Variation In Standard Metabolic Rate Research Articles

Developmental plasticity of melanisation in a beetle reveals sex‐specific responses and performance costs

Abstract Dark pigmentation of adults resulting from cold exposure during development is ubiquitous among insects. Yet the costs associated with the developmental plasticity of melanisation have been explored only for a handful of species. We explored two hypotheses relating to colour plasticity of insects. First, if melanin pigments are costly to synthesise and require allocation of resources, increased melanisation should be negatively associated with performance traits (the cost hypothesis). Second, given the positive association between melanisation and immune responses or desiccation resistance in some species, melanisation and performance may co‐vary positively (the positive covariance hypothesis). Furthermore, given the typical differences in reproductive effort and activity between females and males, we expect these performance–melanisation relationships to differ between sexes. We assessed whether temperature‐induced melanisation of the globally invasive beetle Harmonia axyridis f. succinea (Pallas) (Coleoptera: Coccinellidae) was related to standard metabolic rate and flight performance while accounting for developmental temperature and family affiliation. Melanisation explained little variation in standard metabolic rate at most temperatures tested (15, 20, 25, 30 and 35°C). For flight performance and after the cold‐rearing regime, lighter coloured female beetles had increased flight speed compared with darker females, whereas the opposite was found in males. In addition, in warm‐reared beetles, darker individuals had lower mean flight speeds. The results suggest that at least some flight performance costs are associated with developmentally induced melanisation, supporting the cost hypothesis. Disparities in these relationships between males and females suggest that the drivers underlying variation in melanisation differ between sexes. Importantly, variation in melanisation was more acute for females reared in the warm regime compared with equally treated males and to cold‐reared individuals. We propose new directions to test hypotheses pertaining to the maintenance of colour variation and performance between sexes in nature.

Sustained swimming exercise training decreases the individual variation in the metabolic phenotype of gilthead sea bream (Sparus aurata)

Cultured fish can be induced to swim, although the suitability and benefits remain to be tested. Sustained swimming exercise (SSE) training and detraining (DET) were applied in juvenile gilthead sea bream (Sparus aurata) and the metabolic rates were investigated. Fish with a total body mass of 80.5 ± 1.5 g and total length 17.2 ± 0.1 cm were maintained untrained (spontaneously swimming activity, UNT), swim-trained (induced sustained swimming activity, SSE) at 1 BL s−1 for 28 days, or detrained (28 days of swimming followed by 10 days of untraining, DET). Standard metabolic rate (SMR), maximum metabolic rate (MMR), and excess post-exercise oxygen consumption (EPOC) were assessed (n = 10). In addition, the effects of SSE training (51 days) on blood and plasma parameters were investigated before and immediately after applying a high-intensity swimming (HIS) protocol. SMR, MMR, and EPOC values were not different between SSE, UNT, or DET fish (143.2, 465.5 mg O2 kg−1 h−1, and 459.1 mg O2 kg−1, respectively). Spite the lack of differences between treatments, the dispersion in the residuals for SMR, MMR, and absolute aerobic scope (AAS) values followed the order UNT > DET > SSE, indicating that swim training decreases the individual variation of these metabolic parameters. Haematological parameters, plasma glucose, lactate, and cortisol levels were similar between SSE and UNT groups before HIS. Plasma glucose and lactate levels increased in both groups after HIS, being higher in the SSE group. Plasma cortisol levels were similar between both groups after HIS. Results suggest that SSE training improves energy use and reduces individual variation in SMR and MMR, an effect that declines with detraining.

Territoriality in Drosophila: indirect effects and covariance with body mass and metabolic rate

Abstract Territoriality (i.e., defense of a resource) is the outcome of behavioral interactions that can result in selective advantages in many vertebrates and invertebrates. Since territoriality is expressed in a social context, an individuals’ territoriality may change according to the phenotype of the opponents that they are confronted with (termed “indirect effects”). Defending a territory may also confer energetic costs to individuals, which could be reflected in their standard metabolic rate (SMR), a key component of an ectotherms’ energy budget. Here, we measured territoriality using dyadic contests, body mass, and SMR using flow-through respirometry, twice in each of 192 adult male Drosophila melanogaster. Territoriality, body mass, and (whole-animal) SMR were all significantly repeatable. However, essentially all the among-individual variation in SMR was shared with body mass, as indicated by a very strong among-individual correlation (rind) between body mass and SMR. The among-individual correlation between territoriality and SMR also tended to be positive, suggesting the presence of underlying metabolic costs to territoriality. Although indirect effects on territoriality were present but weak (accounting for 8.4% of phenotypic variance), indirect effects on territoriality were negatively and significantly correlated with body mass. This indicates that larger individuals tended to suppress their opponents territoriality. Variation among individuals in their ability to suppress territoriality in others was not associated with their own territoriality or SMR.

The standard metabolic rate of a land snail (Cepaea hortensis) is a repeatable trait and influences winter survival

Phenotypic selection on physiological parameters is an underrepresented topic in studies of evolutionary biology. There is especially a lack of studies involving invertebrate organisms. We studied the repeatability of the standard metabolic rate (SMR) and the effect of individual variation in SMR on the subsequent winter survival in a terrestrial shell-bearing mollusc, the white-lipped snail (Cepaea hortensis) in mid-Norway. SMR was measured twice during the autumn and – after an experimental overwintering at controlled conditions – twice during the following spring. We found a significant repeatability of SMR over all three time periods tested, with a clear effect of time, with a high repeatability of 0.56 over 4 days during spring, 0.44 over 12 days in the autumn and 0.17 over 194 days from autumn to spring. That SMR is a repeatable physiological trait across the winter period during which a possible selection might occur, suggests that SMR could be a potential target of natural selection. We indeed found that the autumn SMR significantly influenced the probability of survival during the winter period, with a combination of a positive linear (P = .011) and a quadratic stabilizing (P = .001) effect on SMR. Our results hence support the view that metabolic rate is an important physiological component influencing the fitness of an organism.

Can variation in standard metabolic rate explain context-dependent performance of farmed Atlantic salmon offspring?

Escaped farmed Atlantic salmon interbreed with wild Atlantic salmon, leaving offspring that often have lower success in nature than pure wild salmon. On top of this, presence of farmed salmon descendants can impair production of wild‐type recruits. We hypothesize that both these effects connect with farmed salmon having acquired higher standard metabolic rates (SMR, the energetic cost of self‐maintenance) during domestication. Fitness‐related advantages of phenotypic traits associated with both high SMR and farmed salmon (e.g., social dominance) depend on environmental conditions, such as food availability. We hypothesize that farmed offspring have an advantage at high food availability due to, for example, dominance behavior but suffer increased risks of starvation when food is scarce because this behavior is energy‐demanding. To test these hypotheses, we first compare embryo SMR of pure farmed, farmed‐wild hybrids and pure wild offspring. Next, we test early‐life performance (in terms of survival and growth) of hybrids relative to that of their wild half‐siblings, as well as their competitive abilities, in semi‐natural conditions of high and low food availability. Finally, we test how SMR affects early‐life performance at high and low food availability. We find inconclusive support for the hypothesis that domestication has induced increased SMR. Further, wild and hybrid juveniles had similar survival and growth in the semi‐natural streams. Yet, the presence of hybrids led to decreased survival of their wild half‐siblings. Contrary to our hypothesis about context‐dependency, these effects were not modified by food availability. However, wild juveniles with high SMR had decreased survival when food was scarce, but there was no such effect at high food availability. This study provides further proof that farmed salmon introgression may compromise the viability of wild salmon populations. We cannot, however, conclude that this is connected to alterations in the metabolic phenotype of farmed salmon.

Energy metabolism and personality in wild-caught fall field crickets

Standard metabolic rate (SMR) is known to be highly variable across levels of biological organisation (e.g., species, populations, among individuals, within individuals). Some of the variation in SMR can be attributed to factors such as diet, temperature, and body mass, yet much of the residual variation in SMR remains unexplained. Intuitively, we can expect SMR to co-vary with “personality”, but the rapidly accumulating empirical evidence on this topic remains equivocal. The goal of this study was to test for a link between SMR and a behavioural syndrome at the among-individual level in wild-caught fall field crickets (Gryllus pennsylvanicus). Paired measurements of SMR and two behavioural traits were repeatedly taken over a two-month period, thus allowing to estimate the among-individual correlations (rind) separately from the residual (within-individual) correlations. The two behavioural traits (latency to exit a refuge in a novel environment and “freezing” time following a stressful stimulus) were significantly and moderately repeatable and were found to be part of a syndrome, as indicated by a strong and positive among-individual correlation (rind = 0.82 ± 0.27). Yet, only latency to exit was significantly and positively correlated with SMR (rind = 0.45 ± 0.21), suggesting that the link between boldness and SMR may be driven by individual differences in responses to novelty and not to simulated predatory cues. Since we found that bold individuals (short latency to exit) consistently had a lower SMR than shy individuals (long latency to exit), our results go against the pace-of-life syndrome hypothesis. Instead, our results suggest the presence of constrains in the energy budgets of crickets, which generated an allocation trade-off between energy spent on physical activity vs. maintenance costs (SMR).

High oxidative stress despite low energy metabolism and vice versa: Insights through temperature acclimation in an ectotherm

Reactive oxygen species (ROS) are inescapable byproducts of energy metabolism and may cause costly damage to biomolecules. Organisms have evolved different means to counter oxidative stress, such as modulation of ROS production, neutralization of produced ROS through free radical scavenging and the repair or removal of the damaged structures. A positive relationship between metabolic rate and ROS production is commonly expected, but the oxidative burden of aerobic metabolism remains poorly understood. We investigated whether acclimation to ambient temperatures imposing variation in standard metabolic rate (SMR) is mirrored in the oxidative status of an ectotherm. Grass snakes (Natrix natrix) acclimated for six months to warm (32 °C) conditions revealed seven times higher SMR compared to cold-acclimated (18 °C) individuals. In contrast to SMR, the amount of damage measured as reactive oxygen metabolites test (dROMs) and abundance of micronucleated erythrocytes was significantly lower in warm-acclimated individuals, while non-enzymatic antioxidant capacity of plasma was unaltered by thermal acclimation. Our results support that high SMR may allow snakes to better cope with oxidative stress, possibly through tissue repair or removal of damaged tissues that also requires energy costs. The reversed association between self-maintenance metabolism and oxidative damage to biomolecules provides novel rational for temperature dependent life histories of ectotherms. How oxidative stress may contribute to the known reduced rates of ectotherm growth or reproduction under cold temperatures or if oxidative stress may even drive such life history trait declines are now important challenges to be addressed.

Standard metabolic rate predicts growth trajectory of juvenile Chinese crucian carp (Carassius auratus) under changing food availability.

ABSTRACTPhenotypic traits vary greatly within populations and can have a significant influence on aspects of performance. The present study aimed to investigate the effects of individual variation in standard metabolic rate (SMR) on growth rate and tolerance to food deprivation in juvenile Chinese crucian carp (Carassius auratus) under varying levels of food availability. To address this issue, 19 high and 16 low SMR individuals were randomly assigned to a satiation diet for 3 weeks, whereas another 20 high and 16 low SMR individuals were assigned to a restricted diet (approximately 50% of satiation) for the same period. Then, all fish were completely food-deprived for another 3 weeks. High SMR individuals showed a higher growth rate when fed to satiation, but this advantage of SMR did not exist in food-restricted fish. This result was related to improved feeding efficiency with decreased food intake in low SMR individuals, due to their low food processing capacity and maintenance costs. High SMR individuals experienced more mass loss during food deprivation as compared to low SMR individuals. Our results here illustrate context-dependent costs and benefits of intraspecific variation in SMR whereby high SMR individuals show increased growth performance under high food availability but had a cost under stressful environments (i.e. food shortage).

Variation in Metabolic Rate among Individuals Is Related to Tissue-Specific Differences in Mitochondrial Leak Respiration.

Standard metabolic rate (SMR) and maximum metabolic rate (MMR) typically vary two- or threefold among conspecifics, with both traits assumed to significantly impact fitness. However, the underlying mechanisms that determine such intraspecific variation are not well understood. We examined the influence of mitochondrial properties on intraspecific variation in SMR and MMR and hypothesized that if SMR supports the cost of maintaining the metabolic machinery required for MMR, then the mitochondrial properties underlying these traits should be shared. Mitochondrial respiratory capacity (leak and phosphorylating respiration) and mitochondrial content (cytochrome c oxidase activity) were determined in the liver and white muscle of brown trout Salmo trutta of similar age and maintenance conditions. SMR and MMR were uncorrelated across individuals and were not associated with the same mitochondrial properties, suggesting that they are under the control of separate physiological processes. Moreover, tissue-specific relationships between mitochondrial properties and whole-organism metabolic traits were observed. Specifically, SMR was positively associated with leak respiration in liver mitochondria, while MMR was positively associated with muscle mitochondrial leak respiration and mitochondrial content. These results suggest that a high SMR or MMR, rather than signaling a higher ability for respiration-driven ATP synthesis, may actually reflect greater dissipation of energy, driven by proton leak across the mitochondrial inner membrane. Knowledge of these links should aid interpretation of the potential fitness consequences of such variation in metabolism, given the importance of mitochondria in the utilization of resources and their allocation to performance.

Individuals with higher metabolic rates have lower levels of reactive oxygen species in vivo.

There is increasing interest in the effect of energy metabolism on oxidative stress, but much ambiguity over the relationship between the rate of oxygen consumption and the generation of reactive oxygen species (ROS). Production of ROS (such as hydrogen peroxide, H2O2) in the mitochondria is primarily inferred indirectly from measurements in vitro, which may not reflect actual ROS production in living animals. Here, we measured in vivo H2O2 content using the recently developed MitoB probe that becomes concentrated in the mitochondria of living organisms, where it is converted by H2O2 into an alternative form termed MitoP; the ratio of MitoP/MitoB indicates the level of mitochondrial H2O2 in vivo. Using the brown trout Salmo trutta, we tested whether this measurement of in vivo H2O2 content over a 24 h-period was related to interindividual variation in standard metabolic rate (SMR). We showed that the H2O2 content varied up to 26-fold among fish of the same age and under identical environmental conditions and nutritional states. Interindividual variation in H2O2 content was unrelated to mitochondrial density but was significantly associated with SMR: fish with a higher mass-independent SMR had a lower level of H2O2. The mechanism underlying this observed relationship between SMR and in vivo H2O2 content requires further investigation, but may implicate mitochondrial uncoupling which can simultaneously increase SMR but reduce ROS production. To our knowledge, this is the first study in living organisms to show that individuals with higher oxygen consumption rates can actually have lower levels of H2O2.

Individual (co)variation in standard metabolic rate, feeding rate, and exploratory behavior in wild-caught semiaquatic salamanders.

Repeatability is an important concept in evolutionary analyses because it provides information regarding the benefit of repeated measurements and, in most cases, a putative upper limit to heritability estimates. Repeatability (R) of different aspects of energy metabolism and behavior has been demonstrated in a variety of organisms over short and long time intervals. Recent research suggests that consistent individual differences in behavior and energy metabolism might covary. Here we present new data on the repeatability of body mass, standard metabolic rate (SMR), voluntary exploratory behavior, and feeding rate in a semiaquatic salamander and ask whether individual variation in behavioral traits is correlated with individual variation in metabolism on a whole-animal basis and after conditioning on body mass. All measured traits were repeatable, but the repeatability estimates ranged from very high for body mass (R = 0.98), to intermediate for SMR (R = 0.39) and food intake (R = 0.58), to low for exploratory behavior (R = 0.25). Moreover, repeatability estimates for all traits except body mass declined over time (i.e., from 3 to 9 wk), although this pattern could be a consequence of the relatively low sample size used in this study. Despite significant repeatability in all traits, we find little evidence that behaviors are correlated with SMR at the phenotypic and among-individual levels when conditioned on body mass. Specifically, the phenotypic correlations between SMR and exploratory behavior were negative in all trials but significantly so in one trial only. Salamanders in this study showed individual variation in how their exploratory behavior changed across trials (but not body mass, SMR, and feed intake), which might have contributed to observed changing correlations across trials.

The metabolic determinant underlying the rate of energy use during prolonged fasting for the diamondback watersnake, Nerodia rhombifer (1101.4)

During prolonged bouts of fasting, tissues and organs lose mass as a function of their metabolic expenditure and differences in the allocation of endogenous energy. We previously observed considerable variation in the rate at which individual diamondback watersnakes (Nerodia rhombifer) lose body and fat mass during extended fasts. It has also been found that these snakes exhibit variation in their standard metabolic rate (SMR). We therefore hypothesized that individual variation in SMR is responsible for the observed variation in body mass and body fat loss with fasting. Predictably, snakes with relatively high SMR would metabolize fat stores at a greater rate (therefore loss body mass faster) than snakes that possess relatively low SMR. We explored the causal relationships between SMR and body and fat loss during extended bouts of fasting for the diamondback watersnake by examining variation in SMR and differential loss of body, fat, and organ mass over six months of fasting. Baseline SMR and body mass was measured from 55 snakes and organ masses from 10 snakes. For each following month, 5 snakes were dissected, with 10 snakes sampled on the 6th month. SMR of individuals was measured at 2‐month intervals. On average snakes lost 0.1 grams of fat per day while fasting, with a 5‐fold range in daily fat loss among snakes. After 5 months of fasting, variation in body fat loss ranged from 25% to 90%. There was a 2.5‐fold variation in SMR among these same individuals. As predicted, snakes with higher SMR experienced a greater rate of body fat loss. We have demonstrated that basal rates of metabolism can significant determine rates of body fat loss and therefore impact an individual’s capacity to survive long episodes of fasting.

Influence of respirometry methods on intraspecific variation in standard metabolic rates in newts

Standard metabolic rate (SMR) is both a highly informative and variable trait. Variation in SMR stems not only from diverse intrinsic and extrinsic factors, but also from the use of diverse methods for metabolic measurements. We measured CO2 production (V˙CO2) and oxygen consumption rates (V˙O2) using two flow-through respirometry modes, continuous and intermittent (stop-flow), to evaluate their potential contribution to SMR variation in Alpine newts, Ichthyosaura alpestris. Both respirometry modes yielded similar and repeatable V˙CO2 values. Although V˙O2 was highly repeatable, continuous respirometry produced lower V˙O2 than the intermittent method. During intermittent measurements, the total number of activity bouts was higher than during continuous respirometry trials. Statistical correction for disparate activity levels minimized variation in oxygen consumption between respirometry modes. We conclude that use of either method of flow-through respirometry, if properly applied, introduced less noise to SMR estimates than a variation in activity levels.

Effects of food ration on SMR: influence of food consumption on individual variation in metabolic rate in juvenile coho salmon (Onchorhynchus kisutch)

1. Consistency of differences in standard metabolic rate (SMR) between individual juvenile salmonids and the apparently limited ability of individuals to regulate their SMR has led many researchers to conclude that differences in individual SMR are fixed (i.e. genetic). 2. To test for the effects of food ration on individual performance and metabolism, SMR was estimated by measuring oxygen consumption using flow-through respirometry on individually separated young of the year coho salmon (Oncorhynchus kisutch) placed on varying food rations over a period of 44 days. 3. Results demonstrate that the quantity of food consumed directly affects SMR of juvenile coho salmon, independent of specific dynamic action (SDA, an elevation in metabolic rate from the increased energy demands associated with digestion immediately following a meal) and indicates that higher food consumption is a cause of elevated SMR rather than a consequence of it. Juvenile coho salmon therefore demonstrated an ability to regulate their SMR according to food availability and ultimately food consumption. 4. This study indicates that food consumption may play a pivotal role in understanding individual variation in SMR independent of inherent genetic differences. We suggest that studies involving SMR need to be cautious about the effects of intra-individual differences in food consumption in communal tanks or in different microhabitats in the wild as disproportionate food consumption may contribute more to variation in SMR than intrinsic (genetic) factors. 5. In general, our results suggest that evolutionary changes in SMR are likely a response to selection on food consumption and growth, rather than SMR itself.

Estimated standard metabolic rate interacts with territory quality and density to determine the growth rates of juvenile Atlantic salmon

Summary1. Physiological traits can vary greatly within a species and consequently have a significant impact on other aspects of performance. Many species exhibit substantial variation in basal or standard metabolic rate (SMR), even after controlling for body size and age, yet the ecological consequences of this are little known.2. We examined the relationships between mass‐specific SMR of yearling salmon (estimated from their ventilation rate) and their feeding and growth rates across a range of natural population densities within a semi‐natural stream environment.3. Standard metabolic rate was strongly correlated with dominance rank, and higher ranking fish were more likely to acquire good feeding territories. Despite this, there was no overall relationship between SMR and growth. We show for the first time that this paradox can be explained because within territories of a given quality, there was a negative correlation between SMR and growth rate, presumably owing to the costs of metabolism.4. These effects were also influenced by density: lower densities led to reduced aggression and competition and hence higher average feeding and growth rates. Moreover, at low densities, where availability of good feeding locations was not limiting, there was no relationship between SMR and growth.5. As a result of these processes, there was a context‐dependent trade‐off in energy budgets: the fish achieving the greatest growth were those with the lowest SMR that was necessary to achieve dominance over conspecifics (and hence acquire a good territory), but this minimum threshold SMR increased with population density. These relationships and trade‐offs can explain the persistence of variation in SMR within populations.

Adaptive trade-offs in juvenile salmonid metabolism associated with habitat partitioning between coho salmon and steelhead trout in coastal streams

1. Adaptive trade-offs are fundamental to the evolution of diversity and the coexistence of similar taxa and occur when complimentary combinations of traits maximize efficiency of resource exploitation or survival at different points on environmental gradients. 2. Standard metabolic rate (SMR) is a key physiological trait that reflects adaptations to baseline metabolic performance, whereas active metabolism reflects adaptations to variable metabolic output associated with performance related to foraging, predator avoidance, aggressive interactions or migratory movements. Benefits of high SMR and active metabolism may change along a resource (productivity) gradient, indicating that a trade-off exists among active metabolism, resting metabolism and energy intake. 3. We measured and compared SMR, maximal metabolic rate (MMR), aerobic scope (AS), swim performance (UCrit) and growth of juvenile hatchery and wild steelhead and coho salmon held on high- and low-food rations in order to better understand the potential significance of variation in SMR to growth, differentiation between species, and patterns of habitat use along a productivity gradient. 4. We found that differences in SMR, MMR, AS, swim performance and growth rate between steelhead trout and coho salmon were reduced in hatchery-reared fish compared with wild fish. Wild steelhead had a higher MMR, AS, swim performance and growth rate than wild coho, but adaptations between species do not appear to involve differences in SMR or to trade-off increased growth rate against lower swim performance, as commonly observed for high-growth strains. Instead, we hypothesize that wild steelhead may be trading off higher growth rate for lower food consumption efficiency, similar to strategies adopted by anadromous vs. resident brook trout and Atlantic salmon vs. brook trout. This highlights potential differences in food consumption and digestion strategies as cryptic adaptations ecologically differentiating salmonid species. 5. We hypothesize that divergent digestive strategies, which are common and well documented among terrestrial vertebrates, may be an important but overlooked aspect of adaptive strategies of juvenile salmonids, and fish in general.

Influence of temperature and body mass on standard metabolic rate of eastern red-backed salamanders ( Plethodon cinereus)

(1) We estimated the standard metabolic rate (SMR) of wild-caught Plethodon cinereus across a range of body masses and ecologically relevant temperatures. (2) We collected 16 salamanders, ranging 0.06–1.11 g and quantified oxygen consumption at 10, 15, 20, 25, and 30 °C with a computer-controlled, closed-circuit, indirect respirometer. The multiple regression model indicated that SMR increased with body mass and temperature ( F 2,77=815.59, P<0.001). (3) Our allometric relationship explained >95% of the variation in SMR. The regression equation can be used to investigate questions of applied or basic ecological interest, such as quantifying energetic costs of anthropogenic or natural disturbances on energy budgets.

Temporal and spatial metabolic rate variation in the Antarctic springtail Gomphiocephalus hodgsoni

Spatial and temporal environmental variation in terrestrial Antarctic ecosystems are known to impact species strongly at a local scale, but the ways in which organisms respond (e.g. physiologically, behaviourally) to such variation are poorly understood. Further, very few studies have attempted to assess inter-annual variability of such responses.Building on previous work demonstrating intra-seasonal variation in standard metabolic rate in the springtail Gomphiocephalushodgsoni, we investigated variation in metabolic activity of G. hodgsoni across two austral summer periods at Cape Bird, Ross Island. We also examined the influence of spatial variation by comparing metabolic rates of G. hodgsoni at Cape Bird with those from two other isolated continental locations within Victoria Land (Garwood and Taylor Valleys).We found significant differences between metabolic rates across the 2 years of measurement at Cape Bird. In addition, standard metabolic rates of G. hodgsoni obtained from Garwood and Taylor Valleys were significantly higher than those at Cape Bird where habitats are comparable, but environmental characteristics differ (e.g. microclimatic temperatures are higher).We discuss potential underlying causes of these metabolic rate variation patterns, including those related to differences among individuals (e.g. physiological and genetic differences), locations (e.g. habitat quality and microclimatic regime differences) and populations (e.g. acclimation differences among G. hodgsoni populations in the form of metabolic cold adaptation (MCA)).

Metabolic rate, behaviour and winter performance in juvenile Atlantic salmon

Summary 1Contrasting results of relations between standard metabolic rate, behaviour and performance suggest that the evolutionary trade-offs in energy allocation between metabolisms and other parts of the energy budget depend on environmental conditions. Here, we show how habitat complexity changes the relationship between standard metabolic rate, behaviour and performance (winter energy loss-rates) in juvenile Atlantic salmon (Salmo salar L.). 2Juvenile Salmon, individually screened for standard metabolic rate, were stocked in replicated semi-natural stream channels with and without sheltering possibilities under ledge cover. We recorded the use of ledged areas, energy-loss rates and feeding activity during a 61-day period. 3In the channels with ledge cover, individuals with relatively higher standard metabolic rates used areas outside of the cover more frequently and had higher feeding frequency than individuals with lower standard metabolic rates. No relationship was apparent between areas used, feeding and standard metabolic rate for the individuals within the no-cover treatments. In channels without cover, there was a negative relationship between standard metabolic rate and performance (energy balance), whereas standard metabolic rate did not affect performance in channels with ledge cover. 4The effect of standard metabolic rate on performance depended on habitat complexity expressed through differences in behaviour. In channels with ledge cover, individuals with higher standard metabolic rates were apparently able to compensate for higher metabolic costs by utilizing a riskier habitat outside the cover and attaining a higher feeding success. 5The present study demonstrates that the relationship between standard metabolic rate and performance may be dependent on habitat complexity and vary on small spatial scales, contributing to the persistence of large variation in standard metabolic rate within populations.

Physiological and morphological correlates of among-individual variation in standard metabolic rate in the leopard frog Rana pipiens

Rates of standard metabolism (SMR) are highly variable among individuals within vertebrate populations. Because SMR contributes a substantial proportion of an individual's energy budget, among-individual variation in this trait may affect other energetic processes, and potentially fitness. Here, we examine three potential proximate correlates of variation in SMR: organ mass, serum T4 thyroxine and relative mitochondrial content, using flow cytometry. Body-mass-adjusted kidney mass correlated with SMR, but liver, heart, small intestine and gastrocnemius did not. Thyroxine correlated with SMR, as did mitochondrial content. These results suggest several novel proximate physiological and morphological mechanisms that may contribute to among-individual variation in SMR. Variation in SMR may be maintained by diverse environmental conditions. Some conditions, such as low resource availability, may favor individuals with a low SMR, through small organ size, or low thyroxine or mitochondrial content. Other conditions, such as high resource availability, may favor individuals with a high SMR, through large organ size, or high thyroxine or mitochondrial content.