Biology subjects

Walter, G. M.

Publications and source records attributed to Walter, G. M..

16 recordsLinked to original sources

bioRxiv2026

Why development time matters: temperature influences plasticity while nutritional stress reshapes selection and evolutionary potential

The time it takes organisms to reach adulthood is often considered a central life-history trait that influences fitness and is highly plastic across environments. Despite its importance, direct estimates of how selection on development time varies across combinations of environmental stressors remain rare, limiting our understanding of its role in population responses to environmental change. We reared 68 full-sibling families of the vinegar fly, Drosophila melanogaster, at two temperatures (14{degrees}C and 28{degrees}C) on two diets (standard and low-calorie), and measured development time, body size (mass and wing size), egg-to-adult viability and productivity (number of offspring). Temperature caused the largest plastic response, with development taking 4-5 times longer at 14{degrees}C, whereas the low-calorie diet extended development by 20-30%. Yet diet, not temperature, changed directional selection on development time. On the standard diet, faster development was associated with higher fitness at both temperatures. On the low-calorie diet, this advantage disappeared, and at 14{degrees}C slower-developing females tended to produce more offspring. Viability selection showed consistent patterns. The low-calorie diet also increased evolvability in development time c.10-16-fold at both temperatures, and genetic correlations between diets were close to zero, indicating strong genotype-by-environment interactions. Plasticity in development time was not associated with fitness in any treatment. Overall, we provide evidence that development time is a functional trait whose relationship with fitness depends on the environment. Nutritional stress induced a smaller plastic response in development time than temperature, but had a larger effect on directional selection and evolvability. The stressor causing the greatest plastic change in a functional trait is therefore not necessarily the one that shapes its selection and evolution.

evolutionary biology↗

bioRxiv2026

Interactions between thermal and nutritional stress reveal cryptic adaptive potential

Predicting whether populations can adapt fast enough to persist under global change requires understanding how adaptive potential changes with the environment. Theory predicts that cryptic genetic variation hidden under benign conditions may be released under stress, but empirical tests are rare and typically manipulate single stressors in isolation, which is not realistic under global change. Using a quantitative genetic breeding design with Drosophila melanogaster, we quantified additive genetic variance in fitness (female productivity) across combinations of temperature and nutritional stress. We found that nutritional stress increased genetic variance at cold and hot margins. Temperature and diet also interacted to shape which genotypes performed best: nutritional stress weakened genetic correlations across temperatures at the hot margin, whereas genetic correlations across diets strengthened at the cold margin. Adaptive potential is therefore environment-dependent and multidimensional, meaning single stressor experiments are likely to systematically misrepresent adaptive capacity under realistic global change scenarios.

evolutionary biology↗

bioRxiv2026

Evidence from the field that multiple processes maintain hidden adaptive capacity to a novel environment in a wild daisy

Populations often persist in novel environments despite predictions that adaptive capacity to such conditions should be limited by a lack of genetic variation. A leading hypothesis is that genetic variation for adapting to novel environments is maintained but remains hidden under native conditions. However, direct evidence for the mechanisms that maintain this adaptive potential in natural populations is scarce. Here, we integrate data from four large-scale field experiments to test whether variation in selection across life history and environments, together with genetic architecture, maintains genetic variation important for adapting to novel environments. Using a quantitative genetic breeding design, we generated families of the Sicilian daisy, Senecio chrysanthemifolius (Asteraceae), and planted seeds and cuttings across native and novel elevations on Mount Etna. We tracked fitness across elevations, life stages, seasons and generations. Genotypes with higher survival and flowering success at the novel elevation increased adaptive potential, but were only weakly selected against in the native environment where they had slightly lower fitness at a later life-history stage. A negative genetic correlation in seedling survival across seasons indicated that different genotypes were favoured across temporal variation in native environments. Crosses between genotypes with low and high fitness in the novel environment revealed that genotypes that increased adaptive potential had heritable effects on plasticity and fitness across generations, but were recessive and therefore largely hidden in heterozygotes. Together, these results provide rare field-based evidence that weak selection in native environments, temporal variation in selection and dominance effects act together to maintain cryptic adaptive potential in natural populations.

evolutionary biology↗

bioRxiv2025

Local adaptation in climate tolerance at a small geographic scale contrasts with broad latitudinal patterns

While climate adaptation is typically quantified across broad gradients, the potential for adaptation to the same environmental variables at small scales is rarely tested. If local-scale environmental heterogeneity can generate patterns of adaptation similar to broad gradients, then we currently underestimate adaptive capacity to global change. We quantified population variation in climate tolerance traits and their plasticity for five populations of Drosophila melanogaster from a 3000-km latitudinal gradient, which we contrasted with eight local populations from an environmentally heterogeneous 600x300km area. Population variation in stress tolerance at the local scale was comparable to that across latitude. Consistent with local adaptation, populations from warmer and drier environments showed greater heat and desiccation tolerance, and populations from more predictable environments showed greater plasticity. Climate adaptation at smaller geographic scales can therefore be comparable to adaptation across broad geographic scales. However, patterns of adaptation often changed across geography, which makes predicting responses to global change challenging.

evolutionary biology↗

bioRxiv2025

Variation in temperature but not diet determines the stability of latitudinal clines in tolerance traits and their plasticity

Latitudinal clines are routinely used as evidence of adaptation across broad climatic gradients. However, if environmental variation influences the strength of latitudinal clines, then clinal patterns will be unstable, and using patterns of adaptation to predict population responses to global change will be difficult. To test whether environmental variation influences latitudinal clines, we sampled five populations of Drosophila melanogaster spanning 3000-km of east coast Australia, and measured stress tolerance (heat, cold and desiccation) and body size on flies that developed in six combinations of temperature (13{degrees}C, 25{degrees}C and 29{degrees}C) and diet (standard and low-calorie) treatments. We found latitudinal clines where populations further from the equator had larger wings, higher cold tolerance and lower heat tolerance. For all traits, temperature determined the strength of latitudinal clines, whereas diet had little influence. Steeper clines often emerged in warmer treatments, created by latitudinal clines in plasticity. In the warmest temperature, higher latitude populations showed larger increases in heat tolerance, larger reductions in desiccation tolerance but smaller decreases in cold tolerance. Heat tolerance was the only trait that supported the climate variability hypothesis and a trade-off between plasticity and tolerance. Environment-dependent latitudinal clines are therefore likely to determine variation in population responses to global change.

evolutionary biology↗

bioRxiv2025

Genetic differences in plasticity across environmental scales determine fitness along an ecological gradient

When populations suffer reduced fitness in novel environments, genotypes that better adjust their phenotype to cope with environmental change can aid persistence by reducing the severity of fitness declines. However, we know little about how plastic changes in phenotype allow different genotypes to track environmental variation across ecological gradients, particularly as environments become novel. We transplanted numerous clones of 19 genotypes of a Sicilian daisy, Senecio chrysanthemifolius, at four elevations on Mt Etna. We assessed fitness at native and novel elevations and quantified leaf plasticity among and within elevations. Genotypes with higher fitness at novel elevations showed lower variance in fitness, lower plasticity across elevations, but higher plasticity within elevations compared to those with higher fitness in the native range. Our results suggest that there are genotypes hidden in a population whose plasticity better tracks novel environmental variation at multiple scales, which will be crucial for population persistence under rapid environmental change.

evolutionary biology↗

bioRxiv2023

Variation in plasticity in response to combined thermal-nutritional stress is genetically independent between sexes and across traits

Phenotypic plasticity helps animals to buffer the effects of increasing thermal and nutritional stress created by climate change. Plastic responses to single and combined stressors can vary among genetically diverged populations. However, less is known about how plasticity in response to combined stress varies among individuals within a population or whether such variation changes across life-history traits. This is important because individual variation within populations shapes population-level responses to environmental change. Here, we used isogenic lines of Drosophila melanogaster to assess plasticity of egg-to-adult viability and sex-specific body size for combinations of two temperatures (25{degrees}C or 28{degrees}C) and three diets (standard diet, low caloric diet, or low protein:carbohydrate ratio diet). Our results reveal substantial within-population genetic variation in plasticity for egg-to-adult viability and wing size in response to combined thermal-nutritional stress. This genetic variation in plasticity was a result of cross-environment genetic correlations that were often < 1 for both traits, as well as changes in the expression of genetic variation across environments for egg-to-adult viability. Cross-sex genetic correlations for body size were weaker when the sexes were reared in different conditions, suggesting that the genetic basis of traits may change with the environment. Further, our results suggest that plasticity in egg-to-adult viability is genetically independent from plasticity in body size. Importantly, plasticity in response to diet and temperature individually differed from plastic shifts in response to diet and temperature in combination. By quantifying plasticity and the expression of genetic variance in response to combined stress across traits, our study reveals the complexity of animal responses to environmental change, and the need for a more nuanced understanding of the potential for populations to adapt to ongoing climate change.

evolutionary biology↗

bioRxiv2023

Identifying the proximate mechanisms that generate variation in nutritional plasticity for fecundity in Drosophila melanogaster

Nutrition is an important determinant of an animals survival and fitness. Phenotypic plasticity allows a genotype to adjust life history traits to changes in its nutritional environment, and it varies among individuals. To understand how variation in plasticity is achieved, we made use of a Drosophila melanogaster isogenic panel to characterize nutritional plasticity for fecundity by feeding flies diets differing in their yeast content and counting the number of eggs produced. We then identified lines with the highest and lowest plastic responses to diet and dissected the potential proximate mechanisms responsible for these differences in plasticity, including morphology, behaviour, and physiology. Our results suggest that genetic variation in plasticity is not due to differences in ovariole number, but due to both increased food intake, and higher efficiency at converting food into eggs. Our results show that, in this population of D. melanogaster, variation in behaviour and physiology, but not morphology, underlies differences in plasticity for fecundity. Further, they set the stage for future studies aiming to understand how the proximate mechanisms that generate genetic variation in plasticity contribute to a populations persistence when faced with environmental changes.

evolutionary biology↗

bioRxiv2021

Plasticity in novel environments induces larger changes in genetic variance than adaptive divergence

Adaptive plasticity allows populations to cope with environmental variation but is expected to fail as conditions become unfamiliar. In novel conditions, populations may instead rely on rapid adaptation to increase fitness and avoid extinction. Adaptation should be fastest when both plasticity and selection occur in directions of the multivariate phenotype that contain abundant genetic variation. However, tests of this prediction from field experiments are rare. Here, we quantify how additive genetic variance in a multivariate phenotype changes across an elevational gradient, and test whether plasticity and selection align with genetic variation. We do so using two closely related, but ecologically distinct, sister species of Sicilian daisy (Senecio, Asteraceae) adapted to high and low elevations on Mount Etna. Using a paternal half-sibling breeding design, we generated and then reciprocally planted c.19,000 seeds of both species, across an elevational gradient spanning each species native elevation, and then quantified mortality and five leaf traits of emergent seedlings. We found that genetic variance in leaf traits changed more across elevations than between species. The high-elevation species at novel lower elevations showed changes in the distribution of genetic variance among the leaf traits, which reduced the amount of genetic variance in the directions of selection and the native phenotype. By contrast, the low-elevation species mainly showed changes in the amount of genetic variance at the novel high elevation, and genetic variance was concentrated in the direction of the native phenotype. For both species, leaf trait plasticity across elevations was in a direction of the multivariate phenotype that contained a moderate amount of genetic variance. Together, these data suggest that where plasticity is adaptive, selection on genetic variance for an initially plastic response could promote adaptation. However, large environmental effects on genetic variance are likely to reduce adaptive potential in novel environments.

evolutionary biology↗

bioRxiv2021

Adaptive maternal effects in early life history traits help to maintain ecological resilience in novel environments for two contrasting Senecio species

While many organisms shift their development plastically to maintain fitness as environments change, such plasticity has limits. Population mean fitness is expected to decline when novel environments exceed the limits to plasticity, but the level of fitness costs is expected to vary among genotypes that can increase adaptive potential. We lack fundamental insights into how and when changes in early development traits increase adaptive potential in novel environments, which limits our ability to predict the response of natural populations to global change. To test whether genetic variation in development time is associated with increased adaptive potential in novel environments, we used a breeding design to generate c.20,000 seeds of two ecologically contrasting Sicilian species of daisies (Senecio, Asteraceae) adapted to high and low elevations on Mount Etna. We planted the seeds across four elevations that included the native range of each species, and a novel elevation. We tracked seedling mortality and measured development time as the number of days it took seedlings to establish. As predicted, genetic variance in survival increased at novel elevations. However, genetic variance in development time showed the opposite trend, decreasing at novel elevations. A strong negative genetic correlation between development time in the native range and survival at novel elevations suggested that genotypes with faster development in native environments survived better in novel environments. These results were consistent across the two ecologically contrasting species, suggesting that genetic variance in early development in native environments could be used to predict population responses to novel environments.

evolutionary biology↗

bioRxiv2020

Hidden genetic variation in plasticity increases the potential to adapt to novel environments

Rapid environmental change is forcing populations into novel environments where plasticity will no longer maintain fitness. When populations are exposed to novel environments, evolutionary theory predicts that genetic variation in fitness will increase and should be associated with genetic differences in plasticity. If true, then genetic variation in plasticity can increase adaptive potential in novel environments, and population persistence via rapid adaptation is more likely. To test whether genetic variation in fitness increases in novel environments and is associated with plasticity, we transplanted 8,149 clones of 314 genotypes of a Sicilian daisy (Senecio chrysanthemifolius) within and outside its native range, and quantified genetic variation in fitness, and plasticity in leaf traits and gene expression. Although mean fitness declined by 87% in the novel environment, genetic variance in fitness increased threefold and was correlated with plasticity in leaf traits. High fitness genotypes showed greater plasticity in gene expression, but lower plasticity in most leaf traits. Interestingly, genotypes with greater fitness in the novel environment had the lowest fitness at the native site. These results suggest that standing genetic variation in plasticity could help populations to persist and adapt to novel environments, despite remaining hidden in native environments.

evolutionary biology↗

bioRxiv2020

Ecology and evolutionary history determine plastic responses to environmental variation in two closely related species

The evolution of plastic responses to external cues allows species to track the environmental variation they regularly experience. However, it remains unclear how plasticity evolves during adaptation. To test whether distinct patterns of plasticity is associated with recent adaptive divergence, we quantified plasticity for two closely related but ecologically divergent Sicilian daisy species (Senecio, Asteraceae). We sampled c.40 genotypes of each species from natural populations on and around Mt Etna and then reciprocally transplanted multiple clones of each genotype into four field sites along an elevational gradient representing each species native range, the edge of their range, and conditions outside their native range. At each elevation we quantified survival and measured leaf traits that included investment (specific leaf area), morphology, chlorophyll fluorescence, pigment content and gene expression. As evidence of adaptive divergence, both species performed better at their native site and better than the species from the other habitat. Traits and differentially expressed genes that changed with elevation in one species often showed little change in the other species, or changed in the opposite direction. Adaptive divergence is therefore associated with the evolution of distinct plastic responses to environmental variation, despite these two species sharing a recent common ancestor.

evolutionary biology↗

bioRxiv2019

Divergence in hormone signalling links local adaptation and hybrid failure

Natural selection is a significant driver of speciation. Yet it remains largely unknown whether local adaptation can drive speciation through the evolution of hybrid sterility between populations. Here, we show that adaptive divergence in shoot gravitropism, the ability of a plants shoot to bend upwards in response to the downward pull of gravity, contributes to the evolution of hybrid sterility in an Australian wildflower, Senecio lautus. We find that shoot gravitropism has evolved multiple times in association with plant height between adjacent populations inhabiting contrasting environments, suggesting that these traits have evolved by natural selection. We directly tested this prediction using a hybrid population subjected to eight rounds of recombination and three rounds of selection in the field. It revealed that shoot gravitropism responds to natural selection in the expected direction of the locally adapted population. This provided an ideal platform to test whether genetic differences in gravitropism contribute to hybrid sterility in S. lautus. Using this advanced hybrid population, we discovered that crossing individuals with extreme differences in gravitropism reduce their ability to produce seed by 21%, providing strong evidence that this adaptive trait is genetically correlated with hybrid sterility. Our results suggest that natural selection can drive the evolution of locally adaptive traits that also create hybrid sterility, thus indicating an evolutionary connection between local adaptation and the origin of new species. Significance statementNew species originate as populations become reproductively isolated from one another. Despite recent progress in uncovering the genetic basis of reproductive isolation, it remains unclear whether intrinsic reproductive barriers, such as hybrid sterility, evolve as a by-product of local adaptation to contrasting environments or evolve through non-ecological processes, such as meiotic drive. Here, we show that differences in a plants response to the pull of gravity have repeatedly evolved amongst coastal populations of an Australian wildflower, thus implicating a role of natural selection in their evolution. We found a strong genetic correlation between variation in this adaptive trait and hybrid sterility, suggesting that intrinsic reproductive barriers contribute to the origin of new species as populations adapt to heterogeneous environments.

evolutionary biology↗

bioRxiv2019

Evidence for mutation-order speciation between convergent ecotypes

Natural selection shapes how new species arise, yet the mechanisms that generate reproductive barriers remain debated. Although ecological divergence in contrasting environments and mutation-order processes in similar environments are often viewed as distinct speciation mechanisms, we show they act together as part of a continuum we call parallel mosaic speciation. In the Senecio lautus species complex, Dune and Headland ecotypes have evolved repeatedly along the Australian coastline. Through crossing experiments and field studies, we find that divergent natural selection promotes strong reproductive isolation between the Dune and Headland ecotypes. While uniform selection maintains reproductive compatibility among ecologically similar Dune populations, Headland populations have evolved reproductive barriers despite their convergent prostrate phenotypes, likely driven by adaptation to heterogeneous environments. To understand how habitat heterogeneity contributes to patterns of reproductive isolation, we extend previous theoretical work on the accumulation of hybrid incompatibilities to account for environmental gradients and polygenic adaptation. We show that the probability of reproductive isolation depends on three factors: how similar the environments are, how complex the genetic architecture is, and how selection coefficients are distributed among beneficial mutations. These theoretical findings explain how reproductive isolation arises in systems like Senecio, where multiple forms of selection jointly drive parallel speciation.

evolutionary biology↗

bioRxiv2019

Connecting micro and macroevolution using genetic incompatibilities and natural selection on additive genetic variance

Testing whether local adaptation and intrinsic reproductive isolation share a genetic basis can reveal important connections between adaptation and speciation. Local adaptation arises as advantageous alleles spread through a population, but whether these same advantageous alleles fail on the genetic backgrounds of other populations remains largely unknown. We used a quantitative genetic breeding design to produce a late generation (F4) recombinant hybrid population by equally mating amongst four contrasting ecotypes of a native Australian daisy for four generations. We tracked fitness across generations and measured morphological traits in the glasshouse, and used a reciprocal transplant to quantify fitness in all four parental habitats. In the glasshouse, plants in the second generation showed a reduction in fitness as a loss of fertility, but this was fully recovered in the following generation. The F4 hybrid lacked extreme phenotypes present in the parental ecotypes, suggesting that genes reducing hybrid fitness were linked to traits specific to each ecotype. In the natural habitats, additive genetic variance for fitness was greatest for habitats that showed stronger native-ecotype advantage, suggesting that a loss of genetic variation present in the parental ecotypes were adaptive in the natural habitats. Reductions in genetic variance for fitness were associated with a loss of ecological trade-offs previously described in the parental ecotypes. Furthermore, natural selection on morphological traits differed amongst the parental habitats, but was not predicted to occur towards the morphology of the parental ecotypes. Together, these results suggest that intrinsic reproductive isolation removed adaptive genetic variation present in the parental ecotypes. The evolution of these distinct ecotypes was likely governed by genetic variation that resulted in both ecological trade-offs and intrinsic reproductive isolation among populations adapted to contrasting environments.

evolutionary biology↗

bioRxiv2017

Evolution of genetic variance during adaptive radiation

Genetic correlations between traits can bias adaptation away from optimal phenotypes and constrain the rate of evolution. If genetic correlations between traits limit adaptation to contrasting environments, rapid adaptive divergence across a heterogeneous landscape may be difficult. However, if genetic variance can evolve and align with the direction of natural selection, then abundant allelic variation can promote rapid divergence during adaptive radiation. Here, we explored adaptive divergence among ecotypes of an Australian native wildflower by quantifying divergence in multivariate phenotypes of populations that occupy four contrasting environments. We investigated differences in multivariate genetic variance underlying morphological traits and examined the alignment between divergence in phenotype and divergence in genetic variance. We found that divergence in mean multivariate phenotype has occurred along two major axes represented by different combinations of plant architecture and leaf traits. Ecotypes also showed divergence in the level of genetic variance in individual traits, and the multivariate distribution of genetic variance among traits. Divergence in multivariate phenotypic mean aligned with divergence in genetic variance, with most of the divergence in phenotype among ecotypes associated with a change in trait combinations that had substantial levels of genetic variance in each ecotype. Overall, our results suggest that divergent natural selection acting on high levels of standing genetic variation might fuel ecotypic differentiation during the early stages of adaptive radiation.

evolutionary biology↗