The Bioenergetics of World Class Cycling

Asker E Jeukendrup 1, Nell P Craig 2 & John A Hawley 3

1Department of Sport and ExerciseSciences,The Universityof Birmingham, Edgbaston,
Birmingham, UK. 2AustralianCycling Federation & Australian Institute of Sport (Track Cycling
Unit), Henley Beach, South Australia. 3ExerciseMetabolism Group, Department of Human
Biology and Movement Science, RMtT University, Bundoora, Victoria, Australlia.

Jeukendrup, A.E., Craig, N.P., & Hawley, J.A. (2000). The Bioenergetics of Worl Class Cycling.
Journal of Science and Medicine in Sport 3 (4): 414-433.
Professional cycle racing is one of the most demanding of all sports combining
extremes of exercise duration, intensity and frequency. Riders are required to
perform on a variety of surfaces (track, road, cross-country, mountain), terrains
(level, uphill and downhill) and race situations (criterions, sprints, time trials, mass-
start road races) in events ranging in duration from 10 s to 3 wk stage races covenng
200 m to 4,000 km. Fm~hermore, professional road cyclists typically have ~100 race
d/yr. Because of the diversity of cycle races, there are vastly different physiological
demands associated with the various events. Until recently there was little
information on the demands of professional cycling dunng traitKng or competition.
However, with the advent of reliable, valid bicycle crank dynanometers, it is now
possible to quantify real-time power output, cadence and speed during a variety of
track and road cycling races. This article provides novel data on the physiological
demands of professional and world-class amateur cyclists and characterises some
of the physiological attributes necessary for success in cycling at the 61ite level.

Introduction
Cycling is one of the m o s t efficient forms of h u m a n locomotion requiring less
energy per u n i t m a s s per unit distance t h a n a n y form of land transportation
(Brooks, 1989; Wilson, 1973). However, unlike walking or running, d u I ~ g cycling
m a n is combined with a machine: accordingly, there are n u m e r o u s p e r m u t a t i o n s
b y which p e r f o r m a n c e c a n be modified a n d improved. The power required to cycle
at a given velocity d e p e n d s on a complex interaction of m a n y physiological (ie. a n
athlete's maximal oxygen u p t a k e [VO2max], lactate threshold [LT], e c o n o m y of
movement, gross mechanical efficiency), environmental (wind velocity, temper-
ature, humidity, altitude) a n d m e c h a n i c a l (type of bicycle, wheels, tyres a n d
components) variables. A recent article in this j o u r n a l described the physiological
d e t e r m i n a n t s of e n d u r a n c e cycling p e r f o r m a n c e in moderate to weB-trained
cyclists (Coyle, 1999). Here, we p r e s e n t novel d a t a on the physiology and bio-
energetics of the world class t r a c k a n d road cyclist during competition.
Nomenclature and criteria for classification of a World Class
cyclist
There is no c o n s e n s u s in the scientific literature as to w h a t constitutes a n '61ite',
'world-class', 'trained', 'well trained' or a n 'untrained' cyclist. Subjects in several
published studies have b e e n classified as '61ite cyclists' w h e r e a s subjects with
similar physiological characteristics have b e e n t e r m e d 'moderately-trained' in

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 The Bioenergetics of World Class Cycling

Category Trained cyclists Well-trained Elite World Class

Training and race status

Training frequency 2-3 times a week 3-7 times a week 5-8 times a week 5-8 times a week
Training duration 30-60 min 60.240 rain 60-360 rain 60-360 rain
Training background 1 year 3-5 years 5-15 years 5-30 years
Race days per year 0-10 0-20 50-100 90-110
UCl ranking first 2000 first 200
Physiological variables
WmaX {W) 250"400 300-450 350-500 400"600
Wrnax (W/kg) 4.0-5.0 5.0-6.0 6.0-7.0 6.5-8.0
VO2max (L/min) 4.5-5.0 5.0-5.3 5.2-6.0 5,4-7.0
VO2max (ml/kg/min) 64-70 70-75 72-80 75-90
Economy (W/L/min) 72-74 74-75 76-77 >78

Table 1: Criteria for the classification of trained, well-trained, elite and World Classroad cyclists.

other investigations. The scientific literature is contaminated by investigations

that purport subjects to be '61ite eyelists' b u t who are clearly not 6lite and would
stand no chance of suceess in races against real 6lite cyclists. Because of the
confusion that eurrently exists, we propose a unifying classification and
terminology to be adopted by investigators in subsequent studies when describing
the status of their subjeets (Table 1).
It is important to notice that the classification of untrained, trained, or well-
trained is separate from a classification of novice, ~lite and World Class cyclists.
The first classification refers to the a m o u n t of training undertaken and is
independent of any (maximal or submaximal) physiologieal characteristics. For
example, a World Class cyclist with a very high VO2max can be classified as
'untrained' ff h e / s h e does not perform any training for a long period of time. In
Table 1 we have combined the two classifications and set criteria for each
performance ability eategories. The distinetion between trained and well-trained
cyclists should not be applied to individuals b u t only to groups of cyclists or
subjeets. This is because eertain individuals can be classified as 'highly trained',
b u t still lack the genetic endowment neeessary for superior performance. On the
other hand, those cyclists with excellent genetic endowment m a y achieve very
high physiological ratings without (much) training. The data presented in this
paper are derived from a very small and elite group of professional road cyclists
that ean be defined as World Class.
General Model of Cycling
Cycling can be regarded as an exercise in which mechanical energy is produced
to overcome external resistance. The speed of a rider can be estimated ff the power
available for skeletal muscle contraction from physiological sources (ie. an
individual's aerobic and anaerobic capacity) and the power required to overcome
resistance (eg. riding position and body mass, rolling resistance, air resistance,
gradient) are known. Optimal performance occurs when cycling speed increases
to the point where the power d e m a n d closely matches the power supply from all
available energy sources (Olds, 1993).
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The Bioenergeticsof World ClassCycling

The capacity and power of the aerobic and anaerobic systems for producing
adenosine triphosphate (ATP) has been estimated from a n u m b e r of studies
undertaken on h u m a n s performing maximal cycling exercise (Bogdanis, et al.,
1996; Boobis, et al., 1982; McCartney, et al., 1983) and taking muscle biopsy
samples from the vastus lateralis muscle immediately before and after the
workbout. When combined with measurements of an individual's VO2max,
maximal accumulated oxygen deficit (MAOD), anaerobic or LT a n d / o r VO 2
kinetics for the aerobic energy system and deficit kinetics for the anaerobic
system (Bangsbo, et al., 1990; Craig, et al., 1993; Gastin & Lawson, 1994; Medbo
& Tabata, 1989; Medbo, et al., 1988; Withers, et al., 1993), the power and
capacity of an energy system can be quantified. By measuring the gross
mechanical efficiency (ie. the power output relative to the oxygen uptake [VO2] to
produce that power), it is possible to estimate the external power of a biological
system from the following equation:

Gross mechanical efficiency (%) = 60 * W / 20934 * VO2

where W is the power output; VO2 is the absolute oxygen uptake (L/rain); and
20934 J is the equivalent of 1 L of oxygen.
Another measure of the functional ability of a cyclist can be obtained by
measuring economy of motion: the power output (W) at a submaximal steady-
state workload divided by the oxygen uptake (VO2) to ride at that power per unit
time (ie. W/L/rain). Finally, the recent advent of a reliable and valid bicycle crank
dynanometer (SRM, Schoberer Rad Me~technik, Weldorf, Germany), has made it
possible to accurately determine power output during a variety of track and road
cycling events (Bassett, et al., 1999; Broker, et al., 1999; J e u k e n d r u p & Van
Diemen, 1998; Keen, 1994; M a t i n , et al., 1998). These measures will be
discussed subsequently.
In a general model of cycling, there are two methods by which a rider may go
faster: by decreasing the various sources of resistance, a n d / o r by increasing the
power output. The first concern is how best to combine the cyclist with the bicycle
in order to increase power output, which includes improving the cyclist's
physiological attributes, modifying the rider's position, cadence and other
important physical variables. The second concern is how to reduce the mass and
profile of the bike a n d / o r rider to minimise rolling and air resistance, friction etc.
Changes can be made to the bicycle or its components, aerodynamic profile, the
rider's clothing and any other equipment. Such modifications are all designed to
minimise energy losses.
Many factors can affect the rate of power supply or demand. In terms of power
supply, the greatest changes are achieved by improving the athlete's physiological
attributes by training (Hawley, et al., 1997; Spangler & Hooker, 1990). Indeed, it
has been estimated that a 15% increase in a rider's absolute VO2max (L/min)
would provide a 4.5% improvement in 4,000 m performance time in an individual
cycling pursuit race (Olds et al., 1993). The weight of a rider plus bicycle is
another important factor changing the power demand. Weight slows a rider in
three ways: by retarding acceleration, by adding mass (to be carried uphill) and
by adding to the rolling resistance. Adding a small a m o u n t of weight does not
significantly reduce a cyclist's top speed: an extra bicycle mass of 2.7 kg would
add only 0.60 s to the time of a 4,000 m individual pursuit rider completing the

416
 The Bioenergetics of World Class Cycling

event in a time of 290 s. However, it would adversely affect acceleration speed,

(the time taken to reach a given velocity or power output (Martin, et al., 1998).
When cycling on the fiat at speeds <13 k m / h , tyre rolling friction is the
dominant retarding force acting against a bicycle and rider. However, 6lite cyclists
hardly ever race at such low velocities and rolling resistance is overshadowed by
wind resistance at riding speeds >13 k m / h . For example, at speeds >29 k m / h , up
to 80% of the total power output is used to overcome air resistance (Kyle, 1994;
Martin, et al., 1998). In fact, the power necessary to move the rider and bike
through the air increases as the cube of the riding velocity: hence, a small
increase in speed requires a n exponential increase in power output. In addition to
the requirements of overcoming air resistance, power is also required to overcome
rolling resistance, to impart kinetic energy to the bicycle and rider at the start of
an event and to ride up a gradient, the latter of which is related to the m a s s of the
rider and bicycle and the steepness of the slope.
In a recent examination of the evolution of the 'hour record' Bassett et al. (1999)
speculated that ~60% of the improvement in the record h a s come from
aerodynamic advances and 40% from the ability of a rider to sustain a higher
power output. In t e r m s of cycling performance, the sport scientist and coach are
continually struggling with the question "Where do we spend our time and
resources improving the energy supply (ie. training) or decreasing the energy
demand (ie. equipment modifications and innovations)?"
The physiological demands of track cycling
T r a c k cycling is a generic t e r m for all cycling events that take place outdoor and
indoor on a banked track of a standard distance (usually 333 m). Track cycling
can be divided into two broad categories: sprint and endurance races. Events
range from a 200 m flying sprint lasting ~10 s to the 50 k m points score race
lasting ~1 h. In addition, there is the prestigious one h o u r record where
competitors lime-trial alone and cover the greatest distance possible. At the
Sydney 2000 Olympic G a m e s the world's best track cyclists will compete in the
following events: Olympic sprint, 1,000 m individual time-trial, individual pursuit,
t e a m pursuit, Points race, Madison and Keirin.
Unlike road cycling, where the majority of events are performed at submaximal
exercise intensities and power outputs (described subsequently), the shorter track
cycling events require the rider to tax maximally both the aerobic and anaerobic
metabolic pathways. These events are often termed the 'mystery zone' by both
coaches and sport scientists: the consensus is that in order to be successful, both
the aerobic and anaerobic capacities need to be maximally developed through
appropriate training. Table 2 provides information on current world track cycling
records and the estimated contributions of the energy systems to the different
events. The contributions of the energy systems are based on data from several
studies (Bangsbo, et al., 1990; Craig, et al., 1993; Craig, et al., 1995; Gastin &
Lawson, 1994; Gastin, 1998} which have uUlised the accumulated oxygen deficit
method to estimate the aerobic and anaerobic energy contributions d u n n g
exhaustive cycling exercise. It should be noted that the m e a s u r e s on which the
data in Table 2 were derived were m a d e on moderately trained subjects. It is likely
that the capacity and absolute power of the anaerobic energy systems is far
greater in world class cyclists.
Until recently, scientists and coaches have employed mathematical models to

417
The Bioenergetics of World Class Cycling

Event World record Contributionfrom the powersystems(%)

(min:s) Alactic Anaerobic Aerobic
glycolytic
200 m Sprint
Male 0:09,865 a 40 55 5
Female 0:10.831 40 55 5
Time trial
Male (1,000 m) 1:00.148 a 10 40 50
Female (500 m) 0:34.010 20 45 35
Individual pursuit
Male (4,000 m) 4:11.114a* 1 14 85
Female (3,000 m) 3:30.816 I 24 75
Team pursuit
Male (4,000 m) 3:59.710 1 24 75
1 hr record (kin)
Male 56:375 <1 4 >95
Female 48.159 <1 4 >95
a Ridden at altitude. *Ridden in the "Superman' position on the bicycle.
This position is now banned by the UCl.

Table2: Currentworld record times for selectedtrack cyclingeventsand estimationsof the contributionsfrom the
energysystems.

estimate the mechanical power requirements of the different track cycling events.
While such models have been reasonably accurate (Martin, et al., 1998), the SRM
bicycle crank dynamometer has made it possible to accurately quantify instan-
taneous power output during track cycling. An additional feature of the SRM
dynamometer is that it can also provide useful information to the sport scientist
and coach for assessing a rider's technique and bike handling sldlls (Walsh,
C.B.V. personal conmmnication). As such, there are now data characterising the
demands of 6lite track cyclists during international competition.
4,000 m team pursuit
Broker et al. (1999) were the first to report SRM power output profiles on elite
track cyclists during simulated competition. These workers collected data on
seven members of the United States 4,000 m pursuit team during a 2,000 m
ridden at a speed (60 km/h) which would place a team in first position at most
international competitions. When cycling at this speed, the riders in first, second,
third and fourth position generated average power outputs of 607_+45, 430_+39,
389_+32 and 389_+33 W, respectively. In relative terms, riders in second, third and
fourth positions only needed to generate 71, 64 and 640/0 of the power output
sustained by the lead cyclist. When the data were averaged over all four riding
positions, it ranged from 70-75% (425-455 W) of the power of the lead rider
(Broker et al. 1999). These data are in good agreement with that collected on elite
Australian team pursuit cyclists during the 1998 World Cup. These data, which
are the first to profile power output during 6lite track competition, are displayed
in Table 3.
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 The Bioenergetics of World Class Cycling

Cyclist POWER OUTPUT (W)

Lead All four % of lead
position positions position
A 541 394 72.8

B 584 447 76.5

C 642 545 84.9

D 599 478 79.8

E 541 442 81.7

Mean (SD 581 +43 461 + _ 5 6 79.2+_4.7

Table3". Power output data collectedon the Australianmen's Nationalteam duringa 4000 m team pursuit race in
a 1998WorldCupevent.

1400 250
HR
1200
2OO ~"
A- E E
1000
PO 150
800
t~
08 000 100 "~"
D...
400
~-tj
5o ~
200

0 I , 'l | RPM 0
0 0.5 1 1,5 2 2.5 3 3.5 4 4,5
Time (rain)

Figure 1: Poweroutput (PO),heart rate (HPJ,cadence(RPM)and speedprofile for a cyclist riding a 4000m team
pursuit race duringa WorldCupcompetition.

The average power output of 607 W reported by Broker et al. (1999) for their
lead rider is higher than the 581 W value found for Australian riders under
competition conditions (Table 3). Such a discrepancy can probably be explained
by the differences in riding speed: the data of Broker et al. (1999) were
'normalised' for a riding speed of 60 k m / h , whereas the actual speed during the
World Cup competition was 56-58 k m / h . In addition, riding skill and technique,
riding position, body mass and frontal surface area, equipment design and
environmental and track conditions would all be expected to affect riding speed
and the power output necessary to sustain that speed. Although the average
power output data provide useful information as to the minimum requirements of
a 4,000 m team pursuit, the individual rider profile highlights the stochastic
demands of this event. Figure 1 presents the power profile, cadence, speed and
HR response of a rider during a 4,000 m team pursuit in World Cup competition.
419
The Bioenergetics of World Class Cycling

Instantaneous power output was -1,250 W at the start and only dropped to
<1,000 W after the first 8-10 s of the race. Thereafter, depending on the rider's
position in the team, power output fluctuates between 600-650 W in the lead
position and 350-400 W when riding behind other team members. This clearly
illustrates the stochastic power demands of team pursuit racing and highlights
the need for rapid response times to accelerate and decelerate into team
formation, depending on the rider's position within the race.

4,000 m individual pursuit

In contrast with the oscillating power requirements of the 4,000 m team pursuit
(Figure 1), the power profile for rider's competing in individual 4,000 m pursuit
races is generally m u c h more even. Figure 2 displays the power output profiles
for an elite male riding a 4,000 m and an 61ite female riding a 3,000 m race during
a World Cup event.
Compared with the profile for the 4,000 m team pursuit (Figure 1) there is a
m u c h narrower range of power outputs required to ride competitively at this level.
Perhaps surprisingly, power outputs during the first 5-10 s of both races are
remarkably similar (-1,000 W for male and female riders). However, despite the
shorter race distance, the power output for the female rider is considerably lower
for the remainder of the race (363-381 W) compared to the male rider (475-500
W). Interestingly, the female rider demonstrates superior fatigue resistance (ie.
has less of a decline in relative power output) over the race duration than the male
rider, whose power drop-off is marked, especially during the last 45-60 s of the
event. In a mathematical model of the power requirements of the 4,000 m
individual pursuit, Broker et al. (1999) estimated that a rider with a body mass of
- 7 9 kg who was 1.82 m tall would need an average power output of 479 W for a
race time of 4 min 31 s. This estimate agrees well with average power output data
we have collected for Australian riders competing in international events and
riding comparable times (Craig, N.P., unpublished observations).

1200

1000

800
D.
600

400
n

200 Female Individual 3,000 m Pursuit

0
0 0.5 1 1.5 2 2.5 3 3.5 4 4,5

Time (min)

Figure 2: Poweroutput profile for a male cyclistriding a 4,000mindividualpursuit and a female cyclistridinga 3,000m
individual pursuit race during WorldCup competition.
420
 The Bioenergeticsof World ClassCycling

The phySiOlogical demands of road cycling

Road cycling is a generic t e r m for cycling events that take place outdoor on the
road. This b r a n c h of cycling is unique in that it requires competitors to perform
on a variety of terrains (ie. level, uphill and downhill) and race situations (ie.
individual time-trial, t e a m time-trial, drafting in pack formation in the peloton).
These events vary in duration from time trials lasting 5 min, to stage races like
the Tour de France, which last 3 wk. However, it is important to point out that
most World Class cyclists will participate in a combination of events with the
primary emphasis on their best discipline. Some riders m a y also forsake their own
individual ambitions during specific races and ride for a designated t e a m leader
who h a s more chance of achieving overall individual success in the long stage
races. Accordingly, professional road cyclists spend m a n y hours training and
racing, covering between 30,000 and 35,000 k m / y r (Jeukendrup A.E., Unpub-
lished observations). This riding volume clearly differentiates the professional
cyclist from less genetically endowed, albeit 'well-trained' cyclists previously
described in the literature (Coyle, et al., 1991; Hawley, et al., 1997; Lindsay, et al.,
1996; Stepto, et al., 1999; Stepto, et al., 2000) who ride, on average, less t h a n half
this distance (250 versus 625 km/wk).
Another factor which differentiates the professional cyclists from other cyclists
is the n u m b e r of races they enter: the n u m b e r of race days in a group of 22
professional cyclists from a European t e a m was 101+6 d/yr, with a range of 88-
112 d / y r (Jeukendrup A.E., Unpublished observations). Because of the wide
diversity of road cycling events and because the physiological demands of these
events are totally different, they will each be discussed separately.
Time-trials
In time-trials individual riders try to cover a fixed distance as fast as possible. The
winner is the rider with the shortest time. Time-trials m a y vary in distance from
3 k m up to 6 k m and are often ridden as the first 'stage' of longer stage races (ie.
prologue). However, time-trials m a y be as long as 100 km. In these events,
individual maximal s u s t a i n a b l e power a n d aerodynamics will ultimately
determine performance time. Although there h a s been considerable information
collected from simulated time-trials in the laboratory during the past few years
(Jeukendrup et al. 1996; Lindsay et al. 1996; Palmer et al. 1996) there is little
information currently available on time-trials in actual competition, especially in
~lite cyclists. One of the reasons m a y be the difficulty with measuring the exercise
intensity in a field situation. However, with accurate portable telemetric HR
monitors and instantaneous power measuring devices it is now possible to
quantify the exercise intensity in training and competition (Jeukendrup & Van
Diemen, 1998). The power outputs m e a s u r e d during training/competition are
related b a c k to the maximal power output (Wmax) determined during a maximal
test in the laboratory. Wmax is determined during an incremental cycling test
comprising 3-5 min stages with 25-50 W workload increments.

Wmax = Wfinal + (t/T'W)

Where Wfinal is the last completed stage, W is workload increment (W), t is the
time spent in the final (uncompleted) stage, and T is the time (s) of the stage
duration.

421
The Bioenergetics of World Class Cycling

Recently, Padilla et al. (2000) determined the exercise intensity during time
trials in a group of professional cyclists by monitoring HR. Heart rate w a s
recorded in a number of time-trials, including prologues (<10 km), short time-
trials (<40 km), long time-trials (>40 km) and uphill time-trials. These workers
reported significant difference in HR's between the various time trials, which was
largely a fimction of the different duration of the events. The highest HRs were
recorded durtng the short prologues (~88-90% of HRmax). The short time-trials of
27.0-36.5 k m were raced at 82-88% of HRmax and the longer (49 km) time trials
were raced at a slightly lower intensity of between 79 and 84% of HRmax. Speeds
in these events on relatively level roads averaged 4 6 . 3 k m / h , 4 3 . 1 k m / h and
4 4 . 7 k m / h respectively. As would be expected, speed in the uphill time-trial w a s
significantly lower (40 km/h). Similarly, Lucia et al. (1999) reported that during
a 60 k m time-trial in the Tour de France, cyclists spent 95% of race time (70 min)
at HR between 88% and 100% of HRmax. In that time trial, it w a s estimated that
the winner maintained an intensity of >90% of VO2max for 70 min, averaging 50
k m / h . It should be noted, however, that time-trials within stage races m a y not

1400 100

1200

1000
-',~\ "!, ' ,;,,,i', ,'~,~',,, ~, . . . . . . . . . . . . . .
. .
/", ~z~/'~
I " , ,gv', f 40

800

600

¢ -20
400

200

F ~'t -80
0
0;45 1 ;00 1:15 1:30 1:45

T i m e {h:min}

4O0

8O
200 ~ ~ ~,
60

40 A
.E
20 ~

600

-20 el
4OO
-40
2OO
-60

0 -80
0:00 0:30 1:00 1:30
Time (h:min)

Figure 3: Power output (--) and speed (-----) for a male professionalcyclistriding a 30kin (1A) and a 381o11(1B) uphill
time trial.
422
 The Bioenergeticsof World ClassCycling

always be raced at maximal level. The role of a n individual rider within a team
often determines the intensity at which a time trial is raced. This is illustrated by
the findings of Padilla et al (2000). Riders who were aiming at a high overall
ranking rode the 27-70 k m time trials at a n intensity of 81-88% of Wmax, whereas
riders with a strategic role, rode at a slightly lower intensity of 75%-81% of Wmax.
Riding "slow" is not an option for any rider because if riders do not finish each
days stage within a certain time s p a n from the winner (usually 10%) they are
eliminated from the overall race.
Using a power-measuring device, we have collected data dining time trials in
major international races. The HR's measured during these time-trials is in good
agreement with data from previous studies (Lucia, et al., 1999, Padilla, et al.,
2000). Because power output data are highly individual and dependent on a
cyclist's m a s s and anthropometric profile, equipment and, position on the bicycle,
this data will be presented as a case study. The data of an uphill time-trial of a
professional cyclist are displayed in Figure 3.
The average power output during this 30 k m time trial was 381+61 W for an
average speed of 30_+ 11 k m / h . Whereas most time-trials on a fiat course will be
characterised by a relatively constant power profile, this uphill time trial displays
a highly intermittent profile with power outputs varying from 260 to 460 W, HR
ranging from 135 to 165 b e a t s / m i n and speeds as low as 17 k m / h up to 62
k m / h . A further example of a n uphill time-trial is displayed in Figure 3b. This 38
k m time-trial was predominantly uphill, b u t included a downhill section. Average
power output was 320_+148 W and average speed 33.1 k m / h : this illustrates the
dissociation between speed and power output (Figure 3b). As h a s been noted
previously, (Jeukendrup & Van Diemen, 1998), speed is often a poor indicator of
the exercise intensity during cycling, especially when riding on hilly terrain. In the
mountains, speed and exercise intensity seem almost inversely related: the
highest speed is recorded at the lowest power output in a downhill section of the
race (Figure 3b).
The power output recorded with power measuring devices corresponds to the
power outputs calculated from heart rate via a pre-determined relationship
between power output and HR in a laboratory (Grazzi, et al., 1999; Padilla, et al.,
2000). Padilla et al. (2000) estimated that the power output during uphill time
trials was 331-376 W, representing 75-83% of Wmax. There are also case reports
of world h o u r record holders, indicating the extreme physiological d e m a n d s of
time trials (Hawley & Burke, 1998; Padflla & Mujika, 2000). For example, it h a s
been estimated that the current world record holder, Britain's Christopher
Boardman, had to produce an average power output of 442 W for 60 min in order
to ride at an average speed of 56.375 k m / h . This equates to an average oxygen
uptake of 5.6 L/rain or 81 m l / k g / m i n . Although such a sustainable power output
is extreme, a cyclist with a m u c h larger drag coefficient set a previous world 1 h
record of 53.040 k m by producing an estimated average power output of 510 W
(Padilla and Mujika, 2000). It can be estimated that the VO2 during this ride m u s t
have been around 6.25 L / m i n or close to 80 m l / k g / m i n . This rider had a Wmax of
572 W and was able to sustain close to 90% ofWmax for the duration!
In conclusion, professional cyclists produce exceptional power outputs (>320-
450 W) during time-trials ranging in distance from 5-70 km. The absolute power
output will depend on the duration of the time-trial, the course profile and on the
role of a cyclist within the team.

423
The Bioenergetics of World Class Cycling

200 '] ~ HR OBLA:176

160I HR LT: 162
&
120

~ 8o

40 ~

00:00:00' 00:20:00' O0:4O:OO

1 0; :00:00 ~ O1:20:00
200
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . HR OBLA:173
~,~ HR LT: 146
120

8O
"T"

4O

00:00:00' 00:20:00 ' 0~0:40:00 ' 01:00:00 ' Time/hh:mm:ss

Figure4: Heart rates determined duringindividualtime trial and team time trial races(Figureadapted from Padillaet
al., 2000).

Team time-trials
T e a m time-trials are usually part of longer stage races. T e a m s of 4-9 riders try to
cover a fixed distance (usually 100 km) as fast as possible, similar to the
individual time-trial. The physiological d e m a n d s of t e a m time-trialing differ sub-
stantially from individual time-trials, being more intermittent in nature due to the
alternating d e m a n d s of riding in first position and drafting behind the other
riders. In addition to conventional time-trials, there are also a few one-day races
in which two riders (two-ups) ride a time-trial (for example Grand Prix des Nations
and Grand Prix Eddy Merckx). Unfortunately there is very limited information on
the physiological d e m a n d s during t e a m time trials. A recent study (Padflla et al.,
2000) compared the physiological differences between individual time trials and
t e a m time trials. As can be seen (Figure 4), the t e a m time-trial is m u c h more
intermittent compared to the individual race.
Early studies have reported that drafting behind another rider/riders reduces
the oxygen cost of cycling by 25-40% (Kyle, 1979; McCole, et al., 1990). The
largest effect of drafting is seen at high speeds when a cyclist is surrounded by
other riders. Drafting is therefore a very important sldll in professional road
racing. An excellent example of this drafting skill during a road race is displayed
in Figure 5.
This world class cyclist participated in the Tour de France and while the average
speed in this 6 h level stage was 40 k m / h , this cyclists with excellent drafting
skills managed to reduce his average power output to j u s t 98 W. It can be
424
 The Bioenergetics of World ClassCycling

'im

8O

(3

O:O0:CO 1:0(t00 ~00:00 30(1(]0 4:00:00 5:0~C0 6:00:.On

lime (h:rnin)

Figure5: Poweroutput ( - - ) and speed ( ) duringa 220kinroadrace. Thelow power ouput relative to the high
averagespeedindicatesthis rider wasa sldlleddrafter.

calculated that in optimal conditions with no wind and level roads, and with a
good aerodynamic position, riding at that speed would require ~275 W (Martin, et
al., 1998). Drafting in a team time trial, however, does not result in the same
drafting effect that a rider might obtain in the peloton. Drafting allows for some
recovery, however, as can be seen in the HR trace in Figure 4.
In conclusion, compared to individual road race time-trials, team trials are
characterised by a more intermittent nature due to the longer periods of drafting
and recovery. Average HR and power outputs, however, are comparable to
individual time trials of similar duration.

Road races
Road races typically last 60 km (criteriums) up to about 300 km and are
characterised by a mass (bunch} start. Some road races are as long as 600 km
(Paris-Brest-Paris) but most professional classics such as the Tour of Flanders,
Paris-Roubaix and the Amstel Gold race are generally 250-300 km. All riders
begin together as part of the peloton and the first rider who crosses the finish line
is the ~ n n e r , independent of finish time. Although this is an individual race with
only one rider the winner, individuals are usually members of a team and team
tactics can determine a large part of the developments and the physiological
intensity of a race. The principal concerns affecting road race performance are
team tactics and drafting. Because riders remain grouped in the peloton, only the
few who are in the front at any one time face the full effects of wind resistance,
while others can maintain the peloton's pace by drafting. Usually, breakaway
riders can outdistance the petoton only when the complex dynamics of team
support allow it. The major exception to this rule occurs when the peloton
encounters hills. The average power output during road races over 200 km
typically vary between 150 and 300 W (Jeukendrup A.E. et al. Unpublished
observations).
425
The Bioenergeticsof World ClassCycling

Criterium races are shorter road races held on a small, looped course. Riders
perform multiple circuits of the course, with high-speed sharp turns. Bike
handling skills and sprint speed become more important in this event, b u t from
a physiological point of view criteriums are very similar to other road races.
Stage races
Stage races are multiple day events combining time trials and road races (stages)
and sometimes t e a m time-trials. Stage races m a y last 2 days up to 3 wk s u c h as
the major three European stage races (The Vuelta a Espafia, The Giro d'Italia and
the Tour de France). Each day, overall classification is calculated from the Finish
position and times, and there is an overall classification with the leader of the race
being the rider with the lowest overall time. In professional stage races, s u c h as
the Tour de France, time-trial stages and m o u n t a i n stages are often crucially
important and will separate contenders from support riders (ie. domestiques) in
the overall standings, as well as winners from contenders. Stage races are usually
won or lost by very small margins (ie. 200-400 s), which, over the course of a 3
wk stage race lasting ~300,000 s represents 0.07-0.13% of total race time.
Lucia et al. (1999) recorded the HR's of eight professional cyclists during 22
stages of the Tour de France and calculated the time spent in three arbitrarily
chosen exercise intensity zones. These zones were determined by measuring
ventilatory threshold in a laboratory exercise test before the race. A first
ventilatory threshold (VTI) was determined as an increase in VE/VO 2 with a n
increase of VE/VCO2 and the departure from linearity OfVE. VTII was established
using the criteria of a n increase in b o t h VE/O 2 and VE/VCO 2. While
acknowledging the limitations of this method, and the likelihood that these
thresholds m a y have changed during the 22 d of the Tour de France, the data give
an indication of the physiological d e m a n d s of cycling in stage races. Cyclists spent
71, 23, and 8 h in the low-, medium- and high-intensity zones respectively. The
relative contributions were 70%, 23% and 7% of total race time. However, there
were substantial differences between time-trials, fiat stages, high m o u n t a i n
stages, and m e d i u m mountain stages, and between riders depending on their role
within the team.
Associated with the high power outputs sustained by these cyclists over
prolonged periods are very high daffy energy expenditures. Saris et al. (1989)
studied cyclists during the Tour de France using doubly labelled water techniques
and estimated that energy expenditure was 24 M J / d (5,700 kcal/d) with extremes
of up to 40 M J / d a y (9,500 kcal/d) during very long, m o u n t a i n o u s stages. More
recently power output was m e a s u r e d during stages of the Tour de France and it
was demonstrated that the average power output over a 6 h stage was over 240
W with a n estimated energy expenditure of 24 MJ or 5,700 k c a l / d (Jeukendrup
2000). Garcia-Roves et al (1997) reported average energy intakes during the
Vuelta Ciclista a Espafia of 23.5 MJ/d. These data suggest that energy
expenditures reported for riders during stages of the Tour de France in the late
1990's have not changed considerably from those determined in the late 1980's.
With such extreme and high absolute energy expenditures (Brouns et al., 1989,
J e u k e n d r u p , 2000; Saris, 1997; Saris, et al., 1989;) these cyclists have major
problems maintaining energy balance. While eating large a m o u n t s during the race
is impractical or even impossible, time to eat is mainly limited to breakfast and
dinner. Appetite is suppressed immediately after exhausting exercise, which sets

426
 The Bioenergetics of World Class Cycling

73
72
71
~ 7o
.~ 69
~ 68
67

66
65
5 10 15 20
Time (days)

Figure 6: Bodymassof six cyclists during the 1999Tourde Trance.Valuesare mean+_SEM.

a further hurdle for a large food intake. However, despite conditions that make it
difficult for riders to maintain energy balance, body mass d u i ~ g long stage races
like the Tour de France is remarkably well maintained. The body weight of 6
cyclists participating in the Tour de France was recorded daily for 20 d in the
morning after an overnight fast and after voiding (Jeukendrup A.E. Unpublished
observations). Figure 6 shows that although, there was a trend towards a
decreasing body mass at the end of the 3 wk race, this decrease was modest and
not statistically significant.
PhySiOlogical measurements of world class track and road
CyCliStS
With recent advances in the physiological testing of athletes, along with
monitoring training and racing responses, it is slowly becoming accepted that
sport science can make valid contributions to assist coaches in the preparation of
their athletes. However, only recently have data on some of the physiological
characteristics of 61ite amateur and professional cyclists been published (Craig, et
al., 1998; Hawley & Burke, 1998; Jeukendrup & Van Diemen, 1998; Padilla, et
al., 1999). It is of note that most of these data have been obtained from male
cyclists: there is currently little published information on the physiological profiles
of 61ite female cyclists.
Unfortunately there have been few attempts to identify the key physiological
variables associated with 6lite track cyclists. However, an examination of long-
itudinal data collected from the Australian Institute of Sport's track cycling
laboratory (Craig, N P., unpublished observations), in conjunction with previously
published modelling data (Craig, et al., 1993; Olds, et al., 1993) provide an insight
into the major factors impacting on success in international track cycling. One of
the variables frequently discussed by coaches, sport scientists and cyclists is body
mass, which increases the energy cost of acceleration, rolling resistance and the
projected frontal area of the cyclist (and hence air resistance). Olds et al. (1995)
have estimated that an extra fat mass of 2 kg would increase a 4,000 m individual

427
The Bioenergetics of World ClassCycling

pursuit performance time by 1.5 s or a distance o f - 2 0 m. Longitudinal data from

the Australian Institute of Sport's track cycling laboratory indicates that ~lite track
cyclists consistently achieve sum of six skinfolds <40 m m immediately prior to
major international competition (ie. World Championships or Olympic Games).
A consideration of the relative contributions of the aerobic and anaerobic
metabolic energy systems to track cycling events (Table 2) highlights the import-
ance of a high aerobic capacity for success in the majority of track cycling events
(Craig et a0., 1993). Coupled to a high aerobic capacity (>6 L/min) is a high (-500
W) power output, combined with the ability to attain peak power rapidly (spnnt
events). Although many track cycling events are predominantly aerobic activities,
the anaerobic energy system also plays an important role in the required energy
production (Table 2). This notion is supported by the data of Craig et al. (1993)
who reported a significant correlation between MAOD and 4000-m individual
pursuit time (r = -0.50, P <0.05). That study (Craig, et al., 1993) also found that
s p r i n t trained track cyclists had a significantly greater anaerobic capacity
(66.9+2.2 ml/kg) compared to endurance trained track cyclists (57.4_+6.7 ml/kg).
Like most physiological variables, anaerobic capacity can be improved with appro-
priate training. Medbo and Burgess (1990) reported a 10% increase in MAOD after
6 wk of interval training. Such a change in MAOD could, potentially, improve
4,000 m individual pursuit time by ~12 m (Olds, et al., 1993).
In a recent study, Padilla et al. (1999) summarised some of the physiological
characteristics of professional road cyclists and the reader is referred to that paper
for a comprehensive review of the subject. These workers determined the maximal
and submaximal physiological laboratory responses of members of a Spanish
professional road cycling team (Banesto) whose subjects included five-time Tour
de France winner Miguel Indurain, as well as riders who had won other stage
races (Giro d'Italia, Vuelta a Espafia). According to their role in the team, the
cyclists were categorised as either fiat terrain (FT), time trial (Tr), all terrain (AT)
or uphill (UH) specialists.
Figure 7 displays the maximal values for power output and the VO 2 (L/min)
values determined during an incremental test to exhaustion for the team (]1=24).
The maximal protocol employed utilised 4 min stages with workload increments
of 35 W so that steady-state measures of blood lactate could also be determined
(described subsequently). Unfortunately, respiratory gas exchange measurements
were not taken so the VO2max of subjects was estimated from a previously
published regression equation (Hawley & Noakes, 1992).
As can be seen (Figure 7), the maximal power outputs and aerobic capacities of

2" 651 r * 1
i

g
E

FT TT AT UH FT ~ AT UH

Figure~ The maximal physiological characteristics of professional cyclists (n=24). F[ fiat terrain; T[, time-trial; AT,all
terrain; UH, uphill. See text for details. All values are mean+-SD.Data arefrom Padllla et al. (1999). * P<0.05
428
 The Bioenergetics of World Class Cycling

the FT and TiC riders were significantly higher t h a n the UH riders. The highest
Wmax values were m e a s u r e d in the FT riders (461+39 W), which was higher t h a n
AT and UH (432+27 and 404_+34 W, P<0.05). T r (457_+46 W) was also significantly
higher t h a n UH (P<0.05). Absolute VO2max values were also significantly higher
in FT and TI" t h a n UH (5.67_+0.44 and 5.65_+0.53 versus 5.05_+0.39 L/min;
P<0.05), b u t none of these were different from AT (5.36+0.30 L/min). These
findings are predictable based on the specialised role of the team members, and
several anthropometric m e a s u r e m e n t s (not shown) which reveal that the FT and
T r riders were also taller, heavier, and had greater body surface areas t h a n their
AT and UH counterparts. However, when these s a m e parameters are expressed
relative to a riders body m a s s (BM), a different picture emerges, with the UH riders
gaining the ascendancy. The UH riders have the highest power: BM ratio (6.47
(0.33 W/kg) followed by Tr, AT and FT (6.41_+0.12, 6.35_+0.18 and 6.04-+0.29
W/kg, respectively). These values were significantly different between UH and NF.
VO2maxvalues expressed relative to BM were significantly lower in FT t h a n all
other groups (74.4_+3.0 versus 79.2_+1.1, 78.9-+1.9 and 80.9_+3.9 n l l / k g / m i n for
TF, AT and UH; P<0.05).
Although the m a m m a l data reported on these professional cyclists (Padilla et al.
1999) is impressive, perhaps the single m o s t striking physiological attribute of
these riders is their submaMmal exercise capacity, determined by the power
output (W) at the individual LT, defined as the exercise intensity that elicited a 1
mM increase in blood lactate above baseline concentrations when riding at -50-
60% of VO2max(Coyle, et al., 1983). Both FT and TT riders had higher power
outputs at LT t h a n did AT and UH riders (356_+31,357_+41,322_+43 and 308_+46
W respectively). Perhas to emphasise the point, it h a s previously been reported
that in a group of 6lite cyclists whose average 40 kin time was ~54 min, power
output at LT was 311 W (Coyle, et al., 1991). As m o s t professional cyclists com-
plete time-trial races of this distance at speeds in excess of 50 k m / h , with m a n y
riders able to sustain power outputs over 400 W for the duration (see section on
time trials), sport scientists need to exercise caution when classifying their subject
pool (see subsequent section).
Another interesting observation is t h a t gross efficiency m e a s u r e d in
professional road cyclists is high b u t not markedly different to well-trained riders.
Gross efficiency was recently assessed in a group of 16 professional cyclists
during a n incremental exercise test. Gross efficiency was 21.4_+0.2% (Jeukendrup
A. E. Unpublished observations). This value is slightly higher t h a n values reported
for cyclists of a lower ability level (Coyle, et al., 1992). In that study, Coyle et al.
(1992) suggested that cycling efficiency was related to the percentage of type I
muscle fibres: the more type I fibres a cyclists possessed then the higher their
efficiency. Indeed, these workers observed that at a given power output, two
cyclists could vary by as m u c h as 15-20% in their absolute oxygen cost for cycling
(Coyle, et al., 1992). These workers reported that a group of cyclists with a high
proportion of type I fibres (67_+40/0) were capable of generating higher power output
(10%) t h a n a group whose m u s c u l a t u r e contained a lower (53_+6%) percentage of
type I fibres (Coyle, et al., 1992). It was also concluded that cycling performance
was largely determined by gross efficiency, suggesting that a large percentage type
I muscle fibres is a necessary prerequisite for success as a professional cyclist
(Burke, et al., 1977; Coyle, et al., 1992).
At present, however, a causal relationship between muscle fibre type and gross

429
The Bi0e'nergetics of World ClassCycling

cycling efficiency h a s not been directly

d e m o n s t r a t e d . Longitudinal training
studies of professional cyclists are
O4 C~I C~I 0,10,I 0,1 necessary in order to determine whether
the relationship between the proportion
of type I muscle fibres and cycling
performance is causal or coincidental.
For example, in the study of Coyle et al.
(1992), those cyclists with the highest
proportion of type I fibres h a d also been
cycling longer (8.8+0.9 versus 5.0_+3.0
~.J yr, a difference of 76%) t h a n their
counterparts with fewer type I fibres. It
remains to be determined whether the
n u m b e r of years of endurance training
can alter fibre type a n d / o r efficiency.
Such studies are important because
cycling efficiency h a s the potential to
have large effects on performance. For
example, it can be calculated that for an
elite cyclist with a m a s s of 70 kg and a
l h sustainable power of 400 W, a 1%
improvement in gross efficiency equates
to a 48 s improvement in 40-km time-
trial time.
Lr~ o c3 o o o o To the best of our knowledge, there are
m no reports in the scientific literature
~o
documenting the longitudinal changes
that occur in world class cyclists after
z,
several years of intense training. Here for
E:: I:= '-- "I-
the first time we present data on the
•~ ~'~E.~=~ ..
'~ ~ a ~ =~._~ d e v e l o p m e n t of a world-class t r a c k
o cyclist over a 6 yr period (Table 4).
Over the time in which these measure-
m e n t s were taken, this rider lost 3.6%
gaNo "¢3 body mass, and improved his VO2max
_ a and Wmax by 12% and 10% respectively.
Accordingly, this rider's power: m a s s
(1~ ratio increased from 6.25 W / k g to 7.18
~ F~ ~ aa _~ W/kg. This cyclist's best performances
c a m e after 4-5 yr of high-intensity
training: it is clear to see that the de-
velopment of a World class cyclist takes
NN NN m a n y years (see also Table 1).
Conclusions
Professional cycling is one of the m o s t
~f d e m a n d i n g of all sports combining
extremes of exercise intensity, duration
and frequency. The power requirements
430
 The Bioenergetics of World ClassCycling

of cycling depend on a complex interaction of m a n y physiological, environmental

and mechanical variables. With the recent advent of reliable, valid bicycle crank
dynanometers, it h a s been possible to determine power output profiles during a
variety of track and road cycling events. This data reveal that the physiological
d e m a n d s of cycling are extreme: for track events lasting 3-5 min riders produce
average power outputs of 500-600 W. An average power output of ~440 W,
corresponding to a VO 2 of N5.5. L/min, was sustained for 1 b by the current
world-record holder in that event. In road races, power outputs are more
stochastic in nature and related to the type (time-trial, criterium, m a s s - s t a r t stage
race), length and terrain of the race, as well as t e a m tactics. However, professional
cyclists produce power outputs between 325 and 450 W for events ranging in
distance from 5 to 70 km, and between 150 W (when drafting) to 300 W during
long (~200 km) stage races. The maximal physiological characteristics of pro-
fessional road and track cyclists are impressive: values for VO2max and Wmax are
typically >5.5 L / m i n and 450 W, respectively, corresponding to a power:mass
ratio of >6.5 W/kg. Perhaps the single most striking physiological attribute of
these riders is their submaximal exercise capacity, demonstrated by the high
(~350 W) power output at LT. As professional road cyclists frequently ride >30,000
km/yr, including N100 race d/yr, it is not surprising that longitudinal data
suggest that the development of a world class cyclists takes a m i n i m u m of 4-5 yr.
References
Bangsbo, J., Gollnick, P.D., Graham, T.E., Juel, C., Kiens, B., Mizuno, M., & Salfin, B. (1990).
Anaerobic energy production and 02 deficit-debt relationship during exhaustive exercise in
humans. Journal of Physiology 422: 539-559.
Bassett, D.R., Kyle, C.R., Passfield, L, Broker, J.P., & Burke, E.R. (1999). Comparing cycling
world hour records, 1967-1996: modeling with empirical data. Medicine and Science in
Sports and Exercise 31: 1665-1676.
Bogdanis, G.C., Nevill, M.E., Lakomy, H.K.A., Graham, C.M., & Louis, G. (1996). Effects of active
recovery on power output dm~rlg repeated mammal sprint cycling. European Journal of
Applied Physiology 74:461-469
Boobis, L., Williams, C., & Wootton, S. (1982). Human muscle metabolism during brief maximal
exercise. J o u r n a l of Physiology 3 3 8 : 2 1 - 2 2
Broker, J.P., Kyle, C.R., & Burke, E.R. (1999). Racing cyclist power requirements in the 4000-m
individual and team pursuit. Medicine and Science in Sports and Exercise 31: 1677-1685.
Brooks, A.N. (1989). Energy consumption of high efficiency vehicles. Cycling Science 1:6-9
Brouns, F., W. H. M. Saris, Stroecken, J., Beckers, E., Thijssen, R., Rehrer, N.J., & ten Hoot, F.
(1989). Eating, drinking, and cycling. A controlled Tour de France simulation study, part I.
international Journal of Sports Medicine I0: (Supplement 1): $32-$40.
Burke, E.R., Cerny, F., Costill, D.L., & Fink, W. (1977). Characteristics of skeletal muscle in
competitive cyclists. Medicine and Science in Sports and Exercise 9: 109-i 12.
Coyle, E.F. (1999). Physiological determinants of endurance exercise performance. Journal of
Science and Medicine in Sport 2: 181-189.
Coyle, E.F., Felmer, M.E., Kautz, S.A., Hamilton, M.T., Montain, S.J., Baylor, A.M., Abraham,
L.D., & Petrek, G.W. (1991). Physiological and biomechanical factors associated with elite
endurance cycling performance. Medicine and Science in Sports and Exercise 29: 93-107.
Coyle, E.F., W.H. Martin, A.A. Ehsani, J.M. Hagberg, S.A. Bloomfield, D.R. Sinacore, & J.O.
Holloszy (1983). Blood lactate threshold in some wefi-trained ischemic heart disease patients.
Journal of Applied Physiology 54: 18-23.
Coyle, E. F., L S. Sidossis, J. F. Horowitz, and J. D. Beltz. (1992). Cycling efficiency is related to
the percentage of type I muscle fbers. Medicine and Science in Sports and Exercise
24:782-788.
Craig, N. P., Norton, K.I., Bourdon, P.C., Woolford, S.M., Stanef, T., Squires, B., Olds, T.S.,
Conyers, R.A.J., & Walsh, C.B.V. (1993). Aerobic and anaerobic indices contributing to track
endurance cycling performance. European Journal of Applied Physiology 67: 150-158.
Craig, N.P., Norton, K.I., Conyers, R.A.J., Woolford, S.M., Bourdon, P.C., Stanef, T., & Walsh,

431
The Bioenergetics of World Class Cycling

C.B.V. (1995). Influence of test duration and event specificity on maximal accumulated
oxygen deficit of high performance track cyclists. International J o u r n a l of Sports Medicine
16: 534-540.
Craig, N. P., Walsh, C., Martin, D., Woolford, S., Bourdon, P.C., Stanef, T., Barnes, P. & Savage,
B. (1998). High performance track, road and mountain cyclists. Test m e t h o d s manual:
sport specific guidelines for the physiological a s s e s s m e n t of elite athletes, C. J. Gore,
ed., Australian Sports Commission, Canberra.
Garcia-Roves, P., N. Terrados, et al. (1997}. Macronutrients intake of top level cyclists during
continuous competition-change in the feeding pattern. International Journal of Sports
Medicine 19: 61-67.
Gastin, P. (1998). Energetics of high intensity running. Modern Athlete and Coach 36: 3-9.
Gastin, P.B., & Lawson, D.L. (1994). Variable resistance all-out test to generate accumulated
oxygen deficit and predict anaerobic capacity. European Journal of Applied Physiology 69:
331-336.
Grazzi, G., N. Aifieri, C. Borsetto, I. Casoni, F. Manfredini, G. Mazzoni, & F. Conconi (1999). The
power output/heart rate relationship in cycling: test standardization and repeatability.
Medicine and Science in Sports and Exercise 31: 1478-1483.
Hawley, J.A., and Burke, L.M. (1998). The limits to human performance. In: Peak performance:
Training and nutritional strategies for sport, pp.402-406.
Hawley, J.A., Myburgh, K.H., Noakes, T.D., & Dennis, S.C. (1997). Training techniques to
improve fatigue resistance and enhance endurance performance. Journal of Sports
S c i e n c e s 15: 325-333.
Hawley, J.A., & Noakes, T.D. (1992). Peak power predicts maximal oxygen uptake and
performance time in trained cyclists. European Journal of Applied Physiology 65: 79-83.
Jeukendrup, A. E., Saris, W.H.M., Brouns, F., & Kester, A.D.M. (1996). A new validated
endurance performance test. Medicine and Science in Sport and Exercise 28: 266-270.
Jeukendrup, A.E., & Van Diemen, A. (1998). Heart rate monitoring duling training and
competition in cyclists. Journal of Sports S c i e n c e s 16: $91-$99.
Jeukendrup, A. E. (2000). Cycling. IOC Encyclopaedia of Sports Medicine: Nutrition in Sport.
R. J. Maughan (Editor)., BlackweU Science, Oxford, pp.562-573.
Keen, P. (1994). The truth behind the race of truth. Cycle Sport 7: 46-50.
Kyle, C.R. (1979). Reduction of ~ n d resistance and power output of racing cyclists and runners
travelling in groups. Ergonomics 22: 387-397.
Kyle, C. R. (1994). Energy and aerodynamics in cycling. Clinics in Sports Medicine 13: 39-73.
Lindsay, F. H., Hawley, J.A., Myburgh, K.H., Schomer, H.H., Noakes, T.D., & Dennis, S.C. (1996).
Improved athletic performance in highly trained cyclists after interval training. Medicine and
S c i e n c e in Sports and Exercise 28: 1427-1434.
Lucia, A., J. Hoyos, A. Carvajal, & J. L. Chicharro. (1999) Heart rate response to professional road
cycling: the Tour de France. International Journal of Sports Medicine 20:167-72.
Martin, J.C., D.L. Milliken, J.E. Cobb, K.L. McFadden, & A.R. Coggan. (1998). Validation of a
mathematical model for road-cycling power. J o u r n a l of Applied B i o m e c h a n i c s 14:276-291.
McCamaey, N., Heigenhauser, G.J.F., & Jones, N.L. (1983) Power output and fatigue of human
muscle in maximal cycling exercise. J o u r n a l of Applied Physiology 55:218-224.
McCole, S.D., Claney, K., Conte, J.C., Anderson, R., Hagberg, J.M. (1990). Energy expenditure
during bicycling. Journal of Applied Physiology 68: 748-53.
Medbo, ,J.I., Burgers, S. (1990). Effect of training on the anaerobic capacity. Medicine and
Science in Sports and Exercise 22: 501-507.
Medbo, J. I., Mohn, A.C., Tabata, I., & Sejersted, O.M. (1988). Anaerobic capacity determined by
maximal accumulated 02 deficit. J o u r n a l of Applied Physiology: 64:50-60.
Medbo, J.L., & Tabata, I. (1989). Relative importance of aerobic and anaerobic energy release
during short-term exhausting bicycle exercise. Journal of Applied Physiology 67: 1881-
1886.
Olds, T. S., Norton, K., & Craig, N.P. (1993). Mathematical model of cycling performance. Journal
of Applied Physiology 75: 730-737.
Olds, T.S., Norton, K., Craig, N.P., Olive, S., & Lowe, E. (1995). The limits of the possible: Models
of power supply and demand in cycling. The Australian Journal of Science and Medicine
in Sport 27: 29-33.
Padilla, S., Mujika, I., Cuesta, G., & Goiriena, J. (1999}. Level ground and uphill cycling ability
in professional road cycling. Medicine and S c i e n c e in Sports and Exercise 31: 878-885.
Padilla, S., & Mujika, I. Scientific approach to the 1-h cycling world record. A case study (2000}.

432
 The Bioenergetics of World Class Cycling

Journal of Applied Physiology 89: 1522-1527.

Padilla, S., I. Mujika, J. Orbananos, & F. Angulo (2000). Exercise intensity during competition
time trials in professional road cycling. Medicine and Science in Sports and Exercise 32:
850-856.
Palmer, G.S., Dennis, S.C., Noakes, T.D., & Hawley, J.A. (1996). Assessment of the reproducibility
of performance testing on a n air-braked cycle ergometer. International Jollrnal of Sports
Medicine 17: 293-298.
Saris, W. H. M. (1997). Limits of Human Endurance: lessons from the Tour de France. In:
Physiology, stress and malnutrition: functional correlates, nutritional intervention. J.
M. Kinney and H. N. Tucker (Eds.). Lippincott: Raven Publishers, pp. 51-463.
Saris, W. H. M., van Erp-Baart, M.A., Brouns, F., Westerterp, K.R., & ten Hoor, F. (1989). Study
on food intake and energy expenditure during extreme sustained exercise: the Tour de
France. International Journal of Sports Medicine I 0 (Suppl. I):$26-$3 i.
Spangler, D. & Hooker, G. (1990). Scientific training and fitness testing. Cycling Science 21: 23-
26.
Stepto, N.K., J.A. Hawley, S.C. Dennis, & W.G. Hopkins (1999). Effects of different interval-
training programs on cycling time-trial performance. Medicine and Science in Sports and
Exercise 3 1 : 7 3 6 fl 741.
Stepto, N.K., Martin, D.T., Fallon, K.E., & Hawley, J.A. (2001). Metabolic demands of intense
aerobic interval training in competitive cyclists. Medicine and Science in Sports and
Exercise (In press).
Wilson, S.S. (1973). Bicycling technology. Scientific American 228: 81-91.
Withers, R.T., Van Der Ploeg, G., & Finn, J.P. (1993). Oxygen deficits incurred during 45, 60, 75
and 90 s of maximal cycling on a n air-braked ergometer. European Journal of Applied
Physiology 67: 185-191.

433