Detection vs. selection: integration of genetic,epigenetic and environmental cues in fluctuating environments

There are many inputs during development that influence an organism's fit to current or upcoming environments. These include genetic effects, transgenerational epigenetic influences, environmental cues and developmental noise, which are rarely investigated in the same formal framework. We study an analytically tractable evolutionary model, in which cues are integrated to determine mature phenotypes in fluctuating environments. Environmental cues received during development and by the mother as an adult act as detection‐based (individually observed) cues. The mother's phenotype and a quantitative genetic effect act as selection‐based cues (they correlate with environmental states after selection). We specify when such cues are complementary and tend to be used together, and when using the most informative cue will predominate. Thus, we extend recent analyses of the evolutionary implications of subsets of these effects by providing a general diagnosis of the conditions under which detection and selection‐based influences on development are likely to evolve and coexist.     

The evolution of phenotypic plasticity in spatially structured environments: implications of intraspecific competition, plasticity costs and environmental characteristics

We model the evolution of reaction norms focusing on three aspects: frequency-dependent selection arising from resource competition, maintenance and production costs of phenotypic plasticity, and three characteristics of environmental heterogeneity (frequency of environments, their intrinsic carrying capacity and the sensitivity to phenotypic maladaptation in these environments). We show that (i) reaction norms evolve so as to trade adaptation for acquiring resources against cost avoidance; (ii) maintenance costs cause reaction norms to better adapt to frequent rather than to infrequent environments, whereas production costs do not; and (iii) evolved reaction norms confer better adaptation to environments with low rather than with high intrinsic carrying capacity. The two previous findings contradict earlier theoretical results and originate from two previously unexplored features that are included in our model. First, production costs of phenotypic plasticity are only incurred when a given phenotype is actually produced. Therefore, they are proportional to the frequency of environments, and these frequencies thus affect the selection pressure to avoid costs just as much as the selection pressure to improve adaptation. This prevents the frequency of environments from affecting the evolving reaction norm. Secondly, our model describes the evolution of plasticity for a phenotype determining an individual's capability to acquire resources, and thus its realized carrying capacity. When individuals are distributed randomly across environments, they cannot avoid experiencing environments with intrinsically low carrying capacity. As selection pressures arising from the need to improve adaptation are stronger under such extreme conditions than under mild ones, better adaptation to environments with low rather than with high intrinsic carrying capacity results.     

Optimal germination timing in unpredictable environments: the importance of dormancy for both among‐ and within‐season variation

For organisms living in unpredictable environments, timing important life‐history events is challenging. One way to deal with uncertainty is to spread the emergence of offspring across multiple years via dormancy. However, timing of emergence is not only important among years, but also within each growing season. Here, we study the evolutionary interactions between germination strategies that deal with among‐ and within‐season uncertainty. We use a modelling approach that considers among‐season dormancy and within‐season germination phenology of annual plants as potentially independent traits and study their separate and joint evolution in a variable environment. We find that higher among‐season dormancy selects for earlier germination within the growing season. Furthermore, our results indicate that more unpredictable natural environments can counter‐intuitively select for less risk‐spreading within the season. Furthermore, strong priority effects select for earlier within‐season germination phenology which in turn increases the need for bet hedging through among‐season dormancy.     

Seasonal development and variation in abundance among four annual flight periods in a butterfly: a 20‐year study of the speckled wood (Pararge aegeria)

Insects typically spend the winter in a species‐specific diapause stage. The speckled wood butterfly, Pararge aegeria, is unique in having two alternative diapause stages, hibernating as larvae or pupae. In southern Sweden this creates a seasonal flight pattern with four annual adult flight periods: the first in May (pupal diapause), the second in June (larval diapause), and the third and fourth directly developing offspring generations in July and August, respectively. We address the raison d'être of the two diapause pathways by (1) outdoor rearing of cohorts, and (2) performing transect censuses throughout the season for 20 years. We contend that an early start of next season provides a benefit accruing to pupal diapause; conversely, a large proportion of the offspring from adults of the fourth flight peak are unable to reach the pupal stage before winter, providing a benefit accruing to larval winter diapause. The results obtained show that the two hibernation pathways are unlikely to be genetically distinct because of a strong overlap between the two offspring generations, and because sibling offspring from the third and fourth flight periods are likely to choose either of the two hibernation pathways, thereby resulting in a genetic mixing of the pathways. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 635–649.     

Evolution of phenotypic plasticity: Genetic differentiation and additive genetic variation for induced plant defence in wild arugula Eruca sativa

Phenotypic plasticity is the primary mechanism of organismal resilience to abiotic and biotic stress, and genetic differentiation in plasticity can evolve if stresses differ among populations. Inducible defence is a common form of adaptive phenotypic plasticity, and long‐standing theory predicts that its evolution is shaped by costs of the defensive traits, costs of plasticity and a trade‐off in allocation to constitutive versus induced traits. We used a common garden to study the evolution of defence in two native populations of wild arugula Eruca sativa (Brassicaceae) from contrasting desert and Mediterranean habitats that differ in attack by caterpillars and aphids. We report genetic differentiation and additive genetic variance for phenology, growth and three defensive traits (toxic glucosinolates, anti‐nutritive protease inhibitors and physical trichome barriers) as well their inducibility in response to the plant hormone jasmonic acid. The two populations were strongly differentiated for plasticity in nearly all traits. There was little evidence for costs of defence or plasticity, but constitutive and induced traits showed a consistent additive genetic trade‐off within each population for the three defensive traits. We conclude that these populations have evolutionarily diverged in inducible defence and retain ample potential for the future evolution of phenotypic plasticity in defence.     

The consequences of parental age for development,body mass and resistance to stress in the red flour beetle

The performance of most animals deteriorates with age. Motivated by the inconsistency in the literature regarding the effect of parental age on offspring traits and performance, we studied how parental age affects offspring development time, body mass, and starvation and cold tolerance in the red flour beetle (Tribolium castaneum). Offspring of old parents pupated later and at a higher body mass, and there was a general positive correlation between body mass and starvation tolerance. Despite their higher body mass, offspring of old parents tolerated starvation less well than those of young parents, emphasizing the impaired performance of the former. However, parental age did not affect offspring thermal tolerance and offspring of old parents were not more sensitive to cold shock than those of young parents. We also examined how ageing affects body mass and cold tolerance in the parental generations. By contrast to the effect of ontogeny on thermal tolerance, which is better known, change in thermal tolerance with age is seldom studied and can take different shapes. Old beetles were more sensitive to cold shock than younger beetles. Similar to cold tolerance, body mass decreased with age. In summary, older beetles reflect a worse physiological condition than younger ones. Ageing leads to impaired cold tolerance, lower body mass, lower number of offspring reaching adulthood, and deteriorated performance of the offspring, expressed as a lower starvation tolerance and a longer development time of the offspring. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 305–314.     

Environmental determinants of population divergence in life‐history traits for an invasive species: climate,seasonality and natural enemies

Invasive species cope with novel environments through both phenotypic plasticity and evolutionary change. However, the environmental factors that cause evolutionary divergence in invasive species are poorly understood. We developed predictions for how different life‐history traits, and plasticity in those traits, may respond to environmental gradients in seasonal temperatures, season length and natural enemies. We then tested these predictions in four geographic populations of the invasive cabbage white butterfly (Pieris rapae) from North America. We examined the influence of two rearing temperatures (20 and 26.7 °C) on pupal mass, pupal development time, immune function and fecundity. As predicted, development time was shorter and immune function was greater in populations adapted to longer season length. Also, phenotypic plasticity in development time was greater in regions with shorter growing seasons. Populations differed significantly in mean and plasticity of body mass and fecundity, but these differences were not associated with seasonal temperatures or season length. Our study shows that some life‐history traits, such as development time and immune function, can evolve rapidly in response to latitudinal variation in season length and natural enemies, whereas others traits did not. Our results also indicate that phenotypic plasticity in development time can also diverge rapidly in response to environmental conditions for some traits.     

Egg‐size plasticity in Apis mellifera: Honey bee queens alter egg size in response to both genetic and environmental factors

Social evolution has led to distinct life‐history patterns in social insects, but many colony‐level and individual traits, such as egg size, are not sufficiently understood. Thus, a series of experiments was performed to study the effects of genotypes, colony size and colony nutrition on variation in egg size produced by honey bee (Apis mellifera) queens. Queens from different genetic stocks produced significantly different egg sizes under similar environmental conditions, indicating standing genetic variation for egg size that allows for adaptive evolutionary change. Further investigations revealed that eggs produced by queens in large colonies were consistently smaller than eggs produced in small colonies, and queens dynamically adjusted egg size in relation to colony size. Similarly, queens increased egg size in response to food deprivation. These results could not be solely explained by different numbers of eggs produced in the different circumstances but instead seem to reflect an active adjustment of resource allocation by the queen in response to colony conditions. As a result, larger eggs experienced higher subsequent survival than smaller eggs, suggesting that honey bee queens might increase egg size under unfavourable conditions to enhance brood survival and to minimize costly brood care of eggs that fail to successfully develop, and thus conserve energy at the colony level. The extensive plasticity and genetic variation of egg size in honey bees has important implications for understanding life‐history evolution in a social context and implies this neglected life‐history stage in honey bees may have trans‐generational effects.     

Lake‐specific variation in growth,migration timing and survival of juvenile sockeye salmon Oncorhynchus nerka: separating environmental from genetic influences

Time series on juvenile life‐history traits obtained from sockeye salmon Oncorhynchus nerka were analysed to assess lake‐specific environmental influences on juvenile migration timing, size and survival of fish from a common gene pool. Every year for the past two decades, O. nerka have been spawned at a hatchery facility, and the progeny released into two lakes that differ in average summer temperatures, limnological attributes and growth opportunities. Juveniles reared in the warmer, more productive Crosswind Lake were larger and heavier as smolts compared to those from the cooler, less productive Summit Lake and had higher in‐lake and subsequent marine survival. Crosswind Lake smolts migrated from the lake to sea slightly earlier in the season but the migration timing distributions overlapped considerably across years. Fry stocking density had a negative effect on smolt length for both lakes, and a negative effect on in‐lake survival in Summit Lake. Taken together, the results revealed a strong effect of lake‐rearing environment on the expression of life‐history variation in O. nerka. The stocking of these lakes each year with juveniles from a single mixed‐source population provided a large‐scale reverse common‐garden experiment, where the same gene pool was exposed to different environments, rather than the different gene pools in the same environment approach typical of evolutionary ecology studies. Other researchers are encouraged to seek and exploit similar serendipitous situations, which might allow environmental and genetic influences on ecologically important traits to be distinguished in natural or semi‐natural settings.