Repeatable inter‐individual variation in the thermal sensitivity of metabolic rate

Assessing whether trait variations among individuals are consistent over time and among environmental conditions is crucial to understand evolutionary responses to new selective pressures such as climate change. According to the universal thermal dependence hypothesis, thermal sensitivity of metabolic rate should not vary strongly and consistently among organisms, implying limited evolutionary response for metabolic traits under climate change. However, this hypothesis has been rarely tested at an individual level, leaving a gap in our understanding of climate change impacts on metabolic responses and their potential evolution. Using the amphipod Gammarus fossarum, we investigated the variability and repeatability of individual metabolic thermal reaction norms over time. We found large variations in both the thermal sensitivity (i.e. slope) and expression level (i.e. intercept) of individual metabolic reaction norms. Moreover, differences among individuals were consistent over time, and therefore repeatable. Inter‐individual variations in body mass resulted in a high repeatability of metabolic expression level but had no significant effect on the repeatability of thermal sensitivity. Overall, our results highlight that inter‐individual variability and repeatability of thermal reaction norms can be substantial. We conclude that these consistent differences among individuals should not be overlooked when apprehending the ecological and evolutionary effects of climate change.     

An information-theoretic approach to evaluating the size and temperature dependence of metabolic rate

The effects of body mass and temperature on metabolic rate (MR) are among the most widely examined physiological relationships. Recently, these relationships have been incorporated into the metabolic theory of ecology (MTE) that links the ecology of populations, communities and ecosystems to the MR of individual organisms. The fundamental equation of MTE derives the relation between mass and MR using first principles and predicts the temperature dependence of MR based on biochemical kinetics. It is a deliberately simple, zeroth-order approximation that represents a baseline against which variation in real biological systems can be examined. In the present study, we evaluate the fundamental equation of MTE against other more parameter-rich models for MR using an information-theoretic approach to penalize the inclusion of additional parameters. Using a comparative database of MR measurements for 1359 species, from 11 groups ranging from prokaryotes to mammals, and spanning 16 orders of magnitude in mass and a 59°C range in body temperature, we show that differences between taxa in the mass and temperature dependence of MR are sufficiently large as to be retained in the best model for MR despite the requirement for estimation of 22 more parameters than the fundamental equation of MTE.     

The combined effects of reactant kinetics and enzyme stability explain the temperature dependence of metabolic rates

A mechanistic understanding of the response of metabolic rate to temperature is essential for understanding thermal ecology and metabolic adaptation. Although the Arrhenius equation has been used to describe the effects of temperature on reaction rates and metabolic traits, it does not adequately describe two aspects of the thermal performance curve (TPC) for metabolic rate—that metabolic rate is a unimodal function of temperature often with maximal values in the biologically relevant temperature range and that activation energies are temperature dependent. We show that the temperature dependence of metabolic rate in ectotherms is well described by an enzyme‐assisted Arrhenius (EAAR) model that accounts for the temperature‐dependent contribution of enzymes to decreasing the activation energy required for reactions to occur. The model is mechanistically derived using the thermodynamic rules that govern protein stability. We contrast our model with other unimodal functions that also can be used to describe the temperature dependence of metabolic rate to show how the EAAR model provides an important advance over previous work. We fit the EAAR model to metabolic rate data for a variety of taxa to demonstrate the model's utility in describing metabolic rate TPCs while revealing significant differences in thermodynamic properties across species and acclimation temperatures. Our model advances our ability to understand the metabolic and ecological consequences of increases in the mean and variance of temperature associated with global climate change. In addition, the model suggests avenues by which organisms can acclimate and adapt to changing thermal environments. Furthermore, the parameters in the EAAR model generate links between organismal level performance and underlying molecular processes that can be tested for in future work.     

Diversity in the influence of temperature on the growth rates of freshwater algae, and its ecological relevance

1. Growth rates of seven species of planktonic algae were determined in culture over a range of temperature from 2 to 35 °C. Additional observations on growth and viability were made for 13 species in the temperature range 20–35 °C. 2. There was a wide range of growth rates between species at their optimal temperatures, from 1.7 divisions day^?1 (Asterionella formosa) to 0.3 divisions day^?1 (Ceratium furcoides). 3. There were considerable differences between species for growth at low and high temperature. Certain algae, including the diatom A. formosa and the flagellates Cryptomonas marssonii, Dinobryon divergens and Eudorina unicocca var. unicocca, had growth rates of 0.4 divisions day^?1 or more at 5 °C. The cyanophyte Tychonema (formerly Oscillatoria) bourrellyi, the xanthophyte Tribonema sp., the desmid Staurastrum cingulum and the large dinoflagellate C. furcoides grew poorly or not at all at this temperature. All 21 species tested could grow at 25 °C, but many – including most of the diatoms, some cyanophytes, and all the flagellates – failed to grow persistently at 30 °C. Only Aphanizomenon flos‐aquae survived with moderate increase at 35 °C, a lethal temperature for the other species. 4. The applicability was considered of proposed quantitative formulations of the rate‐temperature relationship. Simple exponential relationships applied only to very limited lower ranges of temperature. The relationship proposed by B?lehrádek was a better fit over a wider temperature range, but still excluded rate‐decline at high temperature. 5. The interspecific differences found are of potential significance for restrictions in natural distributions associated with season, altitude (especially above 500 m) and latitude.     

Consistent temperature dependence of respiration across ecosystems contrasting in thermal history

Ecosystem respiration is a primary component of the carbon cycle and understanding the mechanisms that determine its temperature dependence will be important for predicting how rates of carbon efflux might respond to global warming. We used a rare model system, comprising a network of geothermally heated streams ranging in temperature from 5 °C to 25 °C, to explore the nature of the relationship between respiration and temperature. Using this ‘natural experiment’, we tested whether the natal thermal regime of stream communities influenced the temperature dependence of respiration in the absence of other potentially confounding variables. An empirical survey of 13 streams across the thermal gradient revealed that the temperature dependence of whole‐stream respiration was equivalent to the average activation energy of the respiratory complex (0.6–0.7 eV). This observation was also consistent for in‐situ benthic respiration. Laboratory experiments, incubating biofilms from four streams across the thermal gradient at a range of temperatures, revealed that the activation energy and Q₁₀ of respiration were remarkably consistent across streams, despite marked differences in their thermal history and significant turnover in species composition. Furthermore, absolute rates of respiration at standardised temperature were also unrelated to ambient stream temperature, but strongly reflected differences in biofilm biomass. Together, our results suggest that the core biochemistry, which drives the kinetics of oxidative respiratory metabolism, may be well conserved among diverse taxa and environments, and that the intrinsic sensitivity of respiration to temperature is not influenced by ambient environmental temperature.     

Intraspecific variation in temperature dependence of gas exchange characteristics among Plantago asiatica ecotypes from different temperature regimes

There are large inter- and intraspecific differences in the temperature dependence of photosynthesis, but the physiological cause of the variation is poorly understood. Here, the temperature dependence of photosynthesis was examined in three ecotypes of Plantago asiatica transplanted from different latitudes, where the mean annual temperature varies between 7.5 and 16.8 degrees C. Plants were raised at 15 or 30 degrees C, and the CO(2) response of photosynthetic rates was determined at various temperatures. When plants were grown at 30 degrees C, no difference was found in the temperature dependence of photosynthesis among ecotypes. When plants were grown at 15 degrees C, ecotypes from a higher latitude maintained a relatively higher photosynthetic rate at low measurement temperatures. This difference was caused by a difference in the balance between the capacities of two processes, ribulose-1,5-bisphosphate regeneration (J(max)) and carboxylation (V(cmax)), which altered the limiting step of photosynthesis at low temperatures. The organization of photosynthetic proteins also varied among ecotypes. The ecotype from the highest latitude increased the J(max) : V(cmax) ratio with decreasing growth temperature, while that from the lowest latitude did not. It is concluded that nitrogen partitioning in the photosynthetic apparatus and its response to growth temperature were different among ecotypes, which caused an intraspecific variation in temperature dependence of photosynthesis.     

Nutrients and temperature additively increase stream microbial respiration

Rising temperatures and nutrient enrichment are co‐occurring global‐change drivers that stimulate microbial respiration of detrital carbon, but nutrient effects on the temperature dependence of respiration in aquatic ecosystems remain uncertain. We measured respiration rates associated with leaf litter, wood, and fine benthic organic matter (FBOM) across seasonal temperature gradients before (PRE) and after (ENR1, ENR2) experimental nutrient (nitrogen [N] and phosphorus [P]) additions to five forest streams. Nitrogen and phosphorus were added at different N:P ratios using increasing concentrations of N (~80–650 μg/L) and corresponding decreasing concentrations of P (~90–11 μg/L). We assessed the temperature dependence, and microbial (i.e., fungal) drivers of detrital mass‐specific respiration rates using the metabolic theory of ecology, before vs. after nutrient enrichment, and across N and P concentrations. Detrital mass‐specific respiration rates increased with temperature, exhibiting comparable activation energies (E, electronvolts [eV]) for all substrates (FBOM E = 0.43 [95% CI = 0.18–0.69] eV, leaf litter E = 0.30 [95% CI = 0.072–0.54] eV, wood E = 0.41 [95% CI = 0.18–0.64] eV) close to predicted MTE values. There was evidence that temperature‐driven increased respiration occurred via increased fungal biomass (wood) or increased fungal biomass‐specific respiration (leaf litter). Respiration rates increased under nutrient‐enriched conditions on leaves (1.32×) and wood (1.38×), but not FBOM. Respiration rates responded weakly to gradients in N or P concentrations, except for positive effects of P on wood respiration. The temperature dependence of respiration was comparable among years and across N or P concentration for all substrates. Responses of leaf litter and wood respiration to temperature and the combined effects of N and P were similar in magnitude. Our data suggest that the temperature dependence of stream microbial respiration is unchanged by nutrient enrichment, and that increased temperature and N + P availability have additive and comparable effects on microbial respiration rates.     

中华鳖新孵幼体的热耐受性、体温昼夜变化和运动能力的热依赖性

研究中华鳖新孵幼体的热耐受性、体温及温度对运动能力的影响 .结果表明 ,在干燥和潮湿环境下 ,选择体温分别为 2 8.0℃和 30 .3℃ ;潮湿环境下 ,临界高温和低温分别为 40 .9℃和 7.8℃ .在缺乏温度梯度的热环境中 ,水温对幼鳖体温的影响比气温更直接 ,体温和环境温度的昼夜变化相一致 ,说明幼鳖生理调温能力很弱 .在有温度梯度的热环境中 ,幼鳖能通过行为调温将体温维持到较高且较恒定的水平 ,导致体温昼夜变化不明显 .幼鳖运动能力有显著的热依赖性 ,在一定温度范围内随体温升高而增强 .体温31.5℃时 ,幼鳖的运动表现最好 ,最大续跑距离、单位时间跑动距离和单位时间停顿次数分别为 1.87m、4 92m·min-1和 6 .2次·min-1.体温过高时 ,运动能力下降 .当体温为 33 .0℃时 ,最大续跑距离、单位时间跑动距离和单位时间停顿次数分别为 1.30m、4.2 8m·min-1和 7.7次·min-1.     

Context‐specific influence of water temperature on brook trout growth rates in the field

1. Modelling the effects of climate change on freshwater fishes requires robust field‐based estimates accounting for interactions among multiple factors. 2. We used data from an 8‐year individual‐based study of a wild brook trout (Salvelinus fontinalis) population to test the influence of water temperature on season‐specific growth in the context of variation in other environmental (i.e. season, stream flow) or biotic factors (local brook trout biomass density and fish age and size) in West Brook, a third‐order stream in western Massachusetts, U.S.A. 3. Changes in ambient temperature influenced individual growth rates. In general, higher temperatures were associated with higher growth rates in winter and spring and lower growth rates in summer and autumn. However, the effect of temperature on growth was strongly context‐dependent, differing in both magnitude and direction as a function of season, stream flow and fish biomass density. 4. We found that stream flow and temperature had strong and complex interactive effects on trout growth. At the coldest temperatures (in winter), high stream flows were associated with reduced trout growth rates. During spring and autumn and in typical summers (when water temperatures were close to growth optima), higher flows were associated with increased growth rates. In addition, the effect of flow at a given temperature (the flow‐temperature interaction) differed among seasons. 5. Trout density negatively affected growth rate and had strong interactions with temperature in two of four seasons (i.e. spring and summer) with greater negative effects at high temperatures. 6. Our study provided robust, integrative field‐based estimates of the effects of temperature on growth rates for a species which serves as a model organism for cold‐water adapted ectotherms facing the consequences of environmental change. Results of the study strongly suggest that failure to derive season‐specific estimates, or to explicitly consider interactions with flow regime and fish density, will seriously compromise our ability to predict the effects of climate change on stream fish growth rates. Further, the concordance we found between empirical observations and likely energetic mechanisms suggests that our general results should be relevant at broader spatial and temporal scales.     

Geographic variation and thermal plasticity shape salamander metabolic rates under current and future climates

Predicted changes in global temperature are expected to increase extinction risk for ectotherms, primarily through increased metabolic rates. Higher metabolic rates generate increased maintenance energy costs which are a major component of energy budgets. Organisms often employ plastic or evolutionary (e.g., local adaptation) mechanisms to optimize metabolic rate with respect to their environment. We examined relationships between temperature and standard metabolic rate across four populations of a widespread amphibian species to determine if populations vary in metabolic response and if their metabolic rates are plastic to seasonal thermal cues. Populations from warmer climates lowered metabolic rates when acclimating to summer temperatures as compared to spring temperatures. This may act as an energy saving mechanism during the warmest time of the year. No such plasticity was evident in populations from cooler climates. Both juvenile and adult salamanders exhibited metabolic plasticity. Although some populations responded to historic climate thermal cues, no populations showed plastic metabolic rate responses to future climate temperatures, indicating there are constraints on plastic responses. We postulate that impacts of warming will likely impact the energy budgets of salamanders, potentially affecting key demographic rates, such as individual growth and investment in reproduction.     

Thermal variation near the thermal optimum does not affect the growth,metabolism or swimming performance in wild Atlantic salmon Salmo salar

Typically, laboratory studies on the physiological effects of temperature are conducted using stable acclimation temperatures. Nonetheless, information extrapolated from these studies may not accurately represent wild populations living in thermally variable environments. The aim of this study was to compare the growth rate, metabolism and swimming performance of wild Atlantic salmon exposed to cycling temperatures, 16–21°C, and stable acclimation temperatures, 16, 18.5, 21°C. Growth rate, metabolic rate, swimming performance and anaerobic metabolites did not change among acclimation groups, suggesting that within Atlantic salmon's thermal optimum range, temperature variation has no effect on these physiological properties.     

Assessing latitudinal gradients in speciation rates and biodiversity at the global scale

The mechanisms responsible for latitudinal biodiversity gradients have fascinated and perplexed biologists since the time of Darwin. Ecological theory has yielded two general classes of mechanisms to account for variation in biodiversity: dispersal–assembly mechanisms that invoke differences in stochastic rates of speciation, extinction and dispersal; and niche–assembly mechanisms that invoke species differences, species interactions and environmental heterogeneity. Distinguishing between these two classes of mechanisms requires explicit consideration of macroevolutionary dynamics. Here, we assess the importance of dispersal–assembly mechanisms in the origin and maintenance of biodiversity using fossil data that encompass 30 million years of macroevolution for three distinct groups of ocean plankton: foraminifera, nannoplankton and radiolaria. Applying new methods of analysis to these fossil data, we show here for the first time that latitudinal biodiversity gradients exhibit strong positive correlations with speciation rates even after explicitly controlling for variation in sampling effort and for increases in habitat area towards the equator. These findings provide compelling evidence that geographical variation in macroevolutionary dynamics is a primary determinant of contemporary biodiversity gradients, as predicted by dispersal–assembly theory.     

Temporal and spatial variation in potential and realized growth rates of age‐0 year northern rock sole

The possibility of prey limitations on the growth performance of age‐0 year northern rock sole Lepidopsetta polyxystra was evaluated at three sites along the north‐east coast of Kodiak Island, Alaska, U.S.A., by comparison of observed to potential growth rates. Growth potential was measured in the laboratory across the range of temperatures encountered by this species during the first summer of life. Growth potential ( g _L, mm day⁻¹) increased with water temperature (T) between 2 and 13° C, according to: g _L = 0·0151 + 0·3673·log₁₀(T). There were significant differences in growth rate between the three field sites such that Holiday Beach fish were 7·1 mm longer than Shakmanof Beach fish by mid‐September, with Pillar Creek Cove fish of intermediate size. Temperature differences between sites accounted for less than half of this variation. The remainder may have been related to differences in prey availability among the sites in association with observed differences in sediment characteristics. In addition to the spatial variability, there was significant monthly variation in growth performance. Realized growth rates between July and August were in excess of 85% of potential. Between August and September, however, realized growth fell to 43–71% of potential indicating a decline in conditions for growth. The spatial variation in growth rates was not density‐dependent as the site with the highest fish densities (Holiday Beach) also supported the highest growth rates. The available data indicates that for this subtidal species, interannual variation in growth may be more important than site variation.     

Individual variation in metabolism and thermal tolerance in juvenile qingbo (Spinibarbus sinensis)

Studies of individual variation in the physiological performance of animals and their relationship with metabolism may provide insight into how selection influences diversity in phenotypic traits. Thus, the aims of the present study were to investigate variation in thermal tolerance and its relationship with individual metabolism in juvenile qingbo (Spinibarbus sinensis). To fulfill our goal, we first measured the resting metabolic rate (RMR), maximum metabolic rate (MMR), metabolic scope (MS, MMR–RMR) and excess post-exercise oxygen consumption (EPOC) of 40 fish at 25 °C. We then measured the critical thermal minimum (CT_min), lethal thermal minimum (LT_min), critical thermal maximum (CT_max), and lethal thermal maximum (LT_max) of 20 fish. Both MMR and MS were positively correlated with the metabolic recovery rate (MRR) (p = 0.001), indicating that high aerobic metabolic performance individuals possessed an advantage for the recovery of anaerobic metabolism. However, the negative correlation between EPOC and MRR (p = 0.017) indicated a slow recovery of the metabolism of high anaerobic metabolic capacity individuals. The RMR was positively correlated with CT_min and LT_min, whereas all of the metabolic rate parameters (RMR, MMR, and MS) were negatively correlated with CT_max and LT_max (p < 0.05), indicating that high aerobic metabolic performance individuals have a weakened thermal tolerance. These results suggested that there is a trade-off between aerobic metabolic performance and thermal tolerance.     

Estimation of evacuation rates in the field

Two methods are presented to calculate evacuation rates based on observed diel changes in occurrence and mean mass of prey in predator stomachs. The methods do not require predators to exhibit prolonged non‐feeding periods, but the ingestion of each particular prey type must be restricted to certain diel periods. Data from >7500 whiting Merlangius merlangus collected at five locations in the North Sea were used to demonstrate the methods. The evacuation rates estimated from field data were similar to laboratory results, though a tendency for estimates to exceed literature values slightly was noted. Bias was introduced if a large proportion of the prey was evacuated completely in the interval between subsequent samples and if significant amounts of other food were present in the stomach together with the prey in question. The methods can be used to supplement laboratory estimates of evacuation rates or provide first estimates for species that are not easily maintained in the laboratory.