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Circadian rhythms during cycling exercise and

finger-tapping task
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CHRONOBIOLOGY INTERNATIONAL
Vol. 19, No. 6, pp. 11371149, 2002

CIRCADIAN RHYTHMS DURING CYCLING

EXERCISE AND FINGER-TAPPING TASK
S. Moussay,1 F. Dosseville,1 A. Gauthier,1 J. Larue,1
B. Sesboue,2 and D. Davenne1,*
1

Centre de Recherches en Activites Physiques et Sportives (CRAPS

UPRES EA 2131), UFR Sciences et Techniques des Activites Physiques
et Sportives, Universite de Caen Basse Normandie, Caen, France
2
Institut Regional de Medecine du Sport, CHU Caen Cote de Nacre,
Caen, France

ABSTRACT
The aim of this study was to follow the circadian fluctuation of the
spontaneous pedal rate and the motor spontaneous tempo (MST) in a sample
of highly trained cyclists. Ten subjects performed five test sessions at various
times of day. During each test session, subjects were required to perform (i) a
finger-tapping task, in order to set the MST and (ii) a submaximal exercise on
a cycle ergometer for 15 min at 50% of their Wmax. For this exercise, pedal rate
was freely chosen. Spontaneous pedal rate and heart rate (HR) were measured
continuously.
The results demonstrated a circadian variation for mean oral temperature,
HR, and MST. Under submaximal exercise conditions, HR showed no
significant time-of-day influence although spontaneous pedal rate changed
significantly throughout the day. Circadian rhythm of oral temperature and
pedal rate were strongly correlated. Moreover, a significant positive
correlation was found between MST and pedal rate. Both parameters may
be controlled by a common brain oscillator. MST, rest HR, and pedal rate
changes follow the rhythm of internal temperature, which is considered to be
*Corresponding author. D. Davenne, UFR STAPS Caen, Universite de Caen Basse-Normandie,
Boulevard du Marechal Juin, 14032 Caen Cedex, France. E-mail: davenne@staps.unicaen.fr
1137
DOI: 10.1081/CBI-120015966
Copyright q 2002 by Marcel Dekker, Inc.

0742-0528 (Print); 1525-6073 (Online)

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1138

MOUSSAY ET AL.

the major marker in chronobiology, therefore, if there is a relation between

MST and pedal rate, we cannot rule out partial dependence of both parameters
on body temperature. (Chronobiology International, 19(6), 11371149, 2002)
Key Words: Circadian changes; Cyclic activities; Cycling; Motor spontaneous tempo; Spontaneous pedal rate

INTRODUCTION
The natural or spontaneous cadence adopted in all cyclic activities (running,
walking, or swimming) shows fine temporal coordination. To account for the
required accuracy in time processing, some authors[1 6] have suggested the
existence of an internal clock sending periodic signals and serving as a time
reference. When freely chosen by the subject, these periodic signals occur at a given
frequency, which has been called personal tempo,[7 9] referent period,[10] or
motor spontaneous tempo (MST).[11 13] Thus, internal tempo or MST is
considered to be specific to each individual and may directly reflect a persons own
internal clock frequency. It may be described as a central pace-maker, used as a
parameter to organize different internal cognitive and behavioral processes.[8,14]
The origin of the MST could be genetic[15] or may result from a personal
construction, built upon neurophysiological bases.[16] The average MST period is
around 600 msec and is characterized by strong intra-individual stability.[11,17]
However, Oleron et al.[13] identified a diurnal variation in MST with a peak at
midday with low values observed early in the morning and late in the afternoon.
Kikkawa[18] confirmed the increase in MST from 07:00 to 13:00h and a continuous
decrease from 13:00 to 22:00h, however, these fluctuations did not reach a
significant level.
The influence of MST on preferred pedal rate selection in cycling has never
been studied. Cycling is a strongly stereotypical rhythmical movement, and
numerous studies have addressed the problem of how the spontaneous crank rate is
chosen. Some authors have highlighted the importance of the rating of perceived
exertion (RPE) drawn from central and peripheral origins. For instance, Pandolf and
Noble[19] demonstrated a parabolic relationship between RPE and cadence with the
lowest value recorded at 100 revolutions per minute (rev min21) at a power output
of 200 W. These results were confirmed by Marsh and Martin.[20] In other studies,
the minimal metabolic cost[21] or minimal muscular fatigue was taken into account.
Several studies[22 26] show that muscular moments produced and EMG activity
level (recorded throughout a complete pedaling cycle) are minimized for pedal rates
ranging from 90 to 100 rev min21. Since pedal force decreases as a function of pedal
rate, a frequency of 90 100 rev min21 would minimize the level of muscular
contraction produced, thus reducing muscular fatigue.[27]
In a recent study,[28] the crank rate freely chosen by cyclists to develop a
moderate power output [less than 60% of their maximal aerobic power (MAP)]

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CIRCADIAN RHYTHMS DURING CYCLING

1139

significantly increased from 06:00 to 18:00h]. Similarly, for a light power output
(20 30% of MAP), cyclists spontaneously choose a pedaling rate of 75 rev min21
at 06:00h and 95 rev min21 at 18:00h. These results may be compared to those of
Atkinson et al.,[29] which provided evidence for a time-of-day fluctuation in
working cadence in athletic subjects. Furthermore, these fluctuations were
correlated with the circadian rhythm of the core temperature.
In the study of Moussay et al.,[28] cyclists spontaneously decreased their
pedaling rate in the morning, forcing them to produce a higher pedal torque to
keep up with the imposed work-load. The maximal muscular peak torque observed
in the afternoon (17:55h)[30] is not consistent with the reduction of muscular strain
during the pedaling cycle demonstrated by Patterson and Moreno.[25] Muscular
strain would appear not to be the main factor in choosing a pedaling rate at a low
power output, other factors need to be explored. The influence of MST on
spontaneous pedaling rate adopted during cyclic activities is one of these.[2]
It seems an obvious link as a cyclists spontaneous pedaling rate will be
somewhere between 90 and 100 rev min21 depending on power output;[31 33] this
means a period ranging between 660 and 600 msec, which is similar to the average
MST period.[12] Hence MST could be an important factor for the spontaneous
pedaling rate. The aim of this study was to monitor the circadian variation of
pedaling rate and MST in a population of highly trained cyclists in order to detect a
possible relationship between MST and pedaling rate.

METHODS
Subjects
Ten male cyclists (age: 22.2 ^ 2.4 yr; body mass: 68.6 ^ 5 kg; height:
179.5 ^ 5.4 cm) volunteered to take part in this experiment. This study was
granted ethical approval by the ethics committee, CHU Cote de Nacre, Caen,
France. All subjects provided written informed consent after the study procedures
were explained in detail. All subjects had at least 3yr experience in competitive
cycling. Most trained for 8 10 h a week.
Subjects were selected from among 25 cyclists as either moderately
morning n 6 or neither type n 4; from their responses to the self stberg[34] which determines morningassessment questionnaire of Horne and O
ness eveningness.

Experimental Procedure
First, Wmax was assessed by a maximal continuous ramp test performed at
15:00h. The individual Wmax was later used to set the individual power output for
the submaximal steady state exercise.

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1140

MOUSSAY ET AL.

Two weeks later, oral temperature, heart rate (HR), pedal rate, and MST
were recorded during five test sessions carried out at the following times of day:
06:00, 10:00, 14:00, 18:00, and 22:00h. To avoid the effect of sleep deprivation,
no test session was carried out at 02:00h. Each subject performed five test sessions,
the timing of which was randomized. For each subject, there was only one session
per day with a minimal period of 24 h between each session. As proposed by
Baxter and Reilly,[35] subjects ate a meal 2 h before the tests, at 14:00 and 22:00h,
and they were woken up at 05:00h for the test at 06:00h. Before this particular test
they were allowed to drink a glass of water.
In addition, subjects were asked to adhere as closely as possible to their usual
sleeping habits with a minimum of 6 h (mean 7:30 ^ 0:45h) of sleep taken on the
night preceding each test. Subjects were not engaged in any fatiguing exercise
during the protocol. During the protocol, the laboratory temperature remained
constant (19 ^ 18C).
Data Collection
In the first part of the protocol, the cyclists performed a maximal continuous
ramp test; they rode on a cycle ergometer (Cateyew, CS-1000, Osaka, Japan) for
5 min at a work-rate of 150 W to warm-up. Then the work-rate was increased by
15 W min21 steps until exhaustion. Wmax was determined from the last step
completed by the subject.
During the second part of the protocol, each of the five test sessions was
performed on the same ergometer in order to detect a time-of-day effect on
temperature, HR, MST, and pedaling rate. Each test session started with a 20min
rest period during which subjects remained supine. At the end of this rest period,
HR (Polarw, X-trainer Plus, Kempele, Finland) and oral temperature (Omronw,
Wegalaon, Netherlands, accuracy: 0.058C) were recorded. Then, the subjects sat
comfortably on a chair; they were required to perform a finger-tapping task in
order to evaluate the MST. As described by Fraisse,[12] the subjects had to tap at
the most comfortable pace for 30 sec, and the only constraint was to keep the
pacing as regular as possible. During each session, this test was completed three
times, with a rest interval of 1 min, to check the reliability of data collection.
Finally, subjects performed a submaximal exercise on an electrical braked cycle
ergometer (Cateye). Exercise started with a 5min warm-up period at 70 W, after
which subjects were asked to develop 50% of their Wmax for 15 min. Throughout
the exercise, they were free to choose their own crank rate. Spontaneous pedaling
rate and HR were measured continuously (Polar) and averaged every 5 sec.
Data Analysis
For each test session, the most regular of the three 30sec trials (minimal SD)
was taken for analysis. Out of this 30sec period, only the last 20 sec were

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CIRCADIAN RHYTHMS DURING CYCLING

1141

considered for analysis since it takes a little under 10 sec to stabilize the
rhythm.[12] Instantaneous tap-to-tap frequency was computed and averaged over
the 20sec recording for each subject. Mean MST is reported as taps per minute
(taps min21).
During the submaximal test, pedaling rate and HR were continuously
recorded. Mean values were calculated from the 3rd to the 15th min, considering
that it took around 3 min to reach a stable state which was maintained until the end
of the exercise. All results are presented as mean ^ SD.

Statistical Analysis
In order to identify a circadian rhythm and demonstrate that the rhythm
amplitude differed from zero, each parameter was submitted to a one-way analysis
of variance (ANOVA) for repeated measures, with the factor time as a repeated
measure. A protected least significant difference (PLSD) Fischer test was selected
as a post hoc analysis. A least squares regression analysis, using the cosinor
method, was employed to determine the best-fit of a combined 24h period cosine
function[27] of the form Y ti M A cosvti w, where M is the mesor (i.e., the
mean level), A is the amplitude measured as half of the peak-to-trough variation,
and w is the acrophase (i.e., the time when the maximum level occurs, referenced
to local 00:00h). The existence of a sinusoidal circadian rhythm was confirmed or
rejected on the basis of the confidence interval for each parameter. Cosine
functions were compared using as reference values, samples of the 10 subjects
recorded during the five test sessions. Probability ( p ) values of less than 0.05 were
taken to indicate statistical significance.

RESULTS
Maximal Incremental Test
In the first part of the protocol, when the maximal continuous ramp test was
performed, subjects achieved a mean Wmax of 412 ^ 32 W. Maximal HR
measured at exhaustion was 189 ^ 6 beats min21.

Circadian Functions
Temperature
A significant circadian rhythm was found at rest for oral temperature
(Fig. 1A).
Values ranged from 35.7 ^ 0.18C at 06:00h to 36.3 ^ 0.18C at 14:00h, with
significant time-of-day effect F 4;36 3:9; p 0:031: Post hoc analysis
indicated a significant difference between data collected at 06:00h and those

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1142

MOUSSAY ET AL.

Figure 1. Diurnal rhythms of (A) oral temperature, (B) heart rate at rest, (C) MST, and (D) pedal
rate. Mean values (^ SD) are shown. Best fit curve between the experimental data () and the cosine
function curve ( ) is shown.

collected in the afternoon. From cosinor analysis, the 24h mean temperature
(mesor) was 36.1 ^ 0.28C, with a peak-to-trough amplitude of 0.6 ^ 0.28C,
representing a total fluctuation of 1.6% around the mesor. The highest value
(acrophase) was estimated at 16:44 ^ 1:50h (Table 1).

Heart Rate
Rest HR showed a marked circadian fluctuation F 4;36 7:10; p 0:0003
(Fig. 1B). The lowest value, recorded at 06:00h, was significantly different from
other values recorded during the day. Values ranged from 49 ^ 1.2 beats min21 at
06:00h to 58 ^ 1.3 beats min21 at 14:00h. The fitted cosine curve indicated a
circadian variation with an acrophase at 15:19 ^ 1:52h, a 24h mesor of
54 ^ 1.2 beats min21, and a peak-to-trough amplitude of 7 ^ 1.2 beats min21 or
12.43% of the mesor (Table 1). During submaximal exercise at 50% of Wmax, HR

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CIRCADIAN RHYTHMS DURING CYCLING

1143

Table 1. Results of ANOVA for Repeated Measures with the Factor Time and Presentation of the
Characteristics of Circadian Rhythms (w Acrophase, A Peak-to-Trough Amplitude, M Mesor)
ANOVA

Temperature (8C)
Rest HR (beats min21)
MST (taps min21)
Pedal rate (rev min21)

Characteristics of Circadian Rhythms

F(dl)

w (h)

3.09(4,36)
7.1(4,36)
4.84(4,36)
2.82(4,36)

0.031
0.0003
0.0003
0.038

16:44 ^ 1:50
15:19 ^ 1:52
16:52 ^ 1:52
17:49 ^ 2:16

0.6 ^ 0.2
6.7 ^ 1.2
40.2 ^ 8.7
5.9 ^ 1.2

36.12 ^ 0.2
53.9 ^ 1.2
135.6 ^ 9.8
93.9 ^ 2.7

values ranged from 134 ^ 4.6 beats min21 at 06:00h to 137 ^ 5.2 beats min21 at
14:00h but did not show any significant circadian variation F 4;36 1:74; p
0:16:

Motor Spontaneous Tempo

There was a significant time-of-day effect on MST F 4;36 4:8; p 0:003
(Fig. 1C). Post hoc analysis demonstrated a significant increase in MST from
06:00h (117.4 ^ 9.7 taps min21) to 18:00h (155.1 ^ 15.6 taps min21) and a
decrease in the evening from 18:00 to 22:00h (128.9 ^ 18.8 taps min21). The
mesor for finger-tapping, estimated from the cosine function, was 135.6 ^
9.8 taps min21 and tap-to-tap interval was 442 msec. Peak-to-trough amplitude
was 40.2 ^ 8.7 taps min21 (29.64%) of the mesor and the acrophase was
estimated at 16:52 ^ 1:52h (Table 1).

Pedal Rate
Pedal cadences spontaneously adopted by the cyclists showed a stable rate
during each of the test sessions after 3 min of adaptation at the start of the exercise.
The pedaling rate changed significantly at different times of day F 4;36 2:82;
p 0:038 (Fig. 1D).
Values ranged from 90.1 ^ 2.9 rev min21 at 06:00h to 96.5 ^ 2.8 rev min21
at 18:00h. The lowest frequency was recorded at 06:00h and differed significantly
from other values observed during the day. The mesor was 93.9 ^ 2.7 rev min21,
representing 638 msec for a complete revolution, the acrophase occurring at
17:49 ^ 2:16h. Cosinor analysis demonstrated an amplitude of 5.9 ^
1.2 rev min21 throughout the day (Table 1), representing 6.28% of the mesor
value.

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1144

MOUSSAY ET AL.

Correlation Analysis Between Circadian Rhythms

Temperature cosine variation was significantly correlated with pedaling rate
r 0:96; p 0:009 and rest HR r 0:9; p 0:037: A significant correlation
was also found between MST and pedaling rate r 0:87; p 0:048:
There were no significant differences in the acrophases of oral temperature,
HR, MST, or pedal rate F 3;21 0:89; p 0:46:

DISCUSSION
The peak and time course of the diurnal fluctuation in oral temperature are in
agreement with data in the literature.[36 38] The amplitude of the temperature
fluctuation (0.6 ^ 0.28C) is within the range of those measured by Ilmarinen
et al.[39] and Reilly and Down.[40] Such variations are classically observed in
physically active individuals.[29] The observed biphasic fluctuation characterized
by an asymmetry of the curve (a substantial rise between 06:00 and 10:00h, when
no significant variation was observed between 14:00 and 22:00h) is consistent with
the observations of Gauthier et al.[41]
Furthermore, the observed diurnal variation of HR at rest confirms a number
of studies showing a peak value ranging from 15:00 to 17:00h.[36,42 44] However,
the circadian rhythm of HR at rest was lost completely during submaximal
exercise. This seems to confirm the decrease of amplitude in circadian rhythms of
many physiological parameters such as heart rate, oxygen uptake, and ventilation,
during moderate or heavy submaximal exercise.[45]
As for MST, the mean MST period was 442.5 ^ 31.9 msec whereas the
literature indicates MST periods ranging from 380 to 880 msec with an average of
600 msec.[12] This discrepancy might be due to the fact that our subjects were
highly trained, as it has been reported that the level of physical fitness may affect
MST.[12,46]
High intra-individual stability from one day to another is commonly
reported.[12] However, only two studies have tested a time-of-day effect on MST:
one was a single case study,[13] and the other[18] suggested a trend but failed to
highlight any statistical difference. Our results clearly demonstrate a significant
circadian variation in MST, the large amplitude in MST [29.49%
(40.2 ^ 8.7 taps min21) of the mesor value] possibly explained by the subjects
physical fitness.[29,47]
The population was fairly homogenous and the mean W max
(412.5 ^ 32.5 W) is in line with those already observed in highly trained
cyclists.[48]
Pedaling rate measured throughout the day fluctuated between 90.1 ^
2.9 rev min21 at 06:00h and 96.5 ^ 2.8 rev min21 at 22:00h. The mesor value
(93.9 ^ 2.7 rev min21) is in line with that already observed (85100 rev min21) in
previous studies with expert cyclists.[31 33] This is consistent with a rate that

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CIRCADIAN RHYTHMS DURING CYCLING

1145

minimizes neuromuscular fatigue.[23] This fact might play an important role in

choosing the preferred pedaling rate for a cycling exercise.
Interestingly, the diurnal fluctuation in crank rate is not synchronized with
the circadian rhythm of muscular peak torque,[30] as cyclists chose a lower pedal
rate in the morning when muscular torque is at its lowest values. Thus, factors
other than just muscular force need to be taken into account in order to explain the
decrease in the morning pedaling rate.
One possible explanation is to assume a relationship between rhythmicity of
temperature and pedaling rate throughout the day r 0:96; p 0:009:
Circadian fluctuations in the core temperature may modify the musculo-skeletal
system:[49,29]
In the morning, when body temperature is minimal, articular suppleness is
reduced.[50,51] This may reduce the velocity of movements during a
submaximal cycling exercise.
There is a linear relationship between nerve conduction velocity and body
temperature.[52,53] Nerve conduction velocity increases throughout the
day[54] while the time to peak and the relaxation time decrease by 8.3
and 10.7%, respectively,[55] making for better muscle properties in the
evening. Neuromuscular efficiency changes synchronously with core
temperature fluctuation.[56]
An alternative explanation is that a reduced pedaling rate in the morning
could be of benefit to intra- and inter-muscular coordination requirements. Indeed,
Neptune et al.[57] pointed to the importance of the temporal characteristics in burst
onset/offset on muscles coordination. Another argument for a time-of-day
influence on movement coordination is given by Thor[58] who demonstrated a jetlag effect in the timing and coordination of movements.
The days biggest difference was observed between the 06:00h test session
and other experiment times. This effect could be an artifact since subjects were not
allowed to have a meal before this test session and were woken up before their
habitual waketime (<2 h earlier). Nevertheless, our results are consistent with
previous reports[29] showing a minimum in the temperature curve at 06:00h and a
regular increase in self-chosen work-rate during the day that exactly matches the
pedal rate rhythms observed in our study. In addition, MST is characterized by a
minimum at 06:00h which can be linked to fluctuations in arousal[29] and to
fluctuation in maximal tapping rate.[59]
Taken together, these observations make a pure meal or partial sleep
deprivation artifact unlikely.
Another possible artifact is that the minimum at 06:00h may be caused by no
measurement being taken at 02:00h. This can be ruled out however since Atkinson
et al.[29] showed the same minimum at 06:00h despite including a 02:00h
measurement.
Our study points to a significant correlation between pedaling rate and MST
variation during the day r 0:87; p 0:048: However, a pedaling revolution

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1146

MOUSSAY ET AL.

(638 ^ 18 msec) and a tap-to-tap interval (442 ^ 31.9 msec) are of different
durations. Obviously, the two movements differ in terms of motion amplitude and
applied force. Indeed, a moderate cycling exercise involves larger forces and
amplitudes of motion than those required for a finger-tapping task. However, the
maximal rate of reciprocal movements is not necessarily dependent upon the
inertial properties of the limb. The preferred frequency of intra-limb coordination
is subject to a relaxation oscillator arising from neuromuscular dynamics.[60] Thus,
the overt frequency might differ between effectors. Nevertheless, our results show
a similarity in pedaling and finger-tapping frequencies which could be linked to a
common central clock.
In conclusion, this study confirms that rest HR, MST, and spontaneous
pedaling rate during moderate exercise change during the day. Furthermore, MST,
rest HR, and pedal rate fluctuate synchronously with body temperature, which is
held to be a major marker in chronobiology, although there is no evidence for a
causal relationship between fluctuations. Hence, the significant correlation
between MST and pedal rate fluctuations during the day suggests that both
parameters may be controlled by a common brain oscillator such as the
suprachiasmatic nucleus.

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Received January 28, 2002

Returned for revision March 21, 2002
Accepted June 18, 2002