How anaerobic is the Wingate Anaerobic Test for humans?

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1 Eur J Appl Physiol (2002) 87: DOI /s ORIGINAL ARTICLE R. Beneke Æ C. Pollmann Æ I. Bleif Æ R.M. Leitha user M. Hu tler 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 generally used to evaluate anaerobic cycling performance, but knowledge of the metabolic profile of WAnT is limited. Therefore the energetics of WAnT was analysed with respect to working efficiency and performance. A group of 11 male subjects [mean(sd), age 21.6 (3.8) years, height (6.6) cm, body mass 82.2 (12.1) kg] performed a maximal incremental exercise test and a WAnT. Lactic and alactic anaerobic energy outputs were calculated from net lactate production and the fast component of the kinetics of post-exercise oxygenuptake. Aerobic metabolism was determined from oxygen uptake during exercise. The WAnT mean power of 683 (96.0) W resulted from a total energy output above the value at rest of (23.2) kjæ30 s 1 [mean metabolic power=4.3 (0.8) kw] corresponding to a working efficiency of 16.2 (1.6)%. The WAnT working efficiency was lower (P<0.01) thanthe corresponding value of 24.1 (1.7)% at 362 (41) W at the end of an incremental exercise test. During WAnT the fractions of the energy from aerobic, anaerobic alactic and lactic acid metabolism were 18.6 (2.5)%, 31.1 (4.6)%, and 50.3 (5.1)%, respectively. Energy from metabolism of anaerobic lactic acid explained 83% and 81% of the variance of WAnT peak and mean power, respectively. The results indicate firstly that WAnT R. Beneke (&) Department of Biological Sciences, Central Campus, University of Essex, Wivenhoe Park, Colchester, CO4 3SQ, England rbeneke@essex.ac.uk Tel.: Fax: R. Beneke Æ C. Pollmann Æ I. Bleif Æ R.M. Leithäuser Æ M. Hu tler Sports Medicine, Free University Berlin, Berlin, Germany R.M. Leitha user Laboratory 28, Berlin, Germany M. Hu tler Department of Physical Medicine and Rehabilitation, University of Bergen, Bergen, Norway requires the use of more anaerobically derived energy than previously estimated, secondly that anaerobic metabolism is dominated by glycolysis, thirdly that WAnT mechanical efficiency is lower than that found in aerobic exercise tests, and fourthly that the latter finding partly explains discrepancies between previously published and the present data about the metabolic profile of WAnT. Keywords Exercise test Æ Aerobic metabolism Æ Lactic acid metabolism Æ Alactic metabolism Æ Biomechanical efficiency Introduction The Wingate Anaerobic Test (WAnT) is a maximal intensity cycle ergometer test lasting 30 s. It was developed during the 1970s and serves to evaluate anaerobic performance (Ayalon et al. 1974). Generally accepted performance indices of WAnT are peak power, mean power and fatigue index. Peak power is the highest mechanical power elicited from the test takenas the average power over any 5 s period. Mean power is the average power maintained throughout the six 5 s segments. Fatigue index is the amount of the decline in power during the test expressed as a percentage of peak power (Inbar et al. 1996). Inthe past the validity of WAnT was examined by correlating peak power and mean power with other more or less anaerobic tests in the field (Bar-Or et al. 1977; Kaczkowski et al. 1982; Watsonand Sargeant 1986), in the laboratory (Bar-Or et al. 1977; Inbar et al. 1981; Tamayo et al. 1984), or with histochemical findings (Bar-Or et al. 1980; Inbar et al. 1981; Jacobs et al. 1982; Kaczkowski et al. 1982) or with estimations of the contributions of aerobic and anaerobic metabolism (Inbar et al. 1996; Serresse et al. 1988). However, correlations indicate formal interrelationships of selected measurements but they do not allow for quantitative mechanistic conclusions about the underlying energetics. The above mentioned estimations concerning relative metabolic

2 389 contributions based on muscle biopsies, oxygen uptake ( _V O 2 ) or measurements of constituents in the blood were based on limited quantitative data. Measurements of aerobic, anaerobic alactic or anaerobic lactic acid metabolism were missing (Inbar et al. 1996; Serresse et al. 1988) or inadequately determined with respect to timing (Jacobs et al. 1982). Estimations of overall energetics were based on assumed but not measured biomechanical efficiencies (Inbar et al. 1996; Serresse et al. 1988). Thus, although numerous selected details concerning the metabolic profile of the WAnT have been collected data concerning absolute and relative metabolic contributions to different energetic sources at present remain unknown. The aim of the present study was to examine the WAnT with respect to mechanical output, and anaerobic alactic, anaerobic lactic acid, and aerobic metabolic energy production. This requires a re-evaluation of previously published assumptions about biomechanical efficiency during WAnT and the metabolic contributions to WAnT performance. Methods Subjects A group of 11 male regional to national-class rugby players [mean (SD), age 21.6 (3.8) years, height (6.6) cm, body mass 82.2 (12.1) kg, maximal oxygenuptake ( _V O 2max ) 4,220 (466) mlæmin 1 ] participated in the study. All the subjects were healthy and nonsmokers; none was under pharmacological or special dietetic treatment. Informed consent was obtained from all the subjects after explanation of the nature and risks involved in participation in the experiments. The experiment conformed to internationally accepted policy statements regarding the use of human subjects and was approved by the local Ethics Committee. Protocol The subjects performed an incremental exercise test on an electrodynamically braked (Excalibur Sport, Lode) and a WAnT on a mechanically braked cycle ergometer (834 E, Monark). All tests 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 O 2max. 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 encouragement. 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, performed 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 instructed to pedal as fast as possible. A resistance corresponding to 7.5% of the body mass was applied after anaccelerationphase 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 power and fatigue index were calculated as described above. The _V O 2 and carbon dioxide production were measured and recorded breath-by-breath using a spirometric system (Oxycon gamma, Mijnhard) for the duration of the test. Before each test, the device was calibrated according to the manufacturer s instructions. The time course of the _V O 2 inthe recovery after exercise was interpolated using a bi-exponential function as described by (see Fig. 1): _V O 2 ðtþ ¼ae t=sa þ be t=sb þ c where _V O 2 (t) is the oxygenuptake at time t, a and b are the amplitudes of the fast and slow components, respectively, s a and s b the corresponding time constants and c is the _V O 2 at rest. Blood lactate concentration ([La - ] b ) was determined using the enzymatic amperometric method (Ebio Plus, Eppendorf) from capillary blood samples drawnfrom the hyperaemic ear lobe immediately before, and at 1 min intervals up to the 10th, and at 2 min intervals up to the 30th min post WanT. The [La - ] b was also determined before and during the final 30 s of each stage of the incremental exercise test and after the 1st, 3rd and 5th min post-test. Calculations Net aerobic energy was calculated from the _V O 2 above rest during WAnT, the energy equivalent of O 2 being assumed to be 21.1 kjæl 1. The energy produced from anaerobic alactic metabolism was estimated from the fast component of the post WAnT _V O 2 (Fig. 1) and energy equivalent of O 2 (Knuttgen 1970; Roberts and Morton 1978). Net energy produced from anaerobic lactic acid metabolism was determined from net [La - ] b accumulation, body mass, O 2 -lactate equivalent (Beneke and Meyer 1997; di Prampero 1981; Mader and Heck 1986). The average energy equivalent of peak [La - ] b (O 2 -lactate equivalent) was assumed to be 3 ml O 2 Ækg 1 ÆmmolÆl 1, i.e. to be 63 JÆkg 1 ÆmmolÆl 1 (di Prampero 1981). Mechanical efficiency was calculated from average metabolic power and mean power. Statistics Data are reported as mean values and standard deviations (SD). Differences within subjects were analysed by Friedman and Wilcoxon test. Relationships between variables were examined by simple and multiple linear or nonlinear regression analysis. For all statistics, the significance level was set at P<0.05. Results Peak power, meanpower, fatigue index were 900 (115) W, 683 (96) W, and 43.3 (5.5)%, respectively. Net [La - ] b accumulationwas 12.4 (1.6) mmolæl 1, _V O 2 Fig. 1. Oxygen uptake pre, during and post a Wingate anaerobic test (WAnT). Post WAnT oxygen uptake was analysed using a biexponential equation as described in the text and shown in this figure. The first exponential term describes the fast component of oxygenuptake

3 390 above rest during the test was 1,109 (151) ml. The values of a, time constant of a, b, time constant of b and c (see above and Fig. 1) were 2,777 (445) mlæmin 1, 0.7 (0.2) min, 675 (257) mlæmin 1, 14.1 (13.5) minand 329 (48) mlæmin 1, respectively, resulting in a post WAnT _V O 2 calculated from the fast component of the post WAnT _V O 2 of 1,904 (563) ml. Total energy turnover above the value at rest was (23.2) kjæ30 s 1. Fractions of the energy from aerobic, anaerobic alactic acid, and anaerobic lactic acid metabolism above the values at rest were 18.6 (2.5)%, 31.1 (4.6)%, and 50.3 (5.1)%, respectively (P<0.01). The efficiency of exercise during WAnT was 16.2 (1.6)% which was lower (P<0.01) thanthe corresponding value of 24.1 (1.7)% at 362 (41) W at the end of the incremental exercise test. Energy from anaerobic lactic acid metabolism explained 83% and 81% of the variance of peak power, and mean power, respectively. Neither the inclusion of energy from anaerobic alactic metabolism nor from aerobic metabolism improved the latter correlation. Energy from aerobic metabolism above rest did not correlate significantly with _V O 2max. The fatigue index did not show an interrelationship with any of the metabolic measurements made. Discussion The performance of the subjects in the present study was similar to that reported elsewhere and typical of excellent test results (Inbar et al. 1996). The measured fractions of energy from anaerobic alactic lactic acid and aerobic metabolisms yielded a slightly lower or more or less similar aerobic contribution to metabolism than that previously estimated (Boas et al. 1999; Inbar et al. 1996; Seresse et al. 1988; Smith and Hill 1991). It should be considered that these data were based exclusively on _V O 2 and power output. The fractions from anaerobic and aerobic metabolism were estimated by assuming that until peak power was reached all energy was provided 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 assumptions are of questionable validity. A maximal ATP-generation rate from the ATP-PC system of approximately 2.4 mmolæs 1 Ækg 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 glycolysis. Based on experiment data (Bangsbo et al. 1990; Boobis et al. 1982; Jones et al. 1985; Jacobs et al. 1982, 1983) energy from anaerobic lactic acid metabolism can be assumed to contribute to power output even within the initial 5 s of the WAnT. This hypothesis is supported by the high inter-relationship between the energy from anaerobic lactic acid metabolism and peak power in the present study. Several data from biopsies showed that s of maximal cycle-ergometer exercise did not lead to a total depletionof the muscle high energy phosphates (Bangsbo et al. 1990; Boobis et al. 1982; Jones et al. 1985; Jacobs et al. 1982). In the present experiment the fraction of energy from anaerobic alactic metabolism spent on WAnT performance was 40.2 (10.6) kj. This result may have been slightly influenced by the observationthat insome subjects the decrease of post WAnT _V O 2 was delayed (Fig. 2). This delay was previously described by di Prampero et al. (1973) and may have caused the rather slow constant s a, which was nearly twice as high as that reported for the kinetics of phosphocreatine changes upon the onset of exercise (Binzoni et al. 1992). Nevertheless, recalculationof the fractionof post WAnT _V O 2 contributing to anaerobic alactic metabolism (di Prampero et al. 1973; see Fig. 3) resulted in a value of 43.5 (18.8) kj which was not significantly different from the value of the energy from anaerobic 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 cycle ergometer test (Frauendorf et al. 1986). Due to the comparably high pedalling resistance at higher pedalling rates during WAnT not only the leg muscles but also trunk and arm muscles contribute significantly to the power output. Therefore the active muscle mass canbe assumed to be between60% and 85% of the total muscle mass. The latter value was assumed for rowing ergometry (Mader et al. 1988). If approximately 41 kj of energy from anaerobic alactic acid metabolism is provided by roughly 25 kg of muscle the resulting phosphocreatine breakdown during WAnT would be about 22 mmolækg muscle mass 1. Thus the present results compare well with the above-mentioned findings from Fig. 2. Oxygen uptake pre, during and post a Wingate anaerobic test (WAnT). Post WAnT oxygen uptake was slightly delayed compared to the usual first exponential term of a bi-exponential equation(see text)

4 391 Fig. 3. Oxygen uptake pre, during and post a Wingate anaerobic test (WAnT). Post WAnT oxygen uptake was analysed using a mono-exponential equation interpolating post WAnT oxygen uptake (see text). Relying on the work of di Prampero et al. (1973) all breath by breath data after the end of the initially determined fast component were included. The oxygen uptake contributing to the anaerobic alactic acid energy turnover was obtained from graphical back-extrapolation of the mono-exponential regression and subtraction from the post-want oxygen uptake above the value at rest biopsies and indicate that a useful amount of phosphocreatine is still available in the muscle after WAnT termination (Jacobs et al. 1982). The maximal rate of glycolysis canbe expected to be reached between the 5th and 10th s of WAnT when anaerobic lactic acid metabolism appears to have become the dominant energy source up to the end of WAnT. Within this period a glycolysis induced ph decrease becomes an ever increasing limiting factor of the rate of glycolysis and decreases the power from anaerobic lactic acid metabolism as shownby experiments on exhausting exercise up to 90 s (Boobis et al. 1982; Jacobs et al. 1983). The observed net production of lactate, which was equivalent to approximately 50% of the total energy turnover above that at rest during WAnT, and the high proportionof the variance of meanpower explained by energy from anaerobic lactic acid metabolism inthe present study, gives support to the hypothesis that, inspite of the decreasing rate of glycolysis, glycolysis can be expected to be the dominant contributor to performance between the 5th s and the end of WAnT. As WAnT progresses _V O 2 increases. In the present study, at WAnT termination, the _V O 2 was between60% and 80% of _V O 2max individually measured at the end of the incremental exercise test. However, the calculated fractionof the energy from aerobic metabolism may have been underestimated. Assuming the use of an O 2 reserve, bound to myoglobin and haemoglobin, of about 400 ml (Astrand and Rodahl 1977) this would increase the energy available from aerobic sources by approximately 36% and would reduce the contribution calculated to be attributable to the high energy phosphate system by about 20%. The resulting fractions of energy from aerobic and anaerobic alactic acid metabolism would then be 24% and 26%, respectively. Differences between the previously published and the present values of relative 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 onthe basis of assumed biomechanical efficiency (Inbar et al. 1996; Serresse et al. 1988; Smith and Hill 1991). The effect of such assumptions is shown by the differences between the directly measured _V O 2 above values at rest or net [La - ] b accumulationand those calculated onthe 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, estimated from the meanpower and the _V O 2 or net [La - ] b accumulationwould turnout to be 26% and 74%, respectively. The latter is withinthe range of published data (Inbar et al. 1996; Serresse et al. 1988) but does not agree with the fractionof metabolism measured inthe present study, which may illustrate the importance of using realistic values for mechanical efficiency when interpreting the results of anaerobic exercise tests. With respect to the present results it should be considered that the fractions of the energy from anaerobic alactic acid and anaerobic lactic acid metabolism, as calculated inthe present study, are critically dependent on the value assumed for the O 2 equivalent of net [La - ] b accumulation, and the assumption that the time integral of the fast component of the O 2 debt repayment after exercise is an appropriate estimate of the yield of anaerobic energy. Nevertheless, in a recent review these assumptions have been critically reconsidered, concluding that they are reliable (di Prampero and Ferretti 1999). Therefore, based onthe current state of knowledge, they cannot be expected to introduce substantial systematic errors in the calculations. The present study is the first to provide data which allow for the determination of an average mechanical efficiency during WAnT, based on anaerobic, and aerobic metabolism, and on mechanical power. Only one frequently quoted assumption about mechanical efficiency (Serresse et al. 1988) has been similar to the present result. In the latter study WAnT mechanical efficiency was assumed to be equal to the value measured at the end of an incremental _V O 2max test. However, it was unclear how mechanical efficiency was measured, and why the observed value was extremely low compared to the present and other published data measured at the pedalling rates usually employed during incremental exercise tests (Banister and Jackson 1967; Coast and Welch 1985; Dickinson 1929; Ericson 1988; Francescato et al. 1995; Gaesser and Brooks 1975; Zoladz et al. 1998). The mechanical efficiency was found to be nonlinearly related to pedalling rate and mechanical power, with values between 20% and 25%, which underlines the value presently observed at termination of the incremental exercise test. The optimal pedalling rate increases slightly with increasing exercise intensity. Within the normally used range of pedalling rates between 50 to 80Æmin 1 the mechanical efficiency is approximately

5 392 constant, or decreases slightly with increasing exercise intensity or pedalling rate. Due to there being a constant brake load, the WAnT power is a direct result of the pedalling rate attained. Normally the latter reaches 170Æmin 1 withinthe first few seconds and thendecreases to approximately 80Æmin 1 withinthe last few seconds of the test, resulting in average pedalling rates between 110 and 130Æmin 1. The latter, and the fact that higher brake loads enable higher WAnT power outputs to be achieved (Inbar et al. 1996) give evidence of a lower level of mechanical efficiency during WAnT than that determined at _V O 2max as observed inthe present study. Conclusions This study is the first investigating the energetics of WAnT to be based on all energy contributors such as high energy phosphates, glycolysis, and oxidative metabolism. 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