Ali, M., Nicieza, A., Wootton, R. J. (2003). Compensatory growth in fishes: a response to growth depression. Fish and Fisheries, 4, (2), 147-190. Sponsorship: Royal Society of London – Chinese Academy of Sciences Exchange award

Compensatory growth in fishes: a response to growth depression

Resource competition is thought to play a major role in driving evolutionary diversification. For instance, in ecological character displacement, coexisting species evolve to use different resources, reducing the effects of interspecific competition. It is thought that a similar diversifying effect might occur in response to competition among members of a single species. Individuals may mitigate the effects of intraspecific competition by switching to use alternative resources not used by conspecific competitors. This diversification is the driving force in some models of sympatric speciation, but has not been demonstrated in natural populations. Here, we present experimental evidence confirming that competition drives ecological diversification within natural populations. We manipulated population density of three-spine sticklebacks (Gasterosteus aculeatus) in enclosures in a natural lake. Increased population density led to reduced prey availability, causing individuals to add alternative prey types to their diet. Since phenotypically different individuals added different alternative prey, diet variation among individuals increased relative to low-density control enclosures. Competition also increased the diet-morphology correlations, so that the frequency-dependent interactions were stronger in high competition. These results not only confirm that resource competition promotes niche variation within populations, but also show that this increased diversity can arise via behavioural plasticity alone, without the evolutionary changes commonly assumed by theory.

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Intraspecific competition drives increased resource use diversity within a natural population

While the impacts of high exploitation on fish populations and aquatic ecosystems are well-documented for commercial fishing, particularly in the marine environment, the potential biological impacts of angling received less attention. This paper discusses angling patterns within a framework of basic ecological and evolutionary literature and examines potential biological impacts of angling by focusing on study results associated with high exploitation rates and pronounced selective exploitation. The impacts range from impacts occurring directly on the exploited species (truncation of the natural age and size structure, depensatory mechanisms, loss of genetic variability, evolutionary changes), to those that occur on the aquatic ecosystem (changes in trophic cascades, trait-mediated effects). As a third category, impacts related to the angling activity per se are distinguished (habitat modifications, wildlife disturbance, nutrient inputs, loss of fishing gear). Although the main threats to fish often are l...

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Documented and Potential Biological Impacts of Recreational Fishing: Insights for Management and Conservation

Egg size is a widely-studied trait and yet the causes and consequences of variation in this trait remain poorly understood. Egg size varies greatly within many avian species, with the largest egg in a population generally being at least 50% bigger, and sometimes twice as large, as the smallest. Generally, approximately 70% of the variation in egg mass is due to variation between rather than within clutches, although there are some cases of extreme intra-clutch egg-size variation. Despite the large amount of variation in egg size between females, this trait is highly consistent within individuals between breeding attempts; the repeatability of egg size is generally above 0.6 and tends to be higher than that of clutch size or laying date. Heritability estimates also tend to be much higher for egg size (> 0.5) than for clutch size or laying date (< 0.5). As expected, given the high repeatability and heritability of egg size, supplemental food had no statistically significant effect on this trait in 18 out of 28 (64%) studies. Where dietary supplements do increase egg size, the effect is never more than 13% of the control values and is generally much less. Similarly, ambient temperature during egg formation generally explains less than 15% of the variation in egg size. In short, egg size appears to be a characteristic of individual females, and yet the traits of a female that determine egg size are not clear. Although egg size often increases with female age (17 out of 37 studies), the change in egg size is generally less than 10%. Female mass and size rarely explain more than 20% of the variation in egg size within species. A female's egg size is not consistently related to other aspects of reproductive performance such as clutch size, laying date, or the pair's ability to rear young. Physiological characteristics of the female (e.g. endogenous protein stores, oviduct mass, rate of protein uptake by ovarian follicles) show more promise as potential determinants of egg size. With regards to the consequences of egg-size variation for offspring fitness, egg size is often correlated with offspring mass and size within the first week after hatching, but the evidence for more long-lasting effects on chick growth and survival is equivocal. In other oviparous vertebrates, the magnitude of egg-size variation within populations is often as great or greater than that observed within avian populations. Although there are much fewer estimates of the repeatability of egg size in other taxa, the available evidence suggests that egg size may be more flexible within individuals. Furthermore, in non-avian species (particularly fish and turtles), it is more common for female mass or size to explain a substantial proportion of the variation in egg size. Further research into the physiological basis of egg-size variation is needed to shed light on both the proximate and ultimate causes of intraspecific variation in this trait in birds.

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Avian egg size: variation within species and inflexibility within individuals

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.

The evolution of growth trajectories: what limits growth rate?

The compensatory growth responses of individual juveniles of two co-existing species were compared after identical periods of starvation to determine inter-specific similarities and differences. The carnivorous stickleback Gasterosteus aculeatus was compared with the omnivorous minnow Phoxinus phoxinus. Both species experienced 1 or 2 weeks of starvation before being re-fed ad libitum. The two species differed in their response to the starvation periods, with minnows showing a lower weight-specific loss. Both species showed compensatory responses in appetite, growth and to a lesser extent, growth efficiency. Minnows wholly compensated for 1 and 2 weeks of starvation. At the end of the experiment, sticklebacks starved For 2 weeks were still showing a compensatory response and had nut achieved full compensation. The compensatory responses of the sticklebacks showed a lag of a week before developing in the re-feeding phase, whereas the response of the minnows was immediate. Analysis of lipid and dry matter concentrations suggested that the compensatory response restored reserve lipids while also bringing the fish back to the growth trajectory of continuously fed fish. (C) 2001 The Fisheries Society of the British Isles.

Comparison of compensatory growth responses of juvenile three‐spined stickleback and minnow following similar food deprivation protocols

The study examined the response to variation in food ration within a breeding season on components of the reproductive performance of mature female three-spined sticklebacks Gasterosteus aculeatus, a batch spawner. The main effect of food availability was on the duration of the inter-spawning interval (ISI). Low ration resulted in an increase in the number of days until the next spawning. A low ration during one inter-spawning interval increased the length of the next ISI, even if there was a high ration in the latter. Females that experienced high rations for two successive ISIs showed an increase in batch fecundity over the successive spawnings. There was no tendency for fish on low rations to produce bigger eggs or eggs with higher lipid concentration. Females on low ration had a lower carcass dry matter concentration and a lower liver dry weight immediately after their third spawning. Female sticklebacks responded to a low ration with little change in egg number or size, but a slight reduction in physiological condition. Egg characteristics and, to a lesser extent, batch fecundity are insensitive to current rate of food consumption, but the interval between spawnings is sensitive to both current and previous rates of consumption.

Effect of variable food levels on reproductive performance of breeding female three-spined sticklebacks

Ali, M., Wootton, R. J. (2001). Capacity for growth compensation in juvenile three-spined sticklebacks experiencing cycles of food deprivation. Journal of Fish Biology, 58, (6), 1531-1544.

Capacity for growth compensation in juvenile three‐spined sticklebacks experiencing cycles of food deprivation

The temporal dynamics of appetite (weight of food consumed per day) were analysed for fish fed to satiation after a 1 or 2-week period of feed deprivation. Three species were compared: two omnivores (minnow, Phoxinus phoxinus and gibel carp, Carassius auratus gibelio) and a carnivore (three-spined stickleback, Gasterosteus aculeatus). All species showed compensatory changes in appetite and growth during the re-feeding period but in the stickleback, there was a lag of a week before the compensatory response was detected. The temporal dynamics of appetite differed between the three species but not within a species. Appetite of the minnow declined towards control levels from the onset of re-feeding. In gibel carp, appetite increased to a peak and then declined to control levels. In the stickleback, appetite was initially below control levels, increased to a maximum, then declined towards control levels. The differences between the species might have been artefacts of the experimental protocols used but could also reflect underlying differences in the control of appetite in these species of fish.

M. Ali

Papers

Compensatory growth in fishes: a response to growth depression

Comparison of compensatory growth responses of juvenile three‐spined stickleback and minnow following similar food deprivation protocols

Effect of variable food levels on reproductive performance of breeding female three-spined sticklebacks

Capacity for growth compensation in juvenile three‐spined sticklebacks experiencing cycles of food deprivation

Dynamics of appetite in three fish species (Gasterosteus aculeatus, Phoxinus phoxinus and Carassius auratus gibelio) after feed deprivation