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com
BJSM Online First, published on October 20, 2009 as 10.1136/bjsm.2009.063024

EXERCISE AND HEAT STRESS: PERFORMANCE, FATIGUE AND EXHAUSTION

- A HOT TOPIC.

Zachary J. Schlader, Stephen R. Stannard, Toby Mündel

Institute of Food, Nutrition and Human Health, Massey University, Palmerston North,
New Zealand

Corresponding Author:
Zachary J. Schlader
Institute of Food, Nutrition and Human Health
Private Bag 11 222
Palmerston North, New Zealand, 4442
Telephone: 64 6 356 9099 ext 4848
Fax: 64 6 350 5857
Email: Z.J.Schlader@massey.ac.nz

Word Count (Abstract/Editorial): 93/1654

Running Head: Exercise and Heat Stress

Copyright Article author (or their employer) 2009. Produced by BMJ Publishing Group Ltd under licence.
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ABSTRACT
The attenuation of exercise duration/performance during exercise in the heat has
spawned the use of exercise and heat stress as a model for fatigue. However, the
mechanisms underlying this premature fatigue in the heat remain unclear and have been
debated accordingly. Principle to this debate is the use of both fixed-intensity and self-
paced exercise protocols which have produced seemingly opposing theories. However, a
retrospective analysis of previously published data suggests that these theories are in fact
complementary and act together in order to maintain thermoregulation and prevent the
development of exertional heat stroke.

Keywords: Thermoregulation, Anticipatory Regulation, Critical Core Temperature,

Physiological Strain, Homeostasis

EDITORIAL
Fatigue is suggested to be a general concept which is characterized by an acute
impairment in performance and leads to the eventual inability to produce the necessary
force, i.e. volitional exhaustion 2. Homeostasis is maintained by the physiological
regulatory processes occurring throughout the body 3 and during exercise the
development of fatigue and exhaustion are proposed to prevent homeostatic failure 4.
Principle to the regulatory systems maintaining homeostasis is that of thermoregulation,
and as such the combined stressors of exercise and heat stress are used as a model to
investigate the regulatory processes that influence physical performance, the development
of fatigue, and the onset of exhaustion 5,6. Accordingly, theories have been developed,
healthy (lively) debate has arisen, and various journals have seen fit to devote special
issues e.g. Sports Medicine (vol. 37, issues 4-5), Journal of Applied Physiology (vol. 104,
issue 5) and timely reviews 5-8.

Within the exercise and heat stress model two exercise protocols have been
utilized which both describe performance as a dependent variable of time: i.) Fixed-
intensity (i.e. constant workload) exercise – exercise is undergone until volitional
exhaustion, and ii.) self-paced exercise – which evaluates the time to complete a given
task (e.g. a set amount of work) where the workload can be adjusted by the exerciser.
Both exercise time-to-exhaustion and self-paced exercise performance have been found
to be attenuated under heat stress 9-17. However, confounding these conclusions is that
these exercise protocols evaluate a different combination of physiological processes.
Fixed-intensity exercise provides a measure of exhaustion whereas self-paced exercise
provides an indication of the development of fatigue, and arising from this limitation
seemingly opposing theories have developed.

Thermoregulation is attained by both passive (i.e. non-regulated) and active (i.e.

regulated) systems 18. During exercise in the heat passive heat loss (e.g. evaporation,
conduction, etc.) is maximized by the regulation of active thermoregulatory mechanisms
(i.e. autonomous and/or behavioral). When the heat loss required for heat balance is
exceeded by metabolic heat production (i.e. uncompensable heat stress),

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thermoregulation is dependent upon behavioral adjustments in metabolic heat production,

which is proportional to exercise work-rate 19.

During fixed-intensity exercise in the heat metabolic heat production is constant,

heat loss is limited solely to autonomic mechanisms, and thus a proportional, passive (i.e.
non-regulated) rise in core temperature ensues. Under these conditions thermoregulation
mediates voluntary (i.e. behavioral) exercise exhaustion upon the attainment of a core
temperature of ~40°C 10,11,14,15, although the absolute temperature at which exhaustion
occurs varies with aerobic fitness 9. These findings contribute to the theory that exercise
duration in the heat is limited by a critical core temperature 7,20 with this threshold core
temperature presumably preventing the development of exertional heat stroke, which, in
combination with other sign and symptoms, is characterized by a core temperature >40°C
21
. It is acknowledged, however, that this critical core temperature of ~40°C is primarily
a laboratory based phenomenon as this core temperature threshold can be exceeded
during ‘real world’ situations. For instance, during competitive situations core
temperatures in highly trained runners have been found to exceed 40°C without any
indication of the development of heat illness 22,23, suggesting that thermoregulation was
not compromised by a high core temperature in these athletes.

In contrast, during self-paced exercise the thermal compensability of the

environment can be manipulated by the exerciser through adjustments in work-rate (i.e.
metabolic heat production). These behavioral adjustments in work-rate result in a
regulated rise in core temperature and permit completion of the exercise task while
avoiding the attainment of a critically high core temperature. However, the consequence
of this behavior is a reduction in exercise performance 12,13,16,17 which provides the
theoretical basis for the theory of anticipatory regulation and avoidance. 8

Given that fixed-intensity and self-paced exercise protocols evaluate different

physiological phenomena (e.g. exhaustion and fatigue, respectively), the theories of the
critical core temperature and anticipatory regulation and avoidance have been built up
as mutually exclusive 7 and alternatives of each other 20. However, it has recently been
suggested that these theories actually complement one another and are in fact ‘safety’
mechanisms 7 acting to maintain thermoregulation. Yet, despite the suggestion that
future research should try to incorporate both fixed-intensity and self-paced exercise
protocols 7 an established relationship between anticipatory regulation and a critical core
temperature has remained elusive, suggesting that these theories are conflicting.

It is proposed here that anticipatory regulation and critical core temperature exist
on a continuum. At one end of this continuum is voluntary exhaustion, which occurs
immediately prior to loss of thermoregulation, while fatigue is the progression of
processes that potentially develop into this exhaustion. To test this proposal we
conducted a retrospective analysis of the physiological strain at exhaustion/completion of
both fixed-intensity and self-paced exercise protocols, respectively. Physiological strain
was indicated by the physiological strain index (PSI), modified from Moran et al. 1. The
PSI was chosen for this analysis as it mathematically accounted for the limitation posed
by elevations in body temperature.

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Mean heart rates and core temperatures prior to exercise and at exhaustion (fixed-
intensity exercise) or completion (self-paced exercise) were used as noted in the
published manuscripts. Despite there being a relatively large selection of literature, the
studies used for this analysis were chosen based on the availability of published mean
data. If the necessary data at the conclusion of the exercise bouts were unavailable the
study was excluded. Hence, this resulted in the inclusion of 4 (of the 7 considered) for
fixed-intensity exercise and 4 (of 4) for self-paced exercise.

While data at the termination of exercise was not compromised, if pre-exercise

heart rates or core temperatures were not provided 60 bpm and 37.0 °C, respectively,
were used as estimates. In the original PSI maximal core temperature was ethically
limited to 39.5 °C, but given that core temperature was allowed to approach, and in some
cases exceed 40.0 °C in the literature reviewed, we have modified the original PSI to
allow for a maximal core temperature of 40.0 °C.

Despite the relatively small number of studies used in the analysis, the PSI was
regarded as an appropriate platform for comparing the level of physiological strain at the
conclusion of fixed-intensity and self-paced exercise bouts. However, it is acknowledged
that the PSI is a somewhat crude method of evaluating physiological strain, and that it
does not take into account methodological differences concerning core temperature
measurement location, and exercise mode or intensity. However, given the definition of
the thermal core (“those inner tissues of the body whose temperatures are not changed in
their relationship to each other by circulatory adjustments and changes in heat dissipation
to the environment…3”), that all studies utilized one of the acceptable methods of
measuring core body temperature (i.e. either esophageal or rectal temperature) 3, and that
a maximal exercise effort was assumed in all instances, these limitations were considered
acceptable. Further, given the precise regulation of core temperature, the subjects’
relatively high level of aerobic fitness, and that fluctuations in resting core temperature
and heart rates ± 1.0 °C and 10 bpm, respectively, did not significantly affect the PSI
outcome, 37.0 °C and 60 bpm were deemed sufficient estimates where resting core
temperature and heart rate data were unavailable.

As expected, in the heat physiological strain was higher at the finish of the fixed-
intensity exercise bouts when compared to self-paced (Table 1). During fixed-intensity
exercise, the physiological strain was mediated predominantly by the elevation in core
temperature and not heart rate, as heart rate was higher at completion of the self-paced
exercise. This elevated heart rate, probably due to an ‘all out effort’ at the final stages of
the self-paced exercise bouts, indicates the presence of a physiological ‘reserve’ which
allowed this ‘effort’ to be made without compromising thermoregulation; suggesting that
throughout self-paced exercise in the heat the level of physiological strain was controlled
at a lower level when compared to fixed-intensity exercise. In explanation, during fixed-
intensity exercise the progression of physiological strain, most likely coinciding with the
passive rise in core temperature, passed directly through those levels incurred upon
completion of the self-paced exercise bouts; exercise exhaustion was achieved by
attaining a maximum tolerable level of physiological strain (Figure 1), which is suggested
to occur immediately prior to the loss of thermoregulation.

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Table 1: A comparison of physiological strain upon completion of fixed-

intensity and self-paced exercise in the heat.

Exercise Protocol TCo TCt HRo HRt PSI¹

Fixed-Intensity Exercise
70% VO2 max 10 37.8 40.1 68 174.0 10.0
55 - 77% VO2 max 14 36.4 39.3 70 183.3 9.2
60% VO2 max 11 37.4 40.2 95 197.4 11.4
60% VO2 max 15 37.0 40.0 60 179.0 10.0

Mean 39.9* 183.4* 10.1*

Self-Paced Exercise
20 km time-trial 17 37.5 39.2 60² 184.0 8.6
8 km run 12 37.8 39.4 60² 187.5 8.9
8 km run 13 37.0² 39.5 60² 186.0 9.4
30 min time-trial 16 37.0² 39.2 60² 195.0 9.3

Mean 39.3 188.1 9.1

¹ Physiologic Strain Index 1 as calculated: PSI=5(TCt-TCo)/(40.0-TCo)+5(HRt-

HRo)/(180-HRo) where: TCo, mean resting core temperature (°C); TCt, mean
core temperature at exhaustion/completion (°C); HRo, mean resting heart rate
(bpm); HRt, mean heart rate at exhaustion/completion (bpm). ² TCo and HRo
not provided, estimated as 37.0 °C and 60 bpm, respectively. * Significantly
different from self-paced exercise (P<0.05) as determined via paired t-tests.

The reduction in exercise performance in the heat observed in non-laboratory

situations (see 8 pg. 562) is thus proposed to be mediated by both anticipatory regulation
and a critical core temperature. In these ‘real-life’ settings these ‘safety’ mechanisms act
together to maintain thermoregulation. In the heat, the development of fatigue is
monitored and behavioral adjustments in work-rate are made in order to safely complete
the exercise bout at hand (i.e. anticipatory regulation). However, in some situations (e.g.
competitive circumstances, highly motivated individuals, etc.) these fatigue signals can
be overridden and exercise performance is limited by volitional exhaustion (i.e. critical
core temperature), which by definition is exertional heat exhaustion. 21 However, it is
upon the failure of both of these mechanisms that thermoregulation is lost, homeostasis is
compromised, and the development of potentially life threatening exertional heat stroke is
imminent (Figure 2).

These data support our proposal that during exercise in the heat the process of
anticipatory regulation and the attainment of a critical core temperature complement one
another in order to maintain thermoregulation and in turn homeostasis. As our intentions
for this proposal were to: i.) suggest an alternative way to interpret these seemingly
opposing data, and ii.) stimulate further research in this area, the PSI appears to have
been an appropriate tool. Accordingly, the development of novel and innovative
experimental approaches will further integrate these contrasting theories, helping to
extrapolate laboratory findings to ‘real-life’ situations. The prevailing outcome is not
whether this proposal will be supported or refuted by future research. Ultimately, it is

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hoped that it will lead to a better understanding of the development of exercise-induced

hyperthermic fatigue/exhaustion, the breakdown of these ‘safety’ systems, and a greater
comprehension of the fatigue process and its multitude of causes.

COMPETING INTERESTS
None

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FIGURE LEGENDS
Figure 1:
Proposed schematic of fixed-intensity (A) vs. self-paced (B) exercise under heat stress.
Dashed line indicates the hypothetical maximal physiological strain beyond which
thermoregulatory control will be lost. Under heat stress, A: the rise in physiological
strain during fixed-intensity exercise is mediated predominately by the passive rise in
core temperature, and B: the rise in physiological strain during self-paced exercise is
mediated predominately by a regulated rise in core temperature.

Figure 2:
Physiological homeostasis as maintained by anticipatory regulation and a critical core
temperature during exercise in the heat. Thermoregulation is maintained until exercise is
continued after the attainment of a critical core temperature, beyond which physiological
homeostasis is compromised and the development of exertional heat stroke most likely
ensues.

INFORMATION BOX
What is already known on this topic:
Self-paced exercise performance and fixed-intensity exercise time-to-exhaustion
are attenuated during exercise in the heat. Given exercise modality differences
(i.e. self-paced vs. fixed-intensity), seemingly opposing theories - anticipatory
regulation and avoidance (self-paced) and the critical core temperature (fixed-
intensity) - have been developed to explain the decreased exercise
performance/duration.

What this study shows:

This editorial proposes that the theories of anticipatory regulation and avoidance
and the critical core temperature are ‘safety’ mechanisms synergistically
maintaining thermoregulation, and therefore homeostasis, during ‘real world’
exercise in the heat. These thermoregulatory mechanisms are suggested to
prevent the onset of exertional heat stroke.

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Exercise and Heat Stress: Performance,

Fatigue, and Exhaustion - A HOT Topic
Zachary J Schlader, Stephen R Stannard and Toby Mundel

Br J Sports Med published online October 20, 2009

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http://bjsm.bmj.com/content/early/2009/10/20/bjsm.2009.063024

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