Scand J Med Sci Sports 2010: 20 (Suppl.
3): 80–89 & 2010 John Wiley & Sons A/S
doi: 10.1111/j.1600-0838.2010.01212.x
Review
Different effects of heat exposure upon exercise performance in the
morning and afternoon
S. Racinais
Research and Education Centre, ASPETAR, Qatar Orthopaedic Sports Medicine Hospital, Doha, Qatar
Corresponding author: Se´bastien Racinais, Research and Education Centre, ASPETAR, Qatar Orthopaedic and Sports
Medicine Hospital, PO Box 29222, Doha, Qatar. Tel: 1974 4413 2570, Fax: 1974 4413 2034, E-mail: sebastien.racinais@
aspetar.com
Accepted for publication 22 July 2010
Independent of environmental conditions, rectal tempera-
ture follows a circadian rhythm with an acrophase in the late
afternoon. In neutral environment, this diurnal increase in
temperature is believed to have a passive warm-up effect
improving muscle contractility, and in turn, muscle force,
power and performance. However, a hot environment blunts
the diurnal variation in muscle function by only improving
muscle contractility, and in turn, muscle force, power and
performance in the morning, when body temperature is at its
lowest. Despite this diurnal variation in muscle function,
Research examining the effect of hot environments
on physical abilities have generally focused on cen-
tral temperature (e.g., rectal or esophageal) and the
preservation of the homeostasis. However, even in a
thermally comfortable and neutral environment,
body temperatures are not stable physiological
parameters. As most of the biological variables in
both animal and vegetable species, rectal tempera-
ture follows biological rhythms. The first part of
this review describes the circadian rhythm in rectal
temperature and its effect on performance. Then,
the repercussions of the diurnal variation in
both temperature and performance are addressed
when exercising in hot environments. The other
biological rhythms potentially influencing exercise
capacity in hot environment are briefly presented in
a third part.
Circadian rhythms and time-of-day effect on
performance in neutral environment
Generality on circadian rhythms and time-of-day
Biological rhythms represent cyclic variations in
biological functions. They can be described by their
mesor (i.e., average value), their period, their ampli-
tude or their accrophase (i.e., time of the peak value).
Biological rhythms are part of our genetic pheno-
80
long-duration exercise is only slightly affected by the time-
of-day in neutral environment. However, higher afternoon
body temperatures can reduce the heat storage capacity and
result in a reduction in exercise capacity in hot environ-
ments. In addition, in parallel to the circadian variations in
muscle contractility and central temperature, exercise ca-
pacity in hot environment may also be affected by the
diurnal variations in melatonin concentration and in the
onset of peripheral vasodilatation and sweating.
type, thus endogenous. A biological rhythm is stable
under constant environmental factors (i.e., light,
temperature, food intake) but can change its period
according to environmental synchronizers. In exer-
cise physiology, the most studied biological rhythms
are the circadian rhythms. Their endogenous period
is around 25 h, but there are synchronized by the
solar day on a period of 24 h (circa 5 around,
diem 5 day). The most known circadian rhythm is
the sleep–wake cycle.
When studying the specificities of exercising in hot
environments, the circadian rhythms in body tem-
peratures (e.g., rectal) have to be considered. In
neutral environment, rectal temperature follows a
sinusoidal variation (Fig. 1) with an acrophase
in the late afternoon, close to 18:00 hours (Water-
house et al., 1993). This circadian rhythm is mainly
endogenous and persists in free-living conditions,
without the influence of the zeitgebers (Colin et al.,
1968).
Statistical significance of some circadian rhythms
could be linked to extremely low values during night,
whereas most of the physical activities (i.e., sport
events, leisure or work) are performed during the
day. A functional approach for chronobiology in
sports and physical abilities is to investigate the
existence of fluctuation during day-time rather than
the nocturnal nadir. Studies examining the whole

Fig. 1. Circadian rhythm in body temperature (F}F) and
muscle power (- -&- -). These two rhythms fluctuate in
phase, with an acrophase in the late afternoon. Redrawn
with permission from the Data of Souissi et al. (2004).
solar cycle (i.e., 24 h) generally use the terminology of
‘‘circadian rhythms,’’ whereas studies focusing on the
day-time refer to ‘‘time-of-day.’’
This review considers the diurnal variation in
performance (i.e., time-of-day effect), the diurnal
increase in rectal temperature, the effect of an ex-
posure to a hot environment and their interaction
effect on performance.
Diurnal variation in performance during
short-duration exercise
Performances of short duration (i.e., less than a
minute) present the common characteristic to be better
in the afternoon (e.g., between 16:00 and 20:00 hours)
than in the morning (e.g., between 06:00 and 10:00
hours). Even if some studies did not observe any
differences in power output during a sprint on cycle
ergometer in the morning and the afternoon (Falgair-
ette et al., 2003; Giacomoni et al., 2006), most of the
studies consistently reported an improved performance
in the afternoon when compared with the morning.
This diurnal increment has been observed for: vertical
jump height (Bernard et al., 1998; Racinais et al.,
2004a), outdoor sprint running velocity (Javierre
et al., 1996), maximal cycling power output (Bernard
et al., 1998; Racinais et al., 2005a, c), or total work
completed during 30 (Melhim, 1993) and 60 s (Hill
et al., 1992) of cycle exercise. All this studies reported
statistically significant difference of small-to-moderate
amplitude (generally less than 10%).
Diurnal variation in muscle properties
The maximal voluntary force produced by the arm
flexors (Gauthier et al., 1996) or the knee extensors
(Racinais et al., 2005b) has been observed to be
higher in the afternoon than the morning. Interest-
ingly, as displayed in Fig. 2, this higher muscle force
was observed without modification of the electrical
activity (recorded by surface electromyography and
Chronobiology and environmental physiology
quantified by RMS activity) of the biceps brachialis
(Gauthier et al., 1996) or the vastus lateralis (Raci-
nais et al., 2005b). This suggests that the diurnal
improvement in muscle force is not due to a mod-
ification in neural drive but rather due to an im-
provement of the muscle contractile properties in the
afternoon. Better muscle contractile properties in the
afternoon than in the morning have been evidenced
by a higher mechanical response to an electrical
stimulation of the motor nerve (i.e., Pt) in the after-
noon than in the morning (Martin et al., 1999) as well
as a higher RMS/force ratio during voluntary con-
traction in the afternoon than in the morning (Gau-
thier et al., 1996; Racinais et al., 2005b).
What could explain the diurnal increase in force and
power output during short-duration exercise?
The diurnal increase in power output during cycling
sprints has been attributed to a higher force produc-
tion during the sprints performed in the afternoon
than the morning (Racinais et al., 2005a). As detailed
above, this higher force production has been linked
to better muscle contractile properties in the after-
noon than the morning (Gauthier et al., 1996; Martin
et al., 1999; Racinais et al., 2005b). However, the
physiological basis of this diurnal variation in muscle
contractile properties remains unclear. Based on the
fact that the circadian rhythm in short-duration
performance is in phase with the circadian rhythm
in rectal (Reilly & Down, 1986, 1992) and oral
(Souissi et al., 2004) temperature (Fig. 1), it has
been suggested that the lower body temperatures in
the morning than the afternoon do not match the
optimal conditions for muscle contractility (Racinais
et al., 2005b) and that diurnal increase in tempera-
ture could act as a passive warm-up improving
performance (Melhim, 1993; Bernard et al., 1998;
Racinais et al., 2004a, b, 2005b). However, the diur-
nal variation in muscle temperature has never been
investigated yet and the hypothesis of a passive
warm-up effect cannot be demonstrated.
Long duration exercise in neutral environment
Whereas diurnal variations in short-duration perfor-
mance have been widely described, such variations
seem to dissipate when exercise prolonged. In neutral
environments, the ability to perform in middle- and
long-duration events is partly linked to the athlete’s
maximal oxygen consumption (VO ·
2peak), ventilatory
thresholds and running/cycling economy. However,
there is no clear evidence that these parameters
fluctuate during the day. Even though some studies
have observed a diurnal variation in VO ·
2peak (Hill
et al., 1988, 1989; Williams & Hill, 1995), others
·
concluded that VO 2peak is not time-of-day dependent
81
Racinais
Fig. 2. Muscle force and neural drive in neutral and hot environment. A hot environment improves muscle force in the
morning only, when body temperature is at its lowest. Hot environment and time-of-day can improve muscle force without
modification in neural drive by improving the contractile properties. Reproduced with permission from Racinais et al. (2005b).
(Faria & Drummond, 1982; Reilly & Brooks, 1982,
1990; Jones & Hill, 1995). What is interesting to note
is that a diurnal fluctuation in maximal oxygen
consumption does not translate to any variation in
performance itself (e.g., time to exhaustion) (Hill
et al., 1988). A time-of-day effect has been observed
previously for VO ·
2 kinetics during moderate-
intensity cycling exercise (Brisswalter et al., 2007),
but not during high-intensity running (Carter et al.,
2002) or cycling (Bessot et al., 2006) exercise. Venti-
latory threshold (Hill et al., 1989) and VO ·
2 during
high-intensity exercise (Dalton et al., 1997; Deschenes
et al., 1998) also do not depend of time-of-day. There-
fore, prolonged exercise capacity does not appear to
depend on time-of-day or to follow circadian varia-
tions in a neutral environment (Table 1).
Hot environments differently affect exercise capacity
in the morning and the afternoon
Effect of hot environments on exercise capacity
Hot environment exposure affects exercise capacity
in a different way in the function of exercise duration
and heat stress intensity. Briefly, during short-dura-
tion exercise, increasing muscle temperature im-
proves muscle function and there is a positive
relationship between performance and muscle tem-
perature with a 2–5% modification by 1 1C of change
in muscle temperature (for a review, see Racinais &
Oksa, 2010 in this supplement). However, if the
central temperature increases (i.e., hyperthermia),
82
this positive relation stops and performance becomes
impaired due to central and peripheral failures alter-
ing neural drive (Racinais et al., 2008).
When exercise prolonged, the increase in central
temperature is controlled by different thermoregula-
tory systems in an attempt to preserve homeostasis.
The majority of heat dissipation is facilitated by
evaporation, while convection, conduction and ra-
diation play smaller roles. A part of the heat is also
stored in the body, but the storage possibilities are
limited, and central – and may be cerebral – tem-
perature can be a limiting factor to exercising in
warm environment (Caputa et al., 1986; Brück &
Olschewski, 1987; Nielsen et al., 1993; Gonzalez-
Alonso et al., 1999). There is no unique threshold
(Ely et al., 2009) but cessation of activity in hot
environment seems to occur at a critical central
temperature whatever the exercise intensity, the
heat storage capacity, the acclimatized state, the rate
of heat storage, the initial temperature or the skin
temperature (Fig. 5, Brück & Olschewski, 1987;
Nielsen et al., 1993; Gonzalez-Alonso et al., 1999).
Similarly, rats stop exercising in hot environment at
the same abdominal and cerebral temperature what-
ever the modification made to their initial tempera-
ture (Fuller et al., 1998; Walters et al., 2000).
Hot environment vs diurnal increase in central
temperature as passive warm-ups
Power output during a cycling sprint (Falk et al.,
1998) and jump height (Hue et al., 2003) have been
 Chronobiology and environmental physiology
Table 1. Effect of time-of-day on exercise capacity in neutral and hot environment

Neutral environment Hot environment

(A) Short duration exercise (generally less than 1 min – e.g., muscle force, power)
Effect of time-of-day on MorningoAfternoon Morning 5 Afternoon
performance
Possible physiological The diurnal increase in body temperature could A hot environment can act as a passive warm-up
explanation act as a passive warm-up improving muscle and improve muscle contractility and performance
contractility. in the morning only, when body temperature is at
its lowest.
References Bernard et al. (1998), Gauthier et al. (1996), Hill Racinais et al. (2004b, 2005b)
et al. (1992), Javierre et al. (1996), Melhim
(1993), Racinais et al. (2004b, 2005a, b, c)
(B) Long duration exercise (generally more than 10 min – e.g., cycling, running)
Effect of time-of-day on Morning 5 Afternoon Morning4Afternoon
performance
Possible physiological There is no evidence for diurnal variation in The diurnal increase in central temperature can
explanation maximal oxygen consumption, ventilatory reduce heat storage capacity.
threshold or running/cycling economy.
References Faria and Drummond (1982), Hill et al. (1988), Hobson et al. (2009)
Jones and Hill (1995), Reilly and Brooks
(1982, 1990)

See text for more details.

reported to be higher in hot than in neutral environ-
ment, probably due to the passive warm-up effect of
a hot exposure. Similarly, as detailed above, the
diurnal increase in body temperature has been con-
sidered as a passive warm-up improving the power
output also during a cycling sprint or jump height
(Melhim, 1993; Bernard et al., 1998; Racinais et al.,
2004a). Therefore, time-of-day (i.e., the diurnal in-
crease in body temperature) and a hot environment
seem to represent two factors that increase short-
duration muscle performance through a passive
warm-up effect. Hence, the question is to know
whether an external passive warm-up as a hot en-
vironment possibly overrides the passive warm-up
effect of the time-of-day, and reciprocally.
As displayed in Fig. 3, recent data showed that
maximal power output during a cycling sprint (Ra-
cinais et al., 2004a), jump height (Racinais et al.,
2004a) and maximal voluntary muscle force (Raci-
nais et al., 2005b) are significantly better in hot
environment than in neutral environment if the tests
are performed in the morning (e.g., 07:00–09:00
hours) but not in the afternoon (e.g., 17:00–19:00
hours). Consequently, whereas these parameters dis-
played a diurnal increase in neutral environment, the
values recorded in a hot environment are the same in
the morning and the afternoon (Racinais et al.,
2004a, 2005b) and even during the night (Racinais
et al., 2004b). As for the effect of time-of-day, a hot
environment seems to improve muscle contractility
(Racinais et al., 2005b), which in turn, improved
muscle force (Racinais et al., 2005b) and ultimately
power production (Racinais et al., 2004a), but in the
morning only, when body temperature is at its lowest
(Fig. 3, Table 1).
These findings suggest that adding a hot exposure
to the diurnal increase in body temperature had no
cumulative effect on the improvement in muscle
power. The existence of a threshold above which
additional sources of passive warm-up have no
additional effect is supported by the fact that, in
moderately warm and humid environment, neither
the diurnal increase in body temperature and/or
30 min of exposure at a temperature of 76 1C (sauna)
improve power output during cycling sprint, jump
height nor muscle force (Racinais et al., 2006).
Lastly, it is practically relevant to note that athletes
and coaches can also improve maximal power output
in the morning and blunt its diurnal variation by
immersing the leg for 30 min in a hot (38 1C) water
baths (Racinais et al., 2009). Conversely, an active
warm-up improves the maximal power output during
a cycling sprint both in the morning and afternoon
(Racinais et al., 2005a).
Physiological basis and limitations to consider the
diurnal increase in central temperature as a
passive warm-up
Low muscle temperatures have been shown to in-
crease Ca21 retention by the sarcoplasmic reticulum
(Stein et al., 1982; Segal et al., 1986). This could
potentially explain the lower muscle contractility
observed in the morning in neutral environment
than in hot environment or in the afternoon. In
addition, Ca21 concentration and temperature influ-
enced muscle force at low temperature but less when
muscle temperature increased (Stephenson & Wil-
liams, 1981), therefore potentially explaining the no-
cumulative effect of different passive warm-up.

83
Racinais
Fig. 3. The effect of hot environment on muscle function is
different in the morning and the afternoon. Muscle contrac-
tility (top graph), force (middle graph) and performance
(bottom graph) are better in hot (black bars) than in neutral
(white bars) environment, in the morning only (*Po0.05).
Heat does not improve muscle function in the afternoon
(NS, no significant). There is a diurnal increase in muscle
function in neutral environment only (Po0.05). Redrawn
with permission from the data of Racinais et al. (2004a,
2005b).
Moreover, Ca21 concentration is not likely to influ-
ence muscle contractility during submaximal con-
tractions, when the needs are lower. This would also
explain why muscle contractility is significantly
higher in the afternoon than in the morning during
high-intensity contractions (i.e., from 75% to 100%
of max), but not during contractions at 25% or 50%
of max (Racinais et al., 2005b). However, all the
studies on the effect of Ca21 concentration and
temperature were comparing muscle temperature
over a range of 10 1C, whereas the diurnal variation
in muscle temperature in vivo has never been inves-
tigated yet. There is a very strong limitation in
84
considering the diurnal increase in rectal temperature
as a passive warm-up for the muscle. Actual data on
muscle temperature in the morning and the after-
noon are mandatory to better understand the effect
of time-of-day.
Lastly, if a hot exposure can act as a passive warm-
up and improve muscle contractile properties, neu-
romuscular function becomes impaired when tem-
peratures reach hyperthermia (for a review, see
Racinais & Oksa, 2010 in this issue) but the effect
of time-of-day on these alterations is unknown.
Diurnal increase in central temperature limits long-
duration exercise capacity in hot environment
Although some studies have observed a reduction in
the amplitude of the diurnal variation in central
temperature in response to heat exposure (Little &
Rummel, 1971; Torii et al., 1995) or 30–60 min of
exercise (Reilly & Garrett, 1995; Aldemir et al.,
2000), the absolute increase in core temperature
during exercise seems mainly independent of time-
of-day (Reilly & Brooks, 1982; Stephenson et al.,
1984; Reilly & Garrett, 1995; Deschenes et al., 1998;
Bessot et al., 2006) and the circadian rhythm in rectal
temperature is still present in hot environment
(Fig. 4, Racinais et al., 2004b). In addition, an
increase in body temperature before long-duration
exercise can decrease performance by altering the
heat storage capacity (Nadel, 1987; Gonzalez-Alonso
et al., 1999); whereas pre-cooling can increase the
heat storage capacity and increase the time to ex-
haustion (Lee & Haymes, 1995; Gonzalez-Alonso
et al., 1999). From this perspective, the diurnal
increase in body temperature will modify the heat
storage capacity and thus the ability to exercise.
This is valid only if central temperature is the
limiting factor, i.e. when exercising in a hot environ-
ment. Some limitation to this hypothesis comes from
Fig. 4. Circadian rhythm in body temperature in hot envir-
onment. Central (i.e., rectal) temperature follow a circadian
rhythm in hot environment also (average 37.11C, acrophase
at 16:02 hours). Redrawn with permission from the Data of
Racinais et al. (2004b).
 Chronobiology and environmental physiology
McLellan et al. (1999), who found that the diurnal
increase in central temperature remained during
exercising in the heat, thus leading to a significantly
higher final temperature in the afternoon than in the
morning. Given that the temperature threshold for
the onset of sweating and skin vasodilatation have
been shown to occur at a higher temperature in the
afternoon than the morning (Stephenson et al.,
1984), the authors suggested that the central tem-
perature tolerated at exhaustion could also be regu-
lated at a higher temperature in the afternoon
(McLellan et al., 1999). However, the data of this
experiment have to be interpreted with caution as the
participants were not exercising up to volitional
exhaustion but doing an intermittent walk with
nuclear, biological and chemical protective clothing,
which imposed an uncompensable heat stress
(McLellan et al., 1999). In addition, the trials were
conducted only 4 h apart, in the morning and the
early afternoon, which are times that do not max-
imize the amplitude of the diurnal variation in
central temperature.
Given that there is no other evidence to suggest
that the temperature tolerated at exhaustion varies
during the day, it has been widely hypothesized that Fig. 5. Exercise duration in hot environment is partly de-
the diurnal increase in resting central temperature pendant of the initial temperature. A decrease or an increase
would reduce the exercise capacity in hot environ- in initial temperature by pre-cooling or pre-warming can,
ments before reaching a critical temperature (Atkin- respectively, increase or decrease the capacity to exercise in
hot environment (top graph, redrawn with permission from
son & Reilly, 1995; Reilly & Garrett, 1995; Reilly & Gonzalez-Alonso et al., 1999). Following the same model,
Garret, 1998; McLellan et al., 1999; Atkinson et al., the diurnal increase in resting temperature will reduce the
2005; Racinais, 2008; Hobson et al., 2009). Reilly and capacity to exercise in hot environment (bottom graph,
Garret (1998) were the first to experiment the hy- redrawn with permission from Hobson et al., 2009).
pothesis of an improved exercise capacity due to a
lower basal central temperature in the morning than
in the afternoon. No diurnal differences in exercise the afternoon, but not different at exhaustion due to
capacity were observed in seven football players the longer exercise duration (Fig. 5) and may be also
tested during identical sessions performed in the a tendency to a higher rate of temperature increase.
morning and the afternoon (Reilly & Garret, 1998). These data confirm that, even of relatively small
The absence of diurnal variation can be explained by amplitude, the diurnal variation in central tempera-
the cool environmental conditions used in this ex- ture affect the capacity to exercise in hot environment
periment (17.6 1C and 51% RH). As stated above, a (Table 1).
modification in heat storage capacity due to the
circadian rhythm in rectal temperature is only likely
to affect exercise capacity in hot environment, when
Other biological variations potentially affecting
heat dissipation fails to maintain homeostasis and
thus the heat storage capacity becomes a limiting exercise capacity in hot environment
factor. Circadian variation in sweating and thermoregulatory
Morning vs afternoon exercise capacities were abilities
recently compared during a cycling exercise (65% An interaction between the circadian rhythm of
VO2max) performed to volitional exhaustion in a hot central temperature and changes in temperature
environment (35 1C and 60% RH) (Hobson et al., produced by activity has been observed in rodent
2009). The authors aimed to verify if the lower initial (Weinert & Waterhouse, 1998) and human studies
morning central temperature allowed subjects to (Waterhouse et al., 1999). Aldemir et al. (2000)
cycle for longer before volitional cessation. In line suggested that, in neutral environments, skin vaso-
with this hypothesis, time to exhaustion was 13.8% dilatation for heat dissipation could be faster in the
higher in the morning than in the evening. Rectal late afternoon than the early morning. The thresh-
temperature was lower at rest in the morning than olds for the onset of peripheral vasodilatation and

85
Racinais
sweating have been reported to be lower in the
evening, facilitating heat dissipation (Stephenson
et al., 1984; Torii et al., 1995; Waterhouse et al.,
2007). This could explain the tendency for a lower
rate of central temperature increment during an
exercise performed in the late afternoon as compared
with the same exercise in the morning (Aldemir et al.,
2000; Hobson et al., 2009). However, despite the
tendency for a lower rate of increment in central
temperature in the afternoon, the actual values stay
higher in the afternoon than in the morning at a
given exercise duration and this difference only dis-
appeared due to the longer duration of the morning
session (Fig. 5, Hobson et al., 2009). In addition,
another study failed to observe diurnal differences in
the rate of heat storage (McLellan et al., 1999) and
the circadian rhythm in sweat rate observed during
low-intensity exercise disappeared at higher work-
load (Torii et al., 1995). Taken together, these studies
suggest that the diurnal variation in sweating should
not have an impact on performance.
The melatonin circadian variation
The circadian variation in melatonin secretion is lead
by the light–dark cycle (Geoffriau et al., 1998) with
secretion starting generally when luminosity is below
2500 lux (i.e., around 20:00–21:00 hours), peaking in
the early morning around 03:00–04:00 hours and
then decreasing to reach baseline daylight values at
around 08:00 hours (Cagnacci et al., 1992, 1996,
1997; Dawson & Van den Heuvel, 1998). Melatonin
is considered as a strong pacemaker of circadian
rhythms and the nocturnal fall in central temperature
seems inversely related to the melatonin rise (Cag-
nacci et al., 1992). In addition, oral ingestion of
melatonin can induce a decrease in central tempera-
ture (Dawson et al., 1996; McLellan et al., 2000).
Melatonin has been widely considered to have a
hypothermic effect (Strassman et al., 1991; Cagnacci
et al., 1992, 1994, 1996; Dollins et al., 1994; Deacon
& Arendt, 1995; Dawson et al., 1996; Reid et al.,
1996; McLellan et al., 2000). This effect seems to
occur in response to a threshold concentration rather
than represent a dose response (Cagnacci et al., 1996;
Dawson et al., 1996), and given the sleep-inducing
effect of melatonin (Dollins et al., 1994; Reid et al.,
1996), doses higher than 5 mg should be avoided if
used to improve performance (McLellan et al., 2000).
The hypothermic effect of melatonin ingestion has
been reported for doses of 1–10 mg (Strassman et al.,
1991; Cagnacci et al., 1992, 1994, 1996; Dollins et al.,
1994; Deacon & Arendt, 1995; Dawson et al., 1996;
Reid et al., 1996; McLellan et al., 2000) but seems
variable between individuals (Dawson et al., 1996;
Reid et al., 1996) and environmental condition
(McLellan et al., 2000). For example, a hypothermic
86
effect has been observed at rest in neutral but not hot
environment (McLellan et al., 2000), at rest but
disappearing during exercise (McLellan et al.,
2000), or on the contrary, not present at rest but
appearing after 20–25 min of exercise (Atkinson
et al., 2005). Although still not fully understood,
the hypothermic effect of melatonin may be partly
due to an increased convective heat loss as suggested
by the increase in temperature of the feet (Kräuchi &
Wirz-Justice, 1994) and hands (Van den Heuvel et
al., 1999) in humans or of the metatarsals in chickens
(Rozenboim et al., 1998). However, this increase in
peripheral cutaneous vasodilatation to promote heat
loss appears to be negated by the orthostatic chal-
lenge and the increase in vasoconstrictor tone when
changing from a supine to upright position (Kräuchi
et al., 1997), or by the vasodilation in response to a
hot environment exposure that also blunts the hy-
pothermic effect of melatonin observed in neutral
temperature (McLellan et al., 2000). Consequently,
whereas melatonin ingestion has been observed to
moderate the rise in central temperature when ex-
ercising at 27 1C (Atkinson et al., 2005), it failed to
influence the rate of sweat production, the rate of
heat storage or the central temperature at 40 1C,
during rest or exercise (McLellan et al., 1999, 2000).
Therefore, there is no evidence that melatonin is an
ergogenic aid for endurance performance in hot
environments (Atkinson et al., 2005).
Menstrual cycle
There is a consensus that the menstrual phase does
not affect physical performance (Galliven et al., 1997;
Shephard, 2000) nor its diurnal variation (Galliven
et al., 1997) over the course of a normal menstrual
cycle in neutral environment. For example, the
menstrual cycle phase does not modify the muscle
power production during neither jump nor cycling
tests (Giacomoni et al., 2000). Moreover, it is im-
portant to note that body temperature is higher in
the luteal phase and lower in the preovulatory phase
compared with the follicular phase (Coyne et al.,
2000). Following the same model than presented
above for the effect of the diurnal increase in central
temperature, this menstrual increase may reduce the
safe margin for heat storage when exercising in hot
environment (Shephard, 2000). However, the impact
on physiological responses to submaximal exercise
seems minimal, even when such activity is performed
in the heat (Horvath & Drinkwater, 1982). This
could partly be explained by the fact that the
amplitude of the diurnal increase in central tempera-
ture is reduced during the luteal phase (Coyne et al.,
2000), potentially reducing the negative effect of the
menstrual increase in central temperature during this
phase. It is also important to note that neither

menstrual cycle nor the use of oral contraceptives
modifies post-exercise heat loss responses (Kenny et
al., 2008). However, if the effect of physical training
on heat acclimation is not affected by using contra-
ceptive hormones (Armstrong et al., 2005), the in-
crease in sweating with training may be affected by
the phase of the menstrual cycle during which the
training was performed (Ichinose et al., 2009).
Jetlag
Even if it is not the topic of this review, it should be
acknowledged that many athletes living in temperate
countries are exposed to hot environments when
competing in hot countries and, if more than four
time zones are flown through, the athlete will also
suffer from jet lag. Symptoms are due to an internal
desynchronization between different circadian
rhythms adapting to the local time zone at a different
rate. Adaptation required is around 1 day by the time
zone crossed and is easier when flying westwards
than eastwards. The reason for this is that endogen-
ous circadian rhythms have a period of 25 h making
it easier to adapt to a phase delay than a phase
advance. When considering athletes traveling west to
east, phase advance can be achieved by exercising
and light exposure in the morning. When considering
athletes traveling east to west, phase delay is facili-
tated by light exposure in the evening. It is also
important to facilitate nocturnal sleep by avoiding
naps and providing darkness.
Chronobiology and environmental physiology
Conclusion
In neutral environment, muscle contractility in-
creases from the morning to the afternoon (Table
1A). Consequently, muscle force and even short-
duration performance are better in the afternoon
than the morning. However, hot environments blunt
this diurnal variation by improving muscle contrac-
tility (and in turn muscle force, power and perfor-
mance) in the morning only, when body temperature
is at its lowest.
The capacity to perform prolonged exercise (Table
1B) is the same in the morning and afternoon in
neutral environment. However, when exercising in a
hot environment, the diurnal increase in central
temperature can reduce the heat storage capacity
and therefore exercise capacity.
There is an interaction between the time-of-day
and the effect of a hot environment making
it essential to take into account the time-of-day
when investigating the effect of the environmental
conditions.
Key words: time-of-day, circadian rhythms, thermo-
regulation, muscle, temperature.
Acknowledgements
Conflicts of interest: The author has no potential conflicts of
interest.

References
Aldemir H, Atkinson G, Cable T,
Edwards B, Waterhouse J, Reilly T. A
comparison of the immediate effects of
moderate exercise in the early morning
and late afternoon on core temperature
and cutaneous thermoregulatory
mechanisms. Chronobiol Int 2000: 17:
197–207.
Armstrong LE, Maresh CM, Keith NR,
Elliott TA, VanHeest JL, Scheett TP,
Stoppani J, Judelson DA, De Souza
MJ. Heat acclimation and physical
training adaptations of young women
using different contraceptive
hormones. Am J Physiol Endocrinol
Metab 2005: 288: E868–E875.
Atkinson G, Holder A, Robertson C,
Gant N, Drust B, Reilly T, Waterhouse
J. Effects of melatonin on the
thermoregulatory responses to
intermittent exercise. J Pineal Res 2005:
39: 353–359.
Atkinson G, Reilly T. Effects of age and
time of day on preferred work rates
during prolonged exercise. Chronobiol
Int 1995: 12: 121–134.
Bernard T, Giacomoni M, Gavarry O,
Seymat M, Falgairette G. Time-of-day
effects in maximal anaerobic leg
exercise. Eur J Appl Physiol 1998: 77:
133–138.
Bessot N, Nicolas A, Moussay S,
Gauthier A, Sesboue B, Davenne D.
The effect of pedal rate and time of day
on the time to exhaustion from high-
intensity exercise. Chronobiol Int 2006:
23: 1009–1024.
Brisswalter J, Bieuzen F, Giacomoni M,
Tricot V, Falgairette G. Morning-to-
evening differences in oxygen uptake
kinetics in short-duration cycling
exercise. Chronobiol Int 2007: 24: 495–
506.
Brück K, Olschewski H. Body
temperature related factors diminishing
the drive to exercise. Can J Physiol
Pharmacol 1987: 65: 1274–1280.
Cagnacci A, Elliott JA, Yen SS.
Melatonin: a major regulator of the
circadian rhythm of core temperature
in humans. J Clin Endocrinol Metab
1992: 75: 447–452.
Cagnacci A, Krauchi K, Wirz-Justice A,
Volpe A. Homeostatic versus circadian
effects of melatonin on core body
temperature in humans. J Biol
Rhythms 1997: 12: 509–517.
Cagnacci A, Soldani R, Laughlin GA,
Yen SS. Modification of circadian
body temperature rhythm during the
luteal menstrual phase: role of
melatonin. J Appl Physiol 1996: 80:
25–29.
Cagnacci A, Soldani R, Romagnolo C,
Yen SS. Melatonin-induced decrease of
body temperature in women: a
threshold event. Neuroendocrinology
1994: 60: 549–552.
Caputa M, Feistkorn G, Jessen C.
Effects of brain and trunk tem-
peratures on exercise performance in
goats. Pflugers Arch 1986: 406: 184–
189.
Carter H, Jones AM, Maxwell NS, Doust
JH. The effect of interdian and diurnal
variation on oxygen uptake kinetics
during treadmill running. J Sports Sci
2002: 20: 901–909.

87
Racinais
Colin J, Timbal J, Boutelier C, Houdas Y,
Siffre M. Rhythm of the rectal
temperature during a 6-month free-
running experiment. J Appl Physiol
1968: 25: 170–176.
Coyne MD, Kesick CM, Doherty TJ,
Kolka MA, Stephenson LA. Circadian
rhythm changes in core temperature
over the menstrual cycle: method for
noninvasive monitoring. Am J Physiol
Regul Integr Comp Physiol 2000: 279:
R1316–R1320.
Dalton B, McNaughton L, Davoren B.
Circadian rhythms have no effect on
cycling performance. Int J Sports Med
1997: 18: 538–542.
Dawson D, Gibbon S, Singh P. The
hypothermic effect of melatonin on
core body temperature: is more better?
J Pineal Res 1996: 20: 192–197.
Dawson D, van den Heuvel CJ.
Integrating the actions of melatonin on
human physiology. Ann Med 1998: 30:
95–102.
Deacon S, Arendt J. Melatonin-induced
temperature suppression and its acute
phase-shifting effects correlate in a
dose-dependent manner in humans.
Brain Res 1995: 688: 77–85.
Deschenes MR, Kraemer WJ, Bush JA,
Doughty TA, Kim D, Mullen KM,
Ramsey K. Biorhythmic influences on
functional capacity of human muscle
and physiological responses. Med Sci
Sports Exerc 1998: 30: 1399–1407.
Dollins AB, Zhdanova IV, Wurtman RJ,
Lynch HJ, Deng MH. Effect of
inducing nocturnal serum melatonin
concentrations in daytime on sleep,
mood, body temperature, and
performance. Proc Natl Acad Sci USA
1994: 91: 1824–1828.
Ely BR, Ely MR, Cheuvront SN,
Kenefick RW, DeGroot DW, Montain
SJ. Evidence against a 40 1C core
temperature threshold for fatigue in
humans. J Appl Physiol 2009: 107:
1519–1525.
Falgairette G, Billaut F, Ramdani S.
Effects of recovery duration and time
of day on sprint performance. Can J
Appl Physiol 2003: 28: 213–224.
Falk B, Radom-Isaac S, Hoffmann JR,
Wang Y, Yarom Y, Magazanik A,
Weinstein Y. The effect of heat
exposure on performance of and
recovery from high-intensity,
intermittent exercise. Int J Sports Med
1998: 19: 1–6.
Faria IE, Drummond BJ. Circadian
changes in resting heart rate and body
temperature, maximal oxygen
consumption and perceived exertion.
Ergonomics 1982: 25: 381–386.
Fuller A, Carter RN, Mitchell D. Brain
and abdominal temperatures at fatigue
in rats exercising in the heat. J Appl
Physiol 1998: 84: 877–883.
88
Galliven EA, Singh A, Michelson D, Bina
S, Gold PW, Deuster PA. Hormonal
and metabolic responses to exercise
across time of day and menstrual
cycle phase. J Appl Physiol 1997: 83:
1822–1831.
Gauthier A, Davenne D, Martin A,
Cometti G, Van Hoecke J. Diurnal
rhythm of the muscular performance of
elbow flexors during isometric
contractions. Chronobiol Int 1996: 13:
135–146.
Geoffriau M, Brun J, Chazot G, Claustrat
B. The physiology and pharmacology
of melatonin in humans. Horm Res
1998: 49: 136–141.
Giacomoni M, Bernard T, Gavarry O,
Altare S, Falgairette G. Influence of the
menstrual cycle phase and menstrual
symptoms on maximal anaerobic
performance. Med Sci Sports Exerc
2000: 32: 486–492.
Giacomoni M, Billaut F, Falgairette G.
Effects of the time of day on repeated
all-out cycle performance and short-
term recovery patterns. Int J Sports
Med 2006: 27: 468–474.
Gonzalez-Alonso J, Teller C, Andersen
SL, Jensen FB, Hyldig T, Nielsen B.
Influence of body temperature on the
development of fatigue during
prolonged exercise in the heat. J Appl
Physiol 1999: 86: 1032–1039.
Hill DW, Borden DO, Darnaby KM,
Hendricks DN, Hill CM. Effect of time
of day on aerobic and anaerobic
responses to high-intensity exercise.
Can J Sport Sci 1992: 17: 316–319.
Hill DW, Cureton KJ, Collins MA. Effect
of time of day on perceived exertion at
work rates above and below the
ventilatory threshold. R Q Exerc
Sports 1989: 60: 127–133.
Hill DW, Cureton KJ, Collins MA,
Grisham SC. Diurnal variations in
responses to exercise of ‘‘morning
types’’ and ‘‘evening types’’. J Sports
Med 1988: 28: 213–219.
Hobson RM, Clapp EL, Watson P,
Maughan RJ. Exercise capacity in the
heat is greater in the morning than in
the evening in man. Med Sci Sports
Exerc 2009: 41: 174–180.
Horvath SM, Drinkwater B.
Thermoregulation and the menstrual
cycle. Aviat Space Environ Med 1982:
53: 790–794.
Hue O, Coman F, Blonc S, Hertogh C.
Effect of tropical climate on
performance during repeated jump-
and-reach tests. J Sports Med Phys
Fitness 2003: 43: 475–480.
Ichinose TK, Inoue Y, Hirata M,
Shamsuddin AK, Kondo N. Enhanced
heat loss responses induced by
short-term endurance training in
exercising women. Exp Physiol 2009:
94: 90–102.
Javierre C, Calvo M, Diez A, Garrido E,
Segura R, Ventura JL. Influence of
sleep and meal schedules on
performance peaks in competitive
sprinters. Int J Sports Med 1996: 17:
404–408.
Jones HA, Hill DW. VO2 responses to
incremental and constant power
exercise tests at different times of day.
Med Sci Sports Exerc 1995: 27: S109.
Kenny GP, Leclair E, Sigal RJ, Journeay
WS, Kilby D, Nettlefold L, Reardon
FD, Jay O. Menstrual cycle and oral
contraceptive use do not modify
postexercise heat loss responses. J Appl
Physiol 2008: 105: 1156–1165.
Kräuchi K, Cajochen C, Hofmann M,
Wirz-Justice A. Melatonin and
orthostasis: interactions of posture
with subjective sleepiness, heart rate,
and skin and core temperature
(abstract). Sleep Res 1997: 26: 79.
Kräuchi K, Wirz-Justice A. Circadian
rhythm of heat production, heart rate,
and skin and core temperature under
unmasking conditions in men. Am J
Physiol 1994: 267: R819–R829.
Lee DT, Haymes EM. Exercise duration
and thermoregulatory responses after
whole body precooling. J Appl Physiol
1995: 79: 1971–1976.
Little MA, Rummel JA. Circadian
variations in thermal and metabolic
responses to heat exposure. J Appl
Physiol 1971: 31: 556–561.
Martin A, Carpentier A, Guissard N, Van
Hoecke J, Duchateau J. Effect of time
of day on force variation in a human
muscle. Muscle Nerve 1999: 22:
1380–1387.
McLellan TM, Gannon GA, Zamecnik J,
Gil V, Brown GM. Low doses of
melatonin and diurnal effects on
thermoregulation and tolerance to
uncompensable heat stress. J Appl
Physiol 1999: 87: 308–316.
McLellan TM, Smith IF, Gannon GA,
Zamecnik J. Melatonin has no effect on
tolerance to uncompensable heat stress
in man. Eur J Appl Physiol 2000: 83:
336–343.
Melhim AF. Investigation of circadian
rhythms in peak power and mean
power of female physical education
students. Int J Sports Med 1993: 14:
303–306.
Nadel ER. Prolonged exercise and high
and low ambient temperatures. Can J
Sport Sci 1987: 12: 140S–142S.
Nielsen B, Hales JR, Strange S,
Christensen NJ, Warberg J, Saltin B.
Human circulatory and
thermoregulatory adaptations with
heat acclimation and exercise in a hot,
dry environment. J Physiol 1993: 460:
467–485.
Racinais S. Circadian rhythms and
physical activity (Chapter). In: Leglise

AL, ed. Progress in circadian rhythm
research. New York: Nova Publishers,
2008: 163–176.
Racinais S, Blonc S, Hue O. Effects of
active warm-up and diurnal increase
in temperature on muscular power.
Med Sci Sports Exerc 2005a: 37:
2134–2139.
Racinais S, Blonc S, Jonville S, Hue O.
Time-of-day influences the
environmental effects on muscle force
and contractility. Med Sci Sports Exerc
2005b: 37: 256–261.
Racinais S, Blonc S, Oksa J, Hue O. Does
the diurnal increase in central
temperature interact with pre-cooling
or passive warm-up of the leg? J Sci
Med Sport 2009: 12: 97–100.
Racinais S, Chamari K, Hachana Y,
Bartagi Z, Blonc S, Hue O. Effect of an
acute hot and dry exposure in
moderately warm and humid
environment on muscle performance at
different time-of-day. Int J Sports Med
2006: 27: 49–54.
Racinais S, Connes P, Bishop D, Blonc S,
Hue O. Morning versus evening power
output and repeated-sprint ability.
Chronobiol Int 2005c: 22: 1029–1039.
Racinais S, Gaoua N, Grantham J.
Hyperthermia impairs short-term
memory and peripheral motor drive
transmission. J Physiol 2008: 586:
4751–4762.
Racinais S, Hue O, Blonc S. Time-of-day
effects on anaerobic muscular power in
a moderately warm environment.
Chronobiol Int 2004a: 21: 485–495.
Racinais S, Hue O, Hertogh C, Damiani
M, Blonc S. Time-of-day effects in
maximal anaerobic leg exercise in
tropical environment: a first approach.
Int J Sports Med 2004b: 25: 186–190.
Racinais S, Oksa J. Temperature and
neuromuscular function. Scand J Med
Sci Sports 2010: 20(Suppl. 3): 1–18.
Reid K, Van den Heuvel C, Dawson D.
Day-time melatonin administration:
effects on core temperature and sleep
onset latency. J Sleep Res 1996: 5: 150–
154.
Reilly T, Brooks GA. Investigation of
circadian rhythms in metabolic
Chronobiology and environmental physiology
responses to exercise. Ergonomics
1982: 25: 1093–1107.
Reilly T, Brooks GA. Selective
persistence of circadian rhythms in
physiological responses to exercise.
Chronobiol Int 1990: 7: 59–67.
Reilly T, Down A. Circadian variation in
the standing broad jump. Percept Mot
Skills 1986: 62: 830.
Reilly T, Down A. Investigation of
circadian rhythms in anaerobic
power and capacity of the legs.
J Sports Med Phys Fitness 1992:
32: 343–347.
Reilly T, Garret R. Investigation of
diurnal variation in sustained exercise
performance. Ergonomics 1998: 41:
1085–1094.
Reilly T, Garrett R. Effects of time of day
on self-paced performances of
prolonged exercise. J Sports Med Phys
Fitness 1995: 35: 99–102.
Rozenboim I, Miara L, Wolfenson D.
The thermoregulatory mechanism of
melatonin-induced hypothermia in
chicken. Am J Physiol 1998: 274:
R232–R236.
Segal SS, Faulkner JA, White TP.
Skeletal muscle fatigue in vitro is
temperature dependent. J Appl Physiol
1986: 61: 660–665.
Shephard RJ. Exercise and training in
women, part II: influence of menstrual
cycle and pregnancy on exercise
responses. Can J Apl Physiol 2000: 25:
35–54.
Souissi N, Gauthier A, Sesboue B, Larue
J, Davenne D. Circadian rhythms in
two types of anaerobic cycle leg
exercise: force-velocity and 30-s
Wingate tests. Int J Sports Med 2004:
25: 14–19.
Stein RB, Gordon T, Shriver J.
Temperature dependence of mamma-
lian muscle contractions and ATPase
activities. Biophys J 1982: 40: 97–107.
Stephenson DG, Williams DA. Calcium-
activated force responses in fast- and
slow-twitch skinned muscle fibres of
the rat at different temperatures.
J Physiol Lond 1981: 317: 281–302.
Stephenson LA, Wenger CB, O’Donovan
BH, Nadel ER. Circadian rhythm in
sweating and cutaneous blood
flow. Am J Physiol 1984: 246:
R321–R324.
Strassman RJ, Qualls CR, Lisansky EJ,
Peake GT. Elevated rectal temperature
produced by all-night bright light is
reversed by melatonin infusion
in men. J Appl Physiol 1991: 71:
2178–2182.
Torii M, Nakayama H, Sasaki T.
Thermoregulation of exercising men in
the morning rise and evening fall
phases of internal temperature. Br J
Sports Med 1995: 29: 113–120.
Van den Heuvel CJ, Kennaway DJ,
Dawson D. Thermoregulatory and
soporific effects of very low dose
melatonin injection. Am J Physiol
1999: 276: E249–E254.
Walters TJ, Ryan KL, Tate LM, Mason
PA. Exercise in the heat is limited by a
critical internal temperature. J Appl
Physiol 2000: 89: 799–806.
Waterhouse J, Aizawa S, Nevill A,
Edwards B, Weinert D, Atkinson G,
Reilly T. Rectal temperature, distal
sweat rate, and forearm blood flow
following mild exercise at two phases of
the circadian cycle. Chronobiol Int
2007: 24: 63–85.
Waterhouse J, Weinert D, Minors D,
Atkinson G, Reilly T, Folkard S,
Owens D, Macdonald I, Sytnik N,
Tucker P. The effect of activity on the
waking temperature rhythm in
humans. Chronobiol Int 1999: 16:
343–357.
Waterhouse JM, Minors DS, Folkard S.
Estimating the endogenous component
of the circadian rhythm of the rectal
temperature in humans undergoing
normal sleep/activity schedules. J
Interdisciplin Cycle Res 1993: 24:
197–211.
Weinert D, Waterhouse J. Diurnally
changing effects of locomotor
activity on body temperature in
laboratory mice. Physiol Behav 1998:
63: 837–843.
Williams CS, Hill DW. The effect of time
of day on VO · ·
2 kinetics and VO2 max
during high-intensity exercise. Med Sci
Sports Exerc 1995: 27: S109.
89