131

ARTICLE
Air velocity influences thermoregulation and endurance
exercise capacity in the heat
Hidenori Otani, Mitsuharu Kaya, Akira Tamaki, Phillip Watson, and Ronald J. Maughan

Abstract: This study examined the effects of variations in air velocity on time to exhaustion and thermoregulatory and
perceptual responses to exercise in a hot environment. Eight male volunteers completed stationary cycle exercise trials at
70% peak oxygen uptake until exhaustion in an environmental chamber maintained at 30 °C and 50% relative humidity. Four air
velocity conditions, 30, 20, 10, and 0 km/h, were tested, and the headwind was directed at the frontal aspect of the subject by
2 industrial fans, with blade diameters of 1 m and 0.5 m, set in series and positioned 3 m from the subject’s chest. Mean ± SD time
to exhaustion was 90 ± 17, 73 ± 16, 58 ± 13, and 41 ± 10 min in 30-, 20-, 10-, and 0-km/h trials, respectively, and was different between
all trials (P < 0.05). There were progressive elevations in the rate of core temperature rise, mean skin temperature, and perceived
thermal sensation as airflow decreases (P < 0.05). Core temperature, heart rate, cutaneous vascular conductance, and perceived
exertion were higher and evaporative heat loss was lower without airflow than at any given airflow (P < 0.05). Dry heat loss and
plasma volume were similar between trials (P > 0.05). The present study demonstrated a progressive reduction in time to
exhaustion as air velocity decreases. This response is associated with a faster rate of core temperature rise and a higher skin
temperature and perceived thermal stress with decreasing airflow. Moreover, airflow greater than 10 km/h (2.8 m/s) might
contribute to enhancing endurance exercise capacity and reducing thermoregulatory, cardiovascular, and perceptual strain
during exercise in a hot environment.

Key words: airflow, core temperature, fatigue, heat stress, skin temperature.

Résumé : Cette étude scrute les effets de la variation de la vélocité de l’air sur la durée d’épuisement et les réponses thermoré-
gulatrices et perceptuelles à l’effort dans un environnement chaud. Huit hommes se portent volontaires pour des essais sur un
cycloergomètre à 70 % de la consommation d’oxygène de pointe jusqu’à épuisement, et ce, dans une enceinte environnementale
maintenue à 30 °C et à 50 % d’humidité relative. On teste quatre vélocités de l’air : 30, 20, 10 et 0 km/h; on génère un vent de face
au moyen de deux ventilateurs industriels munis d’hélices de 1 m et 0,5 m. Les ventilateurs sont placés en série à 3 m en face de
la poitrine du sujet. La durée d’épuisement (moyenne ± écart-type) est de 90 ± 17, 73 ± 16, 58 ± 13 et 41 ± 10 min lors des essais à
30, 20, 10 et 0 km/h respectivement et diffère d’un essai à l’autre (P < 0,05). On note une augmentation graduelle du taux
d’élévation de la température centrale, de la température moyenne cutanée et de la sensation thermique à mesure que la vélocité
de l’air diminue (P < 0,05). La température centrale, le rythme cardiaque, la conductance vasculaire cutanée et la sensation de
l’intensité de l’effort sont plus élevés et la perte de chaleur par évaporation est plus faible à vélocité nulle qu’à toute autre
vélocité de l’air donnée (P < 0,05). La perte de chaleur sèche et le volume du plasma sont similaires d’un essai à l’autre (P > 0,05).
La présente étude révèle une diminution graduelle de la durée d’épuisement à mesure que la vélocité de l’air diminue. Cette
réponse est associée à une élévation plus rapide de la température centrale et à une augmentation de la température cutanée et
du stress thermique perçu à mesure que la vélocité de l’air diminue. De plus, une vélocité de l’air supérieure à 10 km/h (2,8 m/s)
pourrait contribuer à l’amélioration de la capacité physique d’endurance et à la diminution du stress thermorégulateur,
cardiovasculaire et perçu au cours d’un exercice dans un environnement chaud. [Traduit par la Rédaction]

Mots-clés : vélocité de l’air, température centrale, fatigue, stress dû à la chaleur, température cutanée.

Introduction
Body temperature and endurance performance in exercising
individuals is significantly influenced by environmental condi-
tions, including ambient temperature (Ta), relative humidity (RH),
air velocity, and solar radiation, especially when exercise is com-
bined with exposure to heat stress (Maughan 2010). Core temper-
ature, mean skin temperature (Tsk), and heart rate (HR) increase
during exercise as Ta increases (4 °C, 11 °C, 21 °C, and 31 °C) and
exercise time to exhaustion is reduced progressively with increas-
ing Ta above the optimum (in this case 11 °C) (Galloway and
Maughan 1997). Increasing humidity (24%, 40%, 60%, and 80% RH)
(Maughan et al. 2012) and solar radiation (0, 250, 500 and 800 W/m2)
(Otani et al. 2016) during exercise in a hot environment (both
30 °C) have been shown to progressively elevate Tsk and reduce
exercise time to exhaustion, though neither core temperature nor
HR was substantially altered. Based on these findings, it is clear
that endurance exercise capacity is impaired and skin tempera-
ture is elevated in a hot environment with increasing Ta, RH, and
solar radiation, whereas the responses of core temperature and
HR are different between these environmental conditions.

Received 16 July 2017. Accepted 23 September 2017.

H. Otani. Faculty of Health Care Sciences, Himeji Dokkyo University, 7-2-1 Kamiono, Himeji, Hyogo 670-8524, Japan.
M. Kaya and A. Tamaki. Hyogo University of Health Sciences, 1-3-6 Minatojima, Chuo-ku, Kobe, Hyogo 650-8530, Japan.
P. Watson. Department of Human Physiology, Vrije Universiteit Brussel, Pleinlaan 2, B-1050 Brussels, Belgium.
R.J. Maughan. School of Medicine, University of St Andrews, St Andrews KY16 9AJ, UK.
Corresponding author: Hidenori Otani (email: hotani@himeji-du.ac.jp).
Copyright remains with the author(s) or their institution(s). Permission for reuse (free in most cases) can be obtained from RightsLink.

Appl. Physiol. Nutr. Metab. 43: 131–138 (2018) dx.doi.org/10.1139/apnm-2017-0448 Published at www.nrcresearchpress.com/apnm on 6 October 2017.
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There is only 1 published study that systematically examined
the effects of gradual variations in air velocity on thermoregula-
tory responses during exercise in a hot environment (Saunders
et al. 2005). This study reported elevated core temperature, Tsk,
and HR and decreased exercise tolerance response as wind speed
decreases (0.2, 9.9, 33, and 50 km/h (0.1, 2.8, 9.2 and 13.9 m/s))
during 120 min of exercise in the heat (33 °C, 59% RH), and airflow
greater than 9.9 km/h could significantly attenuate these re-
sponses compared with a 0.2-km/h airflow. However, given that
this study did not employ time-to-exhaustion or time-trial exer-
cise tests to assess endurance exercise performance, the influence
of different air velocities on endurance exercise capacity was ob-
scure. In addition, most athletes and active individuals rarely per-
form exercise under such high wind speeds (50 km/h). A higher
endurance exercise capacity and a lower Tsk in a hot environment
have been demonstrated in subjects exercising with airflow com-
pared with without airflow during a time-to-exhaustion exercise
test with lower core temperature and HR when receiving wind at
4.8 m/s (17.3 km/h) (Morrison et al. 2014) and during a time-trial
exercise test with no effect of wind at 4 m/s (14.4 km/h) on core
temperature and HR (Teunissen et al. 2013). Meanwhile, it is re-
ported that airflow at 32 km/h (8.9 m/s) has been shown to offset
the detrimental effect of 2% hypohydration on an increase in core
temperature and HR and of both 2% and 3% hypohydration on
endurance exercise capacity during a time-trial exercise test in the
heat (Wall et al. 2015). Since these studies (Morrison et al. 2014;
Teunissen et al. 2013; Wall et al. 2015) investigated the influence of
only 1 level of wind speed on thermoregulatory responses to ex-
ercise and endurance exercise capacity, it remains therefore un-
clear to what extent variations in air velocity influence endurance
exercise capacity in a hot environment.
A number of studies have investigated the effects of wind on
thermal responses to exercise in a hot environment without
measuring endurance exercise performance (Adams et al. 1992;
Mora-Rodriguez et al. 2007; Ravanelli et al. 2015; Shaffrath and
Adams 1984). These studies reported that core temperature
(Adams et al. 1992; Ravanelli et al. 2015; Shaffrath and Adams
1984), Tsk (Adams et al. 1992; Mora-Rodriguez et al. 2007; Shaffrath
and Adams 1984), and HR (Adams et al. 1992; Mora-Rodriguez et al.
2007; Ravanelli et al. 2015; Shaffrath and Adams 1984) during ex-
ercise are lowered at any given airflow compared with no airflow.
Moreover, regardless of measuring endurance exercise perfor-
mance, many studies (Adams et al. 1992; Mora-Rodriguez et al.
2007; Morrison et al. 2014; Ravanelli et al. 2015; Saunders et al.
2005; Shaffrath and Adams 1984) demonstrated a lower whole-
body sweat rate with airflow compared with without airflow dur-
ing exercise in a hot environment.
The application of airflow to individuals exercising in a hot
environment has been reported to result in a lower rating of
perceived exertion (RPE) (Morrison et al. 2014; Saunders et al.
2005) and a lower perceived thermal sensation (TS) (Morrison et al.
2014; Teunissen et al. 2013). Given that reduced endurance exer-
cise capacity in the heat is related to a higher RPE (Otani et al.
2017) and TS (Otani et al. 2017; Stevens et al. 2016), a faster airflow
is likely to attenuate the negative impact of both perceptual dis-
turbances on endurance exercise performance. While increased
RPE during exercise in a hot environment has been reported to be
attenuated when air velocity greater than 9.9 km/h (Saunders
et al. 2005), the effects of airflow on TS response and the relation-
ship between these perceptual responses at different air velocities
when exercising in the heat remain unknown.
The aim of the present study was therefore to investigate the
effects of gradual variations in air velocity on exercise time to
exhaustion and thermoregulatory and perceptual responses to
stationary cycle exercise in a hot environment when all other
factors remain constant. In the current study, it was hypothesized
that exercise time to exhaustion in the heat would be reduced as
wind speed decreases, and that decreasing wind speed would have
Appl. Physiol. Nutr. Metab. Vol. 43, 2018
a negative effect on thermoregulatory and perceptual responses.
Moreover, we also hypothesized that airflow greater than 10 km/h
would significantly increase exercise tolerance time and reduce
thermoregulatory and perceptual strain compared with no air-
flow.
Materials and methods
Subjects
Eight healthy, heat-acclimatized males (mean ± SD; age, 21.9 ±
1.1 years; height, 177 ± 5 cm; body mass, 68 ± 8 kg; peak oxygen
uptake (V̇O2max), 49.4 ± 5.8 mL·min−1·kg−1) participated in this
investigation. All volunteers were habitually active and moder-
ately fit, but were not competitive athletes. Before volunteering,
all volunteers received written information regarding the nature
and purpose of this study. Following an opportunity to ask any
questions, a written statement of consent was signed. The proto-
col employed was approved by the local Ethics Advisory Commit-
tee of Hyogo University of Health Sciences (ref.: 16048) and was
conducted in accordance with principles of the Declaration of
Helsinki.
Experimental protocol
All volunteers completed an initial maximal exercise test, a
familiarization trial, and 4 experimental trials each on a separate
day. In an initial maximal exercise test, V̇O2max was determined in
a temperate environment (24–25 °C Ta and 45%–50% RH) using a
discontinuous incremental exercise test to exhaustion on an up-
right cycle ergometer (Monark 828E, Sweden). During this test,
expiratory gases were analyzed using a breath-by-breath gas ana-
lyzer (AE300S, Minato Medical Science Co. Ltd., Osaka, Japan). The
data from this test were used to determine the power output
required to elicit 70% V̇O2max from the oxygen uptake–work rate
relationship. A familiarization trial was undertaken with the air
velocity at 20 km/h to ensure the volunteers were accustomed to
the procedures employed during the investigation and to mini-
mize learning or anxiety effects. This trial was identical to the
experimental trials in all respects (see below for details). Experi-
mental trials were undertaken in an environmental chamber with
Ta at 30 °C, RH at 50%, corresponding to wet-bulb globe tempera-
ture at 26.0 °C, and the air velocity was maintained at either
30 km/h (8.3 m/s), 20 km/h (5.6 m/s), 10 km/h (2.8 m/s), or 0 km/h
(<0.1 m/s). Facing airflow was directed at the frontal aspect of the
subject from 2 industrial fans, using a 1-m blade diameter fan
(BF-100V, Nakatomi Co. Ltd., Takayama-mura, Nagano, Japan) and
a 0.5-m blade diameter fan (SF-50FS-1VP, Suiden Co. Ltd., Sango-
cho, Osaka, Japan), set in series and positioned 3 m from the
subject’s chest. Airflow was calibrated before the trials and mea-
sured at 15-min intervals during exercise using an anemometer
(AM-4214SD, MotherTool Co. Ltd., Ueda, Japan). Experimental tri-
als were completed in a balanced, randomized order and were
separated by at least 7 days to avoid an order effect. No exercise or
alcohol consumption was permitted in the 24 h before each trial.
Volunteers entered the laboratory in the morning after an over-
night fast, other than plain water ingested ad libitum until 90 min
before the start of the trial. Upon arrival volunteers first emptied
their bladder and nude body mass was measured to the nearest
10 g (AD6205B, A&D Co. Ltd., Tokyo, Japan). Volunteers then in-
serted a rectal thermistor (ITP010-11, Nikkiso-Therm Co. Ltd.,
Musashino, Tokyo, Japan) 10 cm beyond the anal sphincter for
measurement of rectal temperature (Tre) to determine core tem-
perature. Surface skin temperature thermistor probes (ITP082-25,
Nikkiso-Therm Co. Ltd., Musashino, Tokyo, Japan) were attached
to 4 sites (chest, upper arm, thigh, and calf) to enable the calcula-
tion of Tsk (Ramanathan 1964). An HR telemetry band (Polar
WearLink + Hybrid transmitter, Polar Electro Oy, Kempele, Fin-
land) was then positioned. A skin blood flow probe (laser-Doppler
flowmetry; LBF-III, Bio Medical Science Co. Ltd., Tokyo, Japan) was
Published by NRC Research Press
Otani et al.
also secured on the medial aspect of right forearm, approximately
5 cm from the radial styloid process, to determine forearm skin
blood flow. Volunteers were dressed in cycling shorts, socks, and
shoes for all trials. Volunteers rested in a seated position for
15 min in a temperate environment (24–25 °C) with 1 hand im-
mersed in warm water (about 40 °C) for 5 min to allow a capillary
blood sample to be drawn at rest, and then a finger-stick blood
sample was taken as described below. During this period resting
Tre, skin temperatures, and HR were recorded using a thermo-
meter (N543R, Nikkiso-Therm Co. Ltd., Musashino, Tokyo, Japan)
and an HR monitor (RCX5, Polar Electro Oy, Kempele, Finland),
respectively. A 60-s measurement of skin blood flow was made
after 10 min of seated rest and blood pressure was measured using
an automated sphygmomanometer (FB-300S, Fukuda Denshi Co.
Ltd., Tokyo, Japan).
Volunteers then entered the environmental chamber and be-
gan cycle exercise at a power output corresponding to 70% V̇O2max;
this exercise intensity has been used in previous studies investi-
gating the effects of environmental conditions on endurance ex-
ercise capacity (Galloway and Maughan 1997; Maughan et al. 2012;
Otani et al. 2016; Watson et al. 2012). Exercise continued to exhaus-
tion, defined as the point at which volunteers were unable to
maintain a pedal cadence of 60 rpm for 60 s despite verbal encour-
agement. Tre, skin temperatures, HR, and skin blood flow were
recorded at 3-min intervals during exercise. Blood pressure, RPE
(Borg 1982), and TS, using a 9-point scale (International Organization
for Standardization (ISO) 1995), were assessed every 6 min. Partici-
pants received no real-time visual feedback concerning exercise
duration and physiological indices during exercise. Mean body
temperature (Tb) was calculated as 0.8 × Tre + 0.2 × Tsk (Hardy and
DuBois 1938). Convective heat loss (W/m2) in the 0 km/h trial was
calculated as 6.0 × (Tsk – Ta) (Adams et al. 1992), where 6.0 is the
convective heat transfer coefficient in still air in m2·°C/W, Tsk is
the average Tsk and Ta is the ambient temperature. Convective
heat loss (W/m2) in the 30, 20, and 10 km/h trials was calculated as
8.3 × v0.5 × (Tsk – Ta) (Nielsen 1996), where v is the air velocity (m/s),
8.3 × v0.5 is the convective heat transfer coefficient in forced air in
m2·°C/W. Radiative heat loss (W/m2) was calculated as 5.2 ×
(Tsk – Tmrt) (Nielsen 1996), where 5.2 is the radiative heat transfer
coefficient in m2·°C/W and Tmrt is the mean radiant temperature.
Since it is well known that Tmrt is close to Ta in an indoor envi-
ronment, the current study substituted Ta for Tmrt (Parsons 2014).
Dry heat loss (W/m2) was calculated as convective heat loss + radi-
ative heat loss. Evaporative heat loss (W/m2) was calculated as
124 × v0.5 × (Psk – Pa) (Nielsen 1996), where 124 × v0.5 is the evapo-
rative heat transfer coefficient in m2·kPa/W, Psk is the average
saturated water vapour pressure at the skin in kPa (assumed to be
the saturated water vapour pressure (Psa) at Tsk, which was cal-
culated using equation below), and Pa is the water vapour pres-
sure in the ambient air in kPa (calculated using equation below).
Psa (mb) was calculated using Antoine’s equation (Parsons 2014):
Psa = exp [18.956 – 4030.18/(Ta + 235)]. Psa in mb was converted
to Psa in kPa by dividing by 10 (i.e., 10 mb = 1 kPa). Pa (kPa) was
calculated as Psa × RH (Parsons 2014), where RH is the relative
humidity in %. Heat loss data are expressed as heat loss/exercise
time in hours. Mean arterial pressure (MAP) was calculated as
(systolic blood pressure – diastolic blood pressure)/3 + diastolic
blood pressure. Cutaneous vascular conductance (CVC) was calcu-
lated as the ratio of skin blood flow (arbitrary units) to MAP, and
normalized to baseline values. A second finger-stick blood sample
was taken immediately after exhaustion. Following exercise, vol-
unteers returned to a comfortable environment and emptied
their bladder, the probes were removed, and nude body mass was
remeasured to allow the estimation of sweat loss as body mass
loss + the volume of water ingested (Maughan et al. 2007). During
experimental trials, volunteers were free to ingest plain water
maintained at about 34–36 °C to avoid a decrease in core temper-
133
ature by consuming a cold fluid. Whole-body sweat rate was esti-
mated as sweat loss/exercise time in hours.
For each blood sample, the volunteer’s finger was cleaned with
alcohol and pricked with a lancet (Blood collection needle appa-
ratus, Sanwa Kagaku Kenkyusyo Co. Ltd. Nagoya, Aichi, Japan),
and then the first 1–2 drops of blood were wiped away before the
blood sample was taken. Ten-microlitre capillary blood samples
were collected in HemoCue (HemoCue Hb 201+, HemoCue AB,
Ängelholm, Sweden) microcuvettes by capillary action for haemo-
globin measurement (ICSH method), and 60-␮L blood samples
were collected in 2 heparinized capillary tubes (VC-H075H, Terumo
Corp., Tokyo, Japan) for haematocrit measurement (microcen-
trifugation). Haemoglobin and haematocrit values were used to
estimate percentage changes in plasma volume relative to the
resting sample (Dill and Costill 1974).
Statistical analysis
Data are presented as means ± SD. An a priori sample-size cal-
culation was performed (G*Power3.1.9.2; Dusseldolf, Germany) us-
ing data from previous investigations, employing the similar
experimental model (Maughan et al. 2012; Otani et al. 2006, 2016,
2017; Watson et al. 2012) in which investigators reported medium
to large effect sizes for exercise time to exhaustion, and this indi-
cated that we would need 8 subjects per group to find statistical
significance with an effect size of 0.6, a power of 0.75, and alpha
set to 0.05. The IBM SPSS (version 21; IBM Corp., Armonk, N.Y.,
USA) was used for all statistical analyses. The significance level
was set at P < 0.05. The normality of the data and the homogeneity
of variance between the trials were tested using Shapiro–Wilk’s
test and Levene’s test, respectively. When Shapiro–Wilk’s test or
Levene’s test was less than a significant level (P < 0.05), data were
analyzed using nonparametric tests. Nonparametric data (TS and
RPE) were analyzed using Friedman’s 2-way ANOVA. When a sig-
nificant difference was found, the pairwise comparisons were
tested using Wilcoxon’s signed-rank test. In all other cases, data
collected once per trial were analyzed using a 1-way ANOVA with
repeated measures, and data collected over time were analyzed
using a 2-way ANOVA with repeated measures (4 trials [air veloc-
ity] × time). When the sphericity assumption was violated, the
Greenhouse–Geisser correction was employed. Pairwise differ-
ences between trials were evaluated using 1-way ANOVAs with a
Bonferroni adjustment applied for multiple comparisons. Effect
size for nonparametric paired samples were calculated as Pearson’s r
(r) using the average of the cross-products of z scores; an r of 0.1
to <0.3 and ≥0.3 to <0.5 has been suggested to represent a small
and medium treatment effect, respectively, while an r ≥ 0.5 rep-
resents a large treatment effect (Cohen 1988). Cohen’s d (d) was
used as a measure of effect size for parametric paired samples; a d
of 0.2 to <0.5 and ≥0.5 to <0.8 has been suggested to represent a
small and medium treatment effect, respectively, while a d ≥0.8
represents a large treatment effect (Cohen 1988). The 95% confi-
dence intervals for effect sizes (CI) were calculated using the for-
mulas described by Nakagawa and Cuthill (2007).
Results
Exercise time to exhaustion
Exercise time to exhaustion was 90.3 ± 16.5, 72.6 ± 15.7, 58.3 ±
12.6, and 41.4 ± 10.1 min in the 30, 20, 10, and 0 km/h trials,
respectively, and was different (P < 0.0001) between all trials
(30 km/h vs. 20 km/h: P < 0.05, d = 1.03 (CI: –0.02–2.08); 30 km/h vs.
10 km/h: P < 0.01, d = 2.04 (CI: 0.80–3.28); 30 km/h vs. 0 km/h:
P < 0.0001, d = 3.35 (CI: 1.77–4.93); 20 km/h vs. 10 km/h: P < 0.01, d =
0.95 (CI: –0.09–1.99); 20 km/h vs. 0 km/h: P < 0.0001, d = 2.22 (CI:
0.94–3.50); 10 km/h vs. 0 km/h: P < 0.01, d = 1.38 (CI: 0.27–2.49)
(Fig. 1)).
Published by NRC Research Press
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Fig. 1. Effect of air velocity on exercise time to exhaustion (ETE).
Columns and error bars are means ± SD, lines are individual
responses of the 8 participants. *, P < 0.05–0.0001 denote a
significant difference between all trials where significance levels
are described in the text.
Thermoregulatory responses
There was a main effect of trial during exercise on Tre (P < 0.01),
which was higher on the 0 km/h trial than on all the other trials
(30 km/h: P < 0.05, d = 0.37 (CI: 0.07–0.67); 20 km/h: P < 0.05, d =
0.47 (CI: 0.17–0.77); 10 km/h: P < 0.05, d = 0.43 (CI: 0.13–0.73)
(Fig. 2a)). Tre at the point of exhaustion was not different between
trials (P = 0.082) and was 38.5 ± 0.5 °C on the 30, 20, and 10 km/h
trials and 38.8 ± 0.5 °C on the 0 km/h trial. The rate of rise in Tre
during exercise was 0.012 ± 0.004 °C/min, 0.016 ± 0.005 °C/min,
0.022 ± 0.007 °C/min, and 0.037 ± 0.014 °C/min on the 30, 20, 10,
and 0 km/h trials, respectively, and was less (P < 0.0001) on the
30 km/h trial than on all the other trials (20 km/h: P < 0.05, d = 0.88
(CI: –0.15–1.91); 10 km/h: P < 0.05, d = 1.76 (CI: 0.58–2.94); 0 km/h:
P < 0.01, d = 2.43 (CI: 1.10–3.76)) and on the 20 km/h trial than on
the 0 km/h trial (P < 0.05, d = 2.00 (CI: 0.77–3.23)). A 2-way interac-
tion during exercise was observed in Tsk (P < 0.0001), which was
higher on the 0 km/h trial than on the 30 km/h trial from 18 min
to the point of exhaustion (all P < 0.05, d = 1.16–1.34 (CI: 0.09–2.44))
and on the 20 and 10 km/h trials at the point of exhaustion (both
P < 0.05, d = 1.56 (CI: 0.42–2.70) and 1.47 (CI: 0.35–2.59) (Fig. 2b)). A
main effect of trial during exercise was shown for Tb (P < 0.001),
which was higher on the 0 km/h trial than on all the other trials
(30 km/h: P < 0.001, d = 0.53 (CI: 0.23–0.83); 20 km/h: P < 0.001, d =
0.57 (CI: 0.27–0.87); 10 km/h: P < 0.05, d = 0.56 (CI: 0.26–0.86)
(Fig. 2c)). There was an interaction during exercise in Tre-to-Tsk gra-
dient (P < 0.0001), which was narrower on the 0 km/h trial than on
the 30 km/h trial from 15 min to the point of exhaustion (all P < 0.05,
d = 1.16–1.34 (CI: 0.09–2.44)) and on the 20 and 10 km/h trials at the
point of exhaustion (both P < 0.05, d = 1.57 (CI: 0.43–2.71) and 1.61 (CI:
0.46–2.76) (Fig. 2d)).
Convective heat loss was lower (P < 0.0001) on the 0 km/h trial
than on all the other trials (30 km/h: P < 0.0001, d = 2.22 (CI:
0.94–3.50); 20 km/h: P < 0.0001, d =2.93 (CI: 1.47–4.39); 10 km/h:
P < 0.001, d = 2.93 (CI: 1.47–4.39) (Fig. 3)). Radiative heat loss was
higher (P < 0.001) on the 0 km/h trial than on the 30 and 20 km/h
trials (30 km/h: P < 0.01, d = 2.37 (CI: 1.05–3.69); 20 km/h: P < 0.05,
d = 2.08 (CI: 0.83–3.33)), on the 10 km/h trial than on the 30 km/h
trial (30 km/h: P < 0.05, d = 2.14 (CI: 0.88–3.40) (Fig. 3)). However,
dry heat loss was not different between trials (P = 0.168; Fig. 3).
Evaporative heat loss was lower (P < 0.0001) on the 0 km/h trial
than on all the other trials (30 km/h: P < 0.0001, d = 4.91 (CI:
2.84–6.98); 20 km/h: P < 0.0001, d = 4.58 (CI: 2.62–6.54); 10 km/h:
P < 0.0001, d = 4.39 (CI: 2.49–6.29) (Fig. 3)).
Appl. Physiol. Nutr. Metab. Vol. 43, 2018
Cardiovascular responses
There was a main effect of trial during exercise on HR (P < 0.01),
which was higher on the 0 km/h trial than on all the other trials
(30 km/h: P < 0.05, d = 0.32 (CI: 0.02–0.62); 20 km/h: P < 0.05, d =
0.44 (CI: 0.14–0.74); 10 km/h: P < 0.01, d = 0.32 (CI: 0.02–0.62)
(Fig. 4a)). Neither an interaction nor a main effect of trial was
apparent during exercise on MAP (interaction: P = 0.244; main
effect: P = 0.081 (Fig. 4b)). A 2-way interaction during exercise was
shown in CVC (P < 0.01), which was higher on the 0 km/h trial than
on the 30 km/h trial at 24 min and at the point of exhaustion (both
P < 0.05, d = 1.19 (CI: 0.12–2.26) and 1.29 (CI: 0.20–2.38)), on the
20 km/h trial from 18 min to the point of exhaustion (all P < 0.05,
d = 1.39–1.56 (CI: 0.28–2.70)), and on the 10 km/h trial at 24 min
(P < 0.05; d = 1.19 [CI: 0.12–2.26]: Fig. 4c).
Fluid balance
Whole-body sweat rate was 0.82 ± 0.16, 0.91 ± 0.15, 0.97 ± 0.18,
and 1.05 ± 0.24 L/h on the 30, 20, 10, and 0 km/h trials, respectively,
and was higher (P < 0.05) on the 0 km/h trial than on the 30 km/h
trial (P < 0.05, d = 1.13 (CI: 0.06–2.20)). The volume of water in-
gested was 434 ± 202, 444 ± 202, 550 ± 380, and 544 ± 271 mL on the
30, 20, 10, and 0 km/h trials, respectively, and was not different
between trials (P = 0.243). Plasma volume fell by 3.3% ± 2.9%, 3.1% ±
2.9%, 3.5% ± 2.5%, and 3.5% ± 2.3% on the 30, 20, 10, and 0 km/h
trials, respectively, and the change was not different between
trials (P = 0.977).
Perception of thermal stress and exertion
A 2-way interaction during exercise was shown in TS from 6 min
to the point of exhaustion (P < 0.0001), which was higher on the
0 km/h trial than on all the other trials (30 km/h: all P < 0.05, all r =
0.56 (CI: 0.12–0.77); 20 km/h: all P < 0.05, r = 0.51–0.57 (CI: 0.06–
0.78); 10 km/h: all P < 0.05, r = 0.51–0.56 (CI: 0.06–0.77)) and on the
20 and 10 km/h trials than on the 30 km/h trial (20 km/h: all
P < 0.05, r = 0.44–0.51 (CI: –0.03–0.75); 10 km/h: all P < 0.05, r =
0.51–0.54 (CI: 0.06–0.76) (Fig. 5a)). There was an interaction during
exercise for RPE (P < 0.0001), which was higher on the 0 km/h trial
than on the 30 km/h trial from 12 to 24 min (all P < 0.05, all r = 0.56
(CI: 0.12–0.77)) and on the 20 and 10 km/h trials from 12 min to the
point of exhaustion (20 km/h: all P < 0.05, r = 0.51–0.54 (CI: 0.06–
0.76); 10 km/h: all P < 0.05, r = 0.51–0.56 (CI: 0.06–0.77) (Fig. 5b)).
Discussion
The results of the present study clearly demonstrate a reduction
in exercise time to exhaustion with decreasing wind speed and
therefore confirmed our experimental hypothesis. A novel find-
ing in the current study is the finding of a linear association
between air velocity and endurance exercise capacity, with de-
creasing airflow reducing exercise time to exhaustion in a hot
environment by approximately 16 min or 5.8 min for each 10 km/h
or 1 m/s decrease in airflow. This early onset of fatigue with de-
creasing airflow is mainly associated with a faster rate of rise in
Tre and an elevation in Tsk and TS as airflow decreases. Mean-
while, although dry heat loss, MAP, and plasma volume changes
were similar between trials, Tre, Tb, HR, CVC, and RPE were low-
ered and convective and evaporative heat losses were elevated at
any given airflow compared with no airflow. Since it is demon-
strated that receiving airflow greater than 9.9 km/h contributes to
significant decreases in Tre, Tsk, whole-body sweat rate, evapora-
tive heat loss, HR, and RPE compared with a 0.2-km/h airflow
during exercise in the heat (Saunders et al. 2005), the results of the
current study presents the novel finding that there are also signif-
icant increases in exercise time to exhaustion and convective heat
loss and decreases in CVC and TS when airflow greater than
10 km/h compared with no airflow. Based on the results of the
present and previous studies (Adams et al. 1992; Mora-Rodriguez
et al. 2007; Morrison et al. 2014; Ravanelli et al. 2015; Saunders
et al. 2005; Shaffrath and Adams 1984; Teunissen et al. 2013; Wall
Published by NRC Research Press
Otani et al. 135

Fig. 2. Rectal temperature (Tre; a), mean skin temperature (Tsk; b), mean body temperature (Tb; c), and the core-to-skin temperature gradient
(Tre-to-Tsk; d) responses to exercise until exhaustion (Exh) in environments with different air velocities. *, P < 0.05; denotes a significant
interaction between the 0 km/h trial and the 30 km/h trial. †, P < 0.05; denotes a significant interaction between the 0 km/h trial and the
20 and 10 km/h trials. ‡, P < 0.05; denotes a significant main effect of trial between the 0 km/h trial and all the other trials. §, P < 0.001;
denotes a significant main effect of trial between the 0 km/h trial and the 30 and 20 km/h trials. ¶, P < 0.05; denotes a significant main effect
of trial between the 0 km/h trial and the 10 km/h trial.

Fig. 3. Effect of air velocity on convective heat loss (C), radiative
heat loss (R), dry heat loss (D), and evaporative heat loss (E).
*, P < 0.05; denotes a significant difference from the 0 km/h trial.
†, P < 0.05; denotes a significant difference from the 10 km/h trial.
et al. 2015), there would be a great potential for attenuating a
decrease in endurance exercise capacity and reducing thermoreg-
ulatory, cardiovascular, and perceptual strain when receiving air-
flow greater than 10 km/h or 2.8 m/s during exercise in a hot
environment.
Environmental limitations to endurance exercise performance
have been demonstrated with increasing Ta (Galloway and
Maughan 1997), RH (Maughan et al. 2012) and solar radiation
(Otani et al. 2016) and limited airflow (Morrison et al. 2014;
Saunders et al. 2005; Teunissen et al. 2013) in a hot environment.
These environmental interventions are accompanied by addi-
tional load placed on the thermoregulatory system (Galloway and
Maughan 1997; Maughan et al. 2012; Morrison et al. 2014; Otani
et al. 2016; Saunders et al. 2005). It is worth noting that Tsk is
positively correlated with all environmental factors and is ele-
vated as Ta (Galloway and Maughan 1997), RH (Maughan et al.
2012), and solar radiation (Otani et al. 2016) increases and air ve-
locity decreases (Saunders et al. 2005). In line with the findings of
Saunders et al. (2005) who investigated the impact of variations in
air velocity on body temperature regulation during exercise in a
hot environment, the present study observed a gradual elevation
in Tsk during exercise with decreasing air velocity (Fig. 2b). Given
the observed relationships between these environmental factors
and Tsk, the current study strongly supports previous notion that
an elevation in skin temperature is an important sign for predic-
tive risk assessment of climatic heat stress and physical stress in
individuals exercising in a hot environment (Cheuvront et al.
2010; Sawka et al. 2012).
Thermoregulatory limitations for causing hyperthermic fatigue
in individuals exercising in a hot environment have been sug-
gested to be both the attainment of high core temperature
(Gonzalez-Alonso et al. 1999; Nielsen et al. 1993) and skin temper-
ature (Sawka et al. 2011, 2012). While it is known that heat stress
fatigue is associated with no single value of core temperature (Ely
et al. 2009), it is recognized that the attainment of high core
temperature during exercise in the heat causes the early onset of
fatigue owing to a direct impact on the central nervous system
rather than to any muscle, cardiovascular or respiratory limita-
tions (Gonzalez-Alonso et al. 1999; Nielsen et al. 1993). This is
proposed to result in a reduced central neural drive to the exer-
cising muscles (Morrison et al. 2004; Thomas et al. 2006), with
exercise-induced hyperthermia shown to contribute to reductions
in voluntary muscle activation during prolonged maximal knee
extension (Nybo and Nielsen 2001) and to power/speed during
time trials (Watson et al. 2005). Hence, a number of studies re-
ported a deteriorate effect of elevated core temperature on endur-

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136
Fig. 4. Changes in heart rate (HR; a), mean arterial pressure (MAP;
b), and cutaneous vascular conductance (CVC; c) in response to
exercise until exhaustion (Exh) in environments with different air
velocities. *, P < 0.05; denotes a significant interaction between the
0 km/h trial and the 20 km/h trial. †, P < 0.05; denotes a significant
interaction between the 0 km/h trial and all the other trials.
‡, P < 0.05; denotes a significant interaction between the 0 km/h
trial and the 30 and 20 km/h trials. §, P < 0.05; denotes a significant
main effect of trial between the 0 km/h trial and the 30 and 20 km/h
trials. ¶, P < 0.01; denotes a significant main effect of trial between
the 0 km/h trial and the 10 km/h trial.
ance exercise capacity in a hot environment, demonstrating that
individuals reach exhaustion at a core temperature of more than
39 °C in athletes (Gonzalez-Alonso et al. 1999; Nielsen et al. 1993)
and 38–39 °C in nonathletes (Maughan et al. 2012; Otani et al.
2006, 2016, 2017; Watson et al. 2012), which is consistent with the
present study demonstrating Tre at the point of exhaustion of
38.5 ± 0.5 °C (range 38.1–39.6 °C) for the 30, 20, and 10 km/h trials
and 38.8 ± 0.5 °C (range 38.3–39.7 °C) for the 0 km/h trial in non-
athletes (Fig. 2a). Moreover, Maughan and colleagues (2012) dem-
onstrated that a progressive reduction in endurance exercise
capacity in a hot environment is accompanied by a faster rate of
Appl. Physiol. Nutr. Metab. Vol. 43, 2018
Fig. 5. Responses of perceived thermal sensation (TS; a) and rating
of perceived exertion (RPE; b) to exercise until exhaustion (Exh) in
environments with different air velocities. *, P < 0.05; denotes a
significant interaction between the 0 km/h trial and all the other
trials. †, P < 0.05; denotes a significant interaction between the
20 and 10 km/h trials and the 30 km/h trial. ‡, P < 0.05; denotes a
significant interaction between the 20 km/h trial and the 30 km/h
trial. §, P < 0.05; denotes a significant interaction between the
0 km/h trial and the 20 and 10 km/h trials.
rise in Tre with decreasing relative humidity. In agreement with
this, the current study observed a faster rate of rise in Tre during
exercise in the heat with decreasing air velocity. Therefore, in the
current study, a slower rate of rise in Tre as air velocity increases
would contribute to a progressive improvement in endurance ex-
ercise capacity in the heat with increasing wind speed.
It is also recognized that the attainment of skin temperature
above 35 °C results in the early onset of fatigue in individuals
exercising in a hot environment (Cheuvront et al. 2010; Otani et al.
2016; Sawka et al. 2012) owing to the narrowed temperature gra-
dient between the body core and the skin, which requires a
greater skin blood flow and thus elevating cardiovascular strain
(Sawka et al. 2011, 2012). This greater cutaneous blood flow during
exercise has been shown to evoke elevated HR for a given stroke
volume (Trinity et al. 2010) and reducing left ventricular filling
(Stöhr et al. 2011), and to lower blood flow to the exercising mus-
cles (Gonzalez-Alonso et al. 2008) and to the brain (Nybo and
Nielsen 2001), which limits oxygen delivery to, and removal of
heat from, these tissues. Given that Tsk increased during exercise
with decreasing airflow and exceeded 35 °C at the point of exhaus-
tion in most of the volunteers during the 0 km/h trial in the
present study, elevated Tsk likely contribute to a reduced ability
to maintain exercise during this trial (Fig. 2b). The narrowed Tre-
to-Tsk gradient elevated both thermoregulatory and cardiovascu-
lar strain, as evidenced by greater CVC and HR in the 0 km/h trial
compared with the airflow trials (Figs. 2d, 4a, and 4c). Meanwhile,
Published by NRC Research Press
Otani et al.
there was a higher whole-body sweat rate in the 0 km/h trial
compared with the 30 km/h trial in the present study that is in
agreement with these previous studies (Adams et al. 1992;
Mora-Rodriguez et al. 2007; Morrison et al. 2014; Ravanelli et al.
2015; Saunders et al. 2005; Shaffrath and Adams 1984) reporting
high sweat rate in no airflow trial rather than receiving airflow
trial during exercise in a hot environment. Moreover, Saunders
et al. (2005) have demonstrated a progressive lowering of the rate
of evaporative heat loss during exercise as air velocity decreases.
This response was well supported by the result of the present
study, with evaporative heat loss lowering by decreasing wind
speed (Fig. 3). These may indicate that increasing airflow enhances
the ability of evaporative heat loss from the body surface to sur-
rounding air. In the present study, therefore, a gradually lower
Tsk and probably enhanced evaporative heat loss as air velocity
increases would contribute to a progressive improvement in en-
durance exercise capacity in the heat with increasing wind speed.
Psychological limitations to endurance exercise capacity have
been known to be accompanied by the attainment of high per-
ceived exertion in temperate (Pageaux et al. 2013) and hot (Otani
et al. 2017) environments. It is also recognized a reduction in
endurance exercise performance in a hot environment resulting
from a higher TS to exercise (Otani et al. 2016, 2017). Also, receiv-
ing airflow during exercise in the heat has been shown to result in
a lower RPE (Morrison et al. 2014; Saunders et al. 2005) and TS
(Morrison et al. 2014; Teunissen et al. 2013). Although Saunders
et al. (2005) reported a progressive increase in RPE as air velocity
decreases, the present study observed that RPE was higher in the
0 km/h trial than at any given airflow, with no measurable differ-
ences between the airflow trials (Fig. 5b). Such discrepancies in
RPE response to exercise in the study of Saunders et al. (2005) and
the current study might be accompanied by a given air velocity
differences; airflow was compared between 0.2, 9.9, 33, and
50 km/h in the Saunders’s study, which was a greater difference
compared with that of this study (0, 10, 20, and 30 km/h). The
present study also demonstrates a progressive lowering of TS dur-
ing exercise with increasing air velocity (Fig. 5a), suggesting that
TS might be more sensitive to changes in air velocity than RPE in
individuals exercising in a hot environment. Stevens et al. (2016)
clearly demonstrated that a higher TS during exercise can reduce
an ability to perform endurance exercise in a hot environment
without an increase in core temperature. Therefore, a progressive
improvement in endurance exercise capacity in a hot environ-
ment as air velocity increases in the present study would be
largely associated with a gradually lower TS with increasing wind
speed.
The present study has important limitations that should be
pointed out. Given that an evaporative cooling to the torso during
exercise in a hot environment has been demonstrated to lower
Tsk and TS (Filingeri et al. 2015), receiving the headwind at the
frontal aspect of the body without shirt was likely to have a cool-
ing effect on the torso and to lower Tsk and TS in the present
study. Therefore, if our volunteers had worn a shirt to cover torso
during exercise, Tsk and TS could possibly be higher than the
results of the current study. However, since the effects of airflow
with or without a covered torso on thermoregulatory and percep-
tual responses during exercise in the heat is unknown, further
study is warranted. Although our volunteers were habitually ac-
tive and moderately fit, they were not athletes. It is known that
endurance training improves the capacity for dissipating heat
during exercise, including a higher sweat rate (Mora-Rodriguez
et al. 2010) and a higher skin blood flow for a given core temper-
ature (Roberts et al. 1977). These adaptations in trained athletes
contribute to tolerate a higher core temperature compared with
untrained individuals (Mora-Rodriguez et al. 2010). Given these
adaptations in trained athletes, it is likely to demonstrate a longer
exercise time to exhaustion accompanied by a higher capacity for
dissipating heat during exercise in well-trained individuals com-
137
pared with our volunteers in this study. The current study was
conducted in the laboratory on a stationary ergometer in a con-
trolled indoor environment to allow environmental conditions to
be kept constant. Although it would be difficult to carry out the
same investigation as the present study in outdoors, it is well
known that overcoming wind resistance, including not only wind
speed but running or cycling speed and wind direction (headwind
or tailwind), would strongly influence physiological responses ob-
served and the resulting exercise performance of individuals ex-
ercising outdoors (Davies 1980). Moreover, air density has been
reported to partly affect body temperature and endurance exer-
cise performance during outdoor exercise in a hot environment
(Racinais et al. 2015). Thus, the direct application of the results of
laboratory-based experiments to outdoor exercise in a hot envi-
ronment may not be valid. It is worth noting that the present
results are directly applicable to the interpretation of environ-
mental studies undertaken in laboratory settings and any situa-
tion where hard exercise is undertaken on stationary exercise
equipment (e.g., gym-based exercise). Further studies are there-
fore required to systematically investigate the beneficial effect of
airflow on physiological and psychological responses and exercise
performance during outdoor as well as indoor exercise in a hot
environment to prevent excessive heat strain and heat-related
illnesses.
In conclusion, the present study clearly demonstrates that ex-
ercise time to exhaustion on a stationary ergometer in a hot en-
vironment is reduced progressively as air velocity decreases. This
response is associated with a faster rate of rise in core tempera-
ture and a higher skin temperature and perceived thermal stress
with decreasing air velocity. The current study also suggests that
airflow greater than 10 km/h (2.8 m/s) is likely to contribute largely
to attenuating a decrease in endurance exercise capacity and re-
ducing thermoregulatory, cardiovascular, and perceptual strain
during exercise in a hot environment. These findings suggest that
airflow and its velocity are key consideration for individuals exer-
cising in a hot environment, to avoid the deleterious effects of
excessive heat strain as well as the potential to develop heat-
related illnesses.
Conflict of interest statement
The authors declare that they have no conflict of interest.
Acknowledgements
The authors thank the volunteers who donated their time and
effort to participate in the present study. The authors also thank
Yuta Fujinami, Ryota Fukuda, Tomoya Matsuda, and Ryo Mizutani
for assistance.
References
Adams, W.C., Mack, G.W., Langhans, G.W., and Nadel, E.R. 1992. Effects of varied
air velocity on sweating and evaporative rates during exercise. J. Appl.
Physiol. (1985), 73(6): 2668–2674. PMID:1490985.
Borg, G.A. 1982. Psychophysical bases of perceived exertion. Med. Sci. Sports
Exerc. 14(5): 377–381. PMID:7154893.
Cheuvront, S.N., Kenefick, R.W., Montain, S.J., and Sawka, M.N. 2010. Mecha-
nisms of aerobic performance impairment with heat stress and dehydration.
J. Appl. Physiol. (1985), 109(6): 1989–1995. doi:10.1152/japplphysiol.00367.
2010.
Cohen, J. 1988. Statistical power analysis for the behavioral sciences. 2nd ed.
Lawrence Erlbaum Associates, Hillsdale.
Davies, C.T. 1980. Effects of wind assistance and resistance on the forward mo-
tion of a runner. J. Appl. Physiol. Respir. Environ. Exerc. Physiol. 48(4): 702–
709. PMID:7380693.
Dill, D.B., and Costill, D.L. 1974. Calculation of percentage changes in volumes of
blood, plasma, and red cells in dehydration. J. Appl. Physiol. 37(2): 247–248.
PMID:4850854.
Ely, B.R., Ely, M.R., Cheuvront, S.N., Kenefick, R.W., Degroot, D.W., and Montain, S.J.
2009. Evidence against a 40°C core temperature threshold for fatigue in
humans. J. Appl. Physiol. (1985), 107(5): 1519–1525. doi:10.1152/japplphysiol.
00577.2009.
Filingeri, D., Fournet, D., Hodder, S., and Havenith, G. 2015. Mild evaporative
cooling applied to the torso provides thermoregulatory benefits during run-
Published by NRC Research Press
138
ning in the heat. Scand. J Med. Sci. Sports, 25(Suppl. 1): 200–210. doi:10.1111/
sms.12322. PMID:25943671.
Galloway, S.D.R., and Maughan, R.J. 1997. Effects of ambient temperature on the
capacity to perform prolonged cycle exercise in man. Med. Sci. Sports Exerc.
29(9): 1240–1249. doi:10.1097/00005768-199709000-00018. PMID:9309637.
Gonzalez-Alonso, J., Teller, C., Andersen, S.L., Jensen, F.B., Hyldig, T., and
Nielsen, B. 1999. Influence of body temperature on the development of fa-
tigue during prolonged exercise in the heat. J. Appl. Physiol. (1985), 86(3):
1032-1039. PMID:10066720.
Gonzalez-Alonso, J., Crandall, C.G., and Johnson, J.M. 2008. The cardiovascular
challenge of exercising in the heat. J. Physiol. 586(1): 45–53. doi:10.1113/jphysiol.
2007.142158. PMID:17855754.
Hardy, J.D., and DuBois, E.F. 1938. Basal metabolism radiation, convection, and
vaporization at temperature of 22–35°C. J. Nutr. 15: 477–497.
ISO. 1995. Ergonomics of the thermal environment - Assessment of the influence
of the thermal environment using subjective judgement scales. ISO, ISO
10551, Geneva, Switzerland.
Maughan, R.J. 2010. Distance running in hot environments: a thermal challenge
to the elite runner. Scand. J. Med. Sci. Sports, 20(Suppl. 3): 95–102. doi:10.1111/
j.1600-0838.2010.01214.x. PMID:21029196.
Maughan, R.J., Shirreffs, S.M., and Leiper, J.B. 2007. Errors in the estimation of
hydration status from changes in body mass. J. Sports Sci. 25(7): 797–804.
doi:10.1080/02640410600875143. PMID:17454547.
Maughan, R.J., Otani, H., and Watson, P. 2012. Influence of relative humidity on
prolonged exercise capacity in a warm environment. Eur. J. Appl. Physiol.
112(6): 2313–2321. doi:10.1007/s00421-011-2206-7. PMID:22012542.
Mora-Rodriguez, R., Del Coso, J., Aguado-Jimenez, R., and Estevez, E. 2007.
Separate and combined effects of airflow and rehydration during exercise
in the heat. Med. Sci. Sports Exerc. 39(10): 1720–1726. doi:10.1249/mss.
0b013e3180de4dad. PMID:17909398.
Mora-Rodriguez, R., Del Coso, J., Hamouti, N., Estevez, E., and Ortega, J.F. 2010.
Aerobically trained individuals have greater increases in rectal temperature
than untrained ones during exercise in the heat at similar relative intensities.
Eur. J. Appl. Physiol. 109(5): 973–981. doi:10.1007/s00421-010-1436-4. PMID:
20349316.
Morrison, S., Sleivert, G.G., and Cheung, S.S. 2004. Passive hyperthermia reduces
voluntary activation and isometric force production. Eur. J. Appl. Physiol.
91(5–6): 729–736. doi:10.1007/s00421-004-1063-z. PMID:15015001.
Morrison, S.A., Cheung, S., and Cotter, J.D. 2014. Importance of airflow for
physiologic and ergogenic effects of precooling. J. Athl. Train. 49(5): 632–639.
doi:10.4085/1062-6050-49.3.27. PMID:25144598.
Nakagawa, S., and Cuthill, I.C. 2007. Effect size, confidence interval and statis-
tical significance: a practical guide for biologists. Biol. Rev. Camb. Philos. Soc.
82(4): 591–605. doi:10.1111/j.1469-185X.2007.00027.x. PMID:17944619.
Nielsen, B. 1996. Olympics in Atlanta: a fight against physics. Med. Sci. Sports
Exerc. 28(6): 665–668. doi:10.1097/00005768-199606000-00004. PMID:8784753.
Nielsen, B., Hales, J.R., Strange, S., Christensen, N.J., Warberg, J., and Saltin, B.
1993. Human circulatory and thermoregulatory adaptations with heat accli-
mation and exercise in a hot, dry environment. J. Physiol. 460: 467–485.
doi:10.1113/jphysiol.1993.sp019482. PMID:8487204.
Nybo, L., and Nielsen, B. 2001. Middle cerebral artery blood velocity is reduced
with hyperthermia during prolonged exercise in humans. J. Physiol. 534(1):
279–286. doi:10.1111/j.1469-7793.2001.t01-1-00279.x. PMID:11433008.
Otani, H., Kaya, M., Tsujita, J., Hori, K., and Hori, S. 2006. Low levels of hypohy-
dration and endurance capacity during heavy exercise in untrained individ-
uals. J. Therm. Biol. 31(1–2): 186–193. doi:10.1016/j.jtherbio.2005.11.018.
Otani, H., Kaya, M., Tamaki, A., Watson, P., and Maughan, R.J. 2016. Effects of
solar radiation on endurance exercise capacity in a hot environment. Eur. J.
Appl. Physiol. 116(4): 769–779. doi:10.1007/s00421-016-3335-9. PMID:26842928.
Otani, H., Kaya, M., Tamaki, A., and Watson, P. 2017. Separate and combined
effects of exposure to heat stress and mental fatigue on endurance exercise
Appl. Physiol. Nutr. Metab. Vol. 43, 2018
capacity in the heat. Eur. J. Appl. Physiol. 117(1): 119–129. doi:10.1007/s00421-
016-3504-x. PMID:27864637.
Pageaux, B., Marcora, S.M., and Lepers, R. 2013. Prolonged mental exertion does
not alter neuromuscular function of the knee extensors. Med. Sci. Sports
Exerc. 45(12): 2254–2264. doi:10.1249/MSS.0b013e31829b504a. PMID:23698244.
Parsons, K. 2014. Human Thermal Environments. Tayler & Francis, London, UK.
Racinais, S., Périard, J.D., Karlsen, A., and Nybo, L. 2015. Effect of heat and heat
acclimatization on cycling time trial performance and pacing. Med. Sci.
Sports Exerc. 47(3): 601–606. doi:10.1249/MSS.0000000000000428. PMID:
24977692.
Ramanathan, N.L. 1964. A new weighting system for mean surface temperature
of the human body. J. Appl. Physiol. 19: 531–533. PMID:14173555.
Ravanelli, N.M., Hodder, S.G., Havenith, G., and Jay, O. 2015. Heart rate and body
temperature responses to extreme heat and humidity with and without elec-
tric fans. JAMA, 313(7): 724–725. doi:10.1001/jama.2015.153. PMID:25688786.
Roberts, M.F., Wenger, C.B., Stolwijk, J.A., and Nadel, E.R. 1977. Skin blood flow
and sweating changes following exercise training and heat acclimation.
J. Appl. Physiol. 43(1): 133–137. PMID:893254.
Saunders, A.G., Dugas, J.P., Tucker, R., Lambert, M.I., and Noakes, T.D. 2005. The
effects of different air velocities on heat storage and body temperature in
humans cycling in a hot, humid environment. Acta Physiol. Scand. 183(3):
241–255. doi:10.1111/j.1365-201X.2004.01400.x. PMID:15743384.
Sawka, M.N., Leon, L.R., Montain, S.J., and Sonna, L.A. 2011. Integrated physio-
logical mechanisms of exercise performance, adaptation, and maladaptation
to heat stress. Compr. Physiol. 1(4): 1883–1928. doi:10.1002/cphy.c100082.
PMID:23733692.
Sawka, M.N., Cheuvront, S.N., and Kenefick, R.W. 2012. High skin temperature
and hypohydration impair aerobic performance. Exp. Physiol. 97(3): 327–332.
doi:10.1113/expphysiol.2011.061002. PMID:22143882.
Shaffrath, J.D., and Adams, W.C. 1984. Effects of airflow and work load on car-
diovascular drift and skin blood flow. J. Appl. Physiol. 56(5): 1411–1417. PMID:
6725094.
Stevens, C.J., Thoseby, B., Sculley, D.V., Callister, R., Taylor, L., and
Dascombe, B.J. 2016. Running performance and thermal sensation in the
heat are improved with menthol mouth rinse but not ice slurry ingestion.
Scand. J. Med. Sci. Sports, 26(10): 1209–1216. doi:10.1111/sms.12555. PMID:
26408395.
Stöhr, E.J., Gonzalez-Alonso, J., Pearson, J., Pearson, J., Low, D.A., Ali, L., et al.
2011. Dehydration reduces left ventricular filling at rest and during exercise
independent of twist mechanics. J. Appl. Physiol. (1985), 111(3): 891–897. doi:
10.1152/japplphysiol.00528.2011.
Teunissen, L.P., de Haan, A., de Koning, J.J., and Daanen, H.A. 2013. Effects of
wind application on thermal perception and self-paced performance. Eur.
J. Appl. Physiol. 113(7): 1705–1717. doi:10.1007/s00421-013-2596-9. PMID:
23385657.
Thomas, M.M., Cheung, S.S., Elder, G.C., and Sleivert, G.G. 2006. Voluntary mus-
cle activation is impaired by core temperature rather than local muscle
temperature. J. Appl. Physiol. (1985), 100(4): 1361–1369. doi:10.1152/
japplphysiol.00945.2005.
Trinity, J.D., Pahnke, M.D., Lee, J.F., and Coyle, E.F. 2010. Interaction of hyper-
thermia and heart rate on stroke volume during prolonged exercise. J. Appl.
Physiol. (1985), 109(3): 745–751. doi:10.1152/japplphysiol.00377.2010.
Wall, B.A., Watson, G., Peiffer, J.J., Abbiss, C.R., Siegel, R., and Laursen, P.B. 2015.
Current hydration guidelines are erroneous: dehydration does not impair
exercise performance in the heat. Br. J. Sports Med. 49(16): 1077–1083. doi:10.
1136/bjsports-2013-092417. PMID:24055782.
Watson, P., Hasegawa, H., Roelands, B., Piacentini, M.F., Looverie, R., and
Meeusen, R. 2005. Acute dopamine/noradrenaline reuptake inhibition en-
hances exercise performance in warm, but not temperate conditions. J. Physiol.
565(Pt. 3): 873–883. doi:10.1113/jphysiol.2004.079202. PMID:15831540.
Watson, P., Shirreffs, S.M., and Maughan, R.J. 2012. Effect of dilute CHO bever-
ages on performance in cool and warm environments. Med. Sci. Sports Exerc.
44(2): 336–343. doi:10.1249/MSS.0b013e31822dc5ed. PMID:22251924.
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