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Kinetics of VO(2) in professional cyclists

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Kinetics of V̇O2 in professional cyclists
ALEJANDRO LUCÍA, JESÚS HOYOS, ALFREDO SANTALLA, MARGARITA PÉREZ, and JOSÉ L. CHICHARRO
Departamento de Ciencias Morfológicas y Fisiologı́a, Universidad Europea de Madrid, SPAIN; Unidad de Investigación
en Fisiologı́a del Ejercicio, Universidad Computense, Madrid, SPAIN; Asociación Deportiva Banesto, Madrid, SPAIN;
and Departamento de Enfermerı́a, Universidad Complutense, Madrid, SPAIN

ABSTRACT
LUCÍA, A., J. HOYOS, A. SANTALLA, M. PÉREZ, and J. L. CHICHARRO. Kinetics of V̇O2 in professional cyclists. Med. Sci.
Sports Exerc., Vol. 34, No. 2, pp. 320 –325, 2002. Purpose: To analyze the kinetics of oxygen uptake (V̇O2) in professional road
cyclists during a ramp cycle ergometer test and to compare the results with those derived from well-trained amateur cyclists. Methods:
Twelve professional cyclists (P group; 25 ⫾ 1 yr; maximal power output (Wmax), 508.3 ⫾ 9.3 watts) and 10 amateur cyclists (A group;
22 ⫾ 1 y; Wmax, 429.9 ⫾ 8.6 watts) performed a ramp test until exhaustion (power output increases of 25 watts·min⫺1). The regression
lines of the V̇O2:power output (W) relationship were calculated for the following three phases: phase I (below the lactate threshold
(LT)), phase II (between LT and the respiratory compensation point (RCP)), and phase III (above RCP). Results: In group P, the mean
slope (⌬V̇O2:⌬W) of the V̇O2:W relationship decreased significantly (P ⬍ 0.01) across the three phases (9.9 ⫾ 0.1, 8.9 ⫾ 0.2, and
3.8 ⫾ 0.6 mL O2·watts⫺1· min⫺1 for phases I, II, and III, respectively). No significant differences (P ⬎ 0.05) were found between
phases I and II (P ⬎ 0.05) in group A, whereas ⌬V̇O2:⌬W significantly increased in phase III (P ⬍ 0.01), compared with phase II (10.2
⫾ 0.3, 9.2 ⫾ 0.4, and 10.1 ⫾ 1.1 mL O2·watts⫺1· min⫺1 in phases I, II, and III, respectively). The mean value of ⌬V̇O2:⌬W for phase
III was significantly lower in group P than in group A (P ⬍ 0.01). Conclusion: Contrary to the case in amateur riders, the rise in V̇O2 in
professional cyclists is attenuated at moderate to high workloads. This is possibly an adaptation to the higher demands of their training/
competition schedule. Key Words: RAMP TEST, LACTATE, VENTILATION, HEART RATE, SLOW COMPONENT OF V̇O2

O
xygen uptake (V̇O2) tends to slowly rise during any
constant-load exercise of moderate to high intensity,
that is, above the lactate threshold (LT), which is the
point after which blood lactate shows a consistent, nonlinear
increase compared with baseline values (6). This phenomenon,
the “slow component of V̇O2,” has been defined as the con-
tinued rise in V̇O2 beyond the third minute of exercise (6). In
accordance with this phenomenon, the V̇O2:workload relation-
ship has been shown to become curvilinear after the LT has
been exceeded during incremental exercise tests (30). A non-
linear increase in V̇O2 has indeed been identified at high
workloads during this type of exercise protocol in healthy,
non-highly trained adults (2,12,29,30).
Several mechanisms could be involved, at least in part, in the
etiology of the slow component phenomenon, including car-
diorespiratory work, lactate and lactic acidosis, epinephrine,
K⫹, or patterns of muscle fiber recruitment. The metabolic cost
of increased ventilation (V̇E) (i.e., to buffer acidosis at high
workloads) can account for 18 –23% of the total V̇O2 slow
component (6). High values of V̇E (⬎100 L·min⫺1) can also
increase cardiac work through the pumping action of the lungs,
which in turn enhances venous return (15). Factors such as
catecholamines, lactate, and H⫹ can exert a stimulatory effect
at various sites (e.g., lactate may increase metabolic activity in
the liver and possibly in skeletal muscle through stimulation of
glycogenolysis) (6,23). Furthermore, the reduction in pH as-
0195-9131/02/3402-0320/$3.00/0
MEDICINE & SCIENCE IN SPORTS & EXERCISE®
Copyright © 2002 by the American College of Sports Medicine
Submitted for publication February 2001.
Accepted for publication May 2001.
320
sociated with lactic acidosis (as well as increased intramuscular
temperature) causes a rightward shift in the oxyhemoglobin
dissociation curve because of the Bohr effect and may thus
permit the continued V̇O2 increase (6). Rising muscle temper-
ature (i.e., the Q10 effect) and decreased efficiency of force
production because of metabolic accumulation (e.g., K⫹ and
H⫹) can also contribute to V̇O2 “wasting” (6). Finally, the
progressive recruitment of less efficient fast motor units (par-
ticularly those consisting of type IIX fibers) at high workloads
(because of fatigue of previously recruited fibers) has been
postulated as the main determinant of the V̇O2 slow component
(6). Although several authors have also explored the etiology
of the additional, nonlinear increase that occurs in V̇O2 during
incremental exercise (2,12,29,30), there is still some contro-
versy. For instance, Zoladz et al. (30) found no relationship
between lactic acidosis and the “excess” V̇O2 during gradual
cycle ergometer tests, whereas Jones et al. (12) documented a
causal relationship between lactic acidosis and the “excess”
V̇O2 during running tests. In addition, the occurrence of the
V̇O2 slow component phenomenon during heavy constant-load
exercise (80 –90% V̇O2max) has been shown in elite endurance
athletes (i.e., triathletes (3) or professional cyclists (15)),
whereas the occurrence of an additional, nonlinear increase in
V̇O2 during incremental exercise remains to be confirmed in
top-level endurance athletes.
It has been shown that professional road cyclists exhibit
several remarkable physiological adaptations, such as the
ability to perform at high workloads (about 90% V̇O2max)
over long periods of time (60 min or more), or a great
resistance to fatigue of slow motor units (17). In a recent
report, it was stated that the magnitude of the V̇O2 slow
component of professional riders during 20 min of cycling
exercise at 80% V̇O2max was only 130 mL in 17 min or 7.6
mL·min⫺1 (15), clearly below values (around 330 mL in 15
min or 22 mL·min⫺1) previously reported for cyclists work-
ing at similar relative intensities (6). Despite the high power
output sustained by professional cyclists (average of ~400
W), no electromyographic (EMG) evidence was found of
what appears to be the main determinant of the slow com-
ponent phenomenon in non-highly trained humans, that is,
an increased recruitment of fast motor units because of
fatigue of previously recruited slow ones. Such high resis-
tance to fatigue of slow motor units is probably attained
after years of intense training (~35,000 km·yr⫺1).
Three physiological gas exchange phases can be identi-
fied during rapid incremental exercise testing (24,27): phase
I, in which CO2 production (V̇CO2) comes mainly from
oxidative metabolism; phase II (“isocapnic buffering”), dur-
ing which V̇E increases in response to the rise in V̇CO2 from
buffering, with regulation of arterial partial pressure of CO2
(PaCO2); and phase III, in which respiratory compensation
for metabolic acidosis with lowering of PaCO2 (“hypocap-
nic hyperventilation”) occurs. The points that limit these
three phases are denoted ventilatory threshold (VT) or LT
(between phases I and II) and second ventilatory threshold
or respiratory compensation point (RCP) (between phases II
and III). The high workloads at which both VT and RCP
occur in professional cyclists (~65% and ~90% of V̇O2max,
respectively) and the marked difference between these val-
ues and those recorded in amateur cyclists (VT ~60% of
V̇O2max and RCP ~80% of V̇O2max, respectively) suggest
that these two variables might be an important performance
factor in endurance events such as professional road races (17).
Accordingly, exercise intensity is often approached as phases I,
II, and III, both for exercise prescription and for estimating the
physiological demands of cycling competition (17). Moreover,
previous studies dealing with the kinetics of physiological
variables during incremental exercise have also used this three-
segmental approach (9,10,21). It would therefore be of interest
to evaluate the V̇O2 kinetics of professional road cyclists across
these three phases during a rapid incremental protocol (i.e.,
ramp test), to determine if the V̇O2:workload relationship ex-
hibits an upward deflection at moderate to high workloads
(above LT, or phases II and III), as appears to occur in indi-
viduals of a lower fitness level.
The main purpose of this study was twofold: 1) to analyze
the kinetics of V̇O2 in professional cyclists across the work-
loads (phases I, II, and III) of an incremental (ramp) protocol;
and 2) to determine the influence of possible determinants of
this type of kinetics, namely, central factors (cardiorespiratory
work) and blood lactate. An additional goal was to compare the
results with those obtained during the same protocol in a group
of well-trained, amateur cyclists.
METHODS
Subjects
After giving their written informed consent, 12 top-level
professional road cyclists (group P) and 10 well-trained,
V̇O2 KINETICS IN PROFESSIONAL CYCLISTS
amateur cyclists (group A) were selected as subjects for this
investigation. Subjects in group P were required to fulfill the
following criteria at the time of the study: 1) a competition
experience of at least 2 yr in the professional category of the
Union Cycliste Internationale (UCI); 2) to have participated
in at least two of the main 3-wk stage races (Vuelta a
España, Giro d’Italia, or Tour de France); and 3) to have
won at least one event of the UCI professional competition
season within the last 2 yr. Several of the subjects in group
P were among the best cyclists in the world (including a
former world champion and winners of major professional
tour races). Subjects in group A were required to 1) be
enrolled in a licensed amateur team and to have a compe-
tition experience in the “sub23-elite” category of the UCI of
at least 2 yr. All were highly competitive in the amateur
category but had never competed in the professional cate-
gory. Subjects’ mean (⫾ SEM) age, height, and weight were
25 ⫾ 1 yr, 181.4 ⫾ 2.1 cm, and 70.4 ⫾ 1.5 kg, respectively
(group P); and 22 ⫾ 1 yr, 177.0 ⫾ 1.0 cm, and 69.9 ⫾ 1.6
kg, respectively (group A). The study protocol was ap-
proved by the institution’s (Universidad Complutense) re-
search ethics committee.
Exercise Protocol
Exercise tests. Subjects from each group performed an
exercise test (ramp protocol) until exhaustion on a cycle
ergometer (Ergometrics 900, Ergo-line, Barcelona, Spain).
All the tests were performed during the months December
and January under similar environmental conditions (20 –
24°C, 45–55% relative humidity). The subjects were cooled
with a fan throughout the bouts of exercise. This type of
protocol has been used for the physiological evaluation of
professional cyclists in several previous studies
(13,15,16,18). After a 2-min rest, sitting on the cycle er-
gometer, the test was started at 25 watts and the workload
was increased by 25 watts·min⫺1. Subjects adopted the
conventional sitting position during the whole duration of
the tests and were allowed to choose their preferred cadence
within the range 70 –90 rpm. This is known to better sim-
ulate actual cycling conditions compared with tests per-
formed at a fixed cadence. During actual racing (3-wk tour
races), the preferred pedaling cadence of professional riders
ranges from 70 rpm (hill climbs) to 90 rpm (flat terrain or
individual time trials) (16). A pedal-frequency meter was
used by the subjects to maintain this range of cadences, and
the mean pedaling cadence of each test was recorded. The
tests were terminated when pedal cadence could not be
maintained at 70 rpm (at least). Verbal encouragement was
given to the subjects to continue the test until they were
exhausted. All the participants had previous experience in
this type of testing.
During the tests, heart rate (HR, in beats·min⫺1) was
continuously recorded and gas exchange data were collected
continuously using an automated breath-by-breath system
(Mvmax 29C, SensorMedics, Yorba Linda, CA) to determine
V̇O2, V̇CO2, V̇E, ventilatory equivalent for oxygen (V̇E·
V̇O2⫺1) and carbon dioxide (V̇E·V̇CO2⫺1), and end-tidal
Medicine & Science in Sports & Exercise姞 321
 TABLE 1. Physiological variables (mean ⫾ SEM) recorded during the exercise tests
in amateur (group A) and professional (group P) cyclists.
Group P Group A
Maximal variables
Wmax 508 ⫾ 9* 430 ⫾ 9
V̇O2max (L䡠min–1) 5021 ⫾ 105 4855 ⫾ 136
V̇O2max 71.3 ⫾ 1.2 69.5 ⫾ 1.6
(mL䡠kg–1䡠min–1)
BLa (mM) 9.1 ⫾ 0.4 9.5 ⫾ 0.5
HR (beats䡠min–1) 196 ⫾ 3 197 ⫾ 2
V̇E (L䡠min–1) 200.3 ⫾ 5.3* 175.2 ⫾ 6.0
LT and RCP
LT (watts) 312 ⫾ 8* 234 ⫾ 12
LT (%Wmax) 61.4 ⫾ 1.8* 54.4 ⫾ 2.8
RCP (watts) 431 ⫾ 12* 336 ⫾ 10
RCP (%Wmax) 84.8 ⫾ 2.5* 78.2 ⫾ 4.0
Cadence (rpm) 78.7 ⫾ 1.3 77.1 ⫾ 1.3
V̇O2max, maximal oxygen uptake; Wmax, maximal power output in watts; BLa, blood
lactate; HR, heart rate; V̇E, pulmonary ventilation; LT, lactate threshold; RCP, respiratory
compensation point.
V̇O2max and V̇E values represent means for the last 60 s of the tests.
FIGURE 1—Mean regression lines of amateur (A) and professional * P ⬍ 0.05 for group P vs group A.
cyclists (P). Regression equations of A and P are shown in the upper
and lower part of the figure, respectively. LT, lactate threshold; RCP,
respiratory compensation point; Wmax, maximal power output; I, II,
and III, phases I (below LT), II (LT to RCP), and III (above RCP), LT; phase II, LT to RCP; and phase III, above RCP). In each
respectively. group, a one-way repeated-measures ANOVA was applied
to determine if there was a significant difference between 1)
phases I, II, and III in the mean slope (⌬V̇O2:⌬W) of the
V̇O2:W relationship; and 2) mean values of mechanical
partial pressure of oxygen (PETO2) and carbon dioxide
efficiency at LT, RCP, and Wmax, respectively. When a
(PETCO2).
significant difference was found in ANOVA tests, the post
Capillary blood samples (50 ␮L) for the measurement of
hoc Scheffé test was applied. A Student’s t-test for paired
blood lactate (BLa) were taken from fingertips (sampling
data was applied to compare the following variables in
period of 15–20 s) at rest, every 3 min throughout the test
groups A and P: 1) mean values of cadence and physiolog-
(e.g., at the end of the stages 75 W, 150 W, 225 W, etc.), and
ical parameters (e.g., V̇O2max, Wmax); 2) mean values of
immediately after termination of exercise. BLa was deter-
⌬V̇O2:⌬W obtained during phases I, II, and III, respec-
mined using an automated analyzer (YSI 1500, Yellow
tively; and 3) mechanical efficiency at VT, RCP, and Wmax.
Springs Instruments, Yellow Springs, OH). The LT was
Pearson product-moment correlation coefficients were
determined by examining the “lactate concentration-work-
calculated for both groups of subjects to determine whether
load (W)” relationship during the tests according to the
there was a significant relationship within each of the phases
methodology described by Weltman and et al. (28) (Fig. 1).
I, II, and III, between ⌬V̇O2:⌬W and the following slopes:
This method defines the workload (W) corresponding to LT
⌬HR:⌬W, ⌬V̇E:⌬W, ⌬V̇E·V̇O2⫺1:⌬W, and ⌬BLa:⌬W.
as the highest not associated with a rise in lactate concen-
These correlations were calculated to determine the possible
tration above baseline. This always occurred just before the
influence of cardiorespiratory factors and BLa on V̇O2 ki-
curvilinear increase in BLa observed at subsequent exercise
netics. The level of significance was set at 0.05 for all the
intensities. An increase of at least 0.5 mM BLa concentra-
statistical tests and results are expressed as means ⫾ SEM.
tion was required for the determination of the LT.
The power output (W) corresponding to the RCP was also
identified using the criterion of an increase in both RESULTS
V̇E·V̇O2⫺1 and V̇E·V̇CO2⫺1 and a decrease in PETCO2 (19).
Mean cadence and physiological variables recorded dur-
RCP was detected by two independent observers. If there
ing the tests are shown in Table 1. The workload (expressed
was disagreement, the opinion of a third investigator was
in W and in %Wmax) eliciting LT and RCP was significantly
sought. Mechanical efficiency (actual mechanical work ac-
higher in group P than in group A (P ⬍ 0.05).
complished (kg·m)/input of energy (kg·m) ⫻ 100) was cal-
Figure 1 shows the mean regression lines of the V̇O2:W
culated as described elsewhere (20) at the power output (W)
relationship in both groups. In group P, ⌬V̇O2:⌬W de-
corresponding to LT and RCP respectively, and at the max-
creased significantly (P ⬍ 0.01) across the three phases
imal power output (Wmax).
(Table 2). In group A, no significant differences (P ⬎ 0.05)
were found between phases I and II (P ⬎ 0.05), whereas
Data Analysis
⌬V̇O2:⌬W significantly increased in phase III compared
The regression lines of the V̇O2:W, HR:W, V̇E:W, with phase II (P ⬍ 0.01). On the other hand, the mean value
V̇E·V̇O2⫺1:W, and BLa:W relationships were calculated in of ⌬V̇O2:⌬W for phase III was significantly lower in group
each subject for the aforementioned phases (phase I, below P than in group A (P ⬍ 0.01).
322 Official Journal of the American College of Sports Medicine http://www.acsm-msse.org
TABLE 2. Slopes (⌬V̇O2:⌬W, in mL O2䡠watts–1䡠min–1) of the regression lines
corresponding to the tests in amateur (group A) and professional cyclists (group P).
Group P Group A
Phase I (below LT) 9.9 ⫾ 0.1 4 NS 3 10.2 ⫾ 0.3
Phase II (LT to RCP) 8.9 ⫾ 0.2* 4 NS 3 9.2 ⫾ 0.4*
Phase III (above RCP) 3.8 ⫾ 0.6** 4 P ⬍ 0.01 3 10.1 ⫾ 1.1
Data are presented as mean ⫾ SEM.
LT, lactate threshold; RCP, respiratory compensation point; NS, no statistical
significance (P ⬎ 0.05 for group P vs group A).
* P ⬍ 0.01 for phase II vs phase III; ** P ⬍ 0.01 for phase III vs both phases I
and II.
Mean values of mechanical efficiency in both groups are
shown in Table 3. Briefly, mechanical efficiency increased
in group P with increasing workloads (P ⬍ 0.01) in the
following order: VT ⬍ RCP ⬍ Wmax. In group A, no
significant differences were found between VT and RCP (P
⬎ 0.05), and mechanical efficiency significantly increased
at Wmax compared with both VT and RCP (P ⬍ 0.01 and P
⬍ 0.05, respectively). Mean values of mechanical efficiency
at both RCP and Wmax were significantly higher in group P
than in group A (P ⬍ 0.05 and P ⬍ 0.01, respectively).
Finally, no significant correlation was found between
⌬V̇O2:⌬W and ⌬HR:⌬W, ⌬V̇E:⌬W, ⌬V̇E·V̇O2⫺1:⌬W, and
⌬BLa:⌬W in groups P and A for each of the three phases.
DISCUSSION
The purpose of this study was to analyze the kinetics of
V̇O2 in professional cyclists during a ramp protocol and to
determine the possible influence of cardiorespiratory factors
and lactate on this phenomenon. We also analyzed V̇O2
kinetics in a group of well-trained, amateur cyclists of a
lower fitness level. In line with previous research, three
physiological gas exchange phases were considered (i.e.,
below LT, LT to RCP, and above RCP (9,10,21)). Our main
finding was that, in professional riders, the rate of the V̇O2
rise elicited by gradual exercise (ramp protocol) decreases at
moderate to high workloads (i.e., from the LT (~300 watts)
to the maximal attainable power output (~500 watts)). Sim-
ilarly, mechanical efficiency seemed to increase with rising
exercise intensity. To the best of our knowledge, very few
data from top-level endurance athletes have been generated
in this field of research, and neither have findings related to
V̇O2 kinetics at such high power outputs (from ~300 to ~500
watts) been published to date. In contrast, the results of
previous research on subjects of a much lower fitness level
have shown a nonlinear increase in V̇O2 at high workloads
TABLE 3. Mechanical efficiency (%) recorded in amateur (group A) and professional
(group P) cyclists at the power output (W) eliciting the LT and the RCP and
at Wmax.
Group P Group A
LT 24.4 ⫾ 0.7 4 NS 3 23.6 ⫾ 0.6
RCP 26.1 ⫾ 0.5* 4 P ⬍ 0.05 3 24.3 ⫾ 0.5
Wmax 28.8 ⫾ 0.6** 4 P ⬍ 0.01 3 25.6 ⫾ 0.7†
Data are presented as mean ⫾ SEM.
NS, no statistical significance (P ⬎ 0.05 for group P vs group A).
* P ⬍ 0.01 for RCP vs LT; ** P ⬍ 0.01 for Wmax vs both RCP and LT; † P ⬍ 0.05
and P ⬍ 0.01 for Wmax vs RCP and for Wmax vs LT, respectively.
V̇O2 KINETICS IN PROFESSIONAL CYCLISTS
during progressive exercise (from the LT to the maximal
workload) (2,12,29,30). This “excess” V̇O2 has recently
been designated the “slow component of V̇O2” by Jones et
al. (12), on the basis of an analogy with the phenomenon
that occurs during heavy, constant-load exercise. Moreover,
in group P, the slope of the V̇O2 rise below the LT (~10 mL
O2·watts⫺1·min⫺1) was within “typical” limits for healthy,
non-highly trained adults during incremental cycle ergome-
ter tests (i.e., ~9 –11 mL O2·watts⫺1·min⫺1 (22)), whereas it
was below these limits after the LT was reached (mean V̇O2
rise of 8.9 and 3.8 mL O2·watts⫺1·min⫺1 in phases II and III,
respectively). In fact, the V̇O2 kinetics recorded at high
workloads in our group of amateur cyclists was closer to that
expected to occur in healthy, non-highly trained subjects
than to that shown by professional riders; amateur cyclists
showed a mean V̇O2 rise of ⬎ 9 mL O2·watts⫺1· min⫺1 at
each workload, and ⌬V̇O2:⌬W significantly increased
above the RCP. A further finding was that V̇O2 kinetics in
both groups were not significantly influenced by lactate or
cardiorespiratory work.
As mentioned above, the occurrence of a deflection in the
V̇O2:W relationship above the LT in group P differs from
that previously described during gradual exercise in subjects
of a lower fitness level (2,12,29,30). The ramp protocol
chosen for the present investigation (workload increases of
25 watts·min⫺1) and also for numerous studies performed
on elite cyclists (13,15,16,18) differs from that used in
previous research in the area of V̇O2 kinetics (i.e., workload
increases every 3– 4 min (3,12,29,30)). Nevertheless, in a
recent experiment in our laboratory (unpublished data), we
also noted a significant “excess” of V̇O2 in sedentary, young
men during high ramp protocol workloads comparable to
those applied here (workload increases of 5 watts·15 s⫺1,
averaging 25 watts·min⫺1). The results recorded for group P
reflect one of the main adaptations to professional road
cycling compared with amateur cycling—that is, a greater
mechanical efficiency at high workloads—which may ex-
plain their higher performance level. Indeed, the mean
V̇O2max values corresponding to groups A and P were very
similar (~70 mL·kg⫺1·min⫺1) despite the greater perfor-
mance of the latter (i.e., considerably higher values of both
maximal power output and power output at the RCP). We
recently determined V̇O2 kinetics in a group of professional
riders of similar characteristics during a 20-min constant-
load cycle ergometer test at 80% V̇O2max (15). Despite the
high average power output sustained by the professional
riders (~400 watts), the average magnitude of the slow
component (130 mL in 17 min or 7.6 mL· min⫺1) was
considerably lower than that reported by previous research
(22 mL·min⫺1) on the basis of constant-load cycle ergome-
ter tests performed at similar relative intensities (6). This
finding is also suggestive of a great cycling efficiency of
professional riders, which is thought to contribute to their
renowned ability to sustain extremely high workloads over
long periods, as shown by previous field research (e.g.,
average power output ⬎ 400 watts during the 1-h record in
a velodrome over the past decade (17)). Professional riders
show considerable resistance to fatigue of recruited motor
Medicine & Science in Sports & Exercise姞 323
units, at least at high, submaximal intensities (17,18). Such
an adaptation is probably attained after years of highly de-
manding training and competition (i.e., ~35,000 km·yr⫺1 and
~90 competition days) as suggested in a previous study in
which the physiological response of professional cyclists was
compared with that of their elite, amateur counterparts (18).
Ramp tests involving gradual, constant workload in-
creases such as the one used here may be the most suitable
for determining the possible influence of motor unit recruit-
ment patterns on V̇O2 kinetics, since they can reproduce the
hierarchy of muscle fiber recruitment that is likely to occur
with rising exercise intensity in humans, that is, type I
(phase I) 3 type IIA (phase II) 3 type IIX (phase III), as
shown by previous EMG (25) and biopsy studies (26).
Given that 1) the nonlinear increase in V̇O2 after the LT is
mostly linked to the recruitment of type II fibers in non-
highly trained humans (6) and that 2) subjects in group P
failed to show this response, one may speculate that the
latter may have a greater proportion of type I fibers in the
main muscles involved in cycling than their amateur coun-
terparts. Previous research performed on amateur, well-
trained cyclists has indeed shown that cycling efficiency
during heavy exercise (above LT) is positively related to the
percentage of type I fibers in the vastus lateralis muscle
(11). As mentioned before, a higher resistance to fatigue of
slow motor units in group P could have also been involved,
and would allow these professional riders to reach moderate
to high workloads (i.e., between LT and RCP, or ~70 –90%
V̇O2max) before significant recruitment of the less-efficient
type II fibers. However, it must be also kept in mind that the
relationship between the rise in V̇O2 and the rise in power
output described here and in previous research involving
gradual exercise tests (2,12,29,30) does not directly reflect
the efficiency of muscle contraction (i.e., myosin cross-
bridge cycling, which is directly related to force production)
but rather the increase in aerobic metabolism as muscle
power output increases (2). Indeed, other ATP-dependent,
non-crossbridge activities termed “activation heat” (i.e., cal-
cium uptake and release from the sarcoplasmic reticulum)
could also affect V̇O2 kinetics during gradual exercise.
Nevertheless, it remains unclear whether activation heat
varies or remains relatively constant during fatiguing exer-
cise (1). Thus, it is not possible to determine to what extent
the attenuation of the V̇O2 rise exhibited by group P at high
intensities solely reflects a higher efficiency of muscle con-
traction, a smaller contribution of activation heat compared
with subjects of a lower fitness level, or both.
Several potential contributors to V̇O2 kinetics in non-
highly trained humans (i.e., cardiorespiratory work, and
lactate and/or lactic acidosis) seemed to play a minor role in
our subjects. In non-highly trained individuals, the V̇O2 cost
of cardiorespiratory work might partially contribute to the
slow component seen during exercise above the LT (4,7).
Pulmonary ventilation and HR significantly increased in
group P to reach high values at the end of the tests (V̇E ~200
L·min⫺1 and HR ~190 beats·min⫺1), but such high cardio-
respiratory work did not seem to influence V̇O2 kinetics.
Indeed, we found no correlation between ⌬V̇O2:⌬W and
324 Official Journal of the American College of Sports Medicine
any of the variables related to cardiorespiratory work (⌬HR:
⌬W, ⌬V̇E:⌬W, or ⌬V̇E·V̇O2⫺1:⌬W). It may be speculated
that the deflection of the V̇O2 rise recorded in group P as
opposed to both amateur riders and nontrained subjects is
partly attributable to more efficient cardiac function during
moderate- to high-intensity exercise (above LT). Indeed, it
was recently shown that approximately two thirds of pro-
fessional cyclists exhibit a deflection in the HR:W relation-
ship at high intensities (~RCP) during a ramp protocol,
which is partly determined by their great myocardial wall
thickness (14). Professional riders, on the other hand, have
been shown to exhibit a characteristic breathing pattern at
the high workloads of a ramp protocol (i.e., lack of “tachy-
pneic shift”) (13). This breathing adaptation might allow
their respiratory muscles to work more efficiently in both
mechanical and metabolic terms (13), and could thus at least
partly account for the V̇O2 kinetics shown by these subjects.
In fact, the oxygen cost of breathing in highly fit individuals
is not negligible (close to 15% V̇O2max) (8). Furthermore,
the work of breathing during heavy exercise compromises
leg blood flow to working limb muscles, possibly because of
sympathetically mediated vasoconstriction originating in the
respiratory musculature (8). In the event of a more efficient
breathing pattern (i.e., subjects in group P), blood flow to
working limbs may not be significantly reduced by the
aforementioned mechanism. This in turn would lead to a
later recruitment of type II fibers, which predominantly
undertake anaerobic metabolism and are less efficient than
type I fibers (5). Finally, we found no correlation between
V̇O2 and lactate kinetics in any of the three phases. These
results suggest that lactate per se does not enhance whole-
body O2 in highly fit endurance athletes during gradual
exercise. In contrast, findings related to non-highly trained
humans provide evidence for the involvement of lactate
and/or lactic acidosis in the V̇O2 slow component that
occurs at high exercise intensities (above LT) in both con-
stant-load (23) and incremental protocols (12).
In conclusion, the rate of the V̇O2 rise elicited by gradual
exercise (ramp protocol) decreases at moderate to high
workloads (from LT to maximal power output) in profes-
sional cyclists. This response is different to that shown by
most individuals (including well-trained cyclists of a lower
performance level) and probably reflects one of the main
adaptations to professional road cycling, that is, a great
ability to tolerate high workloads (at or above the RCP) over
long periods of time (⬎30 min) before fatigue occurs. Our
findings suggest the possibility of using V̇O2 kinetics during
“routine” incremental testing of elite endurance athletes, as an
indicator of both performance and training adaptation (11).
We thank José Ramón Echevarrı́a for his assistance during the
exercise tests and Ana Burton for her linguistic assistance. This
study was financed by Agrupación Deportiva Banestro. The present
study has been awarded with the Pimer Premio Nacional de Inves-
tigación en Medicina del Deporte 2001 of Universidad de Oviedo,
Spain.
Address for correspondence: Alejandro Lucı́a, M.D., Ph.D.,
Departamento de Ciencias Morfológicas y Fisiologı́a, Universidad
Europea de Madrid, E-28670 Madrid, Spain; E-mail: alejandro.
lucia@mrfs.cisa.uem.es.
http://www.acsm-msse.org
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