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CIRCADIAN RHYTHMS DURING CYCLINGEXERCISE AND FINGER-TAPPING TASK

S. Moussay,1 F. Dosseville,1 A. Gauthier,1 J. Larue,1

B. Sesboue,2 and D. Davenne1,*

1Centre 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, France2Institut 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

1137

DOI: 10.1081/CBI-120015966 0742-0528 (Print); 1525-6073 (Online)Copyright q 2002 by Marcel Dekker, Inc. www.dekker.com

*Corresponding author. D. Davenne, UFR STAPS Caen, Universite de Caen Basse-Normandie,

Boulevard du Marechal Juin, 14032 Caen Cedex, France. E-mail: [email protected]

CHRONOBIOLOGY INTERNATIONAL

Vol. 19, No. 6, pp. 11371149, 2002

2002 Marcel Dekker, Inc. All rights reserved. This material may not be used or reproduced in any form without the express written permission of Marcel Dekker, Inc.

MARCEL DEKKER, INC. 270 MADISON AVENUE NEW YORK, NY 10016

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 spon-

taneous 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 90100 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)]

MOUSSAY ET AL.1138



significantly increased from 06:00 to 18:00h]. Similarly, for a light power output

(2030% 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 810 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-assessment questionnaire of Horne and Ostberg[34] which determines morning-

nesseveningness.

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.

CIRCADIAN RHYTHMS DURING CYCLING 1139



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 satcomfortably 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

MOUSSAY ET AL.1140



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., themean 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, referencedto 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, withsignificant time-of-day effect F4;36 3:9; p 0:031: Post hoc analysisindicated a significant difference between data collected at 06:00h and those




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 F4;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

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 cosinefunction curve ( ) is shown.

MOUSSAY ET AL.1142



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 F4;36 1:74; p 0:16:

Motor Spontaneous Tempo

There was a significant time-of-day effect on MST F4;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 F4;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.

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 Characteristics of Circadian Rhythms

F(dl) p w (h) A M

Temperature (8C) 3.09(4,36) 0.031 16:44 ^ 1:50 0.6 ^ 0.2 36.12 ^ 0.2Rest HR (beats min21) 7.1(4,36) 0.0003 15:19 ^ 1:52 6.7 ^ 1.2 53.9 ^ 1.2

MST (taps min21) 4.84(4,36) 0.0003 16:52 ^ 1:52 40.2 ^ 8.7 135.6 ^ 9.8

Pedal rate (rev min21) 2.82(4,36) 0.038 17:49 ^ 2:16 5.9 ^ 1.2 93.9 ^ 2.7




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 correlationwas 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 F3;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 Ilmarinenet 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 Wmax(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

MOUSSAY ET AL.1144



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 jet-

lag 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 (

(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




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