The timing of seed germination determines the environment experienced by a plant’s most vulnerable life stage—the seedling. Germination is environmentally cued, and genotypes can differ in their sensitivity to environmental cues. When genotypes differ in their response to cues, and when cues accurately predict the postgermination environment, the postgermination environment experienced by seedlings can itself have a genetic basis and potential to evolve. We tested for genetic differences in the postgermination environment using Arabidopsis thaliana genotypes that vary in seed dormancy, a trait known to alter germination time. We dispersed seeds into the field in 5 seasonal cohorts over 1.5 years, observed germination timing for 5297 individuals, and measured the soil temperature and moisture experienced by individuals throughout their life cycle. We found that genotypes differed in the environments they experienced during seedling establishment. This environmental variation occurred because genotypes differed in their environmental sensitivity to germination cues, and pregermination cues were correlated with postgermination environments. Seeds exhibited temporal habitat selection by germinating into a nonrandom subset of environmental conditions available, and seed dormancy increased the consistency of habitat selection. Strikingly, the postgermination environment affected fitness by altering the probability of seedling survival such that genotypes that engaged in stronger habitat selection were less likely to reach reproduction. Our results suggest that environmentally cued development may be a widespread mechanism by which genotypes can differ in the environment they experience, introducing the possibility that the environment itself can be inherited and can evolve.
The feeding ecology of the small-bodied benthic naked goby Gobiosoma bosc, a western Atlantic species that occurs in estuaries and other inshore habitats from Connecticut to Texas U.S.A., was investigated in a total of four estuaries spanning South Carolina, North Carolina, Maryland and New Jersey. Gut content analysis of 391 individuals revealed that G. bosc is a benthic microcarnivore that feeds primarily on polychaetes, gammarid amphipods and harpacticoid copepods. Diet composition varied with body size, tidal creek within an estuary and geographic region. Analyses of gut fullness suggest that G. bosc is a daytime visual predator and that nest and egg guarding during the reproductive season reduce foraging activity in mature males. Additionally, G. bosc infected with adult digenean parasites of the gut foraged more intensely than uninfected individuals, a relationship that was strongest for reproductively mature males. Regionally, significant variation in dietary breadth was documented and may reflect a foraging response to a decrease in prey diversity moving from estuaries of higher salinity and lower latitude to estuaries of lower salinity and higher latitude. These results contribute to an understanding of the life history of G. bosc and the role played by this common species in estuarine food webs.
Evolvable traits of organisms can alter the environment those organisms experience. While it is well appreciated that those modified environments can influence natural selection to which organisms are exposed, they can also influence the expression of genetic variances and covariances of traits under selection. When genetic variance and covariance change in response to changes in the evolving, modified environment, rates and outcomes of evolution also change. Here we discuss the basic mechanisms whereby organisms modify their environments, review how those modified environments have been shown to alter genetic variance and covariance, and discuss potential evolutionary consequences of such dynamics. With these dynamics, responses to selection can be more rapid and sustained, leading to more extreme phenotypes, or they can be slower and truncated, leading to more conserved phenotypes. Patterns of correlated selection can also change, leading to greater or less evolutionary independence of traits, or even causing convergence or divergence of traits, even when selection on them is consistent across environments. Developing evolutionary models that incorporate changes in genetic variances and covariances when environments themselves evolve requires developing methods to predict how genetic parameters respond to environments—frequently multifactorial environments. It also requires a population-level analysis of how traits of collections of individuals modify environments for themselves and/or others in a population, possibly in spatially explicit ways. Despite the challenges of elucidating the mechanisms and nuances of these processes, even qualitative predictions of how environment-modifying traits alter evolutionary potential are likely to improve projections of evolutionary outcomes.
The timing of a developmental transition (phenology) can influence the environment experienced by subsequent life stages. When phenology causes an organism to occupy a particular habitat as a consequence of the developmental cues used, it can act as a form of habitat tracking. Evolutionary theory predicts that habitat tracking can alter the strength, direction, and mode of natural selection on subsequently expressed traits.To test whether germination phenology altered natural selection on postgermination traits, we manipulated germination time by planting seedlings in seven germination cohorts spanning 2 yr. We measured selection on postgermination traits relating to drought, freezing, and heat tolerance using a diverse combination of Arabidopsis thaliana mutants and naturally occurring ecotypes.Germination cohorts experienced variable selection: when dry, cold, and hot environments were experienced by seedlings, selection was intensified for drought, freezing, and heat tolerance, respectively. Reciprocally, postgermination traits modified the optimal germination time; genotypes had maximum fitness after germinating in environments that matched their physiological tolerances.Our results support the theoretical predictions of feedbacks between habitat tracking and traits expressed after habitat selection. In natural populations, whether phenological shifts alter selection on subsequently expressed traits will depend on the effectiveness of habitat tracking through phenology.
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