Eur J Appl Physiol (2002) 87: 388–392

DOI 10.1007/s00421-002-0622-4

O R I GI N A L A R T IC L E

R. Beneke Æ C. Pollmann Æ I. Bleif Æ R.M. Leithäuser

M. Hütler

How anaerobic is the Wingate Anaerobic Test for humans?

Accepted: 11 March 2002 / Published online: 28 May 2002

Ó Springer-Verlag 2002

Abstract The Wingate Anaerobic Test (WAnT) is gen- requires the use of more anaerobically derived energy
erally used to evaluate anaerobic cycling performance, than previously estimated, secondly that anaerobic me-
but knowledge of the metabolic profile of WAnT is tabolism is dominated by glycolysis, thirdly that WAnT
limited. Therefore the energetics of WAnT was analysed mechanical efficiency is lower than that found in aerobic
with respect to working efficiency and performance. exercise tests, and fourthly that the latter finding partly
A group of 11 male subjects [mean (SD), age explains discrepancies between previously published and
21.6 (3.8) years, height 178.6 (6.6) cm, body mass the present data about the metabolic profile of WAnT.
82.2 (12.1) kg] performed a maximal incremental exer-
cise test and a WAnT. Lactic and alactic anaerobic en- Keywords Exercise test Æ Aerobic metabolism Æ
ergy outputs were calculated from net lactate production Lactic acid metabolism Æ Alactic metabolism Æ
and the fast component of the kinetics of post-exercise Biomechanical efficiency
oxygen uptake. Aerobic metabolism was determined
from oxygen uptake during exercise. The WAnT mean
power of 683 (96.0) W resulted from a total energy Introduction
output above the value at rest of 128.1 (23.2) kJÆ30 s–1
[mean metabolic power=4.3 (0.8) kW] corresponding The Wingate Anaerobic Test (WAnT) is a maximal in-
to a working efficiency of 16.2 (1.6)%. The WAnT tensity cycle ergometer test lasting 30 s. It was developed
working efficiency was lower (P<0.01) than the corre- during the 1970s and serves to evaluate anaerobic per-
sponding value of 24.1 (1.7)% at 362 (41) W at the end formance (Ayalon et al. 1974). Generally accepted per-
of an incremental exercise test. During WAnT the formance indices of WAnT are peak power, mean power
fractions of the energy from aerobic, anaerobic alactic and fatigue index. Peak power is the highest mechanical
and lactic acid metabolism were 18.6 (2.5)%, power elicited from the test taken as the average power
31.1 (4.6)%, and 50.3 (5.1)%, respectively. Energy from over any 5 s period. Mean power is the average power
metabolism of anaerobic lactic acid explained 83% and maintained throughout the six 5 s segments. Fatigue
81% of the variance of WAnT peak and mean power, index is the amount of the decline in power during the
respectively. The results indicate firstly that WAnT test expressed as a percentage of peak power (Inbar et al.
1996).
In the past the validity of WAnT was examined by
R. Beneke (&) correlating peak power and mean power with other more
Department of Biological Sciences, Central Campus, or less anaerobic tests in the field (Bar-Or et al. 1977;
University of Essex, Wivenhoe Park,
Colchester, CO4 3SQ, England Kaczkowski et al. 1982; Watson and Sargeant 1986), in
E-mail: [email protected] the laboratory (Bar-Or et al. 1977; Inbar et al. 1981;
Tel.: +44-1206-872530 Tamayo et al. 1984), or with histochemical findings
Fax: +44-1206-872592 (Bar-Or et al. 1980; Inbar et al. 1981; Jacobs et al. 1982;
R. Beneke Æ C. Pollmann Æ I. Bleif Æ R.M. Leithäuser Æ M. Hütler Kaczkowski et al. 1982) or with estimations of the con-
Sports Medicine, Free University Berlin, Berlin, Germany tributions of aerobic and anaerobic metabolism (Inbar
R.M. Leithäuser et al. 1996; Serresse et al. 1988). However, correlations
Laboratory 28, Berlin, Germany indicate formal interrelationships of selected measure-
M. Hütler ments but they do not allow for quantitative mechanistic
Department of Physical Medicine and Rehabilitation, conclusions about the underlying energetics. The above
University of Bergen, Bergen, Norway mentioned estimations concerning relative metabolic
 389

contributions based on muscle biopsies, oxygen uptake was calibrated according to the manufacturer’s instructions. The
(V_ O2) or measurements of constituents in the blood were time course of the V_ O2 in the recovery after exercise was interpo-
lated using a bi-exponential function as described by (see Fig. 1):
based on limited quantitative data. Measurements of
aerobic, anaerobic alactic or anaerobic lactic acid me- V_ O2 ðtÞ ¼ a et=sa þ b et=sb þ c
tabolism were missing (Inbar et al. 1996; Serresse et al. where V_ O2(t) is the oxygen uptake at time t, a and b are the am-
1988) or inadequately determined with respect to timing plitudes of the fast and slow components, respectively, sa and sb
(Jacobs et al. 1982). Estimations of overall energetics were the corresponding time constants and c is the V_ O2 at rest.
based on assumed but not measured biomechanical effi- Blood lactate concentration ([La-]b) was determined using the
ciencies (Inbar et al. 1996; Serresse et al. 1988). Thus, al- enzymatic amperometric method (Ebio Plus, Eppendorf) from
capillary blood samples drawn from the hyperaemic ear lobe
though numerous selected details concerning the immediately before, and at 1 min intervals up to the 10th, and at
metabolic profile of the WAnT have been collected data 2 min intervals up to the 30th min post WanT. The [La-]b was also
concerning absolute and relative metabolic contributions determined before and during the final 30 s of each stage of the
to different energetic sources at present remain unknown. incremental exercise test and after the 1st, 3rd and 5th min post-test.
The aim of the present study was to examine the
WAnT with respect to mechanical output, and anaero- Calculations
bic alactic, anaerobic lactic acid, and aerobic metabolic
Net aerobic energy was calculated from the V_ O2 above rest during
energy production. This requires a re-evaluation of WAnT, the energy equivalent of O2 being assumed to be 21.1 kJÆl–1.
previously published assumptions about biomechanical The energy produced from anaerobic alactic metabolism was esti-
efficiency during WAnT and the metabolic contributions mated from the fast component of the post WAnT V_ O2 (Fig. 1)
to WAnT performance. and energy equivalent of O2 (Knuttgen 1970; Roberts and Morton
1978). Net energy produced from anaerobic lactic acid metabolism
was determined from net [La-]b accumulation, body mass, O2-lac-
tate equivalent (Beneke and Meyer 1997; di Prampero 1981; Mader
Methods and Heck 1986). The average energy equivalent of peak [La-]b
(O2-lactate equivalent) was assumed to be 3 ml O2Ækg–1ÆmmolÆl–1,
Subjects i.e. to be 63 JÆkg–1ÆmmolÆl–1 (di Prampero 1981). Mechanical
efficiency was calculated from average metabolic power and mean
A group of 11 male regional to national-class rugby players [mean power.
(SD), age 21.6 (3.8) years, height 178.6 (6.6) cm, body mass 82.2
(12.1) kg, maximal oxygen uptake (V_ O2max) 4,220 (466) mlÆmin–1] Statistics
participated in the study. All the subjects were healthy and non-
smokers; none was under pharmacological or special dietetic Data are reported as mean values and standard deviations (SD).
treatment. Informed consent was obtained from all the subjects Differences within subjects were analysed by Friedman and
after explanation of the nature and risks involved in participation Wilcoxon test. Relationships between variables were examined by
in the experiments. The experiment conformed to internationally simple and multiple linear or nonlinear regression analysis. For all
accepted policy statements regarding the use of human subjects and statistics, the significance level was set at P<0.05.
was approved by the local Ethics Committee.

Protocol
Results

The subjects performed an incremental exercise test on an elec- Peak power, mean power, fatigue index were
trodynamically braked (Excalibur Sport, Lode) and a WAnT on a 900 (115) W, 683 (96) W, and 43.3 (5.5)%, respectively.
mechanically braked cycle ergometer (834 E, Monark). All tests Net [La-]b accumulation was 12.4 (1.6) mmolÆl–1, V_ O2
were performed at similar times in the morning on separate days at
least 2 h after a light meal. The interval between the tests was
1 week. The subjects were instructed not to engage in strenuous
activity during the day before an exercise test.
The incremental exercise test served for the determination of
V_ O2max. The test started with exercise at 100 W and was increased
by 50 W every 3rd min. It finished at the individual’s maximal
power output as indicated by fatigue despite strong vocal en-
couragement. The WAnT was conducted according to the widely
accepted recommendations for standardization (Inbar et al. 1996).
The WAnT session started with a standardized warming up of
5 min cycling at 50 W including two sprints, each lasting 3 s, per-
formed at the end of the 3rd and the 5th min as preparation for the
sprintlike WAnT. After a 10 min rest the subjects were then in-
structed to pedal as fast as possible. A resistance corresponding to
7.5% of the body mass was applied after an acceleration phase
lasting 3 s. The subjects were verbally encouraged to maintain as
high a pedalling rate as possible throughout the 30 s duration of
the test. After termination of the test the subjects were supervised
during a 30 min rest in a sitting position. The peak power, mean Fig. 1. Oxygen uptake pre, during and post a Wingate anaerobic
power and fatigue index were calculated as described above. The test (WAnT). Post WAnT oxygen uptake was analysed using a bi-
V_ O2 and carbon dioxide production were measured and recorded exponential equation as described in the text and shown in this
breath-by-breath using a spirometric system (Oxycon gamma, figure. The first exponential term describes the fast component of
Mijnhard) for the duration of the test. Before each test, the device oxygen uptake
390

above rest during the test was 1,109 (151) ml. The val- be assumed to contribute to power output even within
ues of a, time constant of a, b, time constant of b and the initial 5 s of the WAnT. This hypothesis is supported
c (see above and Fig. 1) were 2,777 (445) mlÆmin–1, by the high inter-relationship between the energy from
0.7 (0.2) min, 675 (257) mlÆmin–1, 14.1 (13.5) min and anaerobic lactic acid metabolism and peak power in the
329 (48) mlÆmin–1, respectively, resulting in a post present study.
WAnT V_ O2 calculated from the fast component of the Several data from biopsies showed that 10–190 s of
post WAnT V_ O2 of 1,904 (563) ml. maximal cycle-ergometer exercise did not lead to a total
Total energy turnover above the value at rest was depletion of the muscle high energy phosphates
128.1 (23.2) kJÆ30 s–1. Fractions of the energy from (Bangsbo et al. 1990; Boobis et al. 1982; Jones et al.
aerobic, anaerobic alactic acid, and anaerobic lactic acid 1985; Jacobs et al. 1982). In the present experiment the
metabolism above the values at rest were 18.6 (2.5)%, fraction of energy from anaerobic alactic metabolism
31.1 (4.6)%, and 50.3 (5.1)%, respectively (P<0.01). spent on WAnT performance was 40.2 (10.6) kJ. This
The efficiency of exercise during WAnT was result may have been slightly influenced by the obser-
16.2 (1.6)% which was lower (P<0.01) than the corre- vation that in some subjects the decrease of post WAnT
sponding value of 24.1 (1.7)% at 362 (41) W at the end V_ O2 was delayed (Fig. 2). This delay was previously
of the incremental exercise test. described by di Prampero et al. (1973) and may have
Energy from anaerobic lactic acid metabolism ex- caused the rather slow constant sa, which was nearly
plained 83% and 81% of the variance of peak power, twice as high as that reported for the kinetics of phos-
and mean power, respectively. Neither the inclusion of phocreatine changes upon the onset of exercise (Binzoni
energy from anaerobic alactic metabolism nor from et al. 1992). Nevertheless, recalculation of the fraction of
aerobic metabolism improved the latter correlation. post WAnT V_ O2 contributing to anaerobic alactic
Energy from aerobic metabolism above rest did not metabolism (di Prampero et al. 1973; see Fig. 3) resulted
correlate significantly with V_ O2max. The fatigue index in a value of 43.5 (18.8) kJ which was not significantly
did not show an interrelationship with any of the met- different from the value of the energy from anaerobic
abolic measurements made. alactic metabolism calculated initially. The muscle mass
primarily engaged is assumed to be about 60% of the
total muscle mass employed in a traditional incremental
Discussion cycle ergometer test (Frauendorf et al. 1986). Due to the
comparably high pedalling resistance at higher pedalling
The performance of the subjects in the present study was rates during WAnT not only the leg muscles but also
similar to that reported elsewhere and typical of excel- trunk and arm muscles contribute significantly to the
lent test results (Inbar et al. 1996). The measured frac- power output. Therefore the active muscle mass can be
tions of energy from anaerobic alactic lactic acid and assumed to be between 60% and 85% of the total muscle
aerobic metabolisms yielded a slightly lower or more or mass. The latter value was assumed for rowing ergom-
less similar aerobic contribution to metabolism than that etry (Mader et al. 1988). If approximately 41 kJ of en-
previously estimated (Boas et al. 1999; Inbar et al. 1996; ergy from anaerobic alactic acid metabolism is provided
Seresse et al. 1988; Smith and Hill 1991). It should be by roughly 25 kg of muscle the resulting phosphocrea-
considered that these data were based exclusively on tine breakdown during WAnT would be about
V_ O2 and power output. The fractions from anaerobic 22 mmolÆkg muscle mass–1. Thus the present results
and aerobic metabolism were estimated by assuming compare well with the above-mentioned findings from
that until peak power was reached all energy was pro-
vided by the adenosine-triphospate-phosphocreatine
system (ATP-PC system) and that the contribution of
high energy phosphates to work output was 0 after 10 s
of exercise (Serresse et al. 1988). Both of these assump-
tions are of questionable validity.
A maximal ATP-generation rate from the ATP-PC
system of approximately 2.4 mmolÆs–1Ækg wet muscle–1
(Weicker and Strobel 1994) implies that the generation
of energy necessary for the production of the peak
power observed requires supplementing the energy from
anaerobic alactic metabolism from other metabolic
sources. The high rate of anaerobic alactic metabolism
during the initial seconds of WAnT leads to an acute
increase in pH which is known to facilitate the activity of
phosphofructokinase and to increase the rate of glycol-
Fig. 2. Oxygen uptake pre, during and post a Wingate anaerobic
ysis. Based on experiment data (Bangsbo et al. 1990; test (WAnT). Post WAnT oxygen uptake was slightly delayed
Boobis et al. 1982; Jones et al. 1985; Jacobs et al. 1982, compared to the usual first exponential term of a bi-exponential
1983) energy from anaerobic lactic acid metabolism can equation (see text)
 391

energy yield from aerobic and anaerobic metabolism

during WanT may also be due to the fact that in previous
studies the anaerobic and aerobic fractions were not
measured, but estimated on the basis of assumed bio-
mechanical efficiency (Inbar et al. 1996; Serresse et al.
1988; Smith and Hill 1991). The effect of such assump-
tions is shown by the differences between the directly
measured V_ O2 above values at rest or net [La-]b accu-
mulation and those calculated on the basis of a set of
mechanical efficiencies ranging between 12% and 22%
(Beneke et al. 1996; Boas et al. 1999; Inbar et al. 1996;
Serresse et al. 1988; Smith and Hill 1991). For example, in
the present study, assuming an efficiency of 22%, the
yields from aerobic and anaerobic energy metabolism,
Fig. 3. Oxygen uptake pre, during and post a Wingate anaerobic estimated from the mean power and the V_ O2 or net [La-]b
test (WAnT). Post WAnT oxygen uptake was analysed using a accumulation would turn out to be 26% and 74%, re-
mono-exponential equation interpolating post WAnT oxygen spectively. The latter is within the range of published data
uptake (see text). Relying on the work of di Prampero et al.
(1973) all breath by breath data after the end of the initially (Inbar et al. 1996; Serresse et al. 1988) but does not agree
determined fast component were included. The oxygen uptake with the fraction of metabolism measured in the present
contributing to the anaerobic alactic acid energy turnover was study, which may illustrate the importance of using re-
obtained from graphical back-extrapolation of the mono-exponen- alistic values for mechanical efficiency when interpreting
tial regression and subtraction from the post-WAnT oxygen uptake
above the value at rest the results of anaerobic exercise tests. With respect to the
present results it should be considered that the fractions
biopsies and indicate that a useful amount of phos- of the energy from anaerobic alactic acid and anaerobic
phocreatine is still available in the muscle after WAnT lactic acid metabolism, as calculated in the present study,
termination (Jacobs et al. 1982). are critically dependent on the value assumed for the O2
The maximal rate of glycolysis can be expected to be equivalent of net [La-]b accumulation, and the assump-
reached between the 5th and 10th s of WAnT when tion that the time integral of the fast component of the O2
anaerobic lactic acid metabolism appears to have be- debt repayment after exercise is an appropriate estimate
come the dominant energy source up to the end of of the yield of anaerobic energy. Nevertheless, in a recent
WAnT. Within this period a glycolysis induced pH de- review these assumptions have been critically reconsid-
crease becomes an ever increasing limiting factor of the ered, concluding that they are reliable (di Prampero and
rate of glycolysis and decreases the power from anaer- Ferretti 1999). Therefore, based on the current state of
obic lactic acid metabolism as shown by experiments on knowledge, they cannot be expected to introduce sub-
exhausting exercise up to 90 s (Boobis et al. 1982; Jacobs stantial systematic errors in the calculations.
et al. 1983). The observed net production of lactate, The present study is the first to provide data which
which was equivalent to approximately 50% of the total allow for the determination of an average mechanical
energy turnover above that at rest during WAnT, and efficiency during WAnT, based on anaerobic, and aero-
the high proportion of the variance of mean power ex- bic metabolism, and on mechanical power. Only one
plained by energy from anaerobic lactic acid metabolism frequently quoted assumption about mechanical effi-
in the present study, gives support to the hypothesis ciency (Serresse et al. 1988) has been similar to the pre-
that, in spite of the decreasing rate of glycolysis, glyc- sent result. In the latter study WAnT mechanical
olysis can be expected to be the dominant contributor to efficiency was assumed to be equal to the value measured
performance between the 5th s and the end of WAnT. at the end of an incremental V_ O2max test. However, it was
As WAnT progresses V_ O2 increases. In the present unclear how mechanical efficiency was measured, and
study, at WAnT termination, the V_ O2 was between 60% why the observed value was extremely low compared to
and 80% of V_ O2max individually measured at the end of the present and other published data measured at the
the incremental exercise test. However, the calculated pedalling rates usually employed during incremental
fraction of the energy from aerobic metabolism may have exercise tests (Banister and Jackson 1967; Coast and
been underestimated. Assuming the use of an O2 reserve, Welch 1985; Dickinson 1929; Ericson 1988; Francescato
bound to myoglobin and haemoglobin, of about 400 ml et al. 1995; Gaesser and Brooks 1975; Zoladz et al. 1998).
(Astrand and Rodahl 1977) this would increase the en- The mechanical efficiency was found to be nonlinearly
ergy available from aerobic sources by approximately related to pedalling rate and mechanical power, with
36% and would reduce the contribution calculated to be values between 20% and 25%, which underlines the
attributable to the high energy phosphate system by value presently observed at termination of the incre-
about 20%. The resulting fractions of energy from aer- mental exercise test. The optimal pedalling rate increases
obic and anaerobic alactic acid metabolism would then slightly with increasing exercise intensity. Within the
be 24% and 26%, respectively. Differences between the normally used range of pedalling rates between 50 to
previously published and the present values of relative 80Æmin–1 the mechanical efficiency is approximately
392

constant, or decreases slightly with increasing exercise Coast JR, Welch HJ (1985) Linear increase in optimal pedal rate
intensity or pedalling rate. Due to there being a constant with increased power output in cycle ergometry. Eur J of Appl
Physiol 53:339–342
brake load, the WAnT power is a direct result of the Dickinson S (1929) The efficiency of bicycle-pedalling, as affected
pedalling rate attained. Normally the latter reaches by speed and load. J Physiol (Lond) 67:242–255
170Æmin–1 within the first few seconds and then decreases Ericson MO (1988) Mechanical muscular power output and work
to approximately 80Æmin–1 within the last few seconds of during ergometer cycling at different work loads and speeds.
Eur J Appl Physiol 57:382–387
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and 130Æmin–1. The latter, and the fact that higher brake of internal work during cycling. Eur J Appl Physiol 72:51–57
loads enable higher WAnT power outputs to be achieved Frauendorf H, Kobryn U, Gelbrich W, Hoffmann B, Erdmann U
(Inbar et al. 1996) give evidence of a lower level of (1986) Ergometrische Untersuchungen an unterschiedlichen
mechanical efficiency during WAnT than that deter- Muskelgruppen in ihrer Auswirkung auf Herzschlagfrequenz
und Blutdruck. Z Klin Med 41:343–346
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aerobic alactic and lactic acid metabolism dominated by cise. J Appl Physiol 55:365–367
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Heigenhauser GJF, Sutton JR (1985) Muscle performance and
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published literature. The latter partly explains the dis- speeds. J Appl Physiol 59:132–136
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