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Thermal limits may arise through a mismatch between oxygen supply and demand in a range of animal taxa. Whilst this oxygen limitation hypothesis is supported by data from a range of marine fish and invertebrates, its generality remains contentious. In particular, it is unclear whether oxygen limitation determines thermal extremes in tracheated arthropods, where oxygen limitation may be unlikely due to the efficiency and plasticity of tracheal systems in supplying oxygen directly to metabolically active tissues. Although terrestrial taxa with open tracheal systems may not be prone to oxygen limitation, species may be affected during other life-history stages, particularly if these rely on diffusion into closed tracheal systems. Furthermore, a central role for oxygen limitation in insects is envisaged within a parallel line of research focussing on insect gigantism in the late Palaeozoic. Here we examine thermal maxima in the aquatic life stages of an insect at normoxia, hypoxia (14 kPa) and hyperoxia (36 kPa). We demonstrate that upper thermal limits do indeed respond to external oxygen supply in the aquatic life stages of the stonefly Dinocras cephalotes, suggesting that the critical thermal limits of such aquatic larvae are set by oxygen limitation. This could result from impeded oxygen delivery, or limited oxygen regulatory capacity, both of which have implications for our understanding of the limits to insect body size and how these are influenced by atmospheric oxygen levels. These findings extend the generality of the hypothesis of oxygen limitation of thermal tolerance, suggest that oxygen constraints on body size may be stronger in aquatic environments, and that oxygen toxicity may have actively selected for gigantism in the aquatic stages of Carboniferous arthropods.

Aquatic ectotherms face the continuous challenge of capturing sufficient oxygen from their environment as the diffusion rate of oxygen in water is 3 ×× 10 5 times lower than in air. Despite the recognized importance of oxygen in shaping aquatic communities, consensus on what drives environmental oxygen availability is lacking. Physiologists emphasize oxygen partial pressure, while ecologists emphasize oxygen solubility, traditionally expressing oxygen in terms of concentrations. To resolve the question of whether partial pressure or solubility limits oxygen supply in nature, we return to first principles and derive an index of oxygen supply from Fick's classic first law of diffusion. This oxygen supply index (OSI) incorporates both partial pressure and solubility. Our OSI successfully explains published patterns in body size and species across environmental clines linked to differences in oxygen partial pressure (altitude, organic pollution) or oxygen solubility (temperature and salinity). Moreover, the OSI was more accurately and consistently related to these ecological patterns than other measures of oxygen (oxygen saturation, dissolved oxygen concentration, biochemical oxygen demand concentrations) and similarly outperformed temperature and altitude, which covaried with these environmental clines. Intriguingly, by incorporating gas diffusion rates, it becomes clear that actually more oxygen is available to an organism in warmer habitats where lower oxygen concentrations would suggest the reverse. Under our model, the observed reductions in aerobic performance in warmer habitats do not arise from lower oxygen concentrations, but instead through organismal oxygen demand exceeding supply. This reappraisal of how organismal thermal physiology and oxygen demands together shape aerobic performance in aquatic ectotherms and the new insight of how these components change with temperature have broad implications for predicting the responses of aquatic communities to ongoing global climate shifts.

1. A positive interspecific abundance-occupancy relationship is one of the most robust patterns in macroecology. Yet, the mechanisms driving this pattern are poorly understood. Here, we use biological traits of freshwater macroinvertebrates to gain a mechanistic understanding and disentangle the various explanations. We ask whether mechanisms underlying the abundance-occupancy relationship differ between species, and whether information on individual species can be used to explain their contribution to the interspecific relationship. 2. We test the hypothesis that the importance of metapopulation dynamics or niche differences in explaining the relationship differs between species, varying in relation to their habitat breadth. In addition, we analyse how a species' biological traits shape its habitat breadth and its abundance and occupancy. 3. The abundance and occupancy of the 234 different aquatic macroinvertebrate species were strongly and positively related. Marked differences were found between habitat specialists and habitat generalists in the goodness-of-fit of abundance-occupancy relationships. The occupancy-frequency distribution was bimodal for habitat generalists, allowing 'satellite species' to be distinguished from 'core species'. 4. Habitat generalists appeared to be more widespread but less abundant than habitat specialists, suggesting that the jack-of-all-trades may be master-of-none. Species traits (trophic position and other life-history traits) explained a significant part of the variation around the general relationship. Among habitat specialists, more species showed synchronized life cycles, a low dispersal capacity or clustered oviposition, being better adapted to predictable habitats. Among habitat generalists, more species had long-lived adults, spreading reproductive effort in time and space, and were strong dispersers, being better adapted to unpredictable habitats. 5. Interspecific abundance-occupancy relationships can be best understood by examining the contribution of individual species. For habitat specialists, the interplay between niche differences (diet and habitat use) and the underlying spatial distribution of environmental conditions result in competitive displacement and differences in species' success. For habitat generalists, differences in colonization and extinction rates between species are more important. Therefore, both metapopulation dynamics and niche differences can operate simultaneously but apply to different species, thus constituting different endpoints of the same continuum.

In order to predict which species can successfully cope with global warming and how other environmental stressors modulate their vulnerability to climate‐related environmental factors, an understanding of the ecophysiology underpinning thermal limits is essential for both conservation biology and invasion biology. Heat tolerance and the extent to which heat tolerance differed with oxygen availability were examined for four native and four alien freshwater peracarid crustacean species, with differences in habitat use across species. Three hypotheses were tested: (1) Heat and lack of oxygen synergistically reduce survival of species; (2) patterns in heat tolerance and the modulation thereof by oxygen differ between alien and native species and between species with different habitat use; (3) small animals can better tolerate heat than large animals, and this difference is more pronounced under hypoxia. To assess heat tolerances under different oxygen levels, animal survival was monitored in experimental chambers in which the water temperature was ramped up (0.25°C min−1). Heat tolerance (CTmax) was scored as the cessation of all pleopod movement, and heating trials were performed under hypoxia (5 kPa oxygen), normoxia (20 kPa) and hyperoxia (60 kPa). Heat tolerance differed across species as did the extent by which heat tolerance was affected by oxygen conditions. Heat‐tolerant species, for example, Asellus aquaticus and Crangonyx pseudogracilis, showed little response to oxygen conditions in their CTmax, whereas the CTmax of heat‐sensitive species, for example, Dikerogammarus villosus and Gammarus fossarum, was more plastic, being increased by hyperoxia and reduced by hypoxia. In contrast to other studies on crustaceans, alien species were not more heat‐tolerant than native species. Instead, differences in heat tolerance were best explained by habitat use, with species from standing waters being heat tolerant and species from running waters being heat sensitive. In addition, larger animals displayed lower critical maximum temperature, but only under hypoxia. An analysis of data available in the literature on metabolic responses of the study species to temperature and oxygen conditions suggests that oxygen conformers and species whose oxygen demand rapidly increases with temperature (low activation energy) may be more heat sensitive. The alien species D. villosus appeared most susceptible to hypoxia and heat stress. This may explain why this species is very successful in colonizing new areas in littoral zones with rocky substrate which are well aerated due to continuous wave action generated by passing ships or prevailing winds. This species is less capable of spreading to other waters which are poorly oxygenated and where C. pseudogracilis is the more likely dominant alien species. A plain language summary is available for this article. Plain Language Summary

There is renewed interest in the regulation and consequences of cell size adaptations in studies on understanding the ecophysiology of ectotherms. Here we test if induction of triploidy, which increases cell size in zebrafish (Danio rerio), makes for a good model system to study consequences of cell size. Ideally, diploid and triploid zebrafish should differ in cell size, but should otherwise be comparable in order to be suitable as a model. We induced triploidy by cold shock and compared diploid and triploid zebrafish larvae under standard rearing conditions for differences in genome size, cell size and cell number, development, growth and swimming performance and expression of housekeeping genes and hsp70.1. Triploid zebrafish have larger but fewer cells, and the increase in cell size matched the increase in genome size (+ 50%). Under standard conditions, patterns in gene expression, ontogenetic development and larval growth were near identical between triploids and diploids. However, under demanding conditions (i.e. the maximum swimming velocity during an escape response), triploid larvae performed poorer than their diploid counterparts, especially after repeated stimuli to induce swimming. This result is consistent with the idea that larger cells have less capacity to generate energy, which becomes manifest during repeated physical exertion resulting in increased fatigue. Triploidy induction in zebrafish appears a valid method to increase specifically cell size and this provides a model system to test for consequences of cell size adaptation for the energy budget and swimming performance of this ectothermic vertebrate.

Hybridization influences speciation processes, either slowing or reversing species differentiation due to gene flow and recombination, or else accelerating speciation via adaptive introgression and/or polyploidization. One of many consequences of polyploidization is an increase in cell size associated with genome multiplications. Although cell size is regarded as affecting Darwinian fitness across environmental gradients, particularly due to its effects on oxygen transport, fitness effects of changes in cell size associated with hybridization are not well understood. In this study, we examined the effects of ploidy level, our proxy for cell size, and genotypes on metabolic responses to thermal and oxygen conditions in tadpoles of European water frogs ( Pelophylax esculentus complex). Hybrids ( P. esculentus ), originating from the primary hybridization between P. lessonae (genotype LL) and P. ridibundus (RR), were crossed to produce tadpoles with various genotypes (RR, LR, LLR, LRR) and ploidy levels (diploid, triploid). Our results indicate that triploids, particularly LLR, are most susceptible to oxygen limitation in hypoxic water. Additionally, RR progeny with introgressed P. lessonae mtDNA exhibited the lowest metabolic rates under normoxia, suggesting mitochondrial dysfunction due to mitonuclear incompatibility. The greater oxygen limitation in triploids, particularly under hypoxic conditions, may explain their preference for cooler climates. In a time of rapid environmental change, uncovering the physiological trade‐offs associated with hybrid and polyploid genotypes, in connection with cell size changes, is a promising framework for predicting species responses to shifting oxygen and temperature regimes.

Both oxygen and temperature are fundamental factors determining metabolic performance, fitness, ecological niches, and responses of many aquatic organisms to climate change. Despite the importance of physical and physiological constraints on oxygen supply affecting aerobic metabolism of aquatic ectotherms, ecological theories such as the metabolic theory of ecology have focused on the effects of temperature rather than oxygen. This gap currently impedes mechanistic models from accurately predicting metabolic rates (i.e., oxygen consumption rates) of aquatic organisms and restricts predictions to resting metabolism, which is less affected by oxygen limitation. Here, we expand on models of metabolic scaling by accounting for the role of oxygen availability and temperature on both resting and active metabolic rates. Our model predicts that oxygen limitation is more likely to constrain metabolism in larger, warmer, and active fish. Consequently, active metabolic rates are less responsive to temperature than are resting metabolic rates, and metabolism scales to body size with a smaller exponent whenever temperatures or activity levels are higher. Results from a metaanalysis of fish metabolic rates are consistent with our model predictions. The observed interactive effects of temperature, oxygen availability, and body size predict that global warming will limit the aerobic scope of aquatic ectotherms and may place a greater metabolic burden on larger individuals, impairing their physiological performance in the future. Our model reconciles the metabolic theory with empirical observations of oxygen limitation and provides a formal, quantitative framework for predicting both resting and active metabolic rate and hence aerobic scope of aquatic ectotherms.

For aquatic breathers, hypoxia and warming can act synergistically causing a mismatch between oxygen supply (reduced by hypoxia) and oxygen demand (increased by warming). The vulnerability of these species to such interactive effects may differ during ontogeny due to differing gas exchange systems. This study examines respiratory responses to temperature and hypoxia across four life-stages of the intertidal porcelain crab Petrolisthes laevigatus. Eggs, megalopae, juveniles and adults were exposed to combinations of temperatures from 6 to 18 °C and oxygen tensions from 2 to 21 kPa. Metabolic rates differed strongly across life-stages which could be partly attributed to differences in body mass. However, eggs exhibited significantly lower metabolic rates than predicted for their body mass. For the other three stages, metabolic rates scaled with a mass exponent of 0.89. Mass scaling exponents were similar across all temperatures, but were significantly influenced by oxygen tension (the highest at 9 and 14 kPa, and the lowest at 2 kPa). Respiratory responses across gradients of oxygen tension were used to calculate the response to hypoxia, whereby eggs, megalopae and juveniles responded as oxyconformers and adults as oxyregulators. The thermal sensitivity of the metabolic rates (Q10) were dependent on the oxygen tension in megalopae, and also on the interaction between oxygen tension and temperature intervals in adults. Our results thus provide evidence on how the oxygen tension can modulate the mass dependence of metabolic rates and demonstrate changes in respiratory control from eggs to adults. In light of our results indicating that adults show a good capacity for maintaining metabolism independent of oxygen tension, our study highlights the importance of assessing responses to multiple stressors across different life-stages to determine how vulnerability to warming and hypoxia changes during development.

Insufficient oxygen delivery to tissues is hypothesised to limit thermal tolerance, but evidence in ectotherms is mixed. We assessed heat tolerance under hypoxia, normoxia and hyperoxia to test whether the extent in which oxygen can lower or increase heat tolerance differed with mode of respiration, comparing gill-breathing caenogastropods and lung-breathing pulmonates with or without an accessory gill. Hypoxia lowered heat tolerance in three of the four pulmonates (Physa fontinalis, Physa acuta and Planorbis carinatus) by 1.2–2.1°C. Hyperoxia, however, did not increase the heat tolerance in any of the pulmonate species. Thus, heat tolerance limits of these pulmonates does not appear to be oxygen limited under normoxia, possibly because of their high capacity to regulate oxygen consumption associated with aerial gas exchange. Instead, other processes may become limiting at thermal extremes such as loss of protein function, loss of membrane stability or neuronal dysfunction. The caenogastropod species tested (Potamopyrgus antipodarum, Bithynia tentaculata) closed their operculum during the warming experiments. This behavioural response prevented us from obtaining clear results. Nevertheless, our results suggested hyperoxia may increase heat tolerance in B. tentaculata. This could be related to its lower capacity to regulate oxygen, owing to its fully aquatic gas exchange mechanism.

Linking variation in species' traits to large-scale environmental gradients can lend insight into the evolutionary processes that have shaped functional diversity and future responses to environmental change. Here, we ask how heat and cold tolerance vary as a function of latitude, elevation and climate extremes, using an extensive global dataset of ectotherm and endotherm thermal tolerance limits, while accounting for methodological variation in acclimation temperature, ramping rate and duration of exposure among studies. We show that previously reported relationships between thermal limits and latitude in ectotherms are robust to variation in methods. Heat tolerance of terrestrial ectotherms declined marginally towards higher latitudes and did not vary with elevation, whereas heat tolerance of freshwater and marine ectotherms declined more steeply with latitude. By contrast, cold tolerance limits declined steeply with latitude in marine, intertidal, freshwater and terrestrial ectotherms, and towards higher elevations on land. In all realms, both upper and lower thermal tolerance limits increased with extreme daily temperature, suggesting that different experienced climate extremes across realms explain the patterns, as predicted under the Climate Extremes Hypothesis . Statistically accounting for methodological variation in acclimation temperature, ramping rate and exposure duration improved model fits, and increased slopes with extreme ambient temperature. Our results suggest that fundamentally different patterns of thermal limits found among the earth's realms may be largely explained by differences in episodic thermal extremes among realms, updating global macrophysiological ‘rules’. This article is part of the theme issue ‘Physiological diversity, biodiversity patterns and global climate change: testing key hypotheses involving temperature and oxygen’.