The evolution of growth trajectories: what limits growth rate?

According to life‐history theory, growth rates are subject to strong directional selection due to reproductive and survival advantages associated with large adult body size. Yet, growth is commonly observed to occur at rates lower than the maximum that is physiologically possible and intrinsic growth rates often vary among populations. This implies that slower growth is favoured under certain conditions. Realized growth rate is thus the result of a compromise between the costs and advantages of growing rapidly, and the optimal rate of growth is not equivalent to the fundamental maximum rate. The ecological and evolutionary factors influencing growth rate are reviewed, with particular emphasis on how growth might be constrained by direct fitness costs. Costs of accelerating growth might contribute to the variance in fitness that is not attributable to age or size at maturity, as well as to the variation in life‐history strategies observed within and among species. Two main approaches have been taken to study the fitness trade‐offs relating to growth rate. First, environmental manipulations can be used to produce treatment groups with different rates of growth. Second, common garden experiments can be used to compare fitness correlates among populations with different intrinsic growth rates. Data from these studies reveal a number of potential costs for growth over both the short and long term. In order to acquire the energy needed for faster growth, animals must increase food intake. Accordingly, in many taxa, the major constraint on growth rate appears to arise from the trade‐off between predation risk and foraging effort. However, growth rates are also frequently observed to be submaximal in the absence of predation, suggesting that growth trajectories also impact fitness via other channels, such as the reallocation of finite resources between growth and other traits and functions. Despite the prevalence of submaximal growth, even when predators are absent, there is surprisingly little evidence to date demonstrating predator‐independent costs of growth acceleration. Evidence that does exist indicates that such costs may be most apparent under stressful conditions. Future studies should examine more closely the link between patterns of resource allocation to traits in the adult organism and lifetime fitness. Changes in body composition at maturation, for example, may determine the outcome of trade‐offs between reproduction and survival or between early and late reproduction. A number of design issues for studies investigating costs of growth that are imposed over the long term are discussed, along with suggestions for alternative approaches. Despite these issues, identifying costs of growth acceleration may fill a gap in our understanding of life‐history evolution: the relationships between growth rate, the environment, and fitness may contribute substantially to the diversification of life histories in nature.     

Offspring size and parental fitness in Daphnia magna

Variation in offspring size and number has been described for a wide range of organisms. Many theoretical models predict that in a given environment, the production of one single offspring size would yield the highest parental fitness. In most planktonic cladocerans, however, offspring size has been found to increase with size and age of the mother, and as individuals of variable size often co-occur within a population, offspring of variable sizes can be produced simultaneously. In this study, I investigated the relationship between age of the mother and size of her offspring to assess at what age of the mother the optimal offspring size was produced. Optimal offspring size was defined as that size of the offspring yielding the highest parental fitness, which translates to a definition of optimal offspring size as the one having the highest juvenile fitness per unit effort put in these juveniles. I observed that the youngest females produced offspring with the highest juvenile fitness per unit effort, and hence concluded that offspring produced by these females were of optimal size. Larger offspring produced by older females were estimated to yield only 70% of the potential fitness of optimally sized offspring.     

Habitat quality ranking depends on habitat-independent environmental factors: a model and results from Callosobruchus maculatus

1. Survival and reproduction in different habitats of a heterogeneous environment are affected by intrinsic habitat properties as well as by habitat-independent extrinsic factors. It is argued that the ranking of habitats according to quality may depend on extrinsic environmental factors that act with the same intensity in all habitats. This can happen if, across habitats, high reproductive success in the absence of extrinsic mortality is coupled with late reproduction.
2. In a model of a semelparous organism with overlapping generations that uses two habitats, the effect of different patterns of extrinsic mortality on habitat quality are analysed, as measured by the reproductive value of an egg, which estimates its expected contribution to the future gene pool.
3. Measurements of fitness components in Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) on four host species demonstrate that, across hosts, long development can be associated with high reproductive success in the absence of extrinsic mortality. As a result, the quality ranking of three out of the four hosts would be different in a stable population regulated by intraspecific competition and egg parasitoids than in a growing population or a stable population regulated by age-independent larval mortality.
4. It is suggested that habitat-independent mortality may play a role in the evolution of ecological niches.     

Temperature adaptation in a geographically widespread zooplankter,Daphnia magna

Evidence for temperature adaptation in Daphnia magna was inferred from variation in the shape of temperature reaction norms for somatic growth rate, a fitness‐related trait. Ex‐ephippial clones from eight populations across Europe were grown under standardized conditions after preacclimation at five temperatures (17–29 °C). Significant variation for grand mean growth rates occurred both within populations (among clones) and between populations. Genetic variation for reaction norm shape was found within populations, with temperature‐dependent trade‐offs in clone relative fitness. However, the population average responses to temperature were similar, following approximately parallel reaction norms. The among‐population variation is not evidence for temperature adaptation. Lack of temperature adaptation at the population level may be a feature of intermittent populations where environmentally terminated diapause can entrain the planktonic stage of the life‐history within a similar range of temperatures.     

Organic Selection: Proximate Environmental Effects on the Evolution of Morphology and Behaviour

Organic selection (the Baldwin Effect) by which an environmentally elicitedphenotypic adaptation comes under genotypic control following selectionwas proposed independently in 1896 by the psychologists James Baldwinand Conwy Lloyd Morgan and by the paleontologist Henry Fairfield Osborn.Modified forms of organic selection were proposed as autonomization bySchmalhausen in 1938, as genetic assimilation by Waddington in 1942, andas an explanation for evolution in changing environments or for speciationby Matsuda and West-Eberhard in the 1980s. Organic selection as amechanism mediating proximate environmental effects on the evolution ofmorphology and behaviour is the topic of this essay. Discussion includesthe context in which organic selection was proposed, Lamarckian or neo-Lamarckian implications of organic selection, Waddingtons experimentalstudies demonstrating the existence and efficacy of genetic assimilation,stabilizing selection and norms of reaction favoured by Schmalhausen, andMatsudas search for a mechanism of organic selection in endocrine changesand in heterochrony.     

Experimental analysis of an early life‐history stage: avian predation selects for larger body size of hatchling turtles

One common life‐history pattern involves an elevated rate and nonrandom distribution of neonatal mortality. However, the mechanisms causing this pattern and the specific traits that confer a survival benefit are not always evident. We conducted a manipulative field experiment using red‐eared slider turtles to test the hypothesis that diurnal avian predators are a primary cause of size‐specific neonatal mortality. Body size was a significant predictor of recapturing hatchlings alive and of finding hatchlings dead under natural conditions, but was unimportant when diurnal predators were excluded from the field site. Overall recapture rates also more than doubled when predators were excluded compared to natural conditions (72.4 vs. 34.9%). We conclude that birds are an important cause of size‐specific mortality of recently emerged hatchling turtles and that ‘bigger is better’ in this system, which has important implications for life‐history evolution in organisms that experience size‐specific neonatal mortality.     

Does urbanization affect selective pressures and life‐history strategies in the common blackbird (Turdus merula L.)?

Urbanization, one of the most extreme land‐use alterations, is currently spreading, and the number of species confronting these changes is increasing. However, contradictory results of previous studies impede a clear interpretation of which selective pressure (nest predation or food limitation) is more important in urban habitats compared with natural situations, and whether birds can confront them by adjusting their life‐history strategies. We investigated life‐history syndromes of three common blackbird (Turdus merula) populations differing in their human influence (urban, rural, and woodland). We analysed daily nest predation and nestling starvation rates to assess the relative importance of these selection pressures in each habitat. Simultaneously, several life‐history traits were investigated to determine if T. merula seem adapted to their main source of selection. Food limitation was more important in the city, whereas nest predation was the most important selective force in the forest. The rural habitat was characterized by an intermediate influence of these two factors. Life‐history syndromes, as the covariation of a suite of traits, confirmed these results because T. merula seem well adapted to the main cause of selection in each habitat. Our results are consistent with urbanization imposing new challenges on birds, and that they adaptively respond to them. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 759–766.