Oxygen- And Capacity-limited Thermal Tolerance Research Articles

Aerobic scope and its optimum temperature: clarifying their usefulness and limitations – correspondence on J. Exp. Biol. 216, 2771-2782

Clark and colleagues ([Clark et al., 2013][1]) have set people straight on how to make high quality measurements of maximum and minimum oxygen uptake rates in fishes. They correctly state that great care and understanding are needed to properly measure a fish's aerobic scope (=maximum metabolic rate

Aerobic scope measurements of fishes in an era of climate change: respirometry, relevance and recommendations

Measurements of aerobic scope [the difference between minimum and maximum oxygen consumption rate ( and , respectively)] are increasing in prevalence as a tool to address questions relating to fish ecology and the effects of climate change. However, there are underlying issues regarding the array of methods used to measure aerobic scope across studies and species. In an attempt to enhance quality control before the diversity of issues becomes too great to remedy, this paper outlines common techniques and pitfalls associated with measurements of , and aerobic scope across species and under different experimental conditions. Additionally, we provide a brief critique of the oxygen- and capacity-limited thermal tolerance (OCLTT) hypothesis, a concept that is intricately dependent on aerobic scope measurements and is spreading wildly throughout the literature despite little evidence for its general applicability. It is the intention of this paper to encourage transparency and accuracy in future studies that measure the aerobic metabolism of fishes, and to highlight the fundamental issues with assuming broad relevance of the OCLTT hypothesis.

Aerobic scope does not predict the performance of a tropical eurythermal fish at elevated temperatures

Climate warming is predicted to negatively impact fish populations through impairment of oxygen transport systems when temperatures exceed those which are optimal for aerobic scope (AS). This concept of oxygen- and capacity-limited thermal tolerance (OCLTT) is rapidly gaining popularity within climate change research and has been applied to several fish species. Here, we evaluated the relevance of aerobic performance of juvenile barramundi (Lates calcarifer) in the context of thermal preference and tolerance by (1) measuring standard and maximum metabolic rates (SMR and MMR, respectively) and AS of fish acclimated to 29°C and acutely exposed to temperatures from 23 to 38°C, (2) allowing the fish to behaviourally select a preferred temperature between 29 and 38°C, and (3) quantifying alterations to AS after 5 weeks of acclimation to 29 and 38°C. SMR and MMR both increased continuously with temperature in acutely exposed fish, but the increase was greater for MMR such that AS was highest at 38°C, a temperature approaching the upper lethal limit (40-41°C). Despite 38°C eliciting maximum AS, when given the opportunity the fish selected a median temperature of 31.7 ± 0.5°C and spent only 10 ± 3% of their time at temperatures >36°C. Following acclimation to 38°C, AS measured at 38°C was decreased to the same level as 29°C-acclimated fish measured at 29°C, suggesting that AS may be dynamically modulated independent of temperature to accommodate the requirements of daily life. Together, these results reveal limited power of the OCLTT hypothesis in predicting optimal temperatures and effects of climate warming on juvenile barramundi.

A role for haemolymph oxygen capacity in heat tolerance of eurythermal crabs.

Heat tolerance in aquatic ectotherms is constrained by a mismatch, occurring at high temperatures, between oxygen delivery and demand which compromises the maintenance of aerobic scope. The present study analyses how the wide thermal tolerance range of an eurythermal model species, the green crab Carcinus maenas is supported and limited by its ability to sustain efficient oxygen transport to tissues. Similar to other eurytherms, C. maenas sustains naturally occurring acute warming events through the integrated response of circulatory and respiratory systems. The response of C. maenas to warming can be characterized by two phases. During initial warming, oxygen consumption and heart rate increase, while stroke volume and haemolymph oxygen partial pressure decrease. During further warming, dissolved oxygen levels in the venous compartment decrease below the threshold of full haemocyanin oxygen saturation. The progressive release of haemocyanin bound oxygen with further warming follows an exponential pattern, thereby saving energy in oxygen transport and causing an associated leveling off of metabolic rate. According to the concept of oxygen and capacity limited thermal tolerance (OCLTT), this indicates that the thermal tolerance window is widened by the increasing contribution of haemocyanin oxygen transport and associated energy savings in cardiocirculation. Haemocyanin bound oxygen sustains cardiac performance to cover the temperature range experienced by C. maenas in the field. To our knowledge this is the first study providing evidence of a relationship between thermal tolerance and blood (haemolymph) oxygen transport in a eurythermal invertebrate.

Thermal Acclimation Is Not Necessary to Maintain a Wide Thermal Breadth of Aerobic Scope in the Common Killifish (Fundulus heteroclitus)

Loss of aerobic scope at high and low temperatures is a physiological mechanism proposed to limit the thermal performance and tolerance of organisms, a theory known as oxygen- and capacity-limited thermal tolerance (OCLTT). Eurythermal organisms maintain aerobic scope over wide ranges of temperatures, but it is unknown whether acclimation is necessary to maintain this breadth. The objective of this study was to examine changes in aerobic scope in Fundulus heteroclitus, a eurythermal fish, after acclimation and acute exposure to temperatures from 5° to 33°C. The range of temperatures over which aerobic scope was nonzero was similar in acclimated and acutely exposed fish, suggesting that acclimation has modest effects on the thermal breadth of aerobic scope. However, in acclimated fish, there was a clear optimum temperature range for aerobic scope between 25° and 30°C, whereas aerobic scope was relatively constant across the entire temperature range with acute temperature exposure. Therefore, the primary effect of acclimation was to increase aerobic scope between 25° and 30°C, which paradoxically resulted in a narrower temperature range of optimal performance in acclimated fish compared to acutely exposed fish. There was only weak evidence for correlations between the thermal optimum of aerobic scope and the thermal optimum of measures of performance (specific growth rate and gonadosomatic index), and indicators of anaerobic metabolism (lactate accumulation and lactate dehydrogenase activity) only increased at high temperatures. Together these data fit many, but not all, of the predictions made by OCLTT.

Physiological Responses in a Variable Environment: Relationships between Metabolism, Hsp and Thermotolerance in an Intertidal-Subtidal Species

Physiological responses to temperature reflect the evolutionary adaptations of organisms to their thermal environment and the capability of animals to tolerate thermal stress. Contrary to conventional metabolism theory, increasing environmental temperatures have been shown to reduce metabolic rate in rocky–eulittoral-fringe species inhabiting highly variable environments, possibly as a strategy for energy conservation. To study the physiological adaptations of an intertidal-subtidal species to the extreme and unpredictable heat stress of the intertidal zone, oxygen consumption rate and heat shock protein expression were quantified in the sea cucumber Apostichopus japonicus. Using simulate natural temperatures, the relationship between temperature, physiological performance (oxygen consumption and heat shock proteins) and thermotolerance were assessed. Depression of oxygen consumption rate and upregulation of heat shock protein genes (hsps) occurred in sequence when ambient temperature was increased from 24 to 30°C. Large-scale mortality of the sea cucumber occurred when temperatures rose beyond 30°C, suggesting that the upregulation of heat shock proteins and mortality are closely related to the depression of aerobic metabolism, a phenomenon that is in line with the concept of oxygen- and capacity-limited thermal tolerance (OCLTT). The physiologically-related thermotolerance of this sea cucumber should be an adaptation to its local environment.

Climate change effects on fishes and fisheries: towards a cause‐and‐effect understanding

Ongoing climate change is predicted to affect individual organisms during all life stages, thereby affecting populations of a species, communities and the functioning of ecosystems. These effects of climate change can be direct, through changing water temperatures and associated phenologies, the lengths and frequency of hypoxia events, through ongoing ocean acidification trends or through shifts in hydrodynamics and in sea level. In some cases, climate interactions with a species will also, or mostly, be indirect and mediated through direct effects on key prey species which change the composition and dynamic coupling of food webs. Thus, the implications of climate change for marine fish populations can be seen to result from phenomena at four interlinked levels of biological organization: (1) organismal-level physiological changes will occur in response to changing environmental variables such as temperature, dissolved oxygen and ocean carbon dioxide levels. An integrated view of relevant effects, adaptation processes and tolerance limits is provided by the concept of oxygen and capacity-limited thermal tolerance (OCLT). (2) Individual-level behavioural changes may occur such as the avoidance of unfavourable conditions and, if possible, movement into suitable areas. (3) Population-level changes may be observed via changes in the balance between rates of mortality, growth and reproduction. This includes changes in the retention or dispersion of early life stages by ocean currents, which lead to the establishment of new populations in new areas or abandonment of traditional habitats. (4) Ecosystem-level changes in productivity and food web interactions will result from differing physiological responses by organisms at different levels of the food web. The shifts in biogeography and warming-induced biodiversity will affect species productivity and may, thus, explain changes in fisheries economies. This paper tries to establish links between various levels of biological organization by means of addressing the effective physiological principles at the cellular, tissue and whole organism levels.