Distribucion de Potencia

Sports Med 2007; 37 (8): 647-667
0112-1642/07/0008-0647/$44.95/0
LEADING ARTICLE
2007 Adis Data Information BV. All rights reserved.
Distribution of Power Output

During Cycling
Impact and Mechanisms
Greg Atkinson,1 Oliver Peacock,2 Alan St Clair Gibson3 and Ross Tucker4
1
2
3
4
School of Sport and Exercise Sciences, Liverpool John Moores University, Liverpool, England
The School for Health, University of Bath, Bath, England
Psychology and Sport Sciences, Northumbria University, Newcastle upon Tyne, England
Department of Human Biology, MRC UCT Research Unit for Exercise Science and Sports
Medicine, University of Cape Town, Cape Town, South Africa
Abstract
We aim to summarise the impact and mechanisms of work-rate pacing during

individual cycling time trials (TTs). Unlike time-to-exhaustion tests, a TT provides an externally valid model for examining how an initial work rate is chosen
and maintained by an athlete during self-selected exercise.
The selection and distribution of work rate is one of many factors that influence
cycling speed. Mathematical models are available to predict the impact of factors
such as gradient and wind velocity on cycling speed, but only a few researchers
have examined the inter-relationships between these factors and work-rate distribution within a TT.
When environmental conditions are relatively stable (e.g. in a velodrome) and
the TT is >10 minutes, then an even distribution of work rate is optimal. For a
shorter TT (10 minutes), work rate should be increased during the starting effort
because this proportion of total race time is significant. For a very short TT (2
minutes), the starting effort should be maximal, since the time saved during the
starting phase is predicted to outweigh any time lost during the final metres
because of fatigue. A similar time saving rationale underpins the advice that
work rate should vary in parallel with any changes in gradient or wind speed
during a road TT. Increasing work rate in headwind and uphill sections, and vice
versa, decreases the variability in speed and, therefore, the total race time.
It seems that even experienced cyclists naturally select a supraoptimal work
rate at the start of a longer TT. Whether such a start can be blunted through
coaching or the monitoring of psychophysiological variables is unknown. Similarly, the extent to which cyclists can vary and monitor work rate during a TT is
unclear. There is evidence that sub-elite cyclists can vary work rate by 5% the
average for a TT lasting 2560 minutes, but such variability might be difficult
with high-performance cyclists whose average work rate during a TT is already
extremely high (>350 watts).
During a TT, pacing strategy is regulated in a complex anticipatory system that
monitors afferent feedback from various physiological systems, and then regulates the work rate so that potentially limiting changes do not occur before the
648
Atkinson et al.
endpoint of exercise is reached. It is critical that the endpoint of exercise is known

by the cyclist so that adjustments to exercise work rate can be made within the
context of an estimated finish time. Pacing strategies are thus the consequence of
complex regulation and serve a dual role: they are both the result of homeostatic
regulation by the brain, as well as being the means by which such regulation is
achieved.
The pacing strategy algorithm is sited in the brain and would need afferent
input from interoceptors, such as heart rate and respiratory rate, as well as
exteroceptors providing information on local environmental conditions. Such
inputs have been shown to induce activity in the thalamus, hypothalamus and the
parietal somatosensory cortex. Knowledge of time, modulated by the cerebellum,
basal ganglia and primary somatosensory cortex, would also input to the pacing
algorithm as would information stored in memory about previous similar exercise
bouts. How all this information is assimilated by the different regions of the brain
is not known at present.
The importance of an appropriate pacing strategy

during athletic competition has been appreciated for
thousands of years. Aesops classic story of the race
between the tortoise and the hare illustrates why it is
important to consider carefully the duration of the
impending race, select an appropriate pacing strategy and not underestimate the opposition. Nevertheless, the optimal distribution of work rate during
exercise has been studied scientifically only in recent years. For example, it was only in 2005 that the
first themed symposium on pacing strategy was
included on the programme of the annual conference
of the American College of Sports Medicine.
The lack of past research on pacing of selfselected work rate might be due, partly, to the limited availability of ambulatory and continuous measurement methods for physiological variables, such
as heart rate, oxygen consumption and body temperature. In the past, it was difficult to measure some
important physiological responses to a selected
work rate without the measurements being so invasive that they interfered with the work rate itself.
This caveat may have perpetuated the use of timeto-exhaustion (TTE) protocols, which involve exercising at a constant sub-maximal work rate until this
work rate cannot be maintained. Physiological variables can be measured periodically during TTE protocols without interfering significantly with the ability to maintain the stipulated work rate, providing
that the exercise is sub-maximal in intensity. However, such periodic measurements during a TTE test
may allow subjects to estimate elapsed time and this
information could be used to surpass (or match) any
previously recorded time.
It is clear that TTE protocols are not immune
from human reactivity effects, especially if cues are
provided unwittingly about elapsed time. Moreover,
TTE protocols are poor analogues of most sporting
contests[1] and, therefore, provide no information on
work-rate distribution. If one accepts that the definition and measurement of endurance performance
should be relevant to real athletic competitions (and,
indeed, to other aspects of the prolonged work of
humans in, for example, occupational contexts),
then work-rate distribution is a vital component of
endurance performance.
Recent developments in the direct measurement
of power output during actual cycling, as well as
new theories concerning the mechanisms of fatigue,
may have helped to raise the profile of pacing strategy research over the last 10 years. First, it is now
possible to record power output accurately and reliably[2] whilst a cyclist is riding his or her own bicycle
in a velodrome[3] or on the road. Second, following
some criticisms of the catastrophe theory of fatigue,[4,5] there has been an increased interest in
complex systems involving the brain for controlling
self-chosen exercise intensity.[6] The perception of
Sports Med 2007; 37 (8)
Pacing and Cycling
exercise duration and the selection of an appropriate

pacing strategy for this duration have been deemed
important inputs and outputs, respectively, of this
control system.
The developments in both the theoretical mechanisms and applied impact of pacing suggest that it is
important to bring together basic and real-world
research in cycling science. Relevant questions in
this respect are: what are the optimal pacing strategies for cycling competitions of different durations,
such as a 1km time trial (TT) in a velodrome or a
50km TT stage of the Tour de France? How is
pacing strategy in the latter type of races influenced
by inevitable variations in environmental conditions, such as gradient, wind direction and temperature? What are the psychophysiological mechanisms
that mediate a particular cyclists choice of pacing
strategy? What goes wrong in the brain and/or body
when an incorrect pacing strategy is adopted by a
cyclist?
The aim of this review is to consider the realworld impact, as well as the mechanisms, of different work-rate distributions during cycling. In the
first part of the review, we consider the various
factors that influence cycling performance in general. Second, we describe how some of these factors
might influence the optimal pacing strategy for various cycling competitions held in a velodrome and on
the road. Third, we outline the important links between pacing strategy and new theories concerning
the mechanisms of fatigue. Last, we discuss the
pychophysiological factors that might influence the
selection of a given power output during exercise.
1. A Model for Factors Affecting
Cycling Performance
Cycling is a complex sport whereby several
physiological, mechanical and environmental factors can interact to influence power and speed.
These various factors have been diagrammatically
modelled (figure 1) in a recent review by Atkinson
et al.[7] An important grounding factor related to
excellent cycling performance is the physiological
makeup of the athlete (e.g. maximal oxygen uptake
649
Overall race velocity

Bike design
Race-specific
retarding
forces
Pacing strategy
Rider position
Cycling power
Race-specific
training
strategies
Race-specific inherent
physiological ability
Race-specific
nutritional
strategies
Fig. 1. Various factors that influence cycling power output and

speed (reproduced from Atkinson et al.[7] with permission).
2max]). In the present review, it should also

[VO
become clear that inherent psychological factors
may also be important in affecting performance. In
this respect, we group the physiological and psychological traits with the term pyschophysiology.
Training and nutritional strategies are also human
factors that have a strong influence on overall race
power output. It is also important that the position of
the rider on the bicycle is analysed ergonomically
for maximal power output. Besides these human
factors, there are also several physical factors that
influence the degree to which a given power output
translates to cycling velocity. Because cycling
speeds can be high, physical resistive forces present
in the race environment (e.g. hills or wind) greatly
influence the relationship between cycling power
and overall race velocity. The presence of different
resistive forces in a race can also govern the choice
of bicycle design or its components. Besides pacing
strategy, there is one other factor that can influence
both cycling power output and the power-velocity
relationship: a poor rider position can lead to lower
velocities for a given power output (e.g. by not being
particularly aerodynamic) as well as being suboptimal in terms of power output itself (e.g. by the
saddle height being too high or low). The various
factors in figure 1 will now be discussed briefly to
illustrate the possible interactions between pacing
strategy and a myriad of other considerations.
Sports Med 2007; 37 (8)
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Atkinson et al.
1.1 Psychophysiological Ability
The inherent psychophysiological ability of the

cyclist encompasses an important set of attributes
that influence power generation and performance.
The major physiological variables that are correlat 2max, lactate
ed to cycling performance are VO
threshold and cycling efficiency. Certain oxygen
consumption kinetics and the degree of muscle
capillarisation may also be related to success in
cycling.
As cycling is essentially an endurance activity, it
is not surprising that large absolute and relative
2max in the region of 5.06.4 L/min and
values of VO
7085 mL/kg/min, respectively, have been recorded.[8,9] Top-class cycling performance also correlates with the work rate recorded at a certain physiological threshold, which is defined conventionally as
the point when there is a non-linear increase in a
physiological variable. Examples of such thresholds
are the lactate threshold[10] or the first and second
ventilatory threshold.[8] Despite the different labels,
the work rate recorded at these thresholds is generally a superior correlate of good performance than
2max.[8,11,12] Using heart rate to represent work
VO
rate, Lucia et al.[13] and Coyle et al.[12] showed that
elite riders can maintain exercise intensities >90%
2max for up to 60 minutes. Such exercise intensiVO
ties were particularly evident during TTs and mountainous stage races.
Gross mechanical efficiency (work done energy consumed 100) is also related to cycling performance and has been shown to range from 24.4%
to 28.8% in professional cyclists.[14] Lucia et al.[14]
suggested that differences in efficiency between
elite and non-elite cyclists became greater when
they exercised above the lactate threshold. Jeukendrup and Jentjens[15] calculated that a 1% improvement in efficiency for a 70kg cyclist, capable of
sustaining 400 watts (W) for 60 minutes, would
elicit a time saving of 48 seconds, but this improvement has not been confirmed with empirical data.
Coyle et al.[16] reported that cyclists with a
greater proportion of type 1 muscle fibres produced
more power and were more efficient. A high proportion of type 1 muscle fibres appears to be correlated
to the maintenance of high power outputs.[8,12] High

pedal frequencies (>90 rev/min) also appear to be
negatively related to type 2 fibre recruitment.[17]
Despite these reported correlations, little is known
about the muscle fibre types of elite cyclists, probably because of the invasiveness of muscle biopsies.
Recently, Lucia et al.[18] managed to describe the
frequency distribution of -actinin-3 genotypes
from the blood samples of 50 top-class cyclists. The
frequency of a premature stop codon polymorphism
(R577X) in this gene has been shown to be significantly higher in female endurance athletes.[19] However, Lucia et al.[18] found no difference in the
frequency distribution of R577X genotypes between
elite cyclists, runners and sedentary males, suggesting that a cyclist can be excellent, irrespective of
muscle fibre type.
It should be noted that, although there are correlations between some of the previously mentioned
physiological factors and cycling performance, the
causal nexus of these correlations might be elusive.
Maximal or submaximal increases (or decreases) in
some physiological responses to exercise may be
due to changes in other psychophysiological responses to exercise. This point will be returned to
when the impact of pacing strategy of human fatigue
mechanisms is discussed in a later section of this
review.
1.2 Training Strategies
Previous investigators have fully described the

underlying mechanisms and benefits of training to
athletic performance, e.g. in the study by Hawley
and Stepto.[20] Such researchers tend to cover the
key components of volume, intensity and frequency
of training.[21] Of particular interest are the recent
findings of Stepto et al.,[22] who examined the effects of a diverse range of different interval training
programmes on cycling TT performance. Six sessions of the shortest and most intense intervals (12
2max output with 4repetitions of 30 seconds at VO
to 5-minute recovery) were as effective as longer
intervals (8 repetitions of 240 seconds at 85%
2max with 1-minute recovery) at improving
VO
40km TT performance. An increased lactate transSports Med 2007; 37 (8)
Pacing and Cycling
port capacity is known to be one adaptation to highintensity sprint training that could be related to this
improvement in performance.[23] Repeated supramaximal sprints have since been recommended as
essential competitive preparation for all endurance
athletes.[20]
1.3 Nutritional Considerations
Cyclists in the Tour de France can expend up to

9000 kcal of energy per day.[24-26] As glycogen is
the primary energy substrate during such endurance
events, carbohydrate ingestion in the region of
710 g/kg/day has been deemed necessary for adequate cycling performance over such long distances.[27] The benefits of carbohydrate ingestion in
optimising glycogen stores and increasing performance in the week prior to competition,[28] 34 hours
before,[29] during[30] and after exercise,[31] are well
documented. For a more complete review see Hawley et al.[28] and Burke.[31]
Cyclists are also advised to consume fluids prior
to and during exercise, since performance could be
impaired by dehydration.[32] Furthermore, providing
fluid and carbohydrate together improves performance more than either fluid or carbohydrate provided
separately.[33] It should be noted that much of the
research work on rehydration during exercise has
been criticised.[6] One criticism is that researchers
have failed to adequately analogue the real environmental conditions that endurance cyclists might exercise in (e.g. the cooling effects of moving at a high
velocity through the air). Such criticisms serve,
again, to illustrate the importance of combining
basic and applied research work in the sport and
exercise sciences.
The ergogenic effects of caffeine on endurance
exercise are well known and have become particularly relevant since caffeine has recently been removed from the International Olympic Committee
list of banned substances. Kovacs et al.[34] examined
the effects of consuming different doses of caffeine
in a carbohydrate-electrolyte drink on 40km cycling
performance. A 5% increase in power was experienced by those athletes taking the highest caffeine
651
doses. A recent review paper by Graham[35] provides

an excellent summary of this literature.
New ergogenic aids will continue to be proposed
and researched scientifically (e.g. bovine colostrums).[36] Coverage of such supplements is beyond
the scope of this review. Nevertheless, it is worth
noting that much of the research on ergogenic aids
and hydration has involved TTE protocols, possibly
alongside the perception that within-subject variability in endurance performance due to pacing
should be minimised for accurate explorations into
human fatigue. Nevertheless, difficulties arise in
extrapolating the results of studies involving TTE
protocols to real-world endurance events, given that
within-subject variability in work-rate distribution is
an inherent characteristic of almost all such events.
It is possible that an ergogenic aid might be effective
in increasing the time that a constant submaximal
work rate is endured, but is ineffective in increasing
an already varying work rate for some or all of a real
cycling endurance TT.
1.4 Resistive Forces, Bike Design and
Rider Position
The major physical retarding forces during cycling are caused by air resistance and resistance due
to gravity. Rolling resistance and frictional losses
are thought to account for <10% of total energy
expenditure.[37] In brief, the major components of
rolling resistance are total load on the tyre, riding
surface, tyre diameter, construction and pressure.[37,38] Frictional losses are thought to occur from
both the chain drive system and in various roller
bearings, although these losses have been deemed
relatively trivial.[39]
Air resistance is the major retarding force during
cycling on flat terrain at speeds >15 km/h.[40] Air
resistance increases as an approximate squared
function of cycling velocity. Aerodynamic drag
comprises both pressure and friction drag, which
relate to the riders shape and frontal area, and surface smoothness, respectively.[7] The pressure drag
component creates a pressure vortex for a slipstreaming (drafting) cyclist who rides behind another cyclist, enabling up to a 30% reduction in power
Sports Med 2007; 37 (8)
652
Atkinson et al.
output.[41] The potential negative impact of air resistance on overall race velocity has influenced both
bicycle design and rider position. Such optimisation
of aerodynamics was clearly demonstrated when
Chris Boardman broke Tony Romingers world
hour record in 1996, despite Boardman weighing
6kg more and being estimated to average 20W less
power than Rominger.[42] Boardmans success was
helped by a revolutionary bicycle design that allowed adoption of a very aerodynamic superman
position, which has since been banned in cycling
events.[7]
Aerodynamic drag is also determined by relative
air movement and not just horizontal ground velocity. Cycling at 30 km/h into a 10 km/h headwind
requires a power output equivalent to riding at
40 km/h with no headwind. Indeed, Martin et al.[39]
reported that the relationship between wind velocity
and cycling speed is very nearly linear, although
these authors considered just one power output in
their calculation (255W). Fluctuations in air density
caused by changes in air temperature, barometric
pressure and humidity, may also impact upon relative power generation.[7]
The impact of gravity on cycling velocity is
determined largely by the mass of rider and bicycle.
Martin et al.[39] maintained that the relationship between gradient and cycling velocity is nearly linear,
in keeping with the effects of wind. When cycling
on the flat or downhill, the mass of rider and bicycle
is less influential or is even an advantage, since air
resistance relative to body mass is less for larger
than for smaller riders.[43]
1.5 Pacing Strategy: General Considerations
for Cycling
An appropriate distribution of energetic resources is considered vital to athletic performance,[44] particularly when success requires completing a given distance in the shortest possible time. To
achieve optimal performance, all available energy
stores should be used before finishing, without causing fatigue and a significant deceleration late in the
event. This optimisation requires an appropriate
pacing strategy, which has been defined by Atkin 2007 Adis Data Information BV. All rights reserved.
son and Brunskill[45] as the within-race distribution

of work rate (power output). Given the small margins between success and failure in top-level competition, appropriate pacing strategies or pacing mistakes may determine competitive outcomes. The
relative lack of empirical studies on how various
pacing strategies might influence performance is,
therefore, surprising.
Since the results of an early study by Robinson
et al.[46] on 1200m running, a popular assumption is
that an even distribution of effort is optimum. More
recently, researchers have identified a more complex relationship between distribution of pace and
subsequent performance in a diverse range of sports
that include: running,[47] cycling,[45] swimming[48]
and kayaking events.[49] New ideas about pacing are
particularly evident in the cycling literature, where
the significant influence of resistive forces (e.g. hills
and wind) on cycling velocity has prompted the
consideration of alternative pacing strategies.[7]
There are a variety of competitive events in cycling, including criteriums, road races, track events
and TTs. Criteriums and road races are mass start
events where various factors, such as bike handling
skills, race tactics, position of team mates, environmental conditions and drafting, influence performance. Therefore, road cyclists might enter the final
3060 minutes of a race having expended very different amounts of energy.[50] Palmer et al.[51] found
that trained road riders try to avoid stochastic energy
expenditure until the final 50km of a race. Nevertheless, few researchers have examined the work-rate
profiles during professional road races other than
anecdotal descriptions of individual rider power outputs. Moreover, difficulties in accounting for the
various other factors involved in bunch races and,
therefore, isolating the impact of different pacing
strategies on performances in these events has
prompted researchers to focus mainly on TT events.
Cycling TT events can be held on the track or road.
2. Optimal Pacing Strategy for
Track Cycling
Track cycling encompasses all events that take
place indoors or outdoors on a banked, hard surfaced
Sports Med 2007; 37 (8)
Pacing and Cycling
track (wood or cement) that typically has a circumference of 333m.[3] TT events staged on a cycling
velodrome include the 1km sprint or kilo, the 4km
pursuit and the hour record. These events are
naturally subject to fewer variations in the physical
characteristics of the environment (particularly
when the velodrome is enclosed). Therefore, gradient, wind speed, ambient temperature, barometric
pressure, altitude and condition of the riding surface
are all quite stable. The relations between physiological effort, power output and cycling velocity are
consequently relatively strong in such events.[7] This
enables pacing strategies to be imposed and monitored using split times during the TT. Moreover, a
TT that is held in the similar constant environment
of the laboratory is externally valid to track racing.
2.1 Laboratory Studies
In one of the first well designed experiments,

Foster et al.[52] predetermined the pace (very slow,
slow, even paced, fast or very fast) over the first
1km of a 2km cycling TT. The final 1km was
completed in as fast a possible time by the riders.
The best performances were recorded when the pacing strategy was quite even (i.e. first 1km = 50.9%
of the total time). Small variations, irrespective of
direction, from this strategy elicited negative consequences. No detectable changes in oxygen uptake
2), accumulated oxygen deficit or post-exercise
(VO
blood lactate concentration could account for the
pacing-related variation in performance. These findings are often used to support an even-paced strategy in longer cycling TTs on the track, such as the
hour record, which from a metabolic viewpoint,
requires the cyclist to perform in steady-state conditions at an extremely high percentage of their
2max.[53]
VO
In a follow-up study, Foster et al.[54] examined
the effects of different pacing strategies over 1500m
of speed skating and cycling. In contrast to previous
findings, the faster starting strategy led to better
performances. By incorporating data on the pattern
of energy expenditure during Wingate type tests,
computer-simulated models have been constructed
that predict track cycling performance with reasona 2007 Adis Data Information BV. All rights reserved.
653
ble success.[44] One factor that may be important in

such models is adequate simulation of the contribution of the standing start to overall race time.
2.2 Modelling the Influence of the
Standing Start
De Koning et al.[55] investigated the effect of

three pacing strategies (all-out effort, fast start
followed by an even pace and even pace for the
entire test) on modelled 1000m and 4000m track
cycling. The superior performance for the 1000m
simulation was obtained with an all-out strategy,
whilst a fast start followed by an even pace was
optimal for the 4000m simulation. These findings
were corroborated by Van Ingen Schenau et al.,[56]
as short-duration supramaximal exercise performance (<90 seconds) was found to require an all-out
strategy, whilst slightly longer events (<260
seconds) needed a fast start prior to a more constant
pace. The effectiveness of maximal rates of energy
release during the first few seconds of a short TT
was supported by Hirvonen et al.,[57] who observed
greater rates of creatine phosphate breakdown during this period in elite sprint runners. Bishop et al.[49]
reported an improved 2-minute kayak ergometer
performance after an all-out start strategy, which
2 kinetics.
appeared to be accompanied by faster VO
[44]
Foster et al.
also found a high initial power
output and anaerobic energy use, which reduced to
approximately 25% of peak power over the remainder of the 1500m event that was studied. In contrast
to the predictions using mathematical models,[55,56]
participants did not achieve a zero anaerobic energetic output at any point, and appeared to distribute
their energetic resources to provide for anaerobic
power output during the closing stages of the event.
Foster et al.[44] considered that these models failed to
incorporate data on how athletes spontaneously pace
themselves, when the target was to finish in the
quickest time. Atkinson et al.[7] also maintained that
further research is required in this area to elucidate
the exact balance between biophysical optimisation
of energy and physiological production of that energy.
Sports Med 2007; 37 (8)
654
Atkinson et al.
3. Optimal Pacing Strategy for

Road Cycling
A road TT can range from shorter prologue
distances (68km) to events lasting up to 24 hours.
A TT during elite stage races and in the Olympics is
typically 4070km in length. In such events, the
standing start is less influential to overall performance, since it represents a much smaller portion of
the total race time. Therefore, maximising the rate of
energy liberation at the onset of exercise is unnecessary and could be potentially damaging to power
output later in the race. Therefore, several researchers have attempted to examine how cyclists naturally pace the first few minutes of endurance TTs
simulated in the laboratory.
3.1 Laboratory Studies
Despite the lack of importance of the standing

start in a longer TT, there is evidence to suggest that
cyclists choose a power output at the beginning of
such events that is substantially greater than the
mean power for the race.[45,58,59] This fast starting
strategy is demonstrated in figure 2 for a group
(n = 8) of sub-elite cyclists[45] during a simulated
16.1km TT. Some improvements in TT performance
have been found when individual cyclists are
coached via sport science support work to blunt
this supraoptimal starting effort.[60] Such coaching
may be helped by the use of power measuring equipment, such as the SRM Powermeter (Schoberer Rad
400
Power (W)
350
300
250
Messtechnik, Julich, Germany). Alternative measurements, such as heart rate and ratings of perceived exertion,[61] were shown by Atkinson and
Brunskill[45] to be relatively insensitive to subtle but
important changes in power output, especially at the
start of an exercise period, although further research
is required to confirm these observations.
In support of the notion that an evenly paced
strategy should be adopted during a longer TT on the
road, as well as on the track, Palmer et al.[51] observed that Chris Boardman won an elite 80km TT,
whilst measurements of heart rate deviated by only 5
beats/min from the 178 beats/min average during the
race. Nevertheless, Swain[43] suggested that the variable effects of wind, gradient and other ambient
conditions on road TT performance may complicate
the choice of pacing strategy.
Fluctuations in gradient and wind almost always
occur during a road TT. It is extremely unlikely that
a TT course is completely flat, is in completely still
conditions and/or orientated so that there is a 90
crosswind from the right or left throughout the
whole race. Such fluctuations in wind direction and
gradient inevitably lead to variations in cycling
speed, even when the cyclist maintains a constant
power output. For example, Martin et al.[39] demonstrated, using a mathematical model, that a hypothetical cyclist who is able to sustain 255W would
travel at 8 m/s in a 5 m/s headwind compared with
14 m/s in a 5 m/s tailwind, and at 6 m/s during a 5%
ascent compared with 17 m/s during a 5% descent.
Since these effects of wind and gradient can be
modelled, it is possible to predict how variations in
power output, wind speed and gradient interact to
affect cycling speeds during a TT.
3.2 Modelling the Influence of Gradient and
Wind Speed
200
150
100
0
100
200
300 400 500 600

Work completed (kJ)
700
800
Fig. 2. Power output measured during a self-paced 800kJ time trial

for individual participants (n = 8) and the sample mean (). Participants tended to begin the trial with a power output that could not be
sustained, although power increased towards the end of the trial
(reproduced from Atkinson et al.,[62] with permission).
Several mathematical models of the power-speed

relationship have been presented in the literature.[39,63-67] These models have been based primarily on anthropometric, physiological and environmental parameters and have been used to (i) predict
cycling TT performance; (ii) identify individuals
who have the physiological requirements for sucSports Med 2007; 37 (8)
Pacing and Cycling
655
cess; (iii) quantify how changes in modelling parameters might affect performance; and (iv) quantify
other aspects of performance, such as the relative
contribution of energy from aerobic and anaerobic
sources.
During cycling, the velocity of the bike and rider
depends on the balance between the propulsive
forces applied to the pedals and the sum of all the
resistive forces acting on the bike. These factors
include some obvious (wind and rolling resistance)
and some less obvious (efficiency of the chain drive
system and inertia of the wheels) forms of resistance. These physical factors were accounted for by
Di Prampero et al.,[63] who used biomechanical data
to calculate the equation of motion of a cyclist.
Swain[43] utilised this equation to examine theoretical time savings by varying power on hills and wind.
2 in mL/min and velocity km/h, the
Modified for VO
equation of Di Prampero et al.[63] is (equation 1):
.
VO2 = 0.143 M s + 0.0108 A (PB T-1) (s + h)2 s
+ 31.2 M i s
(Eq. 1)
where M is the mass of the bicycle and rider in kg, s
is the cyclists speed in km/h, A is the riders total
surface area in m2, PB is barometric pressure in mm
Hg, T is ambient temperature in degrees Kelvin, h is
the headwind speed in km/h and i is the fractional
road incline. The first expression (0.143 M s) is
the oxygen cost of overcoming rolling resistance,
the second [0.0108 A (PB T-1) (s + h)2 s] is
the oxygen cost of overcoming air resistance, and
the third (31.2 M i s) is the oxygen cost of
overcoming gravity.[43]
Swain[43] assumed that the mass of rider and
bicycle was 80kg and that total surface area was
1.8m2, whilst barometric pressure was 760mm Hg
2,
and the ambient temperature was 293K. Net VO
hill grade and wind velocity were then entered into
the equation as independent variables to determine
subsequent bicycle speed. Swain[43] then investigated the effect of variations in road grade (15% to
15%), wind speed (24 to 24 km/h) and variations of
2 and, therefore, power (015%) on 10km and
VO
40km TT performance. The results showed that any
variation in power and, therefore, speed on a course

with no hills or wind would result in longer finish
times than would be achieved at a constant power.
These predictions agree with the conclusions of
Foster et al.[52] Swain[43] also demonstrated that on a
course with variable conditions of hills and wind, if
the rider maintains a constant power then these
environmental conditions elicit changes in cycling
speed that result in longer finish times than on a
completely flat course with no wind. Furthermore, if
a cyclist increases power in slower uphill or
headwind sections, and compensates for the total
work done by reducing power in faster downhill or
tailwind sections, faster times would be recorded
than if an even-paced strategy was adopted. If a
2 of 4.0 L/min varied power output
cyclist with a VO
by 10% in a 40km TT with variable conditions of
wind (16 km/h), the time to complete the race
would decrease by 30 seconds. Additionally, for the
same hypothetical cyclist and variable power strategy, a 10km TT with variable sections of uphill and
downhill (10%) would be completed with a time
saving of 90 seconds.
Kyle[65] developed a model of the energy demands of cycling based on biomechanical principles, but did not incorporate physiological terms
(i.e. energy supply). Olds et al.[66] provided additional detail by integrating expressions for both the
energy requirements of cycling (derived primarily
from data presented by Di Prampero et al.[63]) and
the energy that can be supplied by the cyclist. A
revised model also by Olds et al.[67] was shown to
account for 79% of the variation in TT performance.
The most recent model of cycling performance
was proposed by Martin et al.[39] and is based on
similar fundamental engineering and physical principles as investigated by previous authors.[63,65,67]
However, Martin et al.[39] were able to accurately
measure the power output and drag area of the
cyclist, and either measure or estimate all other
components in the model. Power was measured via
the recently developed SRM training system,[68]
which incorporates a combination of strain gauges
mounted within a deformable disc between the
crank arm and chain ring, and enables power, speed
Sports Med 2007; 37 (8)
656
Atkinson et al.
and pedalling rate data to be recorded in a small

handlebar-mounted computer. Aerodynamic parameters were calculated for each subject (riding a racing bicycle in TT position) using wind tunnel drag
data and were incorporated into the model. It was
calculated that >97% of the variation in cycling
power was accounted for by the following model
(equation 2):
ance, and ranked them in order of importance using

the model devised by Martin et al.[39] Hypothetical
2max of 48 mL/kg/min),
cyclists of novice (VO
trained (VO2max of 66 mL/kg/min) and elite

2max of 80 mL/kg/min) standard were modelled,
(VO
whilst the influence of different variables were expressed as changes in 40km TT performance. The
40km race was designed to reflect a standard
PTOT = {Va2VG (CDA + FW) + VGCRRmTgCos[Tan1(GR)] course experienced by time triallists. The course
included 5km sections completed in certain wind
+ VG (91 + 8.7VG)103 + VGGRmTgSin[Tan1(GR)]
conditions (either a 2 m/s headwind or tailwind) and
+ (mT + I/r2)(VGf2 VGi2)/(ti tf)}/EC
(Eq. 2) at certain road gradients (from flat to 1%). TT
performance was predicted to be improved by trainwhere Va is air velocity tangent to the direction of
ing (17 minutes), altitude training (2334 seconds),
travel of the cyclist and is dependent on both the
carbohydrate-electrolyte drinks (342 seconds), cafground velocity of the bicycle and rider, and wind
feine ingestion (5584 seconds) and aerodynamics
velocity; VG is the ground velocity of the bicycle
(912 minutes). It was particularly apparent that the
and rider; is air density; CD is the coefficient of
novice cyclists experienced greater time savings
drag; A is the frontal area of bicycle and rider; FW is
than elite cyclists, since they spent relatively longer
a factor associated with wheel rotation that equates
time on the course and, therefore, benefited from
to the incremental drag area of the spokes (CDA);
increased speed to a greater extent.
CRR is the coefficient of rolling resistance (includJeukendrup and Martin[50] did not consider pacing tyre and ground surface characteristics); mT is
the collective mass of bicycle and rider; g is the ing strategy as a potential performance-enhancing
[43]
acceleration due to gravity; GR is the road surface manipulation, despite the findings of Swain and
[45]
-3
some
observations
of
Atkinson
and
Brunskill.
gradient; VG (91 + 8.7VG)10 accounts for wheel
[39] to simuThese
latter
researchers
used
equation
2
bearing friction; I is the moment of inertia of the
wheels; r is the radius of the bike wheel; VGi is the late the effect of varying wind equivalent to changes
initial ground velocity; VGf is the final ground ve- in power over a 50km TT. In an initial TT, a hypolocity; ti is the initial time; tf is the final time; and EC thetical cyclist rode at 400W into a 10 m/s
headwind, followed by 25km with a 10 m/s tailwind.
accounts for chain efficiency factors.
Martin et al.[39] employed equation 2 to predict By increasing power output by 20W (5%) into the
the effects of road gradient, wind velocity, rolling headwind and decreasing power in the tailwind (to
resistance and drag area, which were all found to maintain an average power of 400W for the 50km
influence cycling velocity in an almost linear man- TT), a time saving of 63 seconds was predicted.
ner (R2 > 0.99) when cycling power was held conRecently, Atkinson et al.[69] replicated the restant. Over the range of values evaluated, rolling search of Swain,[43] using equation 2,[39] and comresistance and drag area were found to influence pared potential time savings with other performcycling velocity by 6%. Furthermore, when a rider ance-enhancing interventions by employing the
cycled at 255W, every 1% change in gradient altered methods described by Jeukendrup and Martin.[50] A
cycling velocity by approximately 1.24 m/s, whilst hypothetical cyclist (mass = 70kg) and bicycle
wind affected cycling velocity by 62% of a given (mass = 10kg) were studied under varying hypothetchange in wind velocity.
ical wind speeds (10 to 10 m/s), gradients (6 to
[50]
6%) and pacing strategies. Average rider power
A recent review by Jeukendrup and Martin
summarised the physiological, mechanical and envi- outputs of 164W, 289W and 394W were chosen to
ronmental factors that influence cycling perform- mirror the baseline performance times proposed by
Sports Med 2007; 37 (8)
Pacing and Cycling
657
previous researchers.[43,50] The three race scenarios

were courses with (i) 10% uphill and downhill sections every 1km for 10km; (ii) 4.4 m/s headwind and
tailwind sections every 5km for 40km[43] and (iii)
the 40km TT delimited by Jeukendrup and Martin.[50]
Varying an average power of 289W by 10%
over Swains[43] hilly and windy courses resulted in
time savings of 126 seconds and 51 seconds, respectively (figure 3). Time savings for most race scenarios were greater than those suggested by Swain.[43]
For the standard 40km TT modelled by Jeukendrup and Martin,[50] and the average power of 289W,
a time saving of 26 seconds was found with a power
variability of 10%. Greater time savings were found
for lower average rider power outputs.
Atkinson et al.[69] confirmed that time savings are
possible in cycling TTs if the rider varies power in
parallel with hill gradient and wind direction. With a
more recent mathematical model,[39] slightly larger
time savings were predicted than those reported by
Swain.[43] These time savings compared favourably
a
1800
1600
1400
Power variability 0%
1200
1000
800
Finish time (s)
600
400
0%
5%
10%
Gradient
b
3900
3800
3700
3600
3500
3400
3300
3200
0
2.2
4.4
Wind velocity (m/s)
6.6
Fig. 3. Times for completion of (a) a 10km time trial at 289W when
gradient and power are varied; and (b) a 40km time trial at 289W
when wind velocity and power are varied (reproduced from Atkinson et al.,[69] with permission).
to the predicted benefits of interventions such as

altitude training or ingestion of carbohydrate-electrolyte drinks.[50]
In summary, it is suggested that the optimal
pacing strategy for road TTs should involve (i)
caution at the start so that a supraoptimal power
output is not selected; and (ii) careful consideration
of the variable conditions of gradient and wind on
the day of the TT. When these conditions do not
vary, power output should also not vary. The effects
of variable gradient and wind can be attenuated by
parallel variations in power output. The greater the
parallel variation in power output, the less the cyclists overall time will be influenced by variable
gradient and wind. Therefore, the question emerges
as to what extent power output variations can be
tolerated by a cyclist during an actual TT.
3.3 Is a Variable Power Strategy
Physiologically Acceptable?
To confirm the benefits of adopting a variable

pacing strategy that are predicted by hypothetical
models, investigators must determine the physiological acceptability of power variations, superimposed
on the already high exercise intensities chosen during a TT. Swain[43] predicted that the more a rider
can vary power in parallel with changes in wind
direction or gradient, the closer the ideal state of
maintaining a constant speed is reached and, therefore, the greater the time saving. However, it would
be physiologically impossible to ride at a constant
speed in extremely variable conditions. For example, a rider would incur great physiological strain by
attempting to climb a steep hill at the same speed
that they can maintain when riding on the flat.
Furthermore, since cyclists can maintain exercise
2max for up to 60 minutes,[13] a
intensities >90% VO
15% increase in power could not be sustained for
long periods of time. Nevertheless, Swain[43]
demonstrated that even modest (5% above or below
a constant paced effort) variation in power would
result in significant time savings.
2 has been shown to rise gradually during
VO
constant load exercise that is performed above the
lactate threshold. This phenomenon is termed the
Sports Med 2007; 37 (8)
658
2 and is typically defined as

slow component of VO
the continual rise in VO2 beyond the third minute of

exercise.[70] To achieve optimal performance, cyclists must perform TTs in this physiological state,
2max.[71] Assuming
which occurs at 7090% VO
such high intensity exercise, an additional increase
in work rate causes lactate to accumulate in an
exponential manor and may be greater than that
which can be cleared following a reduced power.[72]
As suggested by Foster et al.,[52] a small increase in
power output during high-intensity exercise impacts
significantly on muscle lactate concentration. A reduction in muscle pH in unison with additional
proton accumulation may interfere with muscle contractile processes.[73] It has also been reported that an
increase in muscle lactate may compromise performance via the loss of technique that can accompany fatigue.[52] Furthermore, the progressive recruitment of less efficient motor units (specifically
type 2X fibres) at maximal workloads,[74] because of
the fatigue of fibres originally recruited,[70] is correlated to a rise in muscle lactate concentrations. An
additional increase in power output above a high
2 to rise in a
exercise intensity causes VO
monoexponential fashion (as opposed to a square
wave response), which would lead to an accumulat 2 declines in a
ed oxygen deficit.[72] Furthermore, VO
non-linear manner following a drop in power by
remaining elevated for a relatively larger time above
the steady state value.
Palmer et al.[75] suggested that a variable power
output detriments the physiological processes governing overall endurance performance. Participants
cycled for 150 minutes at a constant power (at 58%
of the peak power output attained in an incremental
test) or a variable power (58% 12.2% peak power
output). The authors then compared the effect that
these strategies had on subsequent performance of a
20km TT. All cyclists experienced a decrease in
performance (race time increased by 6%) in the
20km TT following the variable-paced effort. The
authors speculated that varying work rate may have
led to a greater depletion of muscle glycogen stores
than the constant-paced trial. However, since the
final 10 minutes of the variable trial was performed
Atkinson et al.
at 7080% of peak power output (30% higher than

in the constant trial), this factor alone may explain
the decrease in subsequent performance.
Palmer et al.[76] used a similar protocol to evaluate the metabolic responses associated with constant
and variable pacing strategies. Type 2 muscle fibres
were found to be more depleted of glycogen after the
variable trial, whereas type 1 muscle fibres were
more depleted after the constant trial. In this study,
subsequent TT performance was similar after the
variable and constant pacing strategies. With the
exception of skeletal muscle carbohydrate metabolism and fibre recruitment, cardiovascular and meta 2, energy expenditure and heart
bolic responses (VO
rate) were similar in both trials when controlled for
total work done.
Billat et al.[47] compared free versus constant
pace on performance and oxygen kinetics during
running. Runners performed four constant velocity
2max to
runs at 90%, 95%, 100% and 105% VO
determine TTE and the distance limit of each run.
They then performed the calculated distance limit
for each trial at a constant or variable pace. The
results showed that performance, oxygen kinetics
2 were not modified by a variable pace. Liedl
and VO
et al.[72] asked participants to perform an initial
2max), from
1-hour TT (corresponding to 78% VO
which the mean power was used to examine the
subjective and physiological strain of two further
trials. Cyclists adopted either a constant pace (at the
average work rate) or a variable strategy of alternating 5-minute segments of riding 5% above and
below the average work rate. In agreement with
Billat et al.,[47] these researchers found no differ 2, blood lactate concentration,
ences in mean VO
mean heart rate and mean ratings of perceived exertion (RPE) between the constant and variable power
trials. Liedl et al.[72] also reported the distribution of
physiological responses within an exercise bout.
Three of the sections in which power was 5% higher
than the mean led to observations of greater blood
lactate in comparison to equivalent periods during
the constant trial, whilst a contrasting pattern was
2.
reported for VO
Sports Med 2007; 37 (8)
Pacing and Cycling
when travelling uphill (mean duration of each

climb = 714 seconds) and decreasing power in the
downhill sections (mean duration of each descent =
190 seconds), so that overall mean power was
equivalent to that in (i). All participants maintained
this variable power strategy during the first half of
the TT, but two riders could not adhere to the power
variations during the final 400kJ (figure 4). Nevertheless, mean standard deviation finish time for
the variable power trial (3670 589 seconds) was
significantly faster than that for the constant power
TT (3758 645 seconds), the 95% confidence interval for the percentage improvement being 0.44.3%.
Heart rate and lactate responses were highest in the
initial self-paced TT and did not differ between the
subsequent constant and variable power trials. Ratings of perceived exertion were also similar between
trials. In this externally valid TT, Atkinson et al.[62]
concluded that some cyclists cannot fully adhere to a
pacing strategy involving an approximate 5% variation in mean power in parallel with gradient variation. Nevertheless, an important time saving can still
result even if a variable pacing strategy is only
partially adopted during a hilly TT, so that no additional physiological strain is incurred.
4. Pacing Strategy and Mechanisms
of Fatigue
As described in section 3.1, an initial pacing
strategy is selected by the cyclist. This selection
Power (W)
Liedl et al.[72] varied power output according to

time elapsed rather than distance covered. The latter
scenario is more representative of an actual TT
where gradient and wind speed vary over given
distances. The findings of Liedl et al.[72] support the
physiological acceptability of a variable power strategy as proposed by Swain,[43] but did not demonstrate actual performance improvements relative to
cycling TTs because of the nature of the protocol.
Atkinson and Brunskill[45] confirmed the timesaving hypothesis proposed by Swain[43] using a
more applicable strategy than that adopted by Liedl
et al.[72] A Computrainer 1 cycle ergometer was
used to simulate a 16.1km flat TT course with an
8.05 km/h headwind in the first half of the race and
an 8.05 km/h tailwind over the remainder of the
race. Participants first rode a self-paced TT to ascertain mean power output. Cyclists then adopted either
a constant strategy at the mean power output or a
variable strategy where power output was 5% above
the mean in the headwind section and 5% below the
mean in the tailwind section. Time to complete the
variable trial was 2 seconds faster than the constant
trial, which was 10 seconds faster than the initial
self-paced trial. Additionally, the overall mean RPE
and lactate were lowest in the variable trial, whilst
there was no difference in mean heart rate between
trials. Therefore, a power variability of 5% was
physiologically acceptable and elicited small yet
significant time savings when varied in parallel with
wind direction.
Recently, Atkinson et al.[62] investigated the acceptability of power variation during a cycling TT
with simulated uphill and downhill sections. Seven
cyclists first completed a 800kJ self-paced TT on a
simulated flat course. A 800kJ TT course with four
sections of uphill/downhill was then modelled. Each
section involved 100kJ of cycling up a simulated
gradient of 5%, followed by 100kJ of riding down a
simulated gradient of 5%. Participants were required to complete this simulated course using two
pacing strategies: (i) at a constant power equivalent
to the mean power achieved during the initial TT;
and (ii) increasing power by 5% of mean power
659
260
250
240
230
220
210
200
190
Constant
Variable
Up Down Up Down Up Down Up Down

0
100 200 300 400 500 600 700 800

Work completed (kJ)
Fig. 4. Mean power output measured during a simulated hilly time

trial whilst adopting either a constant or variable power strategy. All
participants were able to adhere to the variable pacing strategy only
for the first half of the time trial. There was also deviation from the
stipulated constant power strategy for one participant towards the
end of the trial (reproduced from Atkinson et al.,[62] with permission).
The use of trade names is for product identification purposes only and does not imply endorsement.
Sports Med 2007; 37 (8)
660
Atkinson et al.
process obviously involves the brain, which might

use some mechanism that responds to environmental conditions and internal metabolic reserve capacity, metabolic rate and core temperature at the start of
exercise. Once exercise has commenced, the selection of power output could be based on the cyclists
interpretation of the physiological changes occurring during exercise, together with prior experience
and knowledge of the duration or distance of exercise remaining. In contrast to this intuitive intelligent system[77-79] explanation for the regulation of
pacing strategies (discussed in detail in the next
section), a commonly held theory is that fatigue and
a resultant decrease in power output develops only
after one or more physiological system is stressed
beyond its maximum capacity, leading to a failure to
maintain force. Support for this theory has mostly
come from studies in which the exercise work rate is
fixed, thus forcing athletes to maintain a given work
rate until they are no longer able to sustain the
required force output.[80] These studies clearly do
not make allowances for pacing strategies, and thus
fail to explain adequately what occurs in all forms of
freely chosen exercise. That is, when exercise work
rate is self-selected, there is evidence to suggest that
the pacing strategy, or the selection of work rate, is
regulated specifically to ensure that factors that are
classically implicated as causing fatigue are instead
regulated so that they do not adversely affect physiological variables before the known endpoint of
exercise is reached. An example is provided in section 4.1 for exercise in hot conditions.
4.1 Fatigue During Constant Work-Rate
Exercise in the Heat
The role of the brain in the regulation of pacing

strategy is perhaps best illustrated during exercise in
hot and humid conditions. It has long been known
that when exercise is performed at a constant work
rate to exhaustion in the heat, volitional fatigue
occurs at a body temperature of 40C, irrespective
of the state of prior heat acclimation,[81] manipulation of pre-exercise body temperature[82] and the rate
of heat storage.[82] Based on the observation that

humans became dizzy and were unable to move
their legs at the point of exhaustion, it was proposed
that fatigue during exercise in the heat is associated
with a critical core temperature limiting exercise
performance,[82] in which a high body temperature
directly affects central nervous function.[83,84]
Recent novel studies of the effects of heat stress
on central drive, muscle function and brain activity
support the hypothesis that skeletal muscle motor
unit recruitment and central activation are decreased
after body temperatures reach critical values. It has
been shown that in subjects with hyperthermic temperatures (40C), voluntary activation percentage,
measured by superimposing electrical stimulation
on a maximal voluntary contraction, was significantly lower than subjects showing normothermic
body temperatures (38C).[78] Furthermore, the
same pattern was evident in handgrip contractions,
indicating that central activation of muscle is independent of whether the muscle was active or not. It
was concluded from this study that hyperthermia
causes a form of central fatigue, with reduced
central activation that leads to a lower force production.[84]
Studies of brain function in the heat also implicate the brain as being responsible for the failure to
produce force. For example, Todd et al.[85] found
that reduced force output during hyperthermia occurred because of a failure of voluntary drive during
passive heating despite the availability of additional
motor cortical output, which would, in theory, allow
increased force output. Thus, according to this
model, a failure of body temperature regulation
causes critical brain and/or core temperatures to be
reached. Central fatigue then occurs, as the hot
brain is no longer able to recruit a sufficient number
of motor units to sustain the previous or expected
power output, even in the presence of a cortical
reserve. This model is thus catastrophic,[78,79] since
fatigue is the inevitable consequence of a failure of
the thermoregulatory system to maintain an optimal
body temperature.
Sports Med 2007; 37 (8)
Pacing and Cycling
4.2 Fatigue During Self-Paced Exercise in

the Heat
In contrast to those studies in which the exercise

work rate is fixed and exercise continues until volitional fatigue, self-paced exercise bouts are
characterised by an ability to alter pace in response
to various external and internal physiological cues
and signals. The pacing strategy, as defined earlier,
is the optimal allocation of physiological resources
for an exercise task of known duration. Under these
conditions, a more complex pattern of regulation
emerges. For example, Tatterson et al.[86] showed a
reduction in power output after only 15 minutes of a
30-minute self-paced cycling TT in the heat, even
though core temperatures rose at similar rates in a
hot and temperate environment. They postulated
that the brain was sensitive to the rate of increase in
arterial blood temperature and selected a power output relative to the rate of rise in core temperature.
However, they did not measure any index of brain
function to confirm this.
Subsequently, Marino et al.[87] found that lighter
runners outperformed heavier runners during a selfpaced 8km TT that followed 30 minutes of running
at 70% peak treadmill speed in hot (35C) conditions. A significant correlation was also found between the rate of heat storage and body mass during
the TT, suggesting that the lighter runners stored
less heat at the same running speed. It was concluded that this reduced rate of heat storage allowed the
lighter runners to run faster before reaching a limiting rectal temperature.[87] This suggests that the rate
of heat storage may contribute to the afferent input
responsible for a reduction in work rate in the heat.
Marino et al.[88] have also shown that African
runners, who have a lower rate of heat storage at a
given running speed than Caucasian runners, were
able to outperform the Caucasian runners in hot
(35C) but not in cool (15C) conditions during an
8km TT. The difference in running speed between
the groups in the heat was present from the onset of
the TT, despite rectal temperatures that were only
moderately elevated (38C) and not different between groups. It was suggested that the early reduction in running speed in the heat occurs because of
661
an anticipatory exercise response that would control the exercise work rate by regulating the number
of motor units that are recruited or derecruited during prolonged exercise in the heat.[88]
Finally, Tucker et al.[89] have recently shown that
during self-paced cycling exercise in the heat, power
output and electromyographic (iEMG) activity (an
indirect measure of muscle activation) decrease well
before the core temperature reaches 40C. Further,
the power output and iEMG activity were greatest in
the final kilometre of a 20km cycling TT, when the
rectal temperatures were also the highest. A catastrophic model for fatigue cannot explain this observation, for it holds that the ability of the brain to
recruit motor units to produce a given force deteriorates at higher body temperatures. Thus, no allowance is made for the brain to reduce the level of
motor unit recruitment before the body temperature
reaches critical levels, and for the motor unit recruitment to increase when body temperature is higher
than before.
Collectively, these studies suggest that when
force output or exercise work rate are self-selected
rather than being fixed, an anticipatory mechanism[86,89-92] adjusts the work rate by regulating the
degree of motor unit recruitment to prevent body
temperature from rising to levels that may cause
harm or premature fatigue.[88] The brain thus acts
pre-emptively to ensure that a catastrophic failure of
thermoregulation does not occur. Such regulation
represents homeostatic control, the goal of which
would be to prevent an abnormal rise in body temperature by regulating the rate of heat production.[88,89]
4.3 Homeostatic Control in Other Conditions
Similar regulation may occur under various conditions, suggesting that the brain monitors afferent
inputs detailing information on the oxygen content
of the inspired air,[93-95] energy substrate availability[96,97] and, crucially, the anticipated duration of the
exercise bout.[58,98] This final factor, discussed in the
previous section, is essential for optimal pacing,
since any anticipatory calculation cannot be made
unless duration of exercise is known with some
Sports Med 2007; 37 (8)
662
accuracy. That is, if the adjustments in pacing strategy serve to prevent harmful or limiting disturbances
to homeostasis before the end of exercise, as is
described in this review, then the expected duration
of exercise would serve as the anchor point against
which this regulation would occur.
In summary, pacing strategy is regulated in a
complex anticipatory system that monitors afferent
feedback from various physiological systems, and
then regulates the work rate so that potentially limiting changes in physiological systems do not occur
before the endpoint of exercise is reached. It is
critical that the endpoint of exercise be known, so
that adjustments to exercise work rate can be made
within the context of a known duration of exercise.
Pacing strategies are thus the consequence of complex regulation and serve a dual role: they are both
the result of homeostatic regulation by the brain, as
well as being the means by which such regulation is
achieved.
5. How Does the Brain Control Pace?
Whatever pacing strategy is chosen or is optimal
for a particular exercise bout, the arguments outlined in the previous section suggest that initiation
and modulation of the chosen pacing strategy is
regulated by control processes in the brain. In order
for the brain to choose a particular pacing strategy
for a particular exercise bout, knowledge of the
distance or time to be completed during the exercise
bout is perhaps the critical factor used to make this
choice.[99,100] Memory of prior events of similar
duration or time stored in the brains neuronal circuits will optimise the choice of strategy chosen by
the brain, as will awareness of the external environmental conditions and internal metabolic reserve
capacity, metabolic rate and core temperature at the
start of the exercise bout.[78,100,101] Once the exercise
bout begins, these factors will be used by a pacing
algorithm in the brain to calculate alteration in power output throughout the exercise bout, which result
from unexpected changes in either the external environment or internal physiological milieu, in order to
maintain or alter the initially chosen pacing strate 2007 Adis Data Information BV. All rights reserved.
Atkinson et al.
gy.[102] In order for the pacing algorithm in the brain

to perform these corrections to the power output
appropriately, knowledge of time past or distance
covered is required and, therefore, a timing mechanism is essential in order for the brain to be able to
regulate the power output so that the distance or time
still to be covered to reach the endpoint is completed
at an optimal pace to perform the exercise bout as
fast as possible, while ensuring homeostasis is maintained and catastrophic failure does not occur in any
physiological system during the exercise bout.[103]
5.1 The Controlling Algorithm
In order for all these different functions associated with initiating and modulating a particular pacing
strategy to be performed by the pacing algorithm in
the brain, it is clear that a large number of different
brain systems are required to be active and synchronised throughout an exercise bout. Afferent input from interoceptors, which monitor physiological
activity such as heart rate, respiratory rate, metabolic rate, fuel reserve levels and core temperature,
have been shown to induce activity in the thalamus,
hypothalamus, insula and cingulate cortical regions
during exercise.[104-106] Afferent input from exteroceptors, which measure external environmental conditions, have been shown to activate the parietal
somatosensory cortex.[104,107] Memory formation
and decision making involving sequences of prior
exercise bouts, linked by their common events and
places, occur in the hippocampus, together with the
parahippocampus and surrounding areas of temporal
cortex, and the prefrontal cortex.[108-110] Motivation
and emotional state are also important in generating
and maintaining a desired goal and associated pacing strategy during exercise, and emotional responses are thought to be triggered in the amygdala,
limbic system, orbitofrontal and anterior cingulate
cortex.[104,111] Once a decision has been made by
the brain, the output to the muscles altering or
maintaining a force output originates in the motor
(M1) and premotor supplementary motor area
(SMA) cortex.[112,113]
Sports Med 2007; 37 (8)
Pacing and Cycling
5.2 The Importance of Time Judgement
As previously suggested, the capacity of the brain

for assessing time is thought to be essential for the
overall control of pacing strategy, and the integration of these different information sources in the
different brain regions may be associated with structures and functions in the brain that assess and
generate knowledge of the passing of time. Although circadian rhythms have been shown to be
controlled by neural circuitry in the suprachiasmatic
nuclei in the hypothalamus,[114] lesion and electrophysiological studies have suggested that interval
timing in the millisecond interval by neural circuits
in the cerebellum, and in the second to minute range,
is controlled by the activity of corticostriatal circuits
involving the basal ganglia, M1 and SMA motor
cortex, and primary somatosensory cortex.[114]
A number of theories have been suggested regarding how cortical circuitry is involved in monitoring this timing information. In the pacemakeraccumulator model, pulses are emitted and stored in
an accumulatory manner by pacemaker neurons in
the basal ganglia, and are stored in a reference
memory.[114,115] During an exercise bout, the number
of pulses stored in the reference memory is compared with the number of pulses created by the
current exercise bout, which is stored in active
memory. In contrast, in the striatal beat frequency
model, basal ganglia neurons monitor the synchronous activity of oscillatory cortical neurons that fire
during an activity such as an exercise bout and
which project on to them, and continuously compare
the current pattern of oscillations with the patterns
detected during previous activity.[114,116] In the
striatal beat frequency model, dopaminergic neurons
from the substantia nigra and ventral tegmental area
fire prominently at the start and end of a trial, and
are thought to act as a start gun for the detection of
the onset of measurement of the synchronous activity of the cortical oscillatory neurons.[117]
Whichever of these or other theories is correct,
the timing control mechanisms necessary to control
pacing during exercise are even more complex, as
these suggested timing mechanisms are located in
internal brain structures and need to be compared
663
with peripheral and environmental factors or cues

that will act as a comparative frame of reference to
indicate how much distance has passed during an
event, in order to calculate the duration and time
required to reach the endpoint. Environmental and
peripheral physiological factors that may be involved in this comparison are: (i) any variable that
displays metronomic type activity, such as heart
beat, respiratory rate or stride frequency and length;
or (ii) visual flow cues generated by the continuously changing surrounding environment that are associated with forward movement of the athlete. How
these external timing cues interact with the internal
timing neural circuitry is not presently known.
Therefore, it is clear that that a large number of
brain regions are required to produce all the factors
involved in the brains decision-making processes
during exercise to control pacing strategy. How all
the information generated in these different brain
regions are assimilated by the algorithm that calculates power output and regulates the pacing strategy,
and where this pacing algorithm is found in the
brain, is not known at this point in time. It is difficult
to suggest that neural circuits in a specific brain
region can generate the level of intellectual processing required to perform the computational tasks
required by this pacing algorithm, and it is more
likely that brain processes involving large-scale synchronisation of oscillatory electrical and neurotransmitter activity in the neural circuitry of multiple
brain areas control the temporal sequencing of power output involved in pacing during exercise. Further
research is required by the exercise science community to elaborate on the nature of these brain pacing
algorithmic processes, and how conscious awareness of these pacing algorithms and strategies
emerge from the brain structures during exercise.
6. Conclusion
In this review, we have discussed the applied and
mechanistic issues surrounding pacing of work rate
during cycling exercise. Cycling was chosen since
most of the relevant studies have involved this mode
of exercise and because a cycling TT involves individual athletes competing solely against the clock.
Sports Med 2007; 37 (8)
664
Atkinson et al.
Therefore, cycling provides an appropriate model

for studying the impact and underlying mechanisms
of the pacing of work rate whilst humans exercise.
For most real-world situations in which humans
need to work for prolonged periods of time, a TT is
more externally valid than a TTE protocol.
An even distribution of power output is physiologically optimum but only when the various factors
governing the relationship between cycling power
and speed are stable. For short events with a standing start of longer events in which gradient and wind
velocity vary, a variable power strategy in parallel
with the different changeable phases of the competition is optimal. Nevertheless, little is known about
whether the variable power strategies that are predicted to save time can be tolerated by elite cyclists.
Cyclists seem to select a supraoptimal power
output at the start of a longer TT. More research is
needed to examine if pacing interventions are affective and can be successfully adopted by competitive
cyclists in the long term. Such interventions may
well alter the pacing algorithm in the brain, but
little is known about which input signals are most
significant and how, exactly, such an algorithm
functions. Without the recent in-depth consideration
of externally valid TTs rather than TTE tests, such
alternative mechanisms for human fatigue would
not have emerged. Nevertheless, more research
work is needed to elucidate fully the impact and
causes of a poorly selected work-rate distribution
during cycling.
Acknowledgements
No sources of funding were used to assist in the preparation of this article. The authors have no conflicts of interest
that are directly relevant to the content of this article.
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Correspondence: Dr Greg Atkinson, Research Institute for

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E-mail: G.Atkinson@ljmu.ac.uk
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Sports Med 2007; 37 (8): 647-667

2007 Adis Data Information BV. All rights reserved.

Distribution of Power Output

We aim to summarise the impact and mechanisms of work-rate pacing during

endpoint of exercise is reached. It is critical that the endpoint of exercise is known

The importance of an appropriate pacing strategy

Pacing and Cycling

exercise duration and the selection of an appropriate

Overall race velocity

Fig. 1. Various factors that influence cycling power output and

2max]). In the present review, it should also

1.1 Psychophysiological Ability

The inherent psychophysiological ability of the

to the maintenance of high power outputs.[8,12] High

Previous investigators have fully described the

Pacing and Cycling

Cyclists in the Tour de France can expend up to

doses. A recent review paper by Graham[35] provides

son and Brunskill[45] as the within-race distribution

Pacing and Cycling

In one of the first well designed experiments,

ble success.[44] One factor that may be important in

De Koning et al.[55] investigated the effect of

3. Optimal Pacing Strategy for

Despite the lack of importance of the standing

300 400 500 600

Fig. 2. Power output measured during a self-paced 800kJ time trial

2007 Adis Data Information BV. All rights reserved.

Several mathematical models of the power-speed

Pacing and Cycling

variation in power and, therefore, speed on a course

and pedalling rate data to be recorded in a small

ance, and ranked them in order of importance using

trained (VO2max of 66 mL/kg/min) and elite

Sports Med 2007; 37 (8)

Pacing and Cycling

previous researchers.[43,50] The three race scenarios

2007 Adis Data Information BV. All rights reserved.

to the predicted benefits of interventions such as

To confirm the benefits of adopting a variable

2 and is typically defined as

the continual rise in VO2 beyond the third minute of

at 7080% of peak power output (30% higher than

Pacing and Cycling

when travelling uphill (mean duration of each

Liedl et al.[72] varied power output according to

Up Down Up Down Up Down Up Down

100 200 300 400 500 600 700 800

Fig. 4. Mean power output measured during a simulated hilly time

2007 Adis Data Information BV. All rights reserved.

Sports Med 2007; 37 (8)

process obviously involves the brain, which might

The role of the brain in the regulation of pacing

of heat storage.[82] Based on the observation that

Pacing and Cycling

4.2 Fatigue During Self-Paced Exercise in

In contrast to those studies in which the exercise

gy.[102] In order for the pacing algorithm in the brain

Pacing and Cycling

5.2 The Importance of Time Judgement

As previously suggested, the capacity of the brain

with peripheral and environmental factors or cues

Therefore, cycling provides an appropriate model