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WIREs Clim Change
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The impacts of heat stress on animal cognition: Implications for adaptation to a changing climate

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Abstract With global surface air temperature rising rapidly, extensive research effort has been dedicated to assessing the consequences of this change for wildlife. While impacts on the phenology, distribution, and demography of wild animal populations are well documented, the impact of increasing temperature on cognition in these populations has received relatively little attention. Cognition encompasses the mental mechanisms that allow individuals to process information from the surrounding environment, respond accordingly, and flexibly adjust behavior. Hence, it is likely to be a key factor in allowing animals to adjust adaptively to climate change. Captive studies show that heat stress can negatively affect cognitive performance not only in the short‐term but also in the long‐term, by altering cognitive development at early life stages. Field studies indicate that cognitive performance may affect survival and reproductive success. However, the link between heat stress, cognition, and fitness in wild animals has yet to be formally established. We propose a comprehensive research framework for the collection of robust empirical datasets on heat stress and cognitive performance in the wild. We then suggest how knowledge of heat stress impacts on cognitive performance could be applied to population viability models and wildlife management actions. We believe that a joint research effort encompassing the fields of thermal physiology, behavioral ecology, comparative cognition, and conservation science, is essential to provide timely mitigation measures against the potential impacts of climate change on wildlife. This article is categorized under: Climate, Ecology, and Conservation > Observed Ecological Changes
The proposed relationship between cognition and animal response to climate change. The same cognitive mechanisms (attention, learning, memory, and decision‐making) regulate multiple behaviors in different contexts. Despite cognition and behavior being intrinsically linked, cognition is not the only cause of changes in behavior: The physiological response to increasing temperatures can directly modify behavior, for example by inducing heat dissipation behaviors and changing activity levels. A key piece of missing information is how increasing temperatures may impact animals' ability to process environmental information, that is cognition itself. This knowledge gap is important because cognition determines behavioral plasticity: It allows individuals to adjust their behavioral responses when conditions change or a behavior stops being successful. Ultimately, cognition can affect an animal's survival and reproductive success (proxies of fitness), which in turn determine population demography and the potential for adjustment to climate changeSource: Photo: Southern pied babbler (Turdoides bicolor) panting and wingspreading to dissipate heat (photo credit: Nicholas B. Pattinson)
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Framework for future research. We propose three steps: (1) determine the temperature‐dependence of cognitive traits in the wild, (2) link variation in cognitive performance to behavioral responses and fitness measures, (3) integrate critical temperature thresholds for cognitive decline into models of population viability under future climate scenarios. Finally, we suggest potential applications to wildlife management and conservation (Ashton et al., 2018, 2019; Barroso et al., 2020; Batabyal & Thaker, 2019; Brakes et al., 2019; Cauchoix et al., 2020; Culina et al., 2020; Dzialowski, 2005; Krochmal et al., 2018; Li et al., 2020; McCafferty et al., 2015; Morand‐Ferron et al., 2016; Salvanes et al., 2013; Shaw & Schmelz, 2017; Shilpa et al., 2017)
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Southern pied babbler (Turdoides bicolor) interacting with an associative color discrimination learning task under a condition of heat stress. This cognitive task measures the ability to learn the association between a color cue (lid) and a food reward (mealworm inside the well). The sequence of photos shows from top to bottom: Individual panting and wingspreading to dissipate heat while perching, individual approaching the task on the ground while panting, individual searching a well and thus completing a trial. The photos were taken at the Kuruman River Reserve (Northern Cape, South Africa) during cognitive testing on a wild ringed individual as part of an ongoing research project (Ridley, 2020)Source: Photo credit: Nicholas B. Pattinson
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The proposed relationship between temperature, behavior, and cognition. Temperature affects animal behavior through bottom‐up and top‐down processes (Abram et al., 2017). The top‐down effect of temperature refers to the processing of temperature information from sensory neurons to produce a thermoregulatory response (both physiological and behavioral). The bottom‐up effect of temperature refers to the temperature‐dependent rate of biochemical processes, which directly affects metabolism, growth rates, and neuronal function. Changes in metabolism can translate into behavioral changes, for example, altered feeding rates, without involving cognition. The present review focuses on how temperature may affect cognitive mechanisms (arrows highlighted in yellow). These include attention, learning, memory, and decision‐making mechanisms, which allow selective filtering of environmental stimuli and modify behavior through experience (Shettleworth, 2010). The bottom‐up highlighted arrow indicates that temperature‐induced changes in neuronal function may lead to changes in cognitive performance, for example, impaired learning. The top‐down highlighted arrow indicates that thermal information may reduce the cognitive capacity to process other competing external stimuli. Cognition regulates behavior in different contexts from foraging to predator avoidance, mate choice, and thermoregulation itself. Therefore, temperature‐induced effects on cognition may translate into behavioral changes relevant for survival and reproductive success in the wild
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