Quantifying and understanding reproductive allocation schedules in plants

A plant's reproductive allocation (RA) schedule describes the fraction of surplus energy allocated to reproduction as it increases in size. While theorists use RA schedules as the connection between life history and energy allocation, little is known about RA schedules in real vegetation. Here we review what is known about RA schedules for perennial plants using studies either directly quantifying RA or that collected data from which the shape of an RA schedule can be inferred. We also briefly review theoretical models describing factors by which variation in RA may arise. We identified 34 studies from which aspects of an RA schedule could be inferred. Within those, RA schedules varied considerably across species: some species abruptly shift all resources from growth to reproduction; most others gradually shift resources into reproduction, but under a variety of graded schedules. Available data indicate the maximum fraction of energy allocated to production ranges from 0.1 to 1 and that shorter lived species tend to have higher initial RA and increase their RA more quickly than do longer‐lived species. Overall, our findings indicate, little data exist about RA schedules in perennial plants. Available data suggest a wide range of schedules across species. Collection of more data on RA schedules would enable a tighter integration between observation and a variety of models predicting optimal energy allocation, plant growth rates, and biogeochemical cycles.     

Physiological dynamics,reproduction‐maintenance allocations,and life history evolution

Allocation of resources to competing processes of growth, maintenance, or reproduction is arguably a key process driving the physiology of life history trade‐offs and has been shown to affect immune defenses, the evolution of aging, and the evolutionary ecology of offspring quality. Here, we develop a framework to investigate the evolutionary consequences of physiological dynamics by developing theory linking reproductive cell dynamics and components of fitness associated with costly resource allocation decisions to broader life history consequences. We scale these reproductive cell allocation decisions to population‐level survival and fecundity using a life history approach and explore the effects of investment in reproduction or tissue‐specific repair (somatic or reproductive) on the force of selection, reproductive effort, and resource allocation decisions. At the cellular level, we show that investment in protecting reproductive cells increases fitness when reproductive cell maturation rate is high or reproductive cell death is high. At the population level, life history fitness measures show that cellular protection increases reproductive value by differential investment in somatic or reproductive cells and the optimal allocation of resources to reproduction is moulded by this level of investment. Our model provides a framework to understand the evolutionary consequences of physiological processes underlying trade‐offs and highlights the insights to be gained from considering fitness at multiple levels, from cell dynamics through to population growth.     

Diversity of age‐specific reproductive rates may result from ageing and optimal resource allocation

This paper reports the results of a dynamic programming model which optimizes resource allocation to growth, reproduction and repair of somatic damage, based on the disposable soma theory of ageing. Here it is shown that different age‐dependent patterns of reproductive rates are products of optimal lifetime strategies of resource partitioning. The array of different reproductive patterns generated by the model includes those in which reproduction begins at the maximum rate at maturity and then declines to the end of life, or increases up to a certain age and then drops. The observed patterns reflect optimal resource allocation shaped by the level of extrinsic mortality. A continuous decline in the reproductive rate from the start of reproduction is associated with high extrinsic mortality, and an early increase in the reproductive rate occurs under low extrinsic mortality. A long‐lived organism shows a low reproductive rate early in life, and short‐lived organisms start reproduction at the maximum rate.     

Optimal resource allocation in perennial plants: A continuous-time model

A continuous-time model, similar to W. M. Schaffer's (1983, Amer. Nat. 121, 418–431), of growth and reproduction for a perennial herb with discrete growing seasons is considered. Assuming that metabolic rates of reproductive and storage structures are equal, it was possible, through the reduction of the continuous model to a discrete one, to find the optimal allocations to the vegetative, reproductive, and reserve structures. The main feature of the optimal strategy is the existence of an optimal reserve size. The allocation to vegetative structures is, every growing season, the allocation which maximizes the total of reproductive and reserve structures at the end of the season. The relative allocation between reserve and reproductive structures is given, when reproductive success is a linear function of investment, by the fastest growth to the optimal size: no reproduction until the optimal size is reached, and, afterwards, allocation to reproduction of everything beyond what is needed to maintain size R*. Asymptotic growth to the equilibrium and cycles are possible, when reproductive success is a nonlinear function of investment (A. Pugliese, 1988, in “Biomathematics and Related Computational Problems” (L. M. Ricciardi, Ed.), Reidel, Dordrecht, to appear). It has therefore been possible to solve the “general life history problem” ( Schaffer, 1983) when growth is in general a concave function of body size. In the Discussion discrete and continuous-time models are compared; if the real dynamics is described by a continuous model of the type analyzed here, life history predictions made by analyzing the system with a discrete model are upheld.     

Plastic reproductive allocation as a buffer against environmental stochasticity – linking life history and population dynamics to climate

Empirical work suggest that long‐lived organisms have adopted risk sensitive reproductive strategies where individuals trade the amount of resources spent on reproduction versus survival according to expected future environmental conditions. Earlier studies also suggest that climate affects population dynamics both directly by affecting population vital rates and indirectly through long‐term changes in individual life histories. Using a seasonal and state‐dependent individual‐based model we investigated how environmental variability affects the selection of reproductive strategies and their effect on population dynamics. We found that: (1) dynamic, i.e. plastic, reproductive strategies were optimal in a variable climate. (2) Females in poor and unpredictable climatic regimes allocated fewer available resources in reproduction and more in own somatic growth. This resulted in populations with low population densities, and a high average female age and body mass. (3) Strong negative density dependence on offspring body mass and survival, along with co‐variation between climatic severity and population density, resulted in no clear negative climatic effects on reproductive success and offspring body mass. (4) Time series analyses of population growth rates revealed that populations inhabiting benign environments showed the clearest response to climatic perturbations as high population density prohibited an effective buffering of adverse climatic effects as individuals were not able to gain sufficient body reserves during summer. Regularly occurring harsh winters ‘harvested’ populations, resulting in persistent low densities, and released them from negative density dependent effects, resulting in high rewards for a given resource allocation.     

Differential growth rates and calcium-allocation strategies in the garden snail Cantareus aspersus

An optimal division of a key resource between growth and reproductionis expected to produce consistent life history schedules inhabitats where its supply is highly predictable. However, differentialgrowth rates are found between populations and within broodsof Cantareus aspersus, a simultaneous hermaphrodite for whichthe reproductive benefits of a large body size may favour rapidgrowth. Although energy is usually assumed to be the limitingresource in allocation theory, calcium limits the distribution,growth and reproduction of snails. This is a very consistentresource and populations may have allocation strategies whichreflect availability in their habitats. Three experiments comparedCa allocation in the progeny of six populations from Ca-richand Ca-poor habitats. In the first, 100 d-old juveniles werecompared between populations for their shell/soft-tissue dryweight ratio, their allocation of Ca to each compartment, andthe variability within broods. The second measured growth, foodconsumption and shell ratios in growth trials of three populationson low Ca. Thirdly, five populations were compared on abundantor excess Ca. The relationship of shell Ca with soft-tissuelevels differs between populations, but shell ratios changedwith Ca availability in all populations. Most favoured soft-tissuegrowth when dietary Ca is low, but one population (LE) alwayshad the highest shell ratios in these trials. Ca in the parentalhabitat was not a good predictor of juvenile-allocation strategies,but the consistency of LE shell ratios across several broodssuggests theirs may be an inherited trait. LE has faster growthrates and a preference for shell building, which probably representsa strategy for early reproduction. The robustness of a snail'sshell may thus be more indicative of its reproductive strategyrather than Ca availability in its habitat. (Received 18 May 2006; accepted 21 December 2006)     

Life history strategy of Leptinaria unilamellata (d'Orbigny, 1835) (Mollusca,Pulmonata, Subulinidae)

Abstract

Life history theory predicts that the patterns of resource allocation in animals are associated with different strategies, selected in the course of evolution. In the present study, the life history of Leptinaria unilamellata was characterized under laboratory conditions. We determined the growth, reproduction, and longevity patterns of this species and elucidated the strategy related to the development of embryos, through direct observations and examination of the morphology of the gravid uterus. Furthermore, we attempted to analyze the glycogen and galactogen contents of the albumen gland, digestive gland and cephalopedal mass in order to understand energy allocation to life history traits, for three life stages. Leptinaria unilamellata's life history is characterized by great longevity, a short juvenile phase, early sexual maturity, and repeated reproductive events, with little reproductive effort at each event and some mortality shortly after the first reproduction. In the terraria, we found juveniles but no eggs. However, the results of the anatomical study showed no morphological connection between the embryos and the parental organism. Thus, this species should be described as ovoviviparous rather than viviparous. Egg retention in the parent organism is the primary cause of the release of juveniles, instead of eggs, enabling the offspring to withstand environmental stress. The higher quantity of galactogen found in the adults' albumen gland, as compared to juveniles and senescent individuals, as well as the ratio of glycogen to galactogen, reveal the allocation of energy to reproduction rather than to growth. The remaining energy is directed to the maintenance of omeostasis. Such pattern was confirmed by the low levels of glycogen and galactogen observed in the senescent stage, compared to the juvenile and adult stages. In the life strategy of L. unilamellata, the distribution of the reproductive effort among many events associated with ovoviviparity indicates a long-term investment in reproductive success.     

SEVERE COSTS OF REPRODUCTION PERSIST IN ANOLIS LIZARDS DESPITE THE EVOLUTION OF A SINGLE‐EGG CLUTCH

A central tenet of life‐history theory is that investment in reproduction compromises survival. We tested for costs of reproduction in wild brown anoles (Anolis sagrei) by eliminating reproductive investment via surgical ovariectomy and/or removal of oviductal eggs. Anoles are unusual among lizards in that females lay single‐egg clutches at frequent intervals throughout a lengthy reproductive season. This evolutionary reduction in clutch size is thought to decrease the physical burden of reproduction, but our results show that even a single egg significantly impairs stamina and sprint speed. Reproductive females also suffered a reduction in growth, suggesting that the cumulative energetic cost of successive clutches constrains the allocation of energy to other important functions. Finally, in each of two separate years, elimination of reproductive investment increased breeding‐season survival by 56%, overwinter survival by 96%, and interannual survival by 200% relative to reproductive controls. This extreme fitness cost of reproduction may reflect a combination of intrinsic (i.e., reduced allocation of energy to maintenance) and extrinsic (i.e., increased susceptibility to predators) sources of mortality. Our results provide clear experimental support for a central tenet of life‐history theory and show that costs of reproduction persist in anoles despite the evolution of a single‐egg clutch.     

Pooled energy budgets: Resituating human energy ‐allocation trade‐offs

Across taxa, many life‐history traits vary as a function of differences in body size. 1 - 5 Among primates, including humans, allometric relationships explain many trends in metabolic, growth, reproductive, and mortality rates. 6 - 8 But humans also deviate from nonhuman primates with respect to other developmental, reproductive, and parenting characteristics. 9 - 13 Broad relationships between life‐history traits and body size assume that energy expended in activity (foraging effort) is proportional to body size, and that energy available for growth and reproduction are equivalent. Because human subsistence and parenting are based on food sharing, and cooperation in labor and childrearing, the ways by which energy is acquired and allocated to alternate expenditures are expanded. We present a modification of the general allocation model to include a mechanism for these energy transfers. Our goal is to develop a framework that incorporates this mechanism and can explain the human life‐history paradox; that is, slow juvenile growth and rapid reproduction. We suggest that the central characteristics of human subsistence and energy transfer need to be accounted for in order to more fully appreciate human life‐history variability.     

Stress promotes changes in resource allocation to growth and reproduction in a simultaneous hermaphrodite with indeterminate growth

In iteroparous animals, investment in growth is compromised by investment in reproduction, especially in species with indeterminate growth. Life‐history theory predicts that growth should be favoured over reproduction, assuming size‐related fecundity or survival. Hence, increase body condition represents an increase in reproductive potential. Simultaneous hermaphrodites should adjust their resource allocation to each sex function in response to current conditions but, recently, it has been suggested that, in hermaphrodites, gender allocation should be considered as a three‐way trade‐off, including the investment in somatic growth. Due to the higher costs involved, the female function is affected to a greater extent by environmentally stressful conditions rather than the male function. To examine this, we induced stress in the hermaphroditic earthworm Eisenia fetida (Savigny, 1826) and looked for changes in resource allocation in nonreproductive and reproductive individuals. Experimental stress was induced by using tweezers to elicit contractile escape movements. We predicted that stressed earthworms would preferentially allocate resources to growth. In nonreproductive individuals, however, stress had a negative effect on growth, although weight recovery was rapid once manipulation ceased, indicating the importance of body condition, as well as the existence of mechanisms of compensatory growth for growth trajectories in this earthworm species. The response of reproductive individuals was consistent with our expectation: (1) stressed worms maintained their growth rate at the expense of current reproduction and (2) stressed earthworms laid 25% fewer cocoons, which were 30% lighter than cocoons laid by control earthworms. The present results suggest that E. fetida regulates its reproductive effort and that future reproduction has more impact on its fitness than current reproduction. The trade‐off between current and future reproduction should be taken into consideration in models of sex allocation in simultaneous hermaphrodites. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 593–600.     

Sex‐specific reproductive investment of summer spawners of Illex argentinus in the southwest Atlantic

Energy investment in reproduction and somatic growth was investigated for summer spawners of the Argentinean shortfin squid Illex argentinus in the southwest Atlantic Ocean. Sampled squids were examined for morphometry and intensity of feeding behavior associated with reproductive maturation. Residuals generated from length‐weight relationships were analyzed to determine patterns of energy allocation between somatic and reproductive growth. Both females and males showed similar rates of increase for eviscerated body mass and digestive gland mass relative to mantle length, but the rate of increase for total reproductive organ weight relative to mantle length in females was three times that of males. For females, condition of somatic tissues deteriorated until the mature stage, but somatic condition improved after the onset of maturity. In males, there was no correlation between somatic condition and phases of reproductive maturity. Reproductive investment decreased as sexual maturation progressed for both females and males, with the lowest investment occurring at the functionally mature stage. Residual analysis indicated that female reproductive development was at the expense of body muscle growth during the immature and maturing stages, but energy invested in reproduction after onset of maturity was probably met by food intake. However, in males both reproductive maturation and somatic growth proceeded concurrently so that energy allocated to reproduction was related to food intake throughout the process of maturation. For both males and females, there was little evidence of trade‐offs between the digestive gland and reproductive growth, as no significant correlation was found between dorsal mantle length‐digestive gland weight residuals. The role of the digestive gland as an energy reserve for gonadal growth should be reconsidered. Additionally, feeding intensity by both males and females decreased after the onset of sexual maturity, but feeding never stopped completely, even during spawning.     

植物种群的繁殖对策

植物种群的繁殖对策钟章成（西南师范大学,重庆６３０７１５）ＲｅｐｒｏｄｕｃｔｉｖｅＳｔｒａｔｅｇｉｅｓｏｆＰｌａｎｔＰｏｐｕｌａｔｉｏｎｓ．￥ＺｈｏｎｇＺｈａｎｇｃｈｅｎｇ（ＳｏｕｔｈｗｅｓｔＣｈｉｎａＴｅａｃｈｅｒｓＵ－ｎｉｖｅｒｓｉｔｙ,Ｃｈｏｎ...     

Trade‐offs between life‐history traits at range‐edge and central locations

The allocation of resources to different life‐history traits should represent the best compromise in fitness investment for organisms in their local environment. When resources are limiting, the investment in a specific trait must carry a cost that is expressed in trade‐offs with other traits. In this study, the relative investment in the fitness‐related traits, growth, reproduction and defence were compared at central and range‐edge locations, using the seaweed Ascophyllum nodosum as a model system. Individual growth rates were similar at both sites, whereas edge populations showed a higher relative investment in reproduction (demonstrated by a higher reproductive allocation and extended reproductive periods) when compared to central populations that invested more in defence. These results show the capability of A. nodosum to differentially allocate resources for different traits under different habitat conditions, suggesting that reproduction and defence have different fitness values under the specific living conditions experienced at edge and central locations. However, ongoing climate change may threaten edge populations by increasing the selective pressure on specific traits, forcing these populations to lower the investment in other traits that are also potentially important for population fitness.     

Life history of Bulimulus tenuissimus (D'Orbigny, 1835) (Gastropoda,Pulmonata, Bulimulidae): effect of isolation in reproductive strategy and in resources allocation over their lifetime

Despite the great diversity of tropical land snail species, the life history strategies of the great majority of them are unstudied. We studied reproduction, growth and survival patterns of the Brazilian species Bulimulus tenuissimus (D'Orbigny, 1835), and verified the effect of isolation on such patterns. We analyzed aspects of the life cycle of snails maintained in groups and in isolation. For both treatments, we determined the duration of the juvenile, adult and senescent stages. Growth pattern, life-time reproductive output, reproductive output during adult and senescent stages and longevity, were also verified. Isolation prolonged the duration of the juvenile stage, causing a decrease in life-time reproductive output and longevity. The reproductive pattern of the species is seasonal and, in grouped snails, three breeding periods occurred during their lifetime. The isolated snails reproduced by self-fertilization, and only reproduced once in their lifetime, indicating a significant change in reproductive strategy in the isolated individuals. Thus, isolation resulted in changes in energy allocation to growth, reproduction and survival. The results indicate that in adverse environmental conditions, life history traits can enhance the capacity for adaption.     

Is there a trade‐off between horn growth and survival in adult female chamois?

Life‐history theory predicts trade‐offs in energy allocation between different life‐history traits when resources are limited, i.e. certain traits should be negatively correlated. However, individuals differ in their ability to acquire resources, which can lead to positive correlations between traits at the population level. Here, we investigated the consequences of the allocation in horn growth and body mass on survival in a bovid (Rupicapra rupicapra) with capture‐mark re‐sighting data on 161 females. In female ungulates, body mass often covaries positively with demographic performance and the few studies on horn size suggest that this trait could be a signal of individual quality. Thus, we expected to measure positive correlations between the allocation in these traits and female survival. However, body mass was not correlated to female survival and there was only a negative, though marginal, effect of horn growth. Hence, it seems that the allocation in growth is not an indicator of female quality. Future studies could investigate the importance of growth on female reproduction to evaluate its effect on lifetime reproductive success. Moreover, it is important to confirm in other populations our result that suggests a cost of the allocation in horn growth to better understand the presence of horns in female bovids. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 516–521.     

Between semelparity and iteroparity: Empirical evidence for a continuum of modes of parity

The number of times an organism reproduces (i.e., its mode of parity) is a fundamental life‐history character, and evolutionary and ecological models that compare the relative fitnesses of different modes of parity are common in life‐history theory and theoretical biology. Despite the success of mathematical models designed to compare intrinsic rates of increase (i.e., density‐independent growth rates) between annual‐semelparous and perennial‐iteroparous reproductive schedules, there is widespread evidence that variation in reproductive allocation among semelparous and iteroparous organisms alike is continuous. This study reviews the ecological and molecular evidence for the continuity and plasticity of modes of parity—that is, the idea that annual‐semelparous and perennial‐iteroparous life histories are better understood as endpoints along a continuum of possible strategies. I conclude that parity should be understood as a continuum of different modes of parity, which differ by the degree to which they disperse or concentrate reproductive effort in time. I further argue that there are three main implications of this conclusion: (1) that seasonality should not be conflated with parity; (2) that mathematical models purporting to explain the general evolution of semelparous life histories from iteroparous ones (or vice versa) should not assume that organisms can only display either an annual‐semelparous life history or a perennial‐iteroparous one; and (3) that evolutionary ecologists should base explanations of how different life‐history strategies evolve on the physiological or molecular basis of traits underlying different modes of parity.     

Contrasting life histories in neighbouring populations of a large mammal

Background

A fundamental life history question is how individuals should allocate resources to reproduction optimally over time (reproductive allocation). The reproductive restraint hypothesis predicts that reproductive effort (RE; the allocation of resources to current reproduction) should peak at prime-age, whilst the terminal investment hypothesis predicts that individuals should continue to invest more resources in reproduction throughout life, owing to an ever-decreasing residual reproductive value. There is evidence supporting both hypotheses in the scientific literature.

Methodology/Principal Findings

We used an uncommonly large, 38 year dataset on Alpine chamois (Rupicapra rupicapra) shot at various times during the rutting period to test these two hypotheses. We assumed that body mass loss in rutting males was strongly related to RE and, using a process-based approach, modelled how male relative mass loss rates varied with age. For different regions of our study area, we provide evidence consistent with different hypotheses for reproductive allocation. In sites where RE declined in older age, this appears to be strongly linked to declining body condition in old males. In this species, terminal investment may only occur in areas with lower rates of body mass senescence.

Conclusions/Significance

Our results show that patterns of reproductive allocation may be more plastic than previously thought. It appears that there is a continuum from downturns in RE at old age to terminal investment that can be manifest, even across adjacent populations. Our work identifies uncertainty in the relationship between reproductive restraint and a lack of competitive ability in older life (driven by body mass senescence); both could explain a decline in RE in old age and may be hard to disentangle in empirical data. We discuss a number of environmental and anthropogenic factors which could influence reproductive life histories, underlining that life history patterns should not be generalised across different populations.     

Life‐history strategy varies with the strength of competition in a food‐limited ungulate population

Fluctuating population density in stochastic environments can contribute to maintain life‐history variation within populations via density‐dependent selection. We used individual‐based data from a population of Soay sheep to examine variation in life‐history strategies at high and low population density. We incorporated life‐history trade‐offs among survival, reproduction and body mass growth into structured population models and found support for the prediction that different life‐history strategies are optimal at low and high population densities. Shorter generation times and lower asymptotic body mass were selected for in high‐density environments even though heavier individuals had higher probabilities to survive and reproduce. In contrast, greater asymptotic body mass and longer generation times were optimal at low population density. If populations fluctuate between high density when resources are scarce, and low densities when they are abundant, the variation in density will generate fluctuating selection for different life‐history strategies, that could act to maintain life‐history variation.     

The role of fecundity and reproductive effort in defining life‐history strategies of North American freshwater mussels

Selection is expected to optimize reproductive investment resulting in characteristic trade‐offs among traits such as brood size, offspring size, somatic maintenance, and lifespan; relative patterns of energy allocation to these functions are important in defining life‐history strategies. Freshwater mussels are a diverse and imperiled component of aquatic ecosystems, but little is known about their life‐history strategies, particularly patterns of fecundity and reproductive effort. Because mussels have an unusual life cycle in which larvae (glochidia) are obligate parasites on fishes, differences in host relationships are expected to influence patterns of reproductive output among species. I investigated fecundity and reproductive effort (RE) and their relationships to other life‐history traits for a taxonomically broad cross section of North American mussel diversity. Annual fecundity of North American mussel species spans nearly four orders of magnitude, ranging from < 2000 to 10 million, but most species have considerably lower fecundity than previous generalizations, which portrayed the group as having uniformly high fecundity (e.g. > 200000). Estimates of RE also were highly variable, ranging among species from 0.06 to 25.4%. Median fecundity and RE differed among phylogenetic groups, but patterns for these two traits differed in several ways. For example, the tribe Anodontini had relatively low median fecundity but had the highest RE of any group. Within and among species, body size was a strong predictor of fecundity and explained a high percentage of variation in fecundity among species. Fecundity showed little relationship to other life‐history traits including glochidial size, lifespan, brooding strategies, or host strategies. The only apparent trade‐off evident among these traits was the extraordinarily high fecundity of Leptodea, Margaritifera, and Truncilla, which may come at a cost of greatly reduced glochidial size; there was no relationship between fecundity and glochidial size for the remaining 61 species in the dataset. In contrast to fecundity, RE showed evidence of a strong trade‐off with lifespan, which was negatively related to RE. The raw number of glochidia produced may be determined primarily by physical and energetic constraints rather than selection for optimal output based on differences in host strategies or other traits. By integrating traits such as body size, glochidial size, and fecundity, RE appears more useful in defining mussel life‐history strategies. Combined with trade‐offs between other traits such as growth, lifespan, and age at maturity, differences in RE among species depict a broad continuum of divergent strategies ranging from strongly r‐selected species (e.g. tribe Anodontini and some Lampsilini) to K‐selected species (e.g. tribes Pleurobemini and Quadrulini; family Margaritiferidae). Future studies of reproductive effort in an environmental and life‐history context will be useful for understanding the explosive radiation of this group of animals in North America and will aid in the development of effective conservation strategies.     

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.