Ulrich K. Steiner

The fitness costs of developmental canalization and plasticity.

Organisms are capable of an astonishing repertoire of phenotypic responses to the environment, and these often define important adaptive solutions to heterogeneous and unpredictable conditions. The terms ‘phenotypic plasticity’ and ‘canalization’ indicate whether environmental variation has a large or small effect on the phenotype. The evolution of canalization and plasticity is influenced by optimizing selection‐targeting traits within environments, but inherent fitness costs of plasticity may also be important. We present a meta‐analysis of 27 studies (of 16 species of plant and 7 animals) that have measured selection on the degree of plasticity independent of the characters expressed within environments. Costs of plasticity and canalization were equally frequent and usually mild; large costs were observed only in studies with low sample size. We tested the importance of several covariates, but only the degree of environmental stress was marginally positively related to the cost of plasticity. These findings suggest that costs of plasticity are often weak, and may influence phenotypic evolution only under stressful conditions.

Phenotypic Plasticity, Costs of Phenotypes, and Costs of Plasticity

Why are some traits constitutive and others inducible? The term costs often appears in work addressing this issue but may be ambiguously defined. This review distinguishes two conceptually distinct types of costs: phenotypic costs and plasticity costs. Phenotypic costs are assessed from patterns of covariation, typically between a focal trait and a separate trait relevant to fitness. Plasticity costs, separable from phenotypic costs, are gauged by comparing the fitness of genotypes with equivalent phenotypes within two environments but differing in plasticity and fitness. Subtleties associated with both types of costs are illustrated by a body of work addressing predator‐induced plasticity. Such subtleties, and potential interplay between the two types of costs, have also been addressed, often in studies involving genetic model organisms. In some instances, investigators have pinpointed the mechanistic basis of plasticity. In this vein, microbial work is especially illuminating and has three additional strengths. First, information about the machinery underlying plasticity—such as structural and regulatory genes, sensory proteins, and biochemical pathways—helps link population‐level studies with underlying physiological and genetic mechanisms. Second, microbial studies involve many generations, large populations, and replication. Finally, empirical estimation of key parameters (e.g., mutation rates) is tractable. Together, these allow for rigorous investigation of gene interactions, drift, mutation, and selection—all potential factors influencing the maintenance or loss of inducible traits along with phenotypic and plasticity costs. Messages emerging from microbial work can guide future efforts to understand the evolution of plastic traits in diverse organisms.

Optimizing Time and Resource Allocation Trade-Offs for Investment into Morphological and Behavioral Defense

Prey organisms are confronted with time and resource allocation trade‐offs. Time allocation trade‐offs partition time, for example, between foraging effort to acquire resources and behavioral defense. Resource allocation trade‐offs partition the acquired resources between multiple traits, such as growth or morphological defense. We develop a mathematical model for prey organisms that comprise time and resource allocation trade‐offs for multiple defense traits. Fitness is determined by growth and survival during ontogeny. We determine optimal defense strategies for environments that differ in their resource abundance, predation risk, and defense effectiveness. We compare the results with results of simplified models where single defense traits are optimized. Our results indicate that selection acts in favor of integrated traits. The selective advantage of expressing multiple defense traits is most pronounced at intermediate environmental conditions. Optimizing single traits generally leads to a more pronounced response of the defense traits, which implies that studying single traits leads to an overestimation of their response to predation. Behavioral defense and morphological defense compensate for and augment each other depending on predator densities and the effectiveness of the defense mechanisms. In the presence of time constraints, the model shows peak investment into morphological and behavioral defense at intermediate resource levels.

Investment in defense and cost of predator-induced defense along a resource gradient

An organism’s investment in different traits to reduce predation is determined by the fitness benefit of the defense relative to the fitness costs associated with the allocation of time and resources to the defense. Inherent tradeoffs in time and resource allocation should result in differential investment in defense along a resource gradient, but competing models predict different patterns of investment. There are currently insufficient empirical data on changes in investment in defensive traits or their costs along resource gradients to differentiate between the competing allocation models. In this study, I exposed tadpoles to caged predators along a resource gradient in order to estimate investment in defense and costs of defense by assessing predator-induced plasticity. Induced defenses included increased tail depth, reduced feeding, and reduced swimming activity; costs associated with these defenses were reduced developmental rate, reduced growth, and reduced survival. At low resource availability, these costs predominately resulted in reduced survival, while at high resource availability the costs yielded a reduced developmental rate. Defensive traits responded strongly to predation risk, but did not respond to resource availability (with the exception of feeding activity), whereas traits construed as costs of defenses showed the opposite pattern. Therefore, defensive traits were highly sensitive to predation risk, while traits construed as costs of defense were highly sensitive to resource allocation tradeoffs. This difference in sensitivity between the two groups of traits may explain why the correlation between the expression of defensive traits and the expression of the associated defense costs was weak. Furthermore, my results indicate that genetic linkages and mechanistic integration of multiple defensive traits and their associated costs may constrain time and resource allocation in ways that are not addressed in existing models.

Linking antipredator behaviour, ingestion, gut evacuation and costs of predator-induced responses in tadpoles

Many prey organisms reduce their activity to reduce predation risk. A common argument is that a reduction in activity is one of the highest costs of defence. I exposed predator-induced and predator-naive morphs to a short-term predator environment and recorded behavioural, life-historical, physiological, and morphological responses. In contrast to expectation, reduced activity was not one of the highest costs of responding to predators. Predator-exposed tadpoles ingested the same amount of food with less feeding effort and evacuated food from their guts at a higher rate. Despite these advantages, predator-exposed tadpoles still paid costs in responding to predators, in decreased development rate. They did not have reduced survival or reduced growth. Costs in responding to predators are probably caused by physiological factors, such as reduced conversion rates, increased metabolic rates or by allocation to morphological defences, such as increased tail depth. My results show that feeding activity is not linked to the amount of food ingested and that physiological mechanisms, such as gut evacuation, decouple feeding activity and ingestion from growth, development and survival. There was no adaptive response in gut morphology. My study improves the understanding of the underlying internal and physiological mechanisms mediating the trade-off between activity and costs of predator-induced defences.

Dynamic heterogeneity and life history variability in the kittiwake

1. Understanding the evolution of life histories requires an assessment of the process that generates variation in life histories. Within-population heterogeneity of life histories can be dynamically generated by stochastic variation of reproduction and survival or be generated by individual differences that are fixed at birth. 2. We show for the kittiwake that dynamic heterogeneity is a sufficient explanation of observed variation of life histories. 3. The total heterogeneity in life histories has a small contribution from reproductive stage dynamics and a large contribution from survival differences. We quantify the diversity in life histories by metrics computed from the generating stochastic process. 4. We show how dynamic heterogeneity can be used as a null model and also how it can lead to positive associations between reproduction and survival across the life span. 5. We believe our approach to identifying the nature of among-individual heterogeneity yields important insights into the forces that generate within-population variation of life-history traits. It provides an alternative to claims that fixed individual differences are a major determinant of heterogeneity in life histories.

Evolutionary Change in Continuous Reaction Norms

Understanding the evolution of reaction norms remains a major challenge in ecology and evolution. Investigating evolutionary divergence in reaction norm shapes between populations and closely related species is one approach to providing insights. Here we use a meta-analytic approach to compare divergence in reaction norms of closely related species or populations of animals and plants across types of traits and environments. We quantified mean-standardized differences in overall trait means (Offset) and reaction norm shape (including both Slope and Curvature). These analyses revealed that differences in shape (Slope and Curvature together) were generally greater than differences in Offset. Additionally, differences in Curvature were generally greater than differences in Slope. The type of taxon contrast (species vs. population), trait, organism, and the type and novelty of environments all contributed to the best-fitting models, especially for Offset, Curvature, and the total differences (Total) between reaction norms. Congeneric species had greater differences in reaction norms than populations, and novel environmental conditions increased the differences in reaction norms between populations or species. These results show that evolutionary divergence of curvature is common and should be considered an important aspect of plasticity, together with slope. Biological details about traits and environments, including cryptic variation expressed in novel environmental conditions, may be critical to understanding how reaction norms evolve in novel and rapidly changing environments.

Neutral theory for life histories and individual variability in fitness components

Individuals within populations can differ substantially in their life span and their lifetime reproductive success but such realized individual variation in fitness components need not reflect underlying heritable fitness differences visible to natural selection. Even so, biologists commonly argue that large differences in fitness components are likely adaptive, resulting from and driving evolution by natural selection. To examine this argument we use unique formulas to compute exactly the variance in life span and in lifetime reproductive success among individuals with identical (genotypic) vital rates (assuming a common genotype for all individuals). Such individuals have identical fitness but vary substantially in their realized individual fitness components. We show by example that our computed variances and corresponding simulated distribution of fitness components match those observed in real populations. Of course, (genotypic) vital rates in real populations are expected to differ by small but evolutionarily important amounts among genotypes, but we show that such differences only modestly increase variances in fitness components. We conclude that observed differences in fitness components may likely be evolutionarily neutral, at least to the extent that they are indistinguishable from distributions generated by neutral processes. Important consequences of large neutral variation are the following: Heritabilities for fitness components are likely to be small (which is in fact the case), small selective differences in life histories will be hard to measure, and the effects of random drift will be amplified in natural populations by the large variances among individuals.

Predator-Induced Changes in Metabolism Cannot Explain the Growth/Predation Risk Tradeoff

Defence against predators is usually accompanied by declining rates of growth or development. The classical growth/predation risk tradeoff assumes reduced activity as the cause of these declines. However, in many cases these costs cannot be explained by reduced foraging effort or enhanced allocation to defensive structures under predation risk. Here, we tested for a physiological origin of defence costs by measuring oxygen consumption in tadpoles (Rana temporaria) exposed to predation risk over short and long periods of time. The short term reaction was an increase in oxygen consumption, consistent with the “fight-or-flight” response observed in many organisms. The long term reaction showed the opposite pattern: tadpoles reduced oxygen consumption after three weeks exposure to predators, which would act to reduce the growth cost of predator defence. The results point to an instantaneous and reversible stress response to predation risk. This suggests that the tradeoff between avoiding predators and growing rapidly is not caused by changes in metabolic rate, and must be sought in other behavioural or physiological processes.

ChaLearn Looking at People 2015: Apparent Age and Cultural Event Recognition Datasets and Results

Following previous series on Looking at People (LAP) competitions [14, 13, 11, 12, 2], in 2015 ChaLearn ran two new competitions within the field of Looking at People: (1) age estimation, and (2) cultural event recognition, both in still images. We developed a crowd-sourcing application to collect and label data about the apparent age of people (as opposed to the real age). In terms of cultural event recognition, one hundred categories had to be recognized. These tasks involved scene understanding and human body analysis. This paper summarizes both challenges and data, as well as the results achieved by the participants of the competition. Details of the ChaLearn LAP competitions can be found at http://gesture.chalearn.org/.