Received: 21 May 2020 Accepted: 30 June 2020

DOI: 10.1113/EP088783

R E S E A R C H PA P E R

Effect of aerobic fitness on the relation between age and

whole-body heat exchange during exercise-heat stress: a
retrospective analysis
Sean R. Notley Robert D. Meade Glen P. Kenny

Human and Environmental Physiology

Research Unit, School of Human Kinetics,
University of Ottawa, Ottawa, Canada
Correspondence
Glen P. Kenny, University of Ottawa, School of
Human Kinetics, 125 University, Room 367,
Montpetit Hall, Ottawa, Ontario, Canada, K1N
6N5
Email: gkenny@uottawa.ca
Abstract
Ageing is associated with decrements in cutaneous vasodilatation and sweating
that attenuate whole-body total heat loss (evaporative + dry heat exchange) during
exercise-heat stress. However, it remains uncertain whether increased aerobic fitness,
as indexed by peak oxygen uptake (V̇ O peak ), slows that age-related decline. To evaluate
2
this possibility, we conducted a retrospective analysis of data from aerobically fit
(n = 38; V̇ O peak : (mean (SD)) 49 (4) ml kg−1 min−1 ) and less fit (n = 35; V̇ O peak : 32 (3)
2 2

Edited by: Michael White ml kg−1 min−1 ) adults spanning a broad age range (18–65 vs. 18–66 years). Participants
performed three, 30 min bouts of cycling at metabolic heat productions of 150, 200
Funding information
Canadian Institutes of Health Research, and 250 W m−2 , each separated by 15 min recovery, in dry heat (40˚C, ∼15% relative
Grant/Award Number: 399434
humidity). Metabolic heat production and whole-body total heat loss were measured
using indirect and direct calorimetry, respectively. Total heat loss (mean (95% CI))
declined at a rate of 5 (2, 8), 6 (3, 8) and 5 (3, 10) W m−2 per decade during exercise at
metabolic heat productions of 150, 200 and 250 W m−2 , respectively, in less aerobically
fit individuals (all P ≤ 0.002), due primarily to reductions in evaporative heat loss. In
contrast, no significant associations between age and total heat loss were observed in
aerobically fit individuals (all P ≥ 0.146). As such, the slope of the age-related reduction
in total heat loss was steeper in less fit compared to fit individuals across all three
exercise bouts (all P ≤ 0.029). These outcomes indicate that increased aerobic fitness
attenuates the age-related decline in exercise thermoregulation.

KEYWORDS
ageing, aerobic fitness, calorimetry, heat exchange, sweating

1 INTRODUCTION
Older adults are among the most vulnerable to heat-related illnesses
and other adverse effects during heat exposure and physical activity
in the heat (Kenny, Yardley, Brown, Sigal, & Jay, 2010; Petitti, Harlan,
Chowell-Puente, & Ruddell, 2013). This is due, at least in part, to
age-related decrements in cutaneous vasodilatation and sweating
(Balmain, Sabapathy, Louis, & Morris, 2018), which attenuate whole-
body total heat loss (evaporative + dry heat loss) by ∼4% per decade
in both men and women during exercise-heat stress (D’Souza, Notley,
& Kenny, 2020; McGinn et al., 2017). However, while our under-
standing of these age effects has grown substantially in recent years
(Meade, Notley, & Kenny, 2019), the factors that modulate the rate at
which thermoregulatory function declines with increasing age remain
underexplored.
Aerobic fitness, as indexed by peak oxygen uptake (V̇ O2 peak ), also
declines progressively with increasing age (Hawkins & Wiswell, 2003).
Further, aerobic fitness is an important determinant of individual

© 2020 The Authors. Experimental Physiology © 2020 The Physiological Society

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NOTLEY ET AL .
variation in heat loss (Havenith & van Middendorp, 1990; Notley,
Lamarche, Meade, Flouris, & Kenny, 2019b), with recent studies
demonstrating that adults with a V̇ O peak 10–20 ml kg−1 min−1 higher
2
than their less fit counterparts display thermoregulatory adaptations
that augment heat loss during moderate-to-vigorous exercise in the
heat (Lamarche, Notley, Louie, Poirier, & Kenny, 2018a; Lamarche,
Notley, Poirier, & Kenny, 2018b; Stapleton et al., 2015a). It is possible,
therefore, that the rate of thermoregulatory decline with increasing
age may be slower in more aerobically fit individuals. However,
while several investigators have provided support for this hypothesis
by demonstrating that age-related thermoregulatory impairments
disappear when assessed in groups matched for V̇ O peak (Best, Caillaud,
2 66 years, a steeper decline in whole-body total
& Thompson, 2012; Davies, 1979; Drinkwater & Horvath, 1979;
Pandolf et al., 1988; Stapleton et al., 2015a; Tankersley, Smolander,
Kenney, & Fortney, 1991), others have shown those impairments
persist between young and older adults with a similar V̇ O peak (Larose
2
et al., 2013; Notley et al., 2017; Stapleton et al., 2015b). Additionally,
since those studies compared groups of young and older adults, rather
than individuals spanning a broad age range, it was not possible to
quantify the relation between increasing age and thermoregulatory
function. Thus, the effect of aerobic fitness on the age-related decline
in whole-body heat loss during exercise-heat stress remains unclear.
The purpose of this work was therefore to assess the interplay
between the effects of age and aerobic fitness on thermoregulatory
function during exercise-heat stress. To achieve this, we conducted a
retrospective analysis of data from aerobically fit (n = 38; V̇ O peak :2
∼50 ml kg−1 min−1 ) and less aerobically fit adults (n = 35; V̇ O2 peak :
∼30 ml kg−1 min−1 ) spanning a broad age range (18–66 years) from
our previous work (D’Souza et al., 2020; Notley et al., 2019b; Notley
et al., 2019c; Stapleton et al., 2015a). Participants performed exercise
at increasing, fixed rates of metabolic heat production of 150, 200,
and 250 W m−2 in dry heat (40◦ C, ∼15% relative humidity), while
whole-body total heat loss (evaporative + dry heat loss) was measured
continuously using direct calorimetry. We hypothesized that whole-
body total heat loss would decline with increasing age in both groups,
albeit at a lesser rate in aerobically fit individuals.
2 METHODS
2.1 Ethical approval
This study was approved by the University of Ottawa Health Sciences
and Science Research Ethics Board (no. H10-04-04B) and agrees with
the latest version of the Declaration of Helsinki, except for registration
in a database. Written and informed consent was obtained from all
volunteers prior to their participation.
2.2 Participants
Data from seventy-three adults (Table 1) spanning a broad age range
were extracted from our previous work (D’Souza et al., 2020; Notley
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1551
New Findings
∙ What is the central question of this study?
Does higher aerobic fitness, indexed by peak oxygen
uptake (V̇ O peak ), attenuate the age-related decline
2
in thermoregulatory function during exercise in the
heat?
∙ What is the main finding and its importance?
When assessed in aerobically fit and less fit adults
(V̇ O peak : ∼30 vs. ∼50 ml kg−1 min−1 ) aged 18–
2
heat loss (evaporative + dry heat exchange) was
observed with increasing age in less aerobically
fit adults. These outcomes indicate that increased
aerobic fitness may attenuate the age-related
decline in thermoregulatory function during
exercise-heat stress.
et al., 2019b, 2019c; Stapleton et al., 2015a) based on their area-
specific metabolic heat production during exercise (150 ± 10, 200 ± 10
and 250 ± 10 W m−2 ) and mass-specific V̇ O peak (ml kg−1 min−1 ), and
2
analysed retrospectively. Participants were assigned to the aerobically
fit group if they possessed a V̇ O peak of ≥45 ml kg−1 min−1 (n = 38;
2
16 women, 22 men), while participants were included in the less
aerobically fit group if their V̇ O2 peak was ≤35 ml kg−1 min−1 (n = 35;
20 women, 15 men). These inclusion criteria were determined prior
to analysis and were chosen to ensure a minimum between-group
separation in V̇ O2 peak of 10 ml kg−1 min−1 given that fitness-related
differences in heat loss have been shown to occur between groups with
an average ∼10–20 ml kg−1 min−1 difference in V̇ O peak (Lamarche
2
et al., 2018a, 2018b; Stapleton et al., 2015a). Participants were non-
smokers, and did not report a history of cardiovascular, respiratory,
or metabolic disease, and were not taking prescription medication.
The only exceptions were three women in the less fit group, who were
taking oral contraceptives (n = 1) or undergoing hormone replacement
therapy (n = 2). However, since excluding these individuals did not
modify our preliminary analysis, they were retained in the final sample.
2.3 Procedures
2.3.1 Overview
Participants completed one preliminary session and one experimental
trial separated by >48 h. Prior to each session, participants were asked
to refrain from exercise, alcohol or caffeine and non-steroidal anti-
inflammatory drugs for >24 h and were instructed to arrive at the
laboratory well hydrated.
 1469445x, 2020, 9, Downloaded from https://physoc.onlinelibrary.wiley.com/doi/10.1113/EP088783 by Nat Prov Indonesia, Wiley Online Library on [25/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
1552 NOTLEY ET AL .

TA B L E 1 Physical characteristics of the aerobically fit (n = 38) and less fit groups (n = 35)

Fit Less fit

Characteristic Mean (SD) Min–max Mean (SD) Min–max P
Age (years) 38 (15) 18–65 42 (17) 18–66 0.271
Body mass (kg) 70.5 (11.6) 50.3–96.0 74.1 (11.6) 54.0–104.3 0.233
Height (cm) 173 (8) 158–190 171 (10) 152–198 0.316
AD (m2 ) 1.9 (0.2) 1.5–2.2 1.8 (0.2) 1.5–2.2 0.675
Body fat (%) 17 (5) 10–25 27 (8) 9–46 <0.001
V̇ O2 peak (ml kg−1 min−1 ) 49 (4) 45–62 32 (3) 26–35 <0.001
−1
Physical activity (min week ) 417 (162) 135–840 123 (80) 0–240 <0.001

P values denote the results from an unpaired t test comparing aerobically fit to less fit individuals. AD , body surface area (DuBois & DuBois, 1915); V̇ O2 peak :
peak oxygen uptake.

2.3.2 Preliminary session
Physical activity, body height, mass, surface area and fat percentage, as
well as V̇ O peak were determined during the preliminary visit. Physical
2
activity was assessed using a 3-month activity recall questionnaire
(Kohl, Blair, Paffenbarger, Macera, & Kronenfeld, 1988), whereby
participants were asked to document their average weekly volume
of aerobic activity (e.g. running, cycling, swimming). Body surface
area was derived from measures of standing height (model 2391,
Detecto, Webb City, MO, USA) and body mass (IND560, Mettler
Toledo Inc., Mississauga, ON, Canada) (DuBois & DuBois, 1915).
Hydrostatic weighing was used to estimate body fat percentage
(Siri, 1956). Indirect calorimetry was used to quantify V̇ O peak (MCD
2
Medgraphics Ultima Series, MGC Diagnostics, St Paul, MN, USA) during
an incremental exercise protocol (1 min stages) to volitional exhaustion
(CSEP, 1986). Exercise was performed on a semi-recumbent cycle
ergometer (Corival, Lode B.V., Groningen, Netherlands) in non-heat
stress conditions (∼23◦ C), with V̇ O2 peak being taken as the highest
oxygen uptake measured over the final 30 s of the protocol.
2.3.3 Experimental trial
Upon arrival to the laboratory, participants changed into shorts,
sleeveless sports top (women only) and sandals, and provided a urine
sample for the measure of urine specific gravity (Reichert TS 400 total
solids refractometer, Reichert, Depew, NY, USA). After confirming
euhydration (urine specific gravity <1.025) (Kenefick & Cheuvront,
2012), participants were instrumented in a temperate room (∼25 ◦ C),
before entering the modified Snellen direct air calorimeter (40◦ C,
∼15% relative humidity). Participants then completed 15 min seated
rest (baseline), followed by three, 30 min bouts of semi-recumbent
cycling at increasing, fixed rates of metabolic heat production, each
separated by 15 min recovery. During each exercise bout, external
work was controlled to elicit metabolic heat productions of 150, 200
and 250 W m−2 (equivalent to external work rates of ∼40, 70 and
95 W, respectively). Given that the effects of both ageing and aerobic
fitness become more pronounced with increases in metabolic heat
production and the subsequent requirement for heat loss to attain
heat balance (Lamarche et al., 2018a; Lamarche et al., 2018b; Stapleton
et al., 2015a), this protocol made it possible to assess their relation at
increasing requirements for heat loss. No food or fluid consumption
was permitted during the experimental protocol.
2.4 Measurements
During each experimental trial, the modified Snellen direct air
calorimeter was used to measure time-dependent changes in whole-
body evaporative and dry heat exchange (Reardon et al., 2006).
Absolute humidity was measured using dew point hygrometry (RH
Systems model 373H, Albuquerque, NM, USA), while air temperature
was measured using resistance temperature detectors (Black Stack
model 1560, Fluke Corporation, Everett, WA, USA). Evaporative heat
loss was calculated using the calorimeter outflow – inflow difference
in absolute humidity, multiplied by the air mass flow (kg s−1 ) through
the system (equivalent to ∼0.3 m s−1 where the participant is seated)
(Snellen, Chang, & Smith, 1983) and the latent heat of vaporization of
sweat (2426 J g−1 ). For readers interested in the associated whole-
body sweat rate (g m−2 s−1 ), those data can be calculated by dividing
evaporative heat loss (W m−2 ) by the latent heat of vaporization of
sweat (2426 J g−1 ). Dry heat loss was calculated using the calorimeter
outflow – inflow difference in air temperature, multiplied by the air
mass flow and specific heat capacity (1005 J kg−1 ◦ C−1 ). Since air
temperature was above skin temperature (i.e. permitting only dry heat
gain), dry heat exchange is referred to hereafter as dry heat gain.
Oxygen consumption and carbon dioxide production were
measured using indirect calorimetry (Moxus modular metabolic
system, AEI Technologies, Bastrop, TX, USA), and used to approximate
metabolic rate (Kenny & Jay, 2013). To account for respiratory heat
exchange, expired air was recycled back into the calorimeter. External
work rate was controlled to maintain metabolic heat production
(metabolic rate – external work rate) during light, moderate and
vigorous exercise at (mean (SD)) 150 (7), 200 (8) and 250 (8) W m−2

NOTLEY ET AL .
for the fit group, and 151 (6), 200 (8) and 251 (8) W m−2 for the less
fit group. Body heat storage was calculated as the difference between
heat production and whole-body total heat loss (evaporative heat loss
– dry heat gain).
Heart rate was recorded at 1 s intervals using either a Polar
M400 or RS400 monitor (Polar Electro Oy, Kempele, Finland).
Body core temperature was measured within the lower gut using
a capsule thermometer (Vital Sense; Mini Mitter, Bend, OR, USA)
ingested ∼1 h prior to data collection (n = 3), or using a thermo-
couple probe (Mon-a-therm General Purpose Temperature Probe,
Mallinckrodt Medical, St Louis, MO, USA) inserted 40 cm past the
nostril entrance (oesophageal temperature; n = 25) or ∼12 cm past
the anal sphincter (rectal temperature; n = 45). These differences
occurred due to the selection of different indices of core temperature
within the previous experiments comprising this analysis (D’Souza
et al., 2020; Notley et al., 2019b; Notley et al., 2019c; Stapleton
et al., 2015a). While the temporal profile of rectal and ingestible
capsule temperature are similar, oesophageal temperature displays a
lower absolute temperature and faster response to thermal transients
(Taylor, Tipton, & Kenny, 2014). We therefore performed our analysis
of core temperature using only rectal (fit: n = 24; less fit: n = 21)
and ingestible capsule temperature (fit: n = 1; less fit: n = 2). Both
heart rate and core temperature were also expressed as a change
from pre-exercise rest (baseline). Skin temperature was measured at
the bicep, chest, thigh, and calf using T-type thermocouples (Concept
Engineering, Old Saybrook, CT, USA), and used to approximate mean
skin temperature (Ramanathan, 1964).
2.5 Data and statistical analysis
All data were expressed as minute averages, with an average of the
final 5 min of baseline and each exercise period used for statistical
analyses. Physical characteristics as well as baseline and end-exercise
responses were compared between-groups using unpaired, two-tailed
Student’s t test. Pearson’s correlation coefficient and least-squares
linear regression were used to assess the relation between age
and all end-exercise calorimetric responses within each exercise
bout for each group. For variables that demonstrated a significant
relation with age, analysis of covariance (ANCOVA) with factors of
group (aerobically fit and less fit), age (covariate), and the interaction
between group and age was then employed. In instances where a
significant group-by-age interaction was detected, this indicated
the slope of the linear regression relationship between age and
the dependent variable of interest (e.g. total heat loss) differed
between the aerobically fit and less fit groups (Zar, 1999). A power
analysis conducted prior to data analysis indicated that to detect a
between-group difference in the slopes of the regression lines of a
large effect size (F value: 0.40), with at least 80% statistical power, ≥52
individuals were required (Faul, Erdfelder, Lang, & Buchner, 2007).
Thus, with the current sample (n = 73), this analysis was adequately
powered (>80%). Test assumptions of normality, linearity and homo-
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1553
scedasticity were determined by inspecting quantile-comparison,
scatter and residual plots. Data were screened for influential
cases using Cook’s distances, leverage values and Studentized
residuals (no cases were removed). α was set at 0.05 for all statistical
comparisons, with data being reported as mean (SD) unless stated
otherwise as mean (95% CI). All analyses were performed using SPSS
24.0 (IBM Corp., Armonk, NY, USA).
3 RESULTS
Participant characteristics are presented in Table 1. Compared to the
less fit group, the aerobically fit group possessed a higher V̇ O peak
2
and volume of physical activity, but lower body fat percentage (all
P < 0.001). However, age, height, body mass and body surface area did
not differ significantly between groups (all P ≥ 0.233).
Average responses at baseline and during each exercise bout are
presented in Table 2, with key outcomes summarized here. Dry heat
gain was higher in the fit compared to less fit group during exercise
eliciting a metabolic heat production of 250 W m−2 (P = 0.020). Across
all exercise bouts, evaporative and total heat loss were higher in the
fit compared to less fit group (all P ≤ 0.027). Consequently, body heat
storage was higher in the less fit relative to the fit group during all
exercise bouts (all P ≤ 0.027). However, while mean skin temperature
was higher in the less fit relative to fit group during exercise eliciting
a metabolic heat production of 250 W m−2 (P = 0.008), neither
core temperature nor its change from baseline differed significantly
between groups (all P ≥ 0.102). Heart rate was higher in the less fit
compared to fit group at baseline and during each exercise bout (all
P ≤ 0.001).
Results from the linear regression analysis are provided in Table 3,
with the results for dry heat gain, evaporative heat loss and total heat
loss highlighted in Figure 1. In the less fit group, dry heat gain was not
significantly related to age (all P ≥ 0.209), whereas it increased with age
during exercise eliciting a heat production of 250 W m−2 in the fit group
(r = 0.469; P = 0.003). However, the slope of the relation between age
and dry heat gain during exercise at a heat production of 250 W m−2
did not differ significantly between groups (P = 0.095). Evaporative and
total heat loss were negatively associated with age during all exercise
bouts in the less fit group (r = −0.358 to −0.588; all P ≤ 0.035).
However, except for exercise at a heat production of 150 W m−2 , where
evaporative was positively associated with age (r = 0.325; P = 0.046),
evaporative and total heat loss were not significantly related to age
in the aerobically fit group (all P ≥ 0.121). As such, the age-related
decline in both evaporative and total heat loss was steeper in the less
fit compared to the fit group during all exercise bouts (all P ≤ 0.029).
Further, body heat storage was positively associated with age in the
less fit group during exercise eliciting a heat production of 250 W m−2
(r = 0.482 to 0.634; all P ≤ 0.003), but not significantly related to age
in the fit group (P ≥ 0.165). Consequently, the slope of that age-related
increase was steeper in the less fit compared to the fit group across all
exercise bouts (all P ≤ 0.015).
 1469445x, 2020, 9, Downloaded from https://physoc.onlinelibrary.wiley.com/doi/10.1113/EP088783 by Nat Prov Indonesia, Wiley Online Library on [25/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
1554 NOTLEY ET AL .

TA B L E 2 Whole-body heat exchange, thermometry and heart rate in the aerobically fit (n = 38) and less fit (n = 35) groups at baseline and
exercise eliciting metabolic heat productions of 150, 200 and 250 W m−2 in dry heat

Fit Less fit Fit vs. less fit

Mean (SD) Mean (SD) Diff. (95% CI) P
−2
EHL (W m ) Baseline 68 (22) 65 (19) 4 (−6, 13) 0.467
150 W m−2 179 (23) 168 (16) 11 (2, 21) 0.017
−2
200 W m 233 (22) 208 (18) 24 (15, 34) <0.001
−2
250 W m 273 (26) 238 (22) 36 (25, 47) <0.001
DHG (W m−2 ) Baseline 38 (9) 38 (9) 0 (−4, 4) 0.975
−2
150 W m 46 (10) 44 (9) 2 (−3, 6) 0.402
200 W m−2 49 (10) 45 (11) 4 (−1, 9) 0.117
250 W m−2 52 (12) 45 (13) 7 (1, 13) 0.020
−2
THL (W m ) Baseline 31 (22) 27 (18) 3 (−6, 13) 0.455
150 W m−2 134 (18) 123 (15) 9 (1, 18) 0.027
−2
200 W m 184 (17) 162 (15) 20 (12, 28) <0.001
250 W m−2 222 (20) 191 (16) 29 (20, 38) <0.001
S (W m−2 ) 150 W m−2 17 (18) 27 (15) −9 (−17, −1) 0.027
−2
200 W m 16 (17) 38 (15) −21 (−28, −13) <0.001
250 W m−2 29 (20) 58 (15) −28 (−36, −19) <0.001
◦
Tc ( C) Baseline 37.0 (0.3) 37.2 (0.3) −0.2 (−0.4, 0.0) 0.052
−2
150 W m 37.3 (0.4) 37.5 (0.2) −0.2 (−0.4, 0.0) 0.102
200 W m−2 37.7 (0.4) 37.8 (0.3) −0.1 (−0.3, 0.1) 0.296
250 W m−2 38.0 (0.4) 38.2 (0.4) −0.2 (−0.4, 0.1) 0.156
ΔTc (◦ C) 150 W m−2
a
0.3 (0.2) 0.3 (0.2) 0.0 (−0.1, 0.1) 0.757
200 W m−2 0.7 (0.3) 0.6 (0.3) 0.1 (−0.1, 0.2) 0.348
−2
250 W m 1.0 (0.3) 1.0 (0.4) 0.0 (−0.2, 0.2) 0.813
Tsk (◦ C)
a

Baseline 35.5 (0.5) 35.4 (0.4) 0.1 (−0.2, 0.3) 0.561

150 W m−2 35.6 (0.5) 35.8 (0.4) −0.1 (−0.3, 0.1) 0.164
−2
200 W m 35.7 (0.5) 35.9 (0.4) −0.3 (−0.5, 0.0) 0.019
−2
250 W m 35.8 (0.5) 36.2 (0.5) −0.4 (−0.6, −0.2) <0.001
HR (beats min−1 ) Baseline 72 (12) 83 (13) −11 (−17, −5) <0.001
150 W m−2 96 (16) 113 (15) −17 (−24, −9) <0.001
200 W m−2 112 (17) 133 (18) −21 (−29, −13) <0.001
250 W m−2 132 (20) 156 (18) −24 (−33, −14) <0.001
−1 −2
ΔHR (beats min ) 150 W m 24 (8) 29 (7) −3 (−7, 0) 0.054
200 W m−2 40 (10) 50 (10) −10 (−15, −5) <0.001
250 W m−2 60 (14) 67 (30) −13 (−19, −6) <0.001
a
Tc data were recorded in a reduced sample of aerobically fit (n = 25) and less fit individuals (n = 23). P values denote the results from an unpaired t test
comparing aerobically fit to less aerobically fit individuals. DHG, dry heat gain; EHL, evaporative heat loss; HR, heart rate; ΔHR, change in heart rate from
baseline; S, body heat storage; Tc ; core temperature; ΔTc ; change in core temperature from baseline; THL, total heat loss; Tsk , mean skin temperature.

4 DISCUSSION
In this retrospective analysis, we evaluated whole-body heat exchange
in adults spanning a broad age range, but differing markedly in aerobic
fitness, during exercise eliciting metabolic heat productions of 150,
200 and 250 W m−2 in dry heat. Consistent with our hypothesis, the
decline in whole-body evaporative and total heat loss with increasing
age was steeper in less aerobically fit individuals across all exercise
bouts. This was paralleled by a greater age-related increase in body
heat storage in the less fit compared to fit group. Together, these
 TA B L E 3 The relation between age (years) and whole-body heat exchange for the aerobically fit (n = 38) and less fit (n = 35) groups during exercise eliciting metabolic heat productions of 150,
200 and 250 W m−2 in dry heat
NOTLEY ET AL .

Aerobically fit Less aerobically fit Slopes

a a b
Coefficient (95% CI) r R2 P Coefficient (95% CI) r R2 P P
EHL (W m−2 )
150 W m−2 Intercept 161.43 (142.48, 180.37) 0.325 0.106 0.046 183.19 (169.15, 197.24) −0.383 0.147 0.023 0.003
Slope 0.48 (0.01, 0.94) −0.36 (−0.67, −0.05)
−2
200 W m Intercept 219.23 (200.54, 237.92) 0.256 0.065 0.121 229.92 (215.18, 244.66) −0.487 0.237 0.003 0.002
Slope 0.36 (−0.10, 0.82) −0.51 (−0.84, −0.19)
250 W m−2 Intercept 258.02 (235.78, 280.25) 0.245 0.060 0.137 256.56 (237.54, 275.58) −0.358 0.128 0.035 0.013
Slope 0.50 (−0.14, 0.96) −0.45 (−0.87, −0.03)
DHG (W m−2 )
150 W m−2 Intercept 38.29 (29.82, 46.76) 0.300 0.090 0.068 38.84 (30.53, 47.15) 0.217 0.047 0.209 —
Slope 0.19 (−0.01, 0.40) 0.12 (−0.07, 0.30)
200 W m−2 Intercept 42.50 (33.50, 51.49) 0.254 0.064 0.124 42.91 (33.10, 52.72) 0.083 0.017 0.634 —
Slope 0.17 (−0.05, 0.39) 0.05 (−0.17, 0.27)
250 W m−2 Intercept 37.65 (28.04, 47.27) 0.469 0.220 0.003 41.40 (29.55, 53.26) 0.103 0.011 0.555 0.095
Slope 0.37 (0.13, 0.61) 0.08 (−0.19, 0.34)
THL (W m−2 )
150 W m−2 Intercept 123.13 (107.56, 138.71) 0.241 0.058 0.146 144.36 (131.19, 157.52) −0.504 0.254 0.002 0.002
Slope 0.28 (−0.10, 0.66) −0.48 (−0.77, −0.19)
200 W m−2 Intercept 176.73 (161.48, 191.98) 0.167 0.028 0.316 187.01 (174.58, 199.44) −0.588 0.346 <0.001 0.002
Slope 0.19 (−0.19, 0.56) −0.56 (−0.84, −0.29)
250 W m−2 Intercept 220.36 (202.44, 238.29) 0.029 0.001 0.861 215.15 (202.60, 227.71) −0.561 0.315 <0.001 0.029
Slope 0.04 (−0.40, 0.48) −0.53 (−0.81, −0.25)
−2
S (W m )
150 W m−2 Intercept 27.19 (11.53, 42.86) −0.230 0.053 0.165 8.16 (−4.17, 20.50) 0.482 0.232 0.003 0.004
Slope −0.27 (−0.66, 0.12) 0.42 (0.15, 0.70)
200 W m−2 Intercept 24.97 (10.06, 39.88) −0.213 0.045 0.199 18.99 (3.76, 27.87) 0.551 0.303 0.001 0.004
Slope −0.24 (−0.60, 0.13) 0.50 (0.23, 0.76)
−2
250 W m Intercept 30.48 (12.25, 48.71) −0.031 0.001 0.852 32.07 (20.63, 43.52) 0.634 0.403 <0.001 0.015
Slope −0.04 (−0.49, 0.41) 0.59 (0.33, 0.84)
a
P values denote the results of least-squares linear regression analysis on the relation between age and each variable. b P values denote the results from an ANCOVA comparing the slopes of those relations (this
analysis was performed only for variables sharing a significant relation with age). DHG, dry heat gain; EHL, evaporative heat loss; S, body heat storage; THL, total heat loss.
1555

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 1469445x, 2020, 9, Downloaded from https://physoc.onlinelibrary.wiley.com/doi/10.1113/EP088783 by Nat Prov Indonesia, Wiley Online Library on [25/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
1556 NOTLEY ET AL .

M-W (150 W m-2) M-W (200 W m-2) M-W (250 W m-2)

(a) 120 2
Fit (R =0.090; p=0.068) (d) 120 2
Fit (R =0.064; p=0.124) (g) 120 2
Fit (R =0.220; p=0.003)
Less fit (R2=0.047; p=0.209) Less fit (R2=0.017; p=0.634) Less fit (R2=0.011; p=0.555)

Dry heat gain (W m-2) 100 100 100

Dry heat gain (W m-2)

Dry heat gain (W m-2)

80 80 80

60 60 60
Slopes
p=0.095

40 40 40

20 20 20

0 0 0
(b) 380 Fit (R2=0.106; p=0.046)
(e) 380 Fit (R2=0.065; p=0.121)
(h) 380 Fit (R2=0.060; p=0.137)
Less fit (R2=0.146; p=0.023) Less fit (R2=0.237; p=0.003) Less fit (R2=0.128; p=0.015)
Evaporative heat loss (W m-2)

Evaporative heat loss (W m-2)

Evaporative heat loss (W m-2)

340 340 340

300 300 300

260 260 260 Slopes

p=0.013

Slopes
220 220 p=0.002 220

180 Slopes
p=0.023
180 180

140 140 140

100 100 100

(c) 280 Fit (R2=0.058; p=0.146) (f) 280 Fit (R2=0.028; p=0.316) (i) 280 Fit (R 2=0.001; p=0.861)
Less fit (R2=0.254; p=0.002) Less fit (R2=0.346; p<0.001) Less fit (R 2=0.315; p<0.001)

240 240 240

Total heat loss (W m-2)
Total heat loss (W m-2)

Total heat loss (W m-2)

Slopes
200 200 200 p=0.029

Slopes
p=0.002
160 160 160

Slopes
p=0.002
120 120 120

80 80 80
10 20 30 40 50 60 70 10 20 30 40 50 60 70 10 20 30 40 50 60 70
Age (years) Age (years) Age (years)

F I G U R E 1 The linear relation between age and dry heat gain (a, d, g), evaporative heat loss (b, e, h), and total heat loss (evaporative heat loss −
dry heat gain) (c, f, i) in aerobically fit (n = 38) and less fit (n = 35) individuals. Participants performed three, 30 min bouts of cycling eliciting
metabolic heat productions (M-W) of 150, 200 and 250 W m−2 in dry heat (40˚C, ∼15% relative humidity). Data are means obtained over the final
5 min of each exercise bout. The continuous lines represent the least-squares linear regression lines with 95% confidence limits (shaded areas). The
goodness of fit (R2 ) and associated P value for each line are provided in parentheses on the figure, while the regression coefficients are provided in
Table 3. The P values on the main panel represent the results from an analysis of covariance (ANCOVA) comparing the slope of each line between
the fit and less fit groups. Groups were composed of both men and women, with women being indicated by symbols with the dot in the centre

outcomes indicate that increased aerobic fitness is associated with an total heat loss did not differ significantly from zero in fit individuals
attenuated age-related decline in thermoregulatory function during (Table 3). Our findings therefore extend previous reports of similar
exercise-heat stress. exercise thermoregulation among young and older adults matched for
While it is relatively well established that ageing is associated with V̇ O peak (Best et al., 2012; Davies, 1979; Drinkwater & Horvath, 1979;
2

decrements in whole-body total heat loss during exercise-heat stress Pandolf et al., 1988; Stapleton et al., 2015a; Tankersley et al., 1991)
(D’Souza et al., 2020; McGinn et al., 2017), the factors modulating by demonstrating that increased aerobic fitness attenuates the age-
the rate of that decline remain poorly understood. Given that aerobic related decline in whole-body heat loss during exercise-heat stress.
fitness also declines with ageing (Hawkins & Wiswell, 2003), and since The observed decline in total heat loss with increasing age in the
those decrements independently attenuate total heat loss (Lamarche less fit group occurred primarily due to reductions in evaporative heat
et al., 2018a; Lamarche et al., 2018b; Stapleton et al., 2015a), we anti- loss (Table 3), which are thought to stem from age-related alterations
cipated that aerobically fit individuals may demonstrate a lesser age- in the control of sweating (Shibasaki, Okazaki, & Inoue, 2013). Given
related decline in whole-body total heat loss. Our analyses support that no significant age-related impairments were observed in the fit
this hypothesis, with total heat loss declining at a rate of 5 (2, 8), 6 group (Table 3), it is likely that those decrements were offset by the
(3, 8) and 5 (3, 10) W m−2 per decade during exercise at metabolic improvements in sweating associated with increased aerobic fitness
heat productions of 150, 200 and 250 W m−2 in less fit individuals, (Buono & Sjoholm, 1988; Henane, Flandrois, & Charbonnier, 1977).
respectively. In contrast, the slope of the relation between age and The specific mechanism(s) driving those improvements are not fully

NOTLEY ET AL .
understood. However, it is presumed they represent thermoregulatory
adaptations (akin to heat acclimation) associated with performing
aerobic activity that causes sustained elevations in body temperature
(Buono & Sjoholm, 1988; Gisolfi & Robinson, 1969), rather than a
direct consequence of increases in V̇ O peak (Avellini, Shapiro, Fortney,
2
Wenger, & Pandolf, 1982; Ravanelli, Gagnon, Imbeault, & Jay, 2020;
Stapleton, Gagnon, & Kenny, 2010) or aerobic activity per se (Amano,
Koga, Inoue, Nishiyasu, & Kondo, 2013; Lamarche, Notley, Poirier,
& Kenny, 2019). It follows that individuals who frequently engage
in exercise in hot environments may too develop adaptations that
attenuate the age-related decline in thermoregulatory function, albeit
the specific ‘dose’ required to induce those adaptations requires
further exploration. Nonetheless, these outcomes underscore the
importance of regular physical activity for preserving physiological
function with ageing (Lazarus, Lord, & Harridge, 2019), particularly
given the increasing frequency of extreme heat (Watts et al., 2019).
An interesting, albeit somewhat unexpected, additional outcome
was that dry heat gain (i.e. environment-to-skin heat transfer via
radiation and convection) increased with ageing in the aerobically fit,
but not less fit group, during exercise at a metabolic heat production
of 250 W m−2 (Figure 1g). This was paralleled by a lower mean skin
temperature in the aerobically fit compared to less fit group (Table 2),
indicating that the observed age-related increases in dry heat gain
likely occurred due to a widening of the thermal gradient between
the skin surface and surrounding environment. It is possible that this
was the result of an increased reliance on sweat evaporation for heat
loss (Table 2), which subsequently cooled the skin and increased the
thermal gradient for dry heat gain. However, while we observed a
significant positive association between age and evaporative heat loss
during exercise at a heat production of 150 W m−2 in aerobically fit
individuals, this did not reach statistical significance during exercise
eliciting a heat production of either 200 or 250 W m−2 (Table 3). These
outcomes highlight an important gap in our understanding of fitness-
related differences in the integration of dry and evaporative heat loss
to meet a given total heat loss.
Based on the observed decline in whole-body total heat loss with
increasing age in less aerobically fit individuals (Table 3), it would
seem apparent that less aerobically fit older adults may experience
the greatest heat strain during exercise or work in the heat. However,
while these decrements caused a corresponding increase in body
heat storage with advancing age in less fit individuals (Table 3), core
temperature, as assessed via rectal temperature (n = 45) and ingestible
capsule temperature (n = 3), did not differ significantly between the fit
and less fit groups (Table 2). This discrepancy may be explained by the
fact that the gastrointestinal tract is a heavily insulated structure that
responds relatively slowly to changes in body heat storage during short
bouts of exercise (Taylor et al., 2014). Indeed, while the incremental
exercise model employed allows one to identify how exercise intensity
influences age- and fitness-related impairments in heat exchange, it
is poorly suited to quantifying the extent to which those impairments
may exacerbate heat strain. As such, there is a need to examine
these effects during more prolonged exercise and/or work in the
heat. Given that existing physical activity (Armstrong et al., 2007) and
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1557
occupational (ACGIH, 2008) heat exposure guidance does not consider
age- and fitness-related differences in thermoregulatory function, the
outcomes of such research may be particularly important.
4.1 Considerations
Since this experiment was a retrospective analysis, there are several
important limitations to note. First, previous evaluations of the effect
of aerobic fitness on age differences in exercise thermoregulation are
generally equivocal, with some investigators reporting that differences
in heat loss responses between young and older adults disappear when
participants possess similar V̇ O peak (Best et al., 2012; Davies, 1979;
2
Drinkwater & Horvath, 1979; Pandolf et al., 1988; Stapleton et al.,
2015a; Tankersley et al., 1991), whereas others indicate they persist
(Larose et al., 2013; Notley et al., 2017; Stapleton et al., 2015b). In an
attempt to reconcile those conflicting reports, we therefore employed
a similar approach whereby participants were characterized as either
aerobically fit or less fit based on V̇ O peak , irrespective of age (i.e.
2
≥45 vs. ≤35 ml kg−1 min−1 ). However, the key limitation with this
approach is that, since V̇ O peak declines progressively with increasing
2
age (Hawkins & Wiswell, 2003), the older individuals in those pre-
vious reports and in our work would be considered ‘relatively’ more
aerobically fit than their younger counterparts based on age-specific
normative data. For example, men aged 20 and 60 years who both
possess a V̇ O peak of ∼50 ml kg−1 min−1 would be considered to possess
2
a V̇ O2 peak representative of the 70th and 95th percentiles, respectively,
according to age-specific normative data (Hoffmann et al., 2019).
As a more practical alternative, aerobic fitness could be classified
according to age-specific norms (e.g. individuals with a V̇ O2 peak ≥80th
and ≤20th percentiles based on their age). However, one of the major
challenges with this approach is that older individuals in the less fit
group may be unable to complete an exercise protocol such as the
one we employed, as the exercise intensity would approach or exceed
their V̇ O peak . A potential solution is to lower the exercise intensity,
2
but since the effects of both aerobic fitness and ageing on heat loss
increase as a function of the requirement for heat loss (Lamarche et al.,
2018a; Lamarche et al., 2018b; Stapleton et al., 2015a), one may be
unable to detect either effect. That said, restricting our analysis to only
those individuals from the current sample who possess a V̇ O peak ≥80th
2
percentile based on age- and sex-specific normative data (n = 35) failed
to reveal a significant age-related decline in whole-body heat loss in
any exercise bout (all P ≥ 0.154). Further, while we could not perform
such an analysis for a less fit group (e.g. individuals possessing a V̇ O peak
2
≤20th percentile), we would anticipate that the age-related decline
in heat loss may be steeper than that observed in the current study,
given that those individuals would possess even lower aerobic fitness
than our less fit group. It is likely, therefore, that our findings (Figure 1)
would hold among individuals who would be classified as aerobically
fit or less fit according to age-specific norms, although this suggestion
requires further scrutiny.
Second, by using a cross-sectional rather than a longitudinal design,
we cannot be certain that the age-related decrements observed

1558
accurately represent the changes occurring across the lifespan. For
instance, given that V̇ O peak declines progressively with increasing age
2
(Hawkins & Wiswell, 2003), it is possible that less fit young adults
may experience an even steeper decline in thermoregulatory function
with increasing age than that observed here (Table 3). We therefore
acknowledge the need for larger studies employing a longitudinal
design to better characterize individual changes in thermoregulatory
function across the lifespan and the factors modulating those changes.
That research may be particularly important given the potential
benefits of increased aerobic fitness for preserving thermoregulatory
function with ageing.
Finally, to maximize our sample size, both men and women were
included in the aerobically fit and less fit groups. This raises two
important considerations. First, the pre-menopausal women studied
were eumenorrhoeic, but were not tested within a specific menstrual
cycle phase. However, while fluctuations in sex hormones during the
menstrual cycle have long been thought to influence thermoregulatory
function (Stephenson & Kolka, 1985), more recent data indicate that
menstrual phase does not significantly modulate whole-body heat
exchange under comparable exercise and environmental conditions
(Notley, Dervis, Poirier, & Kenny, 2019a). Thus, we feel it is unlikely that
any effects of menstrual phase would modify our observations. Second,
since women are known to display attenuated whole-body heat loss
relative to men, irrespective of age (D’Souza et al., 2020), one could
argue that the inclusion of more older women in the less fit group may
have artificially inflated the observed age-related reductions in heat
loss (Table 3). However, sex does not appear to modulate the rate of
the age-related decline in thermoregulatory function during exercise-
heat stress (D’Souza et al., 2020). Further, the proportion of men and
women above and below the median age for both the fit and less
fit groups did not differ significantly (χ2 : 3.358; P = 0.340). We are
therefore doubtful that the inclusion of men and women within both
groups would have confounded the relationships observed, although
we acknowledge there is a need to better understand the interrelation
between sex, age and aerobic fitness on exercise thermoregulation.
5 CONCLUSION
Outcomes from this retrospective analysis indicate that when assessed
during exercise in dry heat, whole-body total heat loss (evaporative
+ dry heat exchange) declines at a slower rate with increasing age in
aerobically fit compared to less aerobically fit adults. Although there
is a need for larger confirmatory studies, which would ideally evaluate
exercise thermoregulation among men and women who are classified
as aerobically fit or less fit according to age-specific norms or utilize a
longitudinal design, these outcomes highlight the potential importance
of increased aerobic fitness for preserving exercise thermoregulation
with increasing age.
ACKNOWLEDGEMENTS
We thank all the participants as well as Mr Dallon Lamarche, Mr
Andrew D’Souza, Dr Martin Poirier and Dr Jill Stapleton for their
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NOTLEY ET AL .
assistance with data collection. This research was in part supported
by the Canadian Institutes of Health Research (399434) (all funds
held by G.P.K.). S.R.N. is supported by a Postdoctoral Fellowship from
the Human and Environmental Physiology Research Unit. R.D.M. is
supported by an Ontario Graduate Scholarship. G.P.K. is supported by
a University of Ottawa Research Chair.
COMPETING INTERESTS
No conflict of interest, financial or otherwise, are declared by the
author(s). The results of the study are presented clearly, honestly, and
without fabrication, falsification, or inappropriate data manipulation.
AUTHOR CONTRIBUTIONS
S.R.N. and G.P.K. conceptualized and designed the research; S.R.N.
analysed data, prepared figures and drafted manuscript; and all authors
edited and revised the final version of the manuscript. All authors
have read and approved the final version of this manuscript and
agree to be accountable for all aspects of the work in ensuring that
questions related to the accuracy or integrity of any part of the work
are appropriately investigated and resolved. All persons designated as
authors qualify for authorship, and all those who qualify for authorship
are listed.
DATA AVAILABILITY STATEMENT
The data that support the findings of this study are available from the
corresponding author upon reasonable request.
ORCID
Sean R. Notley https://orcid.org/0000-0002-5065-5000
Glen P. Kenny https://orcid.org/0000-0001-8683-6973
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1469445x, 2020, 9, Downloaded from https://physoc.onlinelibrary.wiley.com/doi/10.1113/EP088783 by Nat Prov Indonesia, Wiley Online Library on [25/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
NOTLEY ET AL .
How to cite this article: Notley SR, Meade RD, Kenny GP.
Effect of aerobic fitness on the relation between age and
whole-body heat exchange during exercise-heat stress: a
retrospective analysis. Experimental Physiology.
2020;105:1550–1560. https://doi.org/10.1113/EP088783