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Eur J Appl Physiol (2007) 101:359368

DOI 10.1007/s00421-007-0506-8

ORIGINAL ARTICLE

Effects of intermittent hypoxic training on cycling performance


in well-trained athletes
Belle Roels David J. Bentley Olivier Coste
Jacques Mercier Gregoire P. Millet

Accepted: 9 June 2007 / Published online: 17 July 2007


 Springer-Verlag 2007

Abstract This study aimed to investigate the effects of a program, three cycling tests including an incremental test
short-term period of intermittent hypoxic training (IHT) on to exhaustion in normoxia and hypoxia for determination
cycling performance in athletes. Nineteen participants were of maximal aerobic power VO _ 2 max and peak power out-
randomly assigned to two groups: normoxic (NT, n = 9) put (PPO) as well as a 10-min cycle time trial in normoxia
and intermittent hypoxic training group (IHT, n = 10). A (TT) to measure the average power output (Paver). No
3-week training program (5 1 h1 h 30 min per week) _ 2 max was observed between the
significant difference in VO
was completed. Training sessions were performed in two training groups before or after the training period.
normoxia (~30 m) or hypoxia (simulated altitude of When measured in normoxia, the PPO significantly in-
3,000 m) for NT and IHT group, respectively. Each subject creased (P < 0.05) by 7.2 and 6.6% in NT and IHT groups,
performed before (W0) and after (W4) the training respectively. However, only the IHT group significantly
improved (11.3%; P < 0.05) PPO when measured in hy-
poxia. The NT group improved (P < 0.05) Paver in TT by
B. Roels  O. Coste
8.1%, whereas IHT group did not show any significant
UPRES EA 3759Multidisciplinary Approach of Doping,
700 avenue Pic St Loup, 34090 Montpellier, France difference. Intermittent training performed in hypoxia was
less efficient for improving endurance performance at sea
B. Roels level than similar training performed in normoxia. How-
Centre of Sport Medicine and Human Performance,
ever, IHT has the potential to assist athletes in preparation
Brunel University, School of Sport and Education,
London UB8 3PH, UK for competition at altitude.

D. J. Bentley Keywords Aerobic  Altitude  Endurance  Power 


Department of Human and Health Science,
Adaptation
University of Westminster, London, UK

D. J. Bentley
Health and Exercise Science, School of Medical Sciences, Introduction
University of New South Wales, Sydney, Australia

O. Coste Over the last few decades many athletes and coaches have
Direction Regionale et Departementale de la Jeunesse et des used altitude training in various forms to help improve
Sports, 190 avenue du Pe`re Soulas, 34094 Montpellier, France performance at altitude and/or at sea level. The traditional
approach to altitude training was for athletes to live and
J. Mercier
Laboratoire de physiologie des Interactions EA 701, train at moderate altitude. The effects of this form of
Institut de Biologie, Bvd Henri IV, 34060 Montpellier, France stimulus on endurance performance have been researched
extensively (Fulco et al. 2000; Levine 2002; Wilber 2001).
G. P. Millet (&)
A recent approach has been for athletes to live and sleep at
ASPIRE, Academy for Sports Excellence,
PO Box 2287, Doha, Qatar altitude and train near sea level, the so-called live high-
e-mail: gregoire.millet@aspire.qa train low (LHTL) method or the opposite live low-train

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360 Eur J Appl Physiol (2007) 101:359368

high (LLTH) method, to live and sleep at sea level and (2005) also showed that 4 weeks of interval training under
train at altitude (Wilber 2001). Because the geography of hypoxia conditions, i.e. simulated altitude of 3,000 m for
many countries does not allow LHTL or LLTH, other approximately 114 min per week, did not induce a greater
strategies have been developed for athletes, such as being increase in performance compared to similar normoxic
briefly exposed to hypoxia. Intermittent hypoxic exposure training; moreover, no changes in haematological variables
with (IHE) or without (IHT) exercise training is based on were observed.
the assumption that brief exposure to hypoxia (minutes to Zoll et al. (2006) found that specific muscular transcript
hours) is sufficient to stimulate EPO release, and ultimately level adaptations in the vastus lateralis participated in
increase red blood cell (RBC) concentration (Rusko 1996; enhancing endurance performance after a 6-week IHT with
Wilber 2001) and to induce peripheral modifications in two sessions performed at VT2 per week, in long-distance
skeletal muscle that in turn might increase performance runners. Indeed, combining a hypoxic stimulus with
(Dufour et al. 2006; Roels et al. 2007). Rodriguez et al. endurance training resulted in transcriptional adaptations of
(1999) found that hypoxic exposure (4,0005,000 m) for the vastus lateralis muscle in athletes and in turn a sig-
only 90 min, three times a week over 3 weeks, signifi- nificant increase in time to exhaustion and VO _ 2 max : The
cantly increased hematocrit (Hct), RBC, reticulocytes same research group (Ponsot et al. 2006) also found that
(RET) and haemoglobin (Hb) concentration. However, no this 6-week IHT of 2 sessions at VT2 in well-trained ath-
control group (training in normoxia) was included. Dufour letes, qualitatively improved the mitochondrial function by
et al. (2006) found that a 6-week IHT with two weekly increasing the respiratory control by creatine, providing a
high-intensity (second ventilatory threshold; VT2) and tighter integration between ATP demand and supply.
moderate-duration (2440 min) training sessions, enhanced Therefore, they suggested that adaptations of mitochondrial
endurance performance of competitive distance runners. function accompany the increase in endurance performance
The IHT group significantly improved VO _ 2 max (+5%) both after IHT. Geiser et al. (2001) investigated if IHT induces
in normoxic and hypoxic conditions, without any changes greater improvements than the same intensity training in
in blood O2-carrying capacity. Also, the time to exhaustion normoxia and also if there was a difference in the effect of
increased for the IHT group only (+35%), without signifi- high- versus moderate-intensity training on aerobic per-
cant changes in the VO _ 2 kinetics. This study suggested that formance variables. Previously untrained subjects were
IHT induces peripheral muscle adaptations, which result in assigned to four groups, Norm-high, Norm-low, Hyp-high
an enhanced endurance performance. Aside from this and Hyp-low. The training consisted of 30 min a day of
investigation data regarding the effects of IHE or IHT on cycling, 5 days per week for 6 weeks at an intensity of
haematological and muscular variables and endurance 80% and 67% of VO _ 2 max for the high and low intensity
performance are inconclusive (Dufour et al. 2006; Geiser at groups, respectively, and this under normoxic (600 m) or
al. 2001; Rodriguez et al. 1999, 2000, 2003; Roels et al. hypoxic (3850 m) conditions for the normoxic or hypoxic
2005, 2007; Terrados et al. 1988; Truijens et al. 2003). groups, respectively. They found no significant differences
Katayama et al. (2003) investigated the effect of a 3- in performance enhancement between IHT and the similar
week IHE program of 90 min a day, three times a week at normoxic protocol. However, IHT increased the endurance
simulated hypobaric hypoxia of 4,500 m in trained runners performance to a greater extent when measured under
and found that the sea-level 3,000 m running time and the hypoxic conditions. In addition, the training intensity had
time to exhaustion during a maximal exercise test was no effect on the gain of VO _ 2 max : However, the Hyp-high
significantly enhanced, while there was no change in the group was the most effective group to increase the muscle
resting haematological parameters. Therefore, IHE could oxidative capacity.
improve endurance performance at sea level in trained The effects of high intensity interval training performed
runners. In addition, Hendriksen and Meeuwsen (2003) in hypoxia on endurance performance in hypoxia in trained
observed an improvement in maximal power output and athletes has not yet been clearly established and could be a
mean and peak anaerobic power at sea level 9 days after factor in the adaptive response to simulated altitude train-
the end of a 10-day intermittent hypoxic training period at ing (Geiser et al. 2001).
a simulated altitude of 2,500 m (105 min per day). No Collectively data regarding the effects of IHT on
significant differences were observed after the end of a endurance performance measured in hypoxia and normoxia
similar training performed at sea level. In contrast, 5 weeks are minimal and inconclusive in trained athletes. Therefore,
of intermittent hypoxic training (simulated altitude of the present study reports on the effects of a 3-week inter-
2,500 m) in a swimming flume improved sea level per- mittent hypoxic training, including two high intensity
formance (100- and 400-m freestyle time trials) and interval sessions and three continuous training sessions per
_ 2 max : However, no additional effect of a hypoxia
VO week, on sea level and hypoxic cycling performance. We
stimulus was reported (Truijens et al. 2003). Roels et al. hypothesized that IHT would increase cycling performance

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Eur J Appl Physiol (2007) 101:359368 361

both at sea level and in hypoxia more than a similar were performed in normoxia (PIO2 of 160 mmHg) or in a
normoxic training. hypoxic (PIO2 of 100 mmHg, simulated altitude of
~3000 m) environment for NT and IHT group, respec-
tively. Average duration of the hypoxic stimulus per week
Methods during the training period was 382 min. No supplementary
interval training outside the two supervised interval-train-
Subjects ing sessions was performed. Each training session was
meticulously controlled by the same researcher. Table 2
Nineteen healthy male endurance-trained athletes (cyclists presents the workloads of each group averaged over
and triathletes) participated in the study, which was ap- 3 weeks for the interval training and continuous workload
proved by the institutional ethics committee (Nmes, sessions. No significant differences in (relative) training
France). All subjects gave written informed consent and workloads during the training sessions were observed be-
were sea-level residents. Each subject was familiarized tween the groups.
with the testing protocols and equipment used in the Before (W0) and after (W4) the training program, a
experiment. None of the subjects had a history of recent medical examination and determination of physical char-
travel to altitude, i.e. 3 weeks prior to training. Subjects acteristics was completed (Table 1). Subjects also per-
were initially randomized into two groups: an intermittent formed an incremental exercise test to exhaustion, and a
hypoxic training group (IHT; n = 10), and a normoxic laboratory TT. The incremental test was performed in both
training group (NT, n = 9). Groups were of comparable normoxic and hypoxic conditions. In contrast, TT was
training status in terms of average power output and performed only in normoxia. Blood samples at rest were
average oxygen consumption measured during a laboratory obtained at W0 and W4, and repeated at the end of each
time-trial (TT) prior the experimental period. Subject training week (W1W3).
characteristics for each group are presented in Table 1. One Iron supplementation (Oligo-essentiels, Laboratoires
athlete of the NT group did not complete the study due to Richelet, Paris, France) of 1 ml per day (i.e. 5 mg per day)
illness. was initiated in both groups 2 weeks before the beginning
of the experiment and continued during the training to
prevent low iron state which could limit erythropoiesis. A
Experimental design
physician was in attendance at all times and was respon-
sible for the safety of the subjects during the study.
The endurance-training program consisted of two interval
The weekly training regime, outside the training pro-
and three continuous training sessions per week for
gram was controlled and documented.
3 weeks. The continuous sessions consisted of 60 min at
60% of VO _ 2 max : The interval-training required subjects to
Training outside experimental design
perform two sets of three repetitions of 2 min duration at
an intensity of 100% of PPO (measured in either hypoxia
The training completed outside the experimental design
or normoxia). Two minute of rest was allowed between
was recorded daily using a computerized training diary
each repetition with 6 min rest between each set. Each
during the 3 weeks of training period. Briefly, the type of
training session began and ended with 15 min warm-up and
activity, i.e. cycling, swimming, running, etc., and the
15 min cool-down. Training sessions were performed on
intensity was recorded with this training diary. The training
subjects own bicycle, fixed on an electromagnetic resistant
intensity was divided into five intensity levels (Mujika
home-trainer (Elite Travel, Milan, Italy). Training sessions

Table 2 The averaged percentage of target power output (target PO),


i.e. 100% PPO for interval sessions and 60% of VO2max and the
averaged absolute power output (Wabsolute; W) for continuous training
Table 1 Subjects characteristics sessions, of normoxic training (NT, n = 8) and intermittent hypoxic
W0 W4 training (IHT; n = 10) groups

NT IHT NT IHT Continuous Interval


% of Wabsolute % of Wabsolute
Age (years) 24.2 0.4 24.4 0.3 target PO target PO
Height (cm) 181.6 0.7 180.1 0.5
Weight (kg) 71.3 0.9 73.2 0.8 72.0 0.8 72.6 0.8 NT 91.7 0.6 185.0 7.5 91.0 0.7 310.9 4.5
IHT 91.8 1.6 130.0 0.3 91.8 1.0 259.4 0.3
Mean SE; W0: pre; W4: post; NT: normoxic training group
(n = 8); IHT: intermittent hypoxic training group (n = 10) Mean SE

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362 Eur J Appl Physiol (2007) 101:359368

et al. 1996). All training sessions outside the protocol were oxygen uptake VO _ 2 max ; ml kg1 min1 ; peak power
individually timed and each exercise categorized according output (PPO; W), maximal ventilation VE _ max ; 1 min1 ;
to the five intensity levels. The performed training duration and maximal heart rate (HRmax; bpm). Test began at an
was multiplied by its corresponding multiplying factor, i.e. initial power output of 60 W for 3 min and then the
2, 4, 6, 10 and 16, respectively, and the sum was then workload was increased by 30 W every minute until
divided by the overall duration of the session to calculate exhaustion. Exhaustion coincided with heart rate (HR)
the average intensity of each training session. The recorded approaching an age predicted maximum value (220age); a
parameters were the number and average intensity of the plateau in VO _ 2 despite an increase in mechanical power
sessions, calculated according to Mujika et al. (1996) as it output; and an RER > 1.1. Breath-by-breath data were re-
was not sensible to compare hourly volume as the subjects duced to 30-s averages and VO _ 2 max was determined as the
performed exercise training in different modes (swimming, highest 30-s VO _ 2 average. PPO was defined as the highest
running). Table 3 presents the average intensity of addi- mechanical power maintained for 1 min.
tional training outside the experimental design of each A cycle TT was performed to determine the highest
training week for both groups. No significant differences in power output over a period of 10 min in normoxia. The TT
(relative) training workloads outside the training sessions was performed to assess the influence of training on
were observed between the groups. endurance performance in normoxia by determining the
mechanical power output (averaged power over the 10 min
Environmental stimulus TT: Paver; W), percentage of maximal oxygen consumption
%VO _ 2 max ; i.e. the highest 30 s averaged VO _ 2 measured
A simulated altitude of 3,000 m was used because this during the TT expressed as a percentage of the VO _ 2 max
level of hypoxia was sufficient for erythropoietic stimula- measured during the incremental test to exhaustion in
tion, and still well tolerated by subjects (Villa et al. 2005). normoxia, peak ventilation VE _ peak ; 1 min1 ; and peak
Hypoxic gas mixture was delivered continuously by a heart rate (HRpeak, bpm). Subjects were informed of the
system, which enriches the inspired air by modifying elapsed time but received no feedback on power output or
nitrogen content (Altitrainer 200, SMTEC, Geneva, on performance.
Switzerland). Briefly, O2 content of the mixture is contin- In this 10-min cycle, TT is considered to be long enough
uously displayed and can be expressed either by equivalent to obtain an aerobic contribution for energy provision, can
altitude (~3,000 m) or oxygen partial pressure be easily administered and has been used in other studies
(PIO2~100 mmHg), taking into account barometric pres- (Roels et al. 2005).
sure. Users inhale the mixture contained in the tank All the tests were performed on a bicycle equipped with
through a Hans Rudolph two-way respiratory valve. a SRM road professional powermeter (Schoberer Rad
Respiratory mask could at all times be removed and so Messtechnik, Julich, Welldorf, Germany). Saddle height on
subjects found themselves immediately in normoxic con- cycle ergometer measured during the first test was kept
ditions. A breath-by-breath analyzer can be easily attached identical for the remaining tests. Power output and pedaling
to the system in order to measure respiratory exchange. cadence were recorded with an acquisition frequency of
This device has been used previously by our laboratory and 1 s. Averaged values of 30-s were stored. Calibration of the
further details can be found in Roels et al. (2005). powermeter occurred as for the manufacturers recommen-
dations.
Performance tests
Physiological measurements
An incremental test to exhaustion was performed in
normoxic and hypoxic conditions to determine maximal Pre (W0) and post training (W4), respiratory exchange
measurements were made using a K4b2 (Cosmed, Rome,
Italy). K4b2 setup was changed in order to calculate ven-
Table 3 The average intensity of additional training outside the tilatory variables with the accurate FIO2 rather than with
experimental design of each training week for normoxic training (NT; the default version. Aforementioned physiological vari-
n = 8) and intermittent hypoxic training (IHT; n = 10) groups ables were measured, breath-by-breath, and averaged every
W1 W2 W3 30 s. Before each test, the system was calibrated using
ambient air, whose partial O2 composition was assumed to
NT 3.1 0.05 3.1 0.06 3.2 0.1 be 20.93% and a gas of known CO2 (5%) and O2 (16%)
IHT 2.9 0.09 3.0 0.05 3.0 0.07 concentration. Calibration of turbine flowmeter of the K4b2
Mean SE; W1: first week of training protocol; W2: second week of was performed with a 3-l syringe (Quinton Instruments,
training protocol; W3: third week of training protocol Seattle, WA). During the different tests and training ses-

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Eur J Appl Physiol (2007) 101:359368 363

sions, HR was constantly recorded by means of a HR Results


monitor (S810, Polar, Kempele, Finland).
Incremental test to exhaustion
Haematological variables
There were no significant differences in the initial PPO and
Blood samples at rest were obtained by venapuncture by a _ 2 max measured in normoxia or hypoxia between the two
VO
physician at W0 and W4. Samples were analyzed in the experimental groups. Physiological variables obtained
2 hours following collection using a blood analyzer (Pentra from the incremental test performed in normoxic and
120 Retic, abx, Montpellier, France) for RBC (106 mm3), hypoxic conditions are presented in Table 4. PPO signifi-
Hb (g dl1), Hct (%), mean cell volume (MCV; lm3), cantly increased after training (P < 0.001; F = 15.0) for
averaged globular content of hemoglobin (AGCH; pg), both groups when measured in normoxic conditions but no
averaged corpuscular concentration of haemoglobin differences were found between groups. PPO values were
(ACCH; g dl1), red blood disc (RBD; 103 mm3), reticu- 341.7 3.5 and 339.0 4.9 W at W0 and 366.3 3.2 and
locytes (RET; % and 106 mm3) and averaged reticulocytar 361.5 4.4 W at W4 for NT and IHT group respectively.
volume (ARV, lm3). In hypoxic conditions, only the IHT group increased
(P < 0.05; F = 8.09) PPO (11.3%) from 282.5 3.4 to
Statistical analysis 314.5 3.7 W, respectively, at W0 and W4. No significant
differences were found in the NT group: 301.1 2.5 to
All values are reported as mean standard error (SE). The 313.8 1.9 W at W0 and W4, respectively.
effects of the two training methods (IHT vs. NT) on
measured variables was, after analysis of normality and Cycle time-trial
homogeneity of variance of the tested samples, analyzed
using a two-way (training group time) variance (ANO- At the start of the training period there were no significant
VA) with repeated measures on the second factor. Signif- differences between groups in terms of average power
icant effects were subsequently analyzed using Student output (Paver) during TT. Only the NT group improved
Newman-Keuls post-hoc test. The ratio of true variance (P < 0.05; F = 7.05) Paver by 8.1%. Paver values were
over error variance (F) was calculated. Pearson correlations 259.1 5.0 W at W0, and 280.1 5.8 W at W4, whereas
were also completed to indicate significant correlations IHT group did not show any significant difference
between the performance parameters. All analyzes were (255.3 4.1 to 257.5 4.0 W at W0 and W4, respec-
completed using SigmaStat 2.3 (Jandel Corporation, San tively). There were no significant differences for the other
Rafael, CA) and statistical significance was accepted at physiological variables measured during the cycle TT
P < 0.05. between each training group (Table 5).

Table 4 Incremental test to exhaustion under normoxic and hypoxic condition


Normoxia Hypoxia
NT (n = 8) IHT (n = 10) NT (n = 8) IHT (n = 10)
W0 W4 W0 W4 W0 W4 W0 W4

PPO 341.7 3.48 366.3 3.2** 339.0 0.48 361.5 4.41** 301.1 2.45 313.8 1.89 282.5 3.41 314.5 3.67 *
(W)
_ 2 max
VO 58.1 0.83 61.0 1.15 58.5 0.7 58.3 0.59 47.6 0.81 48.8 1.08 48.3 0.94 48.6 0.64
(ml kg1 min1)
HRmax 190.1 1.1 188.3 1.54 189.7 1.1 189.4 0.97 187.6 1.44 187.4 1.15 188.2 0.96 188.9 1.06
(bpm)
_ max
VE 159.9 2.6 174.3 2.5 154.2 1.4 155.0 1.74 151.2 3.26 167.5 3.38* 140.6 2.27 142.5 1.91

(l min1)

RPE 17.9 0.16 17.9 0.14 16.4 0.15 16.9 0.16 17.7 0.26 18.0 0.21 17.5 0.16 17.8 0.15
Mean SE; W0: pre; W4: post; NT: normoxic training group; IHT: intermittent hypoxic training group; PPO: peak power output; VO _ 2 max :
highest value of the oxygen consumption averaged over 30 s; HRmax: highest value of heart rate averaged over 30 s; VE _ max : highest value of
ventilation averaged over 30 s; RPE: rate of perceived exertion; *: P < 0.05, **: P < 0.001 for the differences within a group versus W0; :
P < 0.05 for the differences between groups at a matched time point

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364 Eur J Appl Physiol (2007) 101:359368

Table 5 Cycle time trial


NT (n = 8) IHT (n = 10)
W0 W4 W0 W4

Paver (W) 259.1 5.0 280.1 5.8 * 255.3 4.1 257.5 4.0
_ 2 max
%VO 99.1 0.9 96.9 1.4 94.6 1.1 95.6 0.8
HRpeak (bpm) 182.9 1.5 179.6 1.8 186.2 1.2 183.7 0.8
_ peak (l min1)
VE 145.2 2.5 136.9 2.4 144.8 2.5 142.4 1.8
RPE 17.7 0.1 17.0 0.2 16.9 0.1 16.5 0.1
Mean SE; W0: pre; W4: post; NT: normoxic training group; IHT: intermittent hypoxic training group; Paver: Average 10-min power output;
%VO_ 2 max : the maximal oxygen consumption averaged over 30 s measured during the time trial expressed as a percentage of the maximal
oxygen consumption averaged over 30 s measured during the incremental test to exhaustion under normoxia; HRpeak: highest value of heart rate
_ peak : highest value of ventilation averaged over 30 s; RPE: rate of perceived exertion; *: P < 0.05 for the differences
averaged over 30 s; VE
within a group versus W0

Haematological variables difference was observed between the experimental hypoxic


group and a control normoxic group, i.e. no additional
Haematological variables measured during the training effect of the hypoxic stimulus. In addition, Roels et al.
period are represented in Table 6. No significant differ- (2005) concluded that IHT of two high-intensity cycling
ences were found between the two training groups, before, sessions per week (10090% relative peak power output,
during or after the experimental period. identical to the present high-intensity sessions) for 7 weeks
at a simulated altitude of 3,000 m for well-trained cyclists
and triathletes did not improve performance at sea level,
Discussion i.e. PPO and 10-min cycle time trial nor VO _ 2 max to a
greater extent than a similar sea level training. Similarly,
The present study showed that two interval and three Truijens et al. (2003) found that 5 weeks of intermittent
continuous training sessions per week performed over a hypoxic training at a simulated altitude of 2,500 m, in a
period of 3 weeks significantly improved PPO measured in swimming flume improved sea-level performance in well-
normoxia and cycle time-trial performance. However, trained swimmers, but no significant additive effect of
these improvements were not further enhanced by per- hypoxic training was observed. Geiser et al. (2001) have
forming the same training in hypoxic conditions. Specifi- also shown, in previously untrained subjects, that IHT of
cally the findings of the present study demonstrate that 6 weeks, with five 30 min sessions per day at 3,850 m,
training in hypoxia and normoxia increased PPO at sea improved normoxic performance to the same extent as a
level (normoxia) to the same extent. Of practical signifi- similar normoxic training while improved to a larger extent
cance, exercise training in hypoxia increased PPO in the performance at altitude. However, this study indicates
hypoxic conditions. In contrast to these findings, the that the training intensity performed may reflect the sub-
improvement in endurance performance following the sequent adaptations observed. The peripheral mechanisms
different exercise training interventions was not associated behind this are largely not understood and could be a
with any changes in VO _ 2 max or in haematological vari- function of differential adaptations in skeletal muscle
ables. oxidative capacity. That aside the data from this study and
others indicated that no additional effect on endurance
Effects of training in hypoxia or normoxia performance occurs when a hypoxic stimulus is superim-
on endurance performance posed on an intensified training stimulus.
The present study did not observe any changes in Paver
In the present study PPO and cycle time-trial performance for the IHT group. The reasons of differences between NT
improved after 3 weeks of training in both hypoxic and and IHT on 10-min time-trial are unclear. In normoxia, the
normoxic conditions. Therefore, the present study is in present improvement after six sessions of interval-training
accordance with Terrados et al. (1988) who investigated at PPO is in agreement with previous studies (Stepto et al.
the effect of intermittent hypobaric hypoxic training 1999; Weston et al. 1997). Stepto et al. (1999) found that
(2,300 m; 45 sessions per week for 34 weeks) in elite interval training at 85% of peak power output in well-
cyclists. These authors found an increase in physiological trained endurance cyclists increased significantly the
work capacity and maximal power output; however, no simulated 40 km TT performance. Moreover, these authors

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Eur J Appl Physiol (2007) 101:359368 365

5.06 0.005

32.0 0.007
1.14 0.005
0.051 0.002

Mean SE; W0: pre; W1: first week of experimental training; W2: second week of experimental training; W3: third week of experimental training W4: post; NT: normoxic training group
(n = 8); IHT: intermittent hypoxic training group (n = 10); RBC: red blood cell; Hb: haemoglobin; Hct: hematocrit; RBD: red blood disc; MCV: mean cell volume; AGCH: averaged globular
found a reduced plasma lactate concentration at the same

14.80 0.1
46.27 0.4
92.0 0.4
246.7 1.4
29.39 0.1

105.6 0.3
absolute work rate after intense interval training. In addi-
IHT tion, Weston et al. (1997) observed a significant increase in
muscle buffering capacity in well-trained cyclists after six
high-intensity training sessions. These results indicated that

0.046 0.001
skeletal muscle buffering capacity may be an important
5.09 0.07

0.88 0.03
14.7 0.2
44.61 0.5
88.14 0.5
241.7 5.4
28.94 0.2
32.90 0.1

99.33 0.9
determinant of relatively short-duration (< 60 min)
endurance cycling activity (Weston et al. 1997). Why the
W4
NT

performance did not improve to the same extent after IHT


might be related to several mechanisms including a lower

0.052 0.002
5.0 0.05

1.07 0.03
absolute power output developed during the hypoxic
14.65 0.1
45.60 0.3
91.83 0.4
257.5 2.4
29.48 0.2
32.17 0.1

106.3 0.6
interval-session (85.7 8.1%; mean % of normoxic PPO).
Indeed, a study in untrained subjects has demonstrated that
IHT

content of haemoglobin; ACCH: averaged corpuscular concentration of haemoglobin; RET: reticulocytes; ARV: averaged reticulocytar volume
the absolute intensity of exercise is more important than the
relative exercise intensity in terms of acute skeletal muscle
0.046 0.003
5.01 0.06

0.90 0.04
14.63 0.2
44.24 0.5
88.50 0.4
241.8 4.7
29.25 0.2
33.01 0.1

99.13 0.4

adaptations to training in hypoxia (Wadley et al. 2005). In


the present study, both groups trained at the same relative
intensity; however, it might be the absolute training
W3
NT

intensity that induces larger performance changes in the


NT group. Furthermore, it has been suggested that hypoxic
0.049 0.001
4.79 0.05

32.07 0.04
1.01 0.02
14.07 0.1
43.89 0.3
91.71 0.4
247.4 3.2
29.43 0.1

107.6 0.7

training induces a decrease in muscular power due to the


reduced training intensity at altitude (detraining) (Boning
IHT

1997). It has also been shown that post-training period


following a return to sea level after an altitude training
0.045 0.003

camp is detrimental in term of specific performance. This


5.06 0.05

32.71 0.09
0.89 0.04
14.75 0.2
45.04 0.5
89.25 0.5
225.9 4.5
29.15 0.2

101.1 1.0

detrimental effect is probably due to the re-adaptations of


performance and physiological parameters to normal O2
W2
NT

content (Dick 1992). Although this loss of adaptation has


not been described after IHT, one may suggest that the
0.053 0.002
4.89 0.04

32.11 0.03
1.09 0.04

post-performance that occurred exactly at this time might


14.37 0.1
44.80 0.3
91.71 0.4
262.1 4.2
29.43 0.1

106.3 0.3

have been affected.


Hypoxia per se affects the structural and biochemical
IHT

properties of skeletal muscle by inducing a change in the


profile of type I (oxidative) to type II (glycolytic) fibres
0.046 0.001
4.95 0.05

0.73 0.01

(Ishihara et al. 1994; Itoh et al. 1995). This adaptation


14.43 0.2
43.70 0.4
88.6 0.4
236.6 4.0
29.15 0.2
33.0 0.1

101.9 0.6

will affect the changes in substrate utilization (Roberts


et al. 1996a, b), by shifting to a more efficient and eco-
W1
NT

nomic mode of oxygen utilization due to the lack of


oxygen in the tissue under hypoxic conditions. However,
0.071 0.004
5.15 0.05
14.96 0.09

31.80 0.07
1.34 0.06
47.06 0.3
91.57 0.4
245.6 1.3
29.14 0.2

104.7 0.6

it was observed that 3 weeks of IHT seemed to alter the


intrinsic properties of mitochondrial function in vastus
IHT

lateralis muscle in well-trained athletes (Roels et al.


2007). They observed a change in the muscle substrate
Table 6 Haematological variables

0.046 0.003

preference with a shift towards less fat oxidation and an


0.92 0.04
5.13 0.1
15.14 0.2
45.71 0.6
89.0 0.5
253.1 5.2
29.58 0.2
33.14 0.1

104.2 0.9

increase in carbohydrate oxidation, whereas the normoxic


control group did not show any significant differences
W0
NT

(Roels et al. 2007). This shift in substrate utilization


might explain the lack of improvement in Paver under
RBD (103 mm3)
RBC (106 mm3)

RET (10 mm )

normoxic conditions and is a detrimental effect of hy-


3

poxia on the peripheral level and upon the immediate


MCV (lm )

ARV (lm3)
ACCH (pg)
AGCH (%)
3
Hb (g dl1)

performance under normoxic conditions. Unfortunately,


RET (%)
Hct (%)

these potential explanations are hypothetical and further


investigation is necessary.

123
366 Eur J Appl Physiol (2007) 101:359368

Effects of training in hypoxia on endurance skeletal muscle adaptations to training in hypoxia as


performance in hypoxia compared to training in normoxia in athletes.

There is a scarcity of studies examining the effect of Effects of training on haematological parameters
endurance training performed in hypoxia on performance
in hypoxia in experienced endurance athletes. In the pres- No changes in RBC, Hb, and Hct nor in the other measured
ent study, we found that PPO measured in an incremental haematological variables were observed in this study.
exercise test to exhaustion in hypoxia significantly im- Thus, data from this study seems to indicate that hypoxic
proved following IHT. In a recent investigation maximal exposure needs to exceed 382 min per week to obtain an
running velocity, measured during an incremental exercise effect on haematological variables. However, Rodriguez
test performed in hypoxia, improved following IHT in et al. (2000) found that hypoxic exposure (4,0005,000 m)
trained athletes (Dufour et al. 2006). This is similar to that of only 90 min, three times a week for 3 weeks, i.e.
observed in the present investigation. Therefore, the data 270 min per week, significantly increased Hct, RBC, RET,
from his study and others confirms that altitude training or and Hb concentration. A continuous exposure of 90 min
simulated altitude training could be useful preparation for seems to represent the minimal stimulus to trigger an acute
performance and/or competition at altitude (Dufour et al. EPO-secretion. However, no control group (training in
2006; Wilber 2001). normoxia) was included in this investigation to establish
A possible mechanism could be in the hypoxic venti- whether the training stress in hypoxia was associated with
latory response (HVR) after hypoxic stimulus and any of these changes. Moreover, it has been shown that
endurance training. HVR is enhanced by intermittent haematological variables are already significantly in-
hypoxic exposure (Katayama et al. 1998) and is a posi- creased after 114 min at 3,000 m (Eckardt et al. 1989).
tive adaptation because of its increased VE _ which im- However, although the short duration of the hypoxic
proves alveolar O2 pressure and increases arterial stimulus used in this study might be responsible for the
oxygenation under hypoxic conditions (Huang et al. lack of changes in the haematological variables, other
1984). In contrast, HVR decreases after endurance studies using longer duration of hypoxic stimulus (Terrados
training (Katayama et al. 1999). In addition, not only et al. 1988; Vallier et al. 1996) have observed no changes
HVR but also cardiovascular responses to hypoxia might in selected haematological variables. The interval training
be an underlying mechanism. HVR and systolic blood completed by our subjects was at a power output of 100%
pressure responses were significantly increased after a 2- of PPO. This form of training induces and elevated blood
week intermittent hypoxic training program (5 sessions lactate and associated metabolic acidosis (Billat 2001).
per week; 30 min per day; 70% VO _ 2 max ; 4,500 m), but Metabolic acidosis inhibits EPO secretion, which might be
decreased significantly in a normoxic control group an explanation for the lack of haematological changes
(Katayama et al. 2004). However, it could be that other found in this study (Eckardt et al. 1990). Beside the
peripheral muscle factors are associated with the changes duration of the hypoxic stimulus, the level of the hypoxic
observed following the training in hypoxia. For example, stimulus might also be a factor involved in the EPO
Hoppeler and Fluck (2003) observed a significant in- secretion and in turn the increased RBC. The present study
crease in vastus lateralis sub-sarcolemmal mitochondria used a simulated altitude of 3,000 m, which was lower than
to compensate decreased O2 availability after an inter- the altitudes used in previous studies (Garcia et al. 2000;
mittent hypoxic training protocol. Other studies in trained Rodriguez et al. 2000).
runners have shown that a 6-week period of low to Inadequate iron stores are a contributing factor in the
moderate intensity training in hypoxia (as compared to lack of changes in haematological variables. Extra iron is
the same training in normoxia) resulted in improvements necessary for the increase in production of red cells stim-
in endurance performance which were associated with ulated by exposure to altitude (Rodriguez et al. 2000).
increases in mitochondrial function as well as increases in However, in this investigation all athletes were given iron
the expression of genes related to lactate shuttling and supplements to offset any iron deficiency. Therefore, it is
glucose transport, all of which could be important in unlikely this factor was influential in the present study.
terms of endurance performance (Ponsot et al. 2006; Zoll Finally, using hypoxic training for enhancement of sea-
et al. 2006). These studies suggest that hypoxic training level performance based on increased EPO, leading to in-
induce muscular adaptations, which are less expressed or creased RBC (Levine 2002) is highly debated as Calbet
even absent after normoxic training and which may ex- et al. (2005) suggested that an increased RBC and in-
plain the observed increase in PPO at altitude. However, creased Hb was only responsible for 16% of the variance in
further studies are required to confirm at a protein level, increased VO_ 2 max and in turn in endurance performance.

123
Eur J Appl Physiol (2007) 101:359368 367

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Acknowledgments This study was supported by the International detraining. J Appl Physiol 86:18051811
Olympic Committee and by the French Ministry of Sport. Additional Katayama K, Matsuo H, Ishida K, Mori S, Miyamura M (2003)
funding for the study was provided by Faculty Research Grants Intermittent hypoxia improves endurance performance and
Westminster University, London, UK and University of New South submaximal exercise efficiency. High Alt Med Biol 4:291304
Wales, Sydney, Australia. Katayama K, Sato Y, Matsuo H, Ishida K, Iwasaki K, Miyamura M
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