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Shade avoidance in Trifolium repens: costs and benefits of plasticity in petiole length and leaf size.

We tested whether the degree of shade-induced plasticity in petiole length and leaf area is related to the mean trait value expressed under high-light conditions, and to what extent trait values expressed under high-light and shaded conditions affect plant performance. Thirty-four Trifolium repens genotypes were used with a wide range of petiole lengths and leaf areas. Plants were subjected to a high-light environment and two shading regimes: homogeneous shading and a vertical light gradient. Absolute petiole elongation in response to both shading treatments and absolute leaf area expansion in response to homogeneous shading were independent of the trait values expressed in high light. Consequently, relative plasticity was higher for genotypes with lower high-light trait values. Plasticity was associated with enhanced plant performance in a vertical light gradient but not in homogeneously shaded conditions. We also found costs associated with the ability to express plasticity. Our results suggest that selection can act separately on trait values expressed under high-light conditions and on the degree of plasticity.

Adaptation, Physiological↗

Phenotypic plasticity, sexual selection and the evolution of colour patterns.

When a population comes to occupy a new environment, phenotypically plastic responses alter the distribution of phenotypes, and hence affect both the direction and the intensity of selection. Rates of evolution can be accelerated or retarded compared to what would happen in the absence of plasticity. Plastic responses in one trait result in novel selection pressures on other traits, and this can lead to evolution in completely different directions than predicted in the absence of plasticity. In this paper I use the concept of the adaptive surface in order to identify conditions under which the various different outcomes are expected. I then discuss differences between sexually and naturally selected traits. Sexually selected traits are often expected to be plastic in their expression, with individuals in high condition developing greater elaboration. As examples of sexually selected traits I review the evolution of colour patterns in birds with a view to assessing the magnitude of plastic responses in their development, and to ask how such responses may have influenced genetic evolution. The common colour pigments in birds are carotenoids and melanins. Both are used in social signaling, and consequently are expected to evolve to be phenotypically plastic indicators of an individual's quality. Perhaps partly because they are condition indicators, the quantity of carotenoids in the plumage can be strongly influenced by diet. Examples are described where alterations of carotenoids in the diet are thought to have altered the phenotype, driving genetic evolution in novel directions. Melanin patterns seem to be less affected by diet, but the intensity of melanization after moult is affected by social interactions during the moult and by raising birds in humid conditions. Hormonal manipulations can have dramatic effects on both the kinds of melanin produced (eumelanin or phaeomelanin) as well as the patterns they form. Differences between species in melanin patterns resemble differences produced by environmental manipulations, as well as those produced by simple modulations of parameters in computer simulations of pattern formation. While phenotypic plasticity is one way that genetic change in plumage patterns (and other traits) could be driven, there are others, including the appearance of major mutations and selection on standing variation whose distribution is not altered in the new environment. I consider some evidence for the different alternatives, and ask when they might lead to qualitatively different evolutionary outcomes.

Adaptation, Physiological↗

Quantitative genetic variation of leaf size and shape in a mixed diploid and triploid population of Populus.

In the interspecific cross of Populus trichocarpa x P. deltoides, unexpected simultaneous occurrence of diploid hybrids and triploid hybrids (with two alleles from the female parent and one from the male parent at each locus) led us to examine the evolutionary genetic significance of this phenomenon. As expected, leaf size and shape of the triploid progeny are closer to the female P. trichocarpa than male P. deltoides parent. Although the pure triploid progeny population did not have higher genetic variance in leaf traits than the pure diploid population, the former appears to hide much non-additive genetic variance and display strong genetic control over the phenotypic plasticity of leaf traits. It is suggested that the cryptic non-additive variance, especially epistasis, can be released when a population is disturbed by changes in the environment. A mixed diploid and triploid progeny population combines phenotypic and genetic characteristics of both pure hybrids and is considered to be of adaptive significance for populars to survive and evolve in a fluctuating environment. The significant effect due to general and specific combining ability differences at the population level suggests that the population divergence of these two species is under additive and non-additive genetic control.

Diploidy↗

The Evolution of Plasticity and Nonplastic Spatial and Temporal Adaptations in the Presence of Imperfect Environmental Cues.

A model for the evolution of plasticity is considered in which the phenotype, undergoing stabilizing selection, is modeled as a linear function of an environmental cue correlated with the phenotypic optimum, with the coefficients z0 and z1 evolving according to standard quantitative genetic theory. In contrast to previous theoretical models, as the rate of migration between demes or the rate of cyclic fluctuations in the optimum increases, the amount of plasticity [Formula: see text] at equilibrium is shown to increase gradually, in part accounting for the effect of reduced nonplastic adaptation and reaching a maximum equal to the squared correlation between the environmental cue and the phenotypic optimum. Given that information available to the organism is limited, this bias of the expressed phenotype toward the global optimum is still optimal, however, in a certain decision-theoretic sense. When genetic variation in the plastic component of the trait is small so that spatial or temporal differentiation in plasticity is small, the effect of plasticity on nonplastic adaptation is to reduce the effects of variation in the phenotypic optimum by a factor [Formula: see text] only. Information acquisition costs and joint evolution of sensory systems are discussed.

environmental cues↗

Evolution of phenotypic plasticity: patterns of plasticity and the emergence of ecotypes.

Phenotypic plasticity itself evolves, as does any other quantitative trait. A very different question is whether phenotypic plasticity causes evolution or is a major evolutionary mechanism. Existing models of the evolution of phenotypic plasticity cover many of the proposals in the literature about the role of phenotypic plasticity in evolution. I will extend existing models to cover adaptation to a novel environment, the appearance of ecotypes and possible covariation between phenotypic plasticity and mean trait value of ecotypes. Genetic assimilation does not sufficiently explain details of observed patterns. Phenotypic plasticity as a major mechanism for evolution--such as, invading new niches, speciation or macroevolution--has, at present, neither empirical nor model support.

Adaptation, Physiological↗

Evolution in stressful environments II: adaptive value and costs of plasticity in response to low light in Sinapis arvensis.

Plants possess a remarkable capacity to alter their phenotype in response to the highly heterogeneous light conditions they commonly encounter in natural environments. In the present study with the weedy annual plant Sinapis arvensis, we (a) tested for the adaptive value of phenotypic plasticity in morphological and life history traits in response to low light and (b) explored possible fitness costs of plasticity. Replicates of 31 half-sib families were grown individually in the greenhouse under full light and under low light (40% of ambient) imposed by neutral shade cloth. Low light resulted in a large increase in hypocotyl length and specific leaf area (SLA), a reduction in juvenile biomass and a delayed onset of flowering. Phenotypic selection analysis within each light environment revealed that selection favoured large SLA under low light, but not under high light, suggesting that the observed increase in SLA was adaptive. In contrast, plasticity in the other traits measured was maladaptive (i.e. in the opposite direction to that favoured by selection in the low light environment). We detected significant additive genetic variance in plasticity in most phenotypic traits and in fitness (number of seeds). Using genotypic selection gradient analysis, we found that families with high plasticity in SLA had a lower fitness than families with low plasticity, when the effect of SLA on fitness was statistically kept constant. This indicates that plasticity in SLA incurred a direct fitness cost. However, a cost of plasticity was only expressed under low light, but not under high light. Thus, models on the evolution of phenotypic plasticity will need to incorporate plasticity costs that vary in magnitude depending on environmental conditions.

Adaptation, Biological↗

Comparative life histories and ecophysiology of Drosophila melanogaster and D. simulans.

Numerous laboratory investigations have compared Drosophila melanogaster and D. simulans for various life history traits and fitness related ecophysiological parameters. From presently available information, it is however difficult to get a general comparative pattern describing the divergence of their ecological niches and understanding their demographic success. Two environmental factors seem however to have played a major role: temperature and alcoholic resources. From an ecophysiological approach, D. simulans may be described as generally more sensitive to stresses; other results point to this species as more cold adapted than its sibling; in some cases, however, D. simulans may appear as better adapted to a warm environment. When investigated, ecophysiological traits show a lesser geographic variability in D. simulans than in D. melanogaster. Presently available information does not explain the ecological prevalence of D. simulans in many places with a mild temperate or subtropical climate. This is presumably due to the fact that most comparisons have been done at a single, standard temperature of 25 degrees C. Comparative studies should be undertaken, spanning the thermal ranges of the two species, and the phenotypic plasticity of ecophysiological traits should now be considered.

Acetic Acid↗

Reaction norm functions and QTL-environment interactions for flowering time in Arabidopsis thaliana.

Many plant traits are phenotypically plastic in response to resource levels that vary continuously among environments. To be able to predict phenotypes in new environments, it is useful to model reaction norms as functions, rather than as a collection of discrete character states. Flowering date and rosette leaf number were measured in 100 recombinant inbred lines of Arabidopsis thaliana, grown on a gradient of light intensity. The results show that there is genetic variation among the recombinant inbred lines for parameters of the reaction norm functions. Genetic variances for leaf number and flowering date are highest under low light conditions. Underlying quantitative trait loci (QTLs) affecting the shape of the reaction norm functions were mapped by modifying Haley & Knott (1992) regressions to include polynomial effects of the environment. Quantitative trait loci of large effect were generally insensitive to the resource gradient. Seven QTLs affecting flowering date and eight QTLs for rosette leaf number were identified, of which only two had significant effects on the linear and quadratic components of the reaction norm function. These results suggest that the genotype-environment interactions for flowering time are controlled by many minor genes, whose effects are below the detection limit in most mapping experiments.

Arabidopsis↗

Variation amongst survivor populations of white clover collected from sites across Europe: growth attributes and physiological responses to low temperature.

Experiments were carried out at IGER, Aberystwyth, UK to investigate traits of direct relevance to the processes of overwintering and spring growth in white clover (Trifolium repens L.). The plant material used was derived from baseline populations of the cultivar AberHerald and survivor populations generated after 2-3 years' growth in Germany (Kiel), Sweden (Uppsala) and Switzerland (Zürich). The aims of the experiments were to measure the level of genetic shift that had occurred in certain traits due to selection in the survivor populations by comparing these with the baseline population. The adaptive significance of traits was assessed by determining the extent to which stabilizing selection had operated to reduce levels of intra-population variation. Significant differences were found in the responses of leaf production to two temperature treatments in the survivor populations from Germany and Sweden compared with the Swiss and baseline material. Plants of the former two populations produced much more leaf than the others at the higher temperature, but leaf production rates at the lower temperature did not differ. As this experiment used cloned genotypes in the two treatments, the result suggests that a higher degree of phenotypic plasticity for this trait had been selected for in the German and Swedish populations. These populations also showed greater rates of regrowth of leaves from terminal buds exposed to sub-zero temperatures, but there were no differences between populations in levels of freezing tolerance, or in stolon carbohydrate content. Genetic shift occurred in the degree of unsaturation of stolon lipids, with all three survivor populations possessing higher proportions of unsaturated fatty acids than the baseline. Stabilizing selection also operated on this trait in the survivor populations, suggesting that it is of adaptive significance in cool climates.

Acclimatization↗

The common quantitative genetic basis of wing morphology and diapause occurrence in the cricket Gryllus veletis.

A covariation between wing morphology and diapause occurrence has been observed in many insect species, but the genetic basis of this covariation has never been established. This study measures the heritability of, and genetic correlation between, these two ecologically important threshold traits in the cricket Gryllus veletis. A total of 81 full-sib families were reared in the laboratory to estimate these parameters. A comparison of laboratory and field samples showed that these two traits are highly plastic. The heritability of wing morphology was 0.25 (0.09), the heritability of diapause occurrence was 0.77 (0.11) and the genetic correlation between them was 0.61 (0.19). These estimates did not differ between males and females. The significance of these quantitative genetic parameters is discussed with reference to the monomorphism of natural populations of G. veletis for diapause occurrence and with reference to the trade-off between the ability to disperse by flight and the ability to diapause found in at least one closely related species. A survey of the literature reveals that genetic correlations between diapause occurrence or wing morphology and various other traits are common in insects, suggesting that these two traits are often genetically integrated in insect life-histories.

Animals↗

Evolutionary trade-off between weapons and testes.

It has long been recognized that male mating competition is responsible for the evolution of weaponry for mate acquisition. However, when females mate with more than one male, competition between males can continue after mating in the form of sperm competition. Theory predicts that males should increase their investment in sperm production as sperm competition is increased, but it assumes that males face a trade-off between sperm production and other life-history traits such as mate acquisition. Here, we use a genus of horned beetle, Onthophagus, to examine the trade-off between investment in testes required for fertilizations and investment in weapons used to obtain matings. In a within-species study, we prevented males from developing horns and found that these males grew larger and invested relatively more in testes growth than did males allowed to grow horns. Among species, there was no general relationship between the relative sizes of horns and testes. However, the allometric slope of horn size on body size was negatively associated with the allometric slope of testes size on body size. We suggest that this reflects meaningful evolutionary changes in the developmental mechanisms regulating trait growth, specifically in the degree of nutrition-dependent phenotypic plasticity versus canalization of traits. Finally, we show how this resource allocation trade-off has influenced the evolutionary diversification of weapons, revealing a rich interplay between developmental trade-offs and both pre- and postmating mechanisms of sexual competition.

Animals↗

Abnormal plasticity of the sensorimotor cortex to slow repetitive transcranial magnetic stimulation in patients with writer's cramp.

Previous studies demonstrated functional abnormalities in the somatosensory system, including a distorted functional organization of the somatosensory cortex (S1) in patients with writer's cramp. We tested the hypothesis that these functional alterations render S1 of these patients more susceptible to the "inhibitory" effects of subthreshold 1 Hz repetitive transcranial magnetic stimulation (rTMS) given to S1. Seven patients with writer's cramp and eight healthy subjects were studied. Patients also received rTMS to the motor cortex hand area (M1). As an outcome measure, short-latency afferent inhibition (SAI) was tested. SAI was studied in the relaxed first dorsal interosseous muscle using conditioning electrical stimulation of the index finger and TMS pulses over the contralateral M1. Baseline SAI did not differ between groups. S1 but not M1 rTMS reduced SAI in patients. rTMS had no effects on SAI in healthy subjects. Because SAI is mediated predominantly at a cortical level in the sensorimotor cortex, we conclude that there is an abnormal responsiveness of this area to 1 Hz rTMS in writer's cramp, which may represent a trait toward maladaptive plasticity in the sensorimotor system in these patients.

Adult↗

Digestive morphology and enzyme activity in the Andean toad Bufo spinulosus: hard-wired or flexible physiology?

Gut plasticity is a trait with implications on animal performance. However, and despite their importance as study models in physiology, research on gut flexibility in amphibians is scarce. In the present work, we analyse digestive adjustments of Bufo spinulosus adult individuals to cope with changes in diet quality and quantity at two organizational levels (i.e., digestive morphology and enzymes). We found that changes in gut size are related to the amount of food ingested, but not to diet composition. This is in agreement with "the gut seasonal change" hypothesis and offers a proximal explanation for this change. Digestive enzymatic activity (maltase and aminopeptidase-N) did not change with diet quality or quantity, which agrees with the hypothesis of "hard-wired physiology in adult amphibians". Both hypotheses are in agreement with the general theoretical framework of gut phenotypic flexibility when interpreted in light of amphibian natural history. In addition, our results indicate that the correlation between feeding frequency and the level of gut up-regulation proposed for interspecific comparisons may also be found at the intraspecific level.

Adaptation, Physiological↗

Performance of 18S rDNA helix E23 for phylogenetic relationships within and between the Rotifera-Acanthocephala clades.

The species diversity of the phylum Rotifera has been largely studied on the basis of morphological characters. However, cladistic relationships within this group are poorly resolved due to extensive homoplasy in morphological traits, substantial phenotypic plasticity and a poor fossil record. We undertook this study to determine if a phylogeny based on partial 18S rDNA, which included the helix E23 of 18S rDNA sequence, was concordant with established taxonomic relationships within the order Ploimida (class: Monogononta). We also estimated the level of polymorphism within clones and populations of Ploimida 'species'. Finally, we included the Cycliophora Symbion pandora as outgroup and the variable helix E23 region to examine the influence of their signal on the evolutionary relationships among Acanthocephala, Bdelloidea and Ploimida. Phylogenetic reconstruction was performed using maximum parsimony, neighbour joining and maximum likelihood methods. We found 1) that morphologically similar Ploimida 'species' show vastly different 18S E23 rDNA sequences; 2) inclusion of the helix E23 of 18S rDNA and its secondary structure analysis results in better resolution of family level relationships within the Ploimida; 3) an impact of Symbion pandora as an outgroup with inclusion of the helix E23 on the relationships between the Rotifera and the Acanthocephala; and 4) partial incongruence and differential substitution rate between conserved region and helix E23 region of the 18S rDNA gene depending on the taxomic group studied.

Animals↗

Long-read, high-coverage reference genome of the nymphalid butterfly Catonephele acontius (Nymphalidae: Biblidinae).

Catonephele acontius (Nymphalidae:Biblidinae:Epicalinii) is a butterfly species with a wide distribution across the Neotropics including the Amazon. Here, we present a long-read high-coverage reference genome for this species to serve as a genomic resource for future studies on Biblidinae butterflies, a group that is the subject of ongoing studies of seasonal adaptation under climate change. We used PacBio HiFi and IsoSeq reads to generate a highly contiguous and well-annotated reference genome. Five libraries were constructed, 4 using RNA from different tissues and 1 using high molecular weight (HMW) DNA from a wild-caught female. The DNA was sequenced using PacBio HiFi technology, and the RNA was sequenced using long read PacBio IsoSeq technology. About 20 Gb of raw HiFi data were generated and assembled to an initial size of 520.7 Mb (39 × homozygous coverage) in 90 contigs. The assembly was then polished and decontaminated into 40 contigs with an N50 of 19.927 Mb (BUSCO completeness: 99.0%; duplication: 0.5%; fragmentation: 0.7%; and missing: 0.3%). Final assembly size was 519.2 Mb. Repeats were annotated, showing that the genome consisted of 40.4% transposable elements. IsoSeq transcriptome data from antennae, leg, ovary, and digestive tissue was then used to structurally and functionally annotate gene models for the softmasked genome, uncovering ∼18,500 genes, with 70% of them given functional annotation. This reference assembly joins many published genomes in the Nymphalidae family but represents one of the first high-quality genomes from the Biblidinae subfamily. It provides a valuable resource to study the evolution of plastic and seasonal traits and will help investigate the genetic processes that may influence these species' responses to rapid climate change.

Animals↗

A parasite-mediated life-history shift in Daphnia magna.

The impact of parasitism on host populations will be modulated by both genetic variation for susceptibility, and phenotypically plastic-life-history traits that are altered to lessen the fitness consequences of infection. In this study we tested for life-history shifts in the crustacean Daphnia magna following exposure to the horizontally transmitted microsporidian, Glugoides intestinalis. In two separate experiments, we exposed hosts to parasite spores and measured their fecundity relative to controls. We show that host exposed G. intestinalis show fecundity compensation, i.e. hosts shift their life-history strategy towards early production. Our experiments included multiple host genotypes, and subtle differences among them indicated that fecundity compensation could be subject to parasite-mediated natural selection.

Analysis of Variance↗

Natural selection and genetic variation for reproductive reaction norms in a wild bird population.

Many morphological and life-history traits show phenotypic plasticity that can be described by reaction norms, but few studies have attempted individual-level analyses of reaction norms in the wild. We analyzed variation in individual reaction norms between laying date and three climatic variables (local temperature, local rainfall, and North Atlantic Oscillation) of 1126 female collared flycatchers (Ficedula albicollis) with a restricted maximum likehood linear mixed model approach using random-effect best linear unbiased predictor estimates for the elevation (i.e., expected laying date in the average environment) and slope (i.e., adjustment in laying date as a function of environment) of females' reaction norms. Variation in laying date was best explained by local temperature, and individual females differed in both the elevation and the slope of their laying date-temperature reaction norms. As revealed by animal model analyses, there was weak evidence for additive genetic variance of elevation (h2 +/- SE = 0.09 +/- 0.09), whereas there was no evidence for heritability of slope (h2 +/- SE = 0.00 +/- 0.01). Selection analysis, using a female's lifetime production of fledglings or recruits as an estimate of her fitness, revealed significant selection for a lower phenotypic value and breeding value for elevation (i.e., earlier laying date at the average temperature). There was selection for steeper phenotypic values of slope (i.e., greater plasticity in the adjustment of laying date to temperature), but no significant selection on the breeding values of slope. Although these results suggest that phenotypic laying date is influenced by additive genetic factors, as well as by an interaction with the environment, selection on plasticity would not produce an evolutionary response.

Animals↗

Mapping determinants of gene expression plasticity by genetical genomics in C. elegans.

Recent genetical genomics studies have provided intimate views on gene regulatory networks. Gene expression variations between genetically different individuals have been mapped to the causal regulatory regions, termed expression quantitative trait loci. Whether the environment-induced plastic response of gene expression also shows heritable difference has not yet been studied. Here we show that differential expression induced by temperatures of 16 degrees C and 24 degrees C has a strong genetic component in Caenorhabditis elegans recombinant inbred strains derived from a cross between strains CB4856 (Hawaii) and N2 (Bristol). No less than 59% of 308 trans-acting genes showed a significant eQTL-by-environment interaction, here termed plasticity quantitative trait loci. In contrast, only 8% of an estimated 188 cis-acting genes showed such interaction. This indicates that heritable differences in plastic responses of gene expression are largely regulated in trans. This regulation is spread over many different regulators. However, for one group of trans-genes we found prominent evidence for a common master regulator: a transband of 66 coregulated genes appeared at 24 degrees C. Our results suggest widespread genetic variation of differential expression responses to environmental impacts and demonstrate the potential of genetical genomics for mapping the molecular determinants of phenotypic plasticity.

Animals↗