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Heat adaptation in humans: the significance of controlled and regulated variables for experimental design and interpretation

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Abstract

Herein, the principles of homoeostasis are re-visited, but with an emphasis upon repeated homoeostatic disturbances that give rise to physiological adaptation. The central focus is human heat adaptation, and how, for experimental purposes, one might standardise successive adaptation stimuli, and then evaluate and compare the resulting adaptations. To provide sufficient background for that discussion, the principles of physiological control and regulation have been reviewed. The case is presented that, since it is the regulated variables that drive both the effector organs and the processes of physiological adaptation, then it is those variables (e.g., body temperature) that should be used to set and standardise the adaptation stimuli. Alternatively, some have proposed that the same outcome can be achieved through standardising a controlled variable (e.g., heart rate), and so the merits of that proposition are evaluated. Indeed, it can be an effective approach, although some experimental pitfalls are described to highlight its limitations with regard to between-group (e.g., able-bodied versus spinal-injured participants) and between-treatment comparisons (e.g., hot-water versus hot-air adaptation stimuli). The concept of setting the adaptation stimulus relative to an anaerobic or lactate threshold is also critically evaluated. Finally, an appraisal is offered concerning the merits of three different strategies for using deep-body and mean body temperature changes for evaluating thermoeffector adaptations.

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Fig. 1

[modified from the schematics of Werner et al. (2008) and Taylor (2014) using BioRender.com]. Five relevant, regulated variables are illustrated, each with its own unique sensors (top and bottom rows). However, whilst some variables are discussed in detail, others are presented here, but not discussed, to highlight the importance of understanding the broader implications of integrated physiological regulation. Humans acquired the ability to regulate those variables within zones that optimise physiological function and that define the characteristics of the milieu intérieur. Sensory feedback ascends to the central integrators which, in turn, will modify the function of the autonomically controlled cells, tissues and organs. The outcome of those control mechanisms is that an array of integrated physiological responses can be activated (the controlled variables), and schematics such as this provide a first-principles framework for investigating and understanding human homoeostasis and heat adaptation

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Abbreviations

ATP:

Adenosine triphosphate

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Acknowledgements

NAST was supported by the Brain Pool Program through the National Research Foundation of Korea (NRF), funded by the Ministry of Science and Information, Communication and Technology (Grant number: 2019H1D3A2A01061171). SRN was supported by a postdoctoral fellowship from the Human and Environmental Physiology Research Unit (University of Ottawa, Canada).

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Taylor, N.A.S., Notley, S.R. & Lindinger, M.I. Heat adaptation in humans: the significance of controlled and regulated variables for experimental design and interpretation. Eur J Appl Physiol 120, 2583–2595 (2020). https://doi.org/10.1007/s00421-020-04489-0

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