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Life history theory predicts that investment per offspring should correlate negatively with the\ud quality of environment offspring are anticipated to encounter; parents may use their own\ud experience as juveniles to predict this environment and may modulate offspring traits such as\ud growth capacity as well as initial size. We manipulated nutrient levels in the juvenile habitat of\ud wild Atlantic salmon Salmo salar to investigate the hypothesis that the egg size maximizing\ud juvenile growth and survival depends on environmental quality. We also tested whether\ud offspring traits were related to parental growth trajectory. Mothers that grew fast when young\ud produced more, smaller offspring than mothers that had grown slowly to reach the same size.\ud Despite their size disadvantage, offspring of faster-growing mothers grew faster than those of\ud slow-growing mothers in all environments, counter to the expectation that they would be\ud competitively disadvantaged. However, they had lower relative survival in environments where\ud the density of older predatory/competitor fish was relatively high. These links between maternal\ud (but not paternal) growth trajectory and offspring survival rate were independent of egg size,\ud underscoring that mothers may be adjusting egg traits other than size to suit the anticipated\ud environment faced by their offspring.

Despite considerable theoretical interest in how the evolution of phenotypic plasticity should be shaped by environmental variability and stochasticity, how individuals actually respond to these aspects of the environment within their own lifetimes remains unclear. We propose that this understanding has been hampered by experimental approaches that expose organisms to fluctuating environments (typically treatments where fluctuations in the environment are cyclical vs. erratic) for a pre‐determined duration while ensuring that the mean environment over that the entire exposure period is invariable. This approach implicitly assumes that responses to the mean and variance/predictability in the environment occur over the same time‐scale. If this assumption is false, one potential outcome is that phenotypic differences among the treatment groups might arise in response to differences in the mean environment that are present over shorter time periods among those same treatment groups. We illustrate an experimental design that (a) creates variation in the level of environmental predictability, (b) allows for estimation of the time‐scale over which the phenotypic response to the mean environment occurs and (c) permits statistical estimation of the effect of predictability in the environmental variable of interest while controlling for any effect of the mean environment over the relevant temporal scale. Using the clonally reproducing zooplankton species Daphnia magna, we test for within‐generation plasticity in the ability to tolerate high temperature following exposure to multiple temperature treatments with the same overall mean, but where the pattern of fluctuations differed among them. This approach revealed that heat tolerance in Daphnia was not influenced by variability in temperature per se nor the predictability of fluctuations in temperature but adjusted in response to the mean temperature they experienced 24 hr prior to measurement. Our results suggest that conclusions arising from studies that employ a single manipulation of environmental predictability and which cannot consider such potentially confounding effects may be premature. A free Plain Language Summary can be found within the Supporting Information of this article.

Using data from wild Atlantic salmon Salmo salar returning to spawn in seven Scottish rivers, we developed a model of fecundity based on individual body size and key developmental traits. We used a novel approach to model selection which maximises predictive accuracy for application to target river stocks to select the best from a suite of Bayesian hierarchical models. This approach aims to ensure the optimal model within the candidate set includes covariates that best predict out-of-sample data to estimate fecundity in areas where no direct observations are available. In addition to body size, the final model included the developmental characteristics of age at smolting and years spent at sea. Using two independent long-term monitoring datasets, the consequences of ignoring these characteristics was revealed by comparing predictions from the best model with models that omitted them.

Widespread declines in the body size of aquatic ectotherms have been attributed to the poorer ability of older, larger individuals to tolerate high temperature. Here, using the thermal death time curve framework, we investigate the relationship between temperature tolerance and size/age by measuring the change in heat tolerance of the keystone zooplankton species Daphnia magna across a range of temperature intensities (and hence exposures of varying duration) among individuals that differed up to 3-fold in size and thus varied in age also. Across the gradient of exposure temperatures, younger, smaller individuals were more tolerant than older, larger individuals. This suggests that the young and the small may be better equipped to withstand temperature challenges that are both intense/brief and more moderate/prolonged. Our study generalizes results obtained from more acute tolerance assays, providing physiological evidence consistent with the observed reductions in ectotherm body size as a response to warming in aquatic systems.

When a change in the environment occurs, organisms can maintain an optimal phenotypic state via plastic, reversible changes to their phenotypes. These adjustments, when occurring within a generation, are described as the process of acclimation. While acclimation has been studied for more than half a century, global environmental change has stimulated renewed interest in quantifying variation in the rate and capacity with which this process occurs, particularly among ectothermic organisms. Yet, despite the likely ecological importance of acclimation capacity and rate, how these traits change throughout life among members of the same species is largely unstudied. Here we investigate these relationships by measuring the acute heat tolerance of the clonally reproducing zooplankter Daphnia magna of different size/age and acclimation status. The heat tolerance of individuals completely acclimated to relatively warm (28°C) or cool (17°C) temperatures diverged during development, indicating that older, larger individuals had a greater capacity to increase heat tolerance. However, when cool acclimated individuals were briefly exposed to the warm temperature (i.e. were ‘heat-hardened'), it was younger, smaller animals with less capacity to acclimate that were able to do so more rapidly because they obtained or came closer to obtaining complete acclimation of heat tolerance. Our results illustrate that within a species, individuals can differ substantially in how rapidly and by how much they can respond to environmental change. We urge greater investigation of the intraspecific relationship between acclimation and development along with further consideration of the factors that might contribute to these enigmatic patterns of phenotypic variation.

A shift from coral to macroalgae dominance of reef systems affected by volcanically acidified waters around Maug (Mariana Islands, North Pacific Ocean) increases fears that reef corals will be displaced by algae as a result of ocean acidification.

Both the environments experienced by a mother as a juvenile and an adult can affect her investment in offspring. However, the implications of these maternal legacies, both juvenile and adult, for offspring fitness in natural populations are unclear. We investigated whether the juvenile growth rate and adult reproductive traits (length, body condition, and reproductive investment at spawning) of female wild Atlantic salmon (Salmo salar) were related to the growth and survival of their offspring. Adult salmon captured on their upstream migration were used to create experimental full-sib clutches of eggs, which were mixed and then placed in artificial nests in a natural stream that lacked salmon due to a migration barrier. Four months later we resampled the stream to obtain family-level estimates of offspring size and survival. Mothers that had grown slowly as juveniles (as determined by scalimetry) but had invested heavily in reproduction (egg production for a given body length) and were in relatively poor body condition (somatic mass for a given body length) at spawning produced the largest eggs. Larger eggs resulted in larger juveniles and higher juvenile survival. However, after controlling for egg size, offspring growth was positively related to maternal juvenile growth rate and reproductive investment. The predictors of offspring survival (i.e., reproductive success) varied with the juvenile growth rate of the mother: If females grew slowly as juveniles, their reproductive success was negatively related to their own body condition. In contrast, the reproductive success of females that grew quickly as juveniles was instead related positively to their own body condition. Our results show that maternal influences on offspring in the wild can be complex, with reproductive success related to the early life performance of the mother, as well as her state at the time of breeding.

We investigated the independent effects of age at smolting and body condition at the time of spawning on egg production by female Atlantic salmon (Salmo salar). For a given body size, female salmon that had smolted as juveniles after 2 years in fresh water produced smaller, more numerous eggs than females that smolted 1 year later. Furthermore, fecundity (but not egg size) was related positively to maternal body condition at spawning. Given that age at smolting is closely related to juvenile growth rate; results from this study suggest that conditions experienced by female Atlantic salmon during both early life and adulthood have implications for the size and number of eggs that they produce.

Individual differences in the energy cost of self-maintenance (resting metabolic rate, RMR) are substantial and the focus of an emerging research area. These differences may influence fitness because self-maintenance is considered as a life-history component along with growth and reproduction. In this review, we ask why do some individuals have two to three times the ‘maintenance costs’ of conspecifics, and what are the fitness consequences? Using evidence from a range of species, we demonstrate that diverse factors, such as genotypes, maternal effects, early developmental conditions and personality differences contribute to variation in individual RMR. We review evidence that RMR is linked with fitness, showing correlations with traits such as growth and survival. However, these relationships are modulated by environmental conditions (e.g. food supply), suggesting that the fitness consequences of a given RMR may be context-dependent. Then, using empirical examples, we discuss broad-scale reasons why variation in RMR might persist in natural populations, including the role of both spatial and temporal variation in selection pressures and trans-generational effects. To conclude, we discuss experimental approaches that will enable more rigorous examination of the causes and consequences of individual variation in this key physiological trait.

This study demonstrated that, irrespective of hormone type or dose, administering cocoa butter implants during egg development affected the growth of female brown trout Salmo trutta and reduced the size of their offspring. Cortisol treatment also increased adult mortality. Caution is urged in the use of implants for studies of maternal hormonal influences on adult fishes and their offspring. (C) 2011 Crown Copyright Journal of Fish Biology (C) 2011 The Fisheries Society of the British Isles

Summary 1. Social status can vary considerably among individuals and has significant implications for performance. In addition to a genetic component, social status may be influenced by environmental factors including maternal effects such as prenatal hormone exposure. Maternal effects on traits determining social status have previously been examined in species where mothers provide parental care for relatively few offspring and therefore directly influence postnatal development. However, the generality of conclusions arising from these investigations is unclear because species that employ different reproductive strategies have not been studied. 2. We investigated the hypothesis that egg steroid hormone levels influence the social status of juvenile brown trout (Salmo trutta). We manipulated intra-clutch levels of cortisol and testosterone in eggs from 15 mothers using dilute hormone baths at the time of fertilization and examined their effects on traits that correlate with social status in juveniles [including standard body size, aggression, competitive ability and standard metabolic rate (SMR)]. 3. Hormone treatment did not affect whole-animal or mass-corrected SMR at the critical developmental stage when juveniles switch from reliance on a maternally provisioned yolk sac to independent feeding. However, juveniles from cortisol-treated eggs were smaller at this developmental stage. They were also less aggressive than, and subordinate to, fish from untreated eggs in socially competitive conditions, even after correcting for the observed effect of cortisol on body size. Egg testosterone treatment resulted in a likely pharmacological or toxicological dose with subsequent effects on both body size and behaviour in independently feeding juveniles. 4. Results from this study demonstrate that variation in the amount of cortisol deposited in eggs by spawning females influences juvenile social status and performance. The effects of elevated egg cortisol in fish are similar to the actions of embryonic glucocorticoids reported in other vertebrate taxa with very different reproductive strategies, suggesting a widespread mechanism for the effects of maternal stress on offspring. Possible adaptive aspects of this relationship are discussed.

Fertilised eggs, stage one and stage two juveniles of the smooth marron, Cherax cainii, are attached to the pleopods of the gravid female via specialised structures. The eggs are fixed to the maternal pleopods by egg stalks, whereas stage one and stage two juveniles remain attached by recurved spines on the dactyls of their fourth and fifth pereopods. Crayfish of the third juvenile stage are independent of the mother and receive no maternal care; their dactyl spines are straight and cannot grasp the mother's pleopods; the female displays agonistic behaviour toward her young commencing with stage III instars. We suggest that eggs and the first two juvenile stages attached to the mother would experience reduced levels of predation and be secured against strong river flow. We conclude by reviewing, briefly, the potential fitness costs and benefits of maternal care, to the mother and young of freshwater crayfish, from a life-history perspective.

We investigated whether among-sibling differences in the phenotypes of juvenile fish were systematically related to the position in the egg mass where each individual developed during oogenesis. We sampled eggs from the front, middle and rear thirds of the egg mass in female brown trout of known dominance rank. In the resulting juveniles, we then measured traits that are related to individual fitness: body size, social status and standard metabolic rate (SMR). When controlling for differences among females in mean egg size, siblings from dominant mothers were initially larger (and had a lower mass-corrected SMR) if they developed from eggs at the rear of the egg mass. However, heterogeneity in the size of siblings from different positions in the egg mass diminished in lower-ranking females. Location of the egg within the egg mass also affected the social dominance of the resulting juvenile fish, although the direction of this effect varied with developmental age. This study provides the first evidence of a systematic basis for among-sibling differences in the phenotypes of offspring in a highly fecund organism.