment interactions affecting life-history traits in Caenorhabditis elegans.

Phenotypic plasticity and genotype-environment interactions (GEI) play an important role in the evolution of life histories. Knowledge of the molecular genetic basis of plasticity and GEI provides insight into the underlying mechanisms of life-history changes in different environments. We used a genomewide single-nucleotide polymorphism map in a recombinant N2 x CB4856 inbred panel of the nematode Caenorhabditis elegans to study the genetic control of phenotypic plasticity to temperature in four fitness-related traits, that is, age at maturity, fertility, egg size and growth rate. We mapped quantitative trait loci (QTL) for the respective traits at 12 and 24 degrees C, as well as their plasticities. We found genetic variation and GEI for age at maturity, fertility, egg size and growth rate. GEI in fertility and egg size was attributed to changes in rank order of reaction norms. In case of age at maturity and growth rate, GEI was caused mainly by differences in the among-line variance. In total, 11 QTLs were detected, five QTL at 12 degrees C and six QTL at 24 degrees C, which were associated with life-history traits. Five QTL associated with age at maturity, fertility and growth rate showed QTL x environment interaction. These colocalized with plasticity QTL for the respective traits suggesting allelic sensitivity to temperature. Further fine mapping, complementation analyses and gene silencing are planned to identify candidate genes underlying phenotypic plasticity for age at maturity, fertility and growth.

Animals↗

Microclinal variation for ovariole number and body size in Drosophila melanogaster in 'Evolution Canyon'.

Sites that display strong environmental contrasts in close proximity, such as 'Evolution Canyon' on Mt. Carmel, Israel, are natural theatres for investigating adaptive evolution in action. We reared Drosophila melanogaster from collection sites along altitudinal transects on the north- and south-facing canyon slopes in each of three temperature environments, and assessed genetic variation in ovariole number and body size between and within collection sites, and temperature plasticity. Both traits exhibited significant genetic variation within collection sites and phenotypic plasticity in response to temperature, but not genetic variation for plasticity. Between-site genetic variation in ovariole number was negatively correlated with altitude on both slopes of the canyon, and collections from the north- and south-facing slopes were genetically differentiated for male, but not female, body size. Genetic variation between sites within easy dispersal range is consistent with the action of strong natural selection, although neither the selective agent(s) nor the direct targets of selection are known.

Adaptation, Biological↗

Reaction norms of Arabidopsis IV. Relationships between plasticity and fitness.

The study of the association between fitness and reaction norms is of primary importance given the hypothesized role for phenotypic plasticity in shaping evolutionary patterns: in microevolution, as one of mechanism for maintaining genetic variation, and in macroevolution, as a means of generating phenotypic novelties. In a glasshouse experiment, we investigated variation in reaction norms to nutrient availability in populations of Arabidopsis thaliana, and the relationship between this variation and reproductive fitness. We found evidence for across-treatment directional selection on the means for leaf number, flowering time, plant height, branching and growth rate; across-treatment stabilizing selection was detected for growth rate; and across-treatment disruptive selection was significant for leaf number. We also uncovered selection on the plasticity of some traits: directional for the plasticity of branching, and stabilizing for the plasticity of both branching and growth rate. When the two environments were considered separately, directional selection for height was detected under low nutrients; under high nutrients, we found evidence for directional selection on leaf number and height, and for disruptive selection on flowering time. The genetic correlation between a trait's expression in one environment and its expression in the alternate environment was positive and highly significant only for flowering time and growth rate. A principle components analysis revealed possible constraints on future selection responses, because of correlations among character means and among character plasticities.

Analysis of Variance↗

Phylogenetic Constraints and Environmental Filtering Jointly Drive Adaptive Evolution in Phragmites australis: From Genetic Structure to Trait Decoupling on the Mongolian Plateau.

The Mongolian Plateau, a typical arid and semi-arid zone in Eurasia, is characterized by highly heterogeneous and fragmented wetland habitats. Phragmites australis, a common wetland species in this region, exhibits remarkable adaptability. Unraveling the coordination between phylogenetic history and local environmental filtering is crucial for elucidating its adaptive mechanisms. Integrating landscape genomics and trait-based phylogenetic analyses, we analyzed transcriptome-wide SNPs, multidimensional functional traits, and environmental variables across 90 individuals from 30 natural P. australis populations. This study aims to reveal the genetic and phenotypic variation patterns underlying population genetic structure and trait variation, specifically distinguishing the roles of geographic isolation, environmental filtering, and phylogenetic history. Results reveal a significant drainage-dependent pattern in genetic structure. Populations in hydrologically connected basins show extensive admixture, whereas those in isolated endorheic basins form distinct lineages. While geographic isolation underpins genetic differentiation, environmental filtering independently explains ~33.84% of the genetic variation, driven primarily by moisture heterogeneity (precipitation seasonality and soil moisture). Crucially, we observed differentiated evolutionary trajectories across functional traits. Structural traits (e.g., plant height, leaf thickness) are phylogenetically conserved; in contrast, physiological traits (e.g., water use efficiency) are decoupled from phylogeny, showing patterns consistent with high plasticity regulated by local environments. This evolutionary decoupling strategy enables P. australis to flexibly adapt to heterogeneous habitats while maintaining structural stability. This study uncovers the synergistic mechanisms by which geographic isolation and environmental filtering jointly shape the genetic patterns of this cosmopolitan species at a regional scale, clarifies that its evolutionary responses may depend heavily on the differentiated plasticity of trait types, and provides valuable regional insights into how widespread wetland species adapt to heterogeneous environments under global change.

Mongolia Plateau↗

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

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.

Animals↗

Inducible reproductive plasticity of the guppy Poecilia reticulata in response to predation cues.

Predation has long been described as one of the major driving forces in evolution. Guppies (Poecilia reticulata) from natural populations exposed to different predation pressures, were found to have different life history traits. Reproductive plasticity in response to direct predation cues has mainly been reported for invertebrates. The goals of the present study were to determine whether exposure to predation cues would induce reproductive phenotypic plasticity in female guppies and to determine whether the effective cues are visual, chemical, or a combination of both. In our first experiment, female guppies exposed to predation cues of the african cichlids Aulonocara nyassae increased their reproductive output by almost two fold, having larger brood-sizes and shorter brood-interval at the first spawn. This effect disappeared in the second spawn in the absence of predators. In the second experiment we found that exposure to the predators induced an increase in the brood-size regardless of whether the cue was: only visual, only chemical, visual and chemical or visual, chemical and tactile. The impacts of these cues were equally powerful on the tested variables and they did not have any cumulative effect. Similar to the results of the first experiment, this effect disappeared in the second spawn, in the absence of predation cues. The present study demonstrates a direct immediate and reversible effect of predation cues on guppy reproduction.

Adaptation, Physiological↗

A chromosomal analysis of the phenotypic plasticity of some life history traits in relation to developmental temperature.

The capacity to delay egg deposition in D. melanogaster females in the absence of a sexual partner is genetically determined and opposite types can be artificially selected. In natural populations, the relative frequency of these genotypes varies geographically and seasonally, with temperature as a selective factor. However, the retention duration of these genotypes can be modified by developmental temperature change. To study the genetic control of this response, chromosome substitution between opposite types of line was carried out in order to produce every possible homozygous chromosomal combination of the three major chromosomes (X,2,3). Eggs of these eight constructed lines were developed at two different temperatures (25 degrees C and 14 degrees C). Low temperature development directly affected the number of ovarioles but also modified the subsequent expression of adult characteristics such as retention duration and fecundity. The comparison of the eight lines revealed that, although the 3 chromosomes were involved in the genetic determinism of each trait, only one or two of them were sensitive to temperature change, and these differed according to the trait. For retention duration and fecundity, the effect of chromosome 3 from the long retention strain was particularly affected by low temperature, showing antagonism between the selective effect detected in natural populations and the effect on phenotypic plasticity studied here.

Animals↗

Maintenance of genetic variation in phenotypic plasticity: the role of environmental variation.

We study genetic variation in phenotypic plasticity maintained by a balance between mutation and weak stabilizing selection. We consider linear reaction norms allowing for spatial and/or temporal variation in the environments of development and selection. We show that the overall genetic variation maintained does not depend on whether the trait is plastic or not. The genetic variances in height and slope of a linear reaction norm, and their covariance, are predicted to decrease with the variation in the environment. Non-pleiotropic loci influencing either height or slope are expected to decrease the genetic variance in slope relative to that in height. Decrease in the ratio of genetic variance in slope to genetic variance in height with increasing variation in the environment presents a test for the presence of loci that only influence the slope, and not the height. We use data on Drosophila to test the theory. In seven of eight pair-wise comparisons genetic variation in reaction norm is higher in a less variable environment than in a more variable environment, which is in accord with the model's predictions.

Alleles↗

Phenotypic plasticity for life-history traits in Drosophila melanogaster. III. Effect of the environment on genetic parameters.

We estimated genetic and environmental variance components for developmental time and dry weight at eclosion in Drosophila melanogaster raised in ten different environments (all combinations of 22, 25 and 28 degrees C and 0.5, 1 and 4% yeast concentration, and 0.25% yeast at 25 degrees C). We used six homozygous lines derived from a natural population for complete diallel crosses in each environment. Additive genetic variances were consistently low for both traits (h2 around 10%). The additive genetic variance of developmental time was larger at lower yeast concentrations, but the heritability did not increase because other components were also larger. The additive genetic effects of the six parental lines changed ranks across environments, suggesting a mechanism for the maintenance of genetic variation in heterogenous environments. The variance due to non-directional dominance was small in most environments. However, there was directional dominance in the form of inbreeding depression for both traits. It was pronounced at high yeast levels and temperatures but disappeared when yeast or temperature were decreased. This meant that the heterozygous flies were more sensitive to environmental differences than homozygous flies. Because dominance effects are not heritable, this suggests that the evolution of plasticity can be constrained when dominance effects are important as a mechanism for plasticity.

Adaptation, Physiological↗

Phenotypic plasticity in bacterial plasmids.

Plasmid pB15 was previously shown to evolve increased horizontal (infectious) transfer at the expense of reduced vertical (intergenerational) transfer and vice versa, a key trade-off assumed in theories of parasite virulence. Whereas the models predict that susceptible host abundance should determine which mode of transfer is selectively favored, host density failed to mediate the trade-off in pB15. One possibility is that the plasmid's transfer deviates from the assumption that horizontal spread (conjugation) occurs in direct proportion to cell density. I tested this hypothesis using Escherichia coli/pB15 associations in laboratory serial culture. Contrary to most models of plasmid transfer kinetics, my data show that pB15 invades static (nonshaking) bacterial cultures only at intermediate densities. The results can be explained by phenotypic plasticity in traits governing plasmid transfer. As cells become more numerous, the plasmid's conjugative transfer unexpectedly declines, while the trade-off between transmission routes causes vertical transfer to increase. Thus, at intermediate densities the plasmid's horizontal transfer can offset selection against plasmid-bearing cells, but at high densities pB15 conjugates so poorly that it cannot invade. I discuss adaptive vs. nonadaptive causes for the phenotypic plasticity, as well as potential mechanisms that may lead to complex transfer dynamics of plasmids in liquid environments.

Analysis of Variance↗

Characterization of Tn917 insertion mutants of Staphylococcus epidermidis affected in biofilm formation.

Biofilm formation is thought to result from the concerted action of primary attachment to a specific surface and accumulation in multilayered cell clusters. Here we describe the isolation and characterization of transposon (Tn917) mutants of Staphylococcus epidermidis O-47 which were biofilm negative in the polystyrene microtiter plate assay. Among 5,000 Tn917 insertion mutants, 4 biofilm-negative mutants were isolated. Each mutant carried one copy of Tn917. The mutants were divided into two phenotypic classes: class A (mut1 and mut1a) and class B (mut2 and mut2a). Mutants of phenotypic class A lacked four cell surface proteins, were less hydrophobic, and were affected in primary attachment to polystyrene, but were still able to form multilayered cell clusters. They were able to form a biofilm on a glass surface, a trait that was even more pronounced than in the wild-type stain O-47. Loss of several surface proteins might have led to the reduced surface hydrophilic structures, thus favoring primary attachment to a glass surface and leading to subsequent biofilm formation. Mutants of phenotype class B were able to attach to polystyrene but were unable to form multilayered cell clusters, had unchanged cell surface proteins and hydrophobicity, and were unable to form a biofilm on a glass surface, mut1 and mut2 could be complemented by wild-type DNA fragments containing the Tn917 insertion sites of mut1 and mut2, respectively. The complemented biofilm-positive clone mut1 (pRC20) produced a 60-kDa protein which is postulated to function as the adhesin for binding to plastic. The traits of binding to polystyrene and the ability to form multilayered cell clusters are phenotypically and genetically distinct.

Animals↗

Progressive bilateral nasal alar collapse: a dominantly inherited trait.

We describe a 47-year-old woman with progressive bilateral collapse of the alae nasi first noted at age 16 years. Her dizygotic twin daughters have similar nasal collapse beginning at age 20 years. This condition appears to be inherited as a dominant trait. Although plastic surgical correction has been successful for a phenotypically similar condition due to trauma, surgical correction must be considered cautiously in individuals with an atraumative, possibly inherited, progressive form of the disorder.

Adult↗

Contribution of relative growth rate to root foraging by annual and perennial grasses from California oak woodlands.

Plants forage for nutrients by increasing their root length density (RLD) in nutrient-rich soil microsites through root morphological changes resulting in increased root biomass density (RBD), specific root length (SRL), or branching frequency (BF). It is commonly accepted that fast-growing species will forage more than slow-growing species. However, foraging responses may be due solely to differences in relative growth rates (RGR). There is little evidence, after the effects of RGR are removed, that the fast versus slow foraging theory is correct. In a pot study, we evaluated foraging of four grass species that differed in RGR: one fast-growing annual species, Bromus diandrus, two intermediate-growing species, annual Bromus hordeaceus and perennial Elymus glaucus, and one slow-growing perennial species, Nassella pulchra. We harvested plants either at a common time (plants varied in size) or at a common leaf number (plants similar size, surrogate for common biomass). By evaluating species at a common time, RGR influenced foraging. Conversely, by evaluating species at a common leaf number, foraging could be evaluated independent of RGR. When RGR was allowed to contribute to foraging (common time harvest), foraging and RGR were positively correlated. B. diandrus (fast RGR) foraged to a greater extent than did E. glaucus (intermediate RGR) and N. pulchra (slow RGR). E. glaucus (intermediate RGR) foraged to a greater extent than N. pulchra (slow RGR). Root growth within nutrient-rich microsites was due to significant increases in RBD, not to modifications of SRL or BF. However, when RGR was not allowed to influence foraging (common leaf number harvest), none of the four species significantly enhanced RLD in nutrient-rich compared to control microsites. This suggests that RGR strongly influenced the ability of these grass species to forage and also supports the need to evaluate plastic root traits independent of RGR.

Biomass↗