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Human Heat Adaptation

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Abstract

In this overview, human morphological and functional adaptations during naturally and artificially induced heat adaptation are explored. Through discussions of adaptation theory and practice, a theoretical basis is constructed for evaluating heat adaptation. It will be argued that some adaptations are specific to the treatment used, while others are generalized. Regarding ethnic differences in heat tolerance, the case is put that reported differences in heat tolerance are not due to natural selection, but can be explained on the basis of variations in adaptation opportunity. These concepts are expanded to illustrate how traditional heat adaptation and acclimatization represent forms of habituation, and thermal clamping (controlled hyperthermia) is proposed as a superior model for mechanistic research. Indeed, this technique has led to questioning the perceived wisdom of body‐fluid changes, such as the expansion and subsequent decay of plasma volume, and sudomotor function, including sweat habituation and redistribution. Throughout, this contribution was aimed at taking another step toward understanding the phenomenon of heat adaptation and stimulating future research. In this regard, research questions are posed concerning the influence that variations in morphological configuration may exert upon adaptation, the determinants of postexercise plasma volume recovery, and the physiological mechanisms that modify the cholinergic sensitivity of sweat glands, and changes in basal metabolic rate and body core temperature following adaptation. © 2014 American Physiological Society. Compr Physiol 4:325‐365, 2014.

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Figure 1. Figure 1. An integrated overview of the homeostatic mechanisms and physiological responses accompanying hot thermal challenges. Five critical variables (bottom row) must be kept within ranges conducive to optimal physiological function while simultaneously avoiding states that are either hazardous or life threatening. Heat‐induced modifications of these variables will be explored within acclimatized, indigenous populations, and in those who have been repeatedly exposed to artificially manipulated thermal stress (heat acclimated). These regulated variables () define, in part, the internal environment over which humans have acquired an ability to maintain considerable stability. To do this, dedicated sensors (top row) respond to changes in these regulated variables (arrows at the bottom link to these sensors), providing feedback to central nervous structures that integrate and evaluate this information, and then modulate the function of one or more effector organs (). As a consequence, specific controlled (dependent) variables (second bottom row) are modified to sustain a relatively constant milieu intérieur (). These separate, yet well‐integrated regulatory processes defend homeostasis in the heat, and form a framework for understanding and investigating human heat adaptation.
Figure 2. Figure 2. Characteristics of physiological adaptation (adapted from ideas of ). With the application of a sufficiently strong stimulus (thermal impulse), homeostasis is disturbed. If this disruption is large enough, the adaptation threshold will be exceeded and physiological adaptation will be initiated, albeit with some delay (latency). Adaptation impulses that do not induce systemic failure will eventually elicit more complete adaptation (plateau), with the difference between the genetically determined physiological maximum and the adaptation plateau defining the potential for further improvements to be realized (adaptation reserve). Variations in the location of this plateau reflect changes in the capacity to tolerate stress (accommodation reserve). Finally, removal of the thermal impulse for a sufficiently long duration results in a decay of these acquired physiological adaptations.
Figure 3. Figure 3. Local sweating responses of matched Asian (Thai; N = 5) and Caucasian women (N = 5) cycling at 35% of their peak power for 90 min in humid heat (35.0°C, 59% relative humidity). Unpublished observations: Thoicharoen and Taylor (2006).
Figure 4. Figure 4. Rectal temperature, sweating, and heart rate responses during successive days of heat adaptation to a fixed exercise forcing function (N = 6; bench stepping 12 steps.min−1, 4 h) and a constant thermal load (34°C, ∼80% relative humidity). Data extracted and redrawn, with permission, from Wyndham et al. ().
Figure 5. Figure 5. Serial changes in blood volume during consecutive days (April, 1922) that involved continuous exposure firstly to cold air (2 days), then to the heat (2 days) and finally back to cold air (data extracted and redrawn, with permission, from : N = 1).
Figure 6. Figure 6. Basal mass‐specific, body‐fluid compartment volumes for sedentary women (white bars), men (hatched bars), and endurance‐trained males (cross‐hatched bars). Sedentary data are gender‐specific reference means () while data for trained individuals are means taken from Maw et al. ().
Figure 7. Figure 7. Basal mass‐specific, body‐fluid compartment volumes before (day 1 [control]: white bars), during (day 8: hatched bars), and immediately after (day 22: cross‐hatched bars) 17 days of heat acclimation during which participants cycled (semirecumbent) in the heat (N = 12; 90 min, 40°C, 60% relative humidity), but with the exercise intensity modified to clamp core temperature at 38.5°C (controlled hyperthermia). A rest day preceded each test day, while all other heat exposures occurred consecutively. Data are means with standard errors of the means extracted from Patterson et al. (), with each asterisk indicating a significant change from day 1. Nonsignificant increases in the extracellular and interstitial volumes on day 22 were ascribed to intersubject variability, since subsequent data from the same laboratory, collected using more sensitive techniques, revealed these increments to be significant ().
Figure 8. Figure 8. A theoretical explanation for reductions in the basal mean body temperature following heat adaptation (adapted, with permission, from ). The gray lines show the dependence of mean body temperature on cutaneous blood flow, with lower flows resulting in higher temperatures, and with heat adaptation increasing cutaneous perfusion at any given body temperature (upward displacement to the broken line). The black lines illustrate sudomotor activity prior to (solid), and following heat adaptation, as driven by changes in mean body temperature. Adaptation lowers the body temperature threshold for initiating sweating (leftward displacement to the broken line) as well as increasing both sudomotor sensitivity and sweat flows. The intersections of the solid and the broken (postadaptation) lines determine the basal mean body temperature for each state, which is reduced following heat adaptation.
Figure 9. Figure 9. Core temperature responses (solid line) of preheated individuals (38°C; N = 16) placed within an air‐tight and heated chamber (1.2 m3, 45°C, 40% relative humidity). Beyond 10 min, the ambient water vapor pressure increased to the point that effective evaporative cooling became negligible. Data taken from Haberley et al. ().
Figure 10. Figure 10. Local sweat rates during an exercising heat stress test conducted before (day 1 [control]: white bars), during (day 8: hatched bars), and immediately after (day 22: cross‐hatched bars) 17 days of heat acclimation. During heat acclimation, participants cycled (semirecumbent) in the heat (N = 11; 90 min, 40°C, 60% relative humidity), alternately exercising and resting to clamp core temperature at 38.5°C. Data are means with standard errors of the means. Percentage changes for day 22 are expressed relative to the sweat rates observed on day 1. Modified from Patterson et al. ().


Figure 1. An integrated overview of the homeostatic mechanisms and physiological responses accompanying hot thermal challenges. Five critical variables (bottom row) must be kept within ranges conducive to optimal physiological function while simultaneously avoiding states that are either hazardous or life threatening. Heat‐induced modifications of these variables will be explored within acclimatized, indigenous populations, and in those who have been repeatedly exposed to artificially manipulated thermal stress (heat acclimated). These regulated variables () define, in part, the internal environment over which humans have acquired an ability to maintain considerable stability. To do this, dedicated sensors (top row) respond to changes in these regulated variables (arrows at the bottom link to these sensors), providing feedback to central nervous structures that integrate and evaluate this information, and then modulate the function of one or more effector organs (). As a consequence, specific controlled (dependent) variables (second bottom row) are modified to sustain a relatively constant milieu intérieur (). These separate, yet well‐integrated regulatory processes defend homeostasis in the heat, and form a framework for understanding and investigating human heat adaptation.


Figure 2. Characteristics of physiological adaptation (adapted from ideas of ). With the application of a sufficiently strong stimulus (thermal impulse), homeostasis is disturbed. If this disruption is large enough, the adaptation threshold will be exceeded and physiological adaptation will be initiated, albeit with some delay (latency). Adaptation impulses that do not induce systemic failure will eventually elicit more complete adaptation (plateau), with the difference between the genetically determined physiological maximum and the adaptation plateau defining the potential for further improvements to be realized (adaptation reserve). Variations in the location of this plateau reflect changes in the capacity to tolerate stress (accommodation reserve). Finally, removal of the thermal impulse for a sufficiently long duration results in a decay of these acquired physiological adaptations.


Figure 3. Local sweating responses of matched Asian (Thai; N = 5) and Caucasian women (N = 5) cycling at 35% of their peak power for 90 min in humid heat (35.0°C, 59% relative humidity). Unpublished observations: Thoicharoen and Taylor (2006).


Figure 4. Rectal temperature, sweating, and heart rate responses during successive days of heat adaptation to a fixed exercise forcing function (N = 6; bench stepping 12 steps.min−1, 4 h) and a constant thermal load (34°C, ∼80% relative humidity). Data extracted and redrawn, with permission, from Wyndham et al. ().


Figure 5. Serial changes in blood volume during consecutive days (April, 1922) that involved continuous exposure firstly to cold air (2 days), then to the heat (2 days) and finally back to cold air (data extracted and redrawn, with permission, from : N = 1).


Figure 6. Basal mass‐specific, body‐fluid compartment volumes for sedentary women (white bars), men (hatched bars), and endurance‐trained males (cross‐hatched bars). Sedentary data are gender‐specific reference means () while data for trained individuals are means taken from Maw et al. ().


Figure 7. Basal mass‐specific, body‐fluid compartment volumes before (day 1 [control]: white bars), during (day 8: hatched bars), and immediately after (day 22: cross‐hatched bars) 17 days of heat acclimation during which participants cycled (semirecumbent) in the heat (N = 12; 90 min, 40°C, 60% relative humidity), but with the exercise intensity modified to clamp core temperature at 38.5°C (controlled hyperthermia). A rest day preceded each test day, while all other heat exposures occurred consecutively. Data are means with standard errors of the means extracted from Patterson et al. (), with each asterisk indicating a significant change from day 1. Nonsignificant increases in the extracellular and interstitial volumes on day 22 were ascribed to intersubject variability, since subsequent data from the same laboratory, collected using more sensitive techniques, revealed these increments to be significant ().


Figure 8. A theoretical explanation for reductions in the basal mean body temperature following heat adaptation (adapted, with permission, from ). The gray lines show the dependence of mean body temperature on cutaneous blood flow, with lower flows resulting in higher temperatures, and with heat adaptation increasing cutaneous perfusion at any given body temperature (upward displacement to the broken line). The black lines illustrate sudomotor activity prior to (solid), and following heat adaptation, as driven by changes in mean body temperature. Adaptation lowers the body temperature threshold for initiating sweating (leftward displacement to the broken line) as well as increasing both sudomotor sensitivity and sweat flows. The intersections of the solid and the broken (postadaptation) lines determine the basal mean body temperature for each state, which is reduced following heat adaptation.


Figure 9. Core temperature responses (solid line) of preheated individuals (38°C; N = 16) placed within an air‐tight and heated chamber (1.2 m3, 45°C, 40% relative humidity). Beyond 10 min, the ambient water vapor pressure increased to the point that effective evaporative cooling became negligible. Data taken from Haberley et al. ().


Figure 10. Local sweat rates during an exercising heat stress test conducted before (day 1 [control]: white bars), during (day 8: hatched bars), and immediately after (day 22: cross‐hatched bars) 17 days of heat acclimation. During heat acclimation, participants cycled (semirecumbent) in the heat (N = 11; 90 min, 40°C, 60% relative humidity), alternately exercising and resting to clamp core temperature at 38.5°C. Data are means with standard errors of the means. Percentage changes for day 22 are expressed relative to the sweat rates observed on day 1. Modified from Patterson et al. ().
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Nigel A.S. Taylor. Human Heat Adaptation. Compr Physiol 2014, 4: 325-365. doi: 10.1002/cphy.c130022