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Changes in surface electromyography (sEMG) frequency during maximal voluntary contractions following a 30-s Wingate anaerobic test (WAnT) using discrete wavelet transform (DWT) changes in sEMG were also compared between DWT and mean frequency (MNF) obtained by fast Fourier transform (fft) 17 healthy men performed a WAnT with a 7. % of body mass load.
Changes in surface electromyography (sEMG) frequency during maximal voluntary contractions following a 30-s Wingate anaerobic test (WAnT) using discrete wavelet transform (DWT) changes in sEMG were also compared between DWT and mean frequency (MNF) obtained by fast Fourier transform (fft) 17 healthy men performed a WAnT with a 7. % of body mass load.
Changes in surface electromyography (sEMG) frequency during maximal voluntary contractions following a 30-s Wingate anaerobic test (WAnT) using discrete wavelet transform (DWT) changes in sEMG were also compared between DWT and mean frequency (MNF) obtained by fast Fourier transform (fft) 17 healthy men performed a WAnT with a 7. % of body mass load.
Changes in surface EMG assessed by discrete wavelet transform
during maximal isometric voluntary contractions following supramaximal cycling Luis Pen ailillo
Rony Silvestre
Kazunori Nosaka Received: 18 April 2012 / Accepted: 11 September 2012 / Published online: 23 September 2012 Springer-Verlag 2012 Abstract To better understand characteristics of neuro- muscular fatigue in supramaximal cycling exercise, this study examined changes in surface electromyography (sEMG) frequency during maximal voluntary isometric contractions (MVC) following a 30-s Wingate anaerobic test (WAnT) using discrete wavelet transform (DWT). The changes in sEMG were also compared between DWT and mean frequency (MNF) obtained by fast Fourier transform (FFT). 17 healthy men performed a WAnT with a 7.5 % of body mass load. Knee extensor MVC torque was measured before and 1, 3, 6, 9, 12 and 15 min following WAnT, and sEMG was recorded from vastus lateralis muscle during the torque measures. sEMG was analysed for (RMS), MNF by FFT and frequency domains of DWT (divided into six domains). MVC torque decreased 2123 % at 315 min, RMS increased 2634 % at 115 min, and MNF decreased 810 % from baseline (76.3 3.2 Hz) at 13 min post- cycling (P\0.05). The DWT frequency domains showed that the changes lasted longer than MNF such that the intensity increased at 12 and 15 min for domain 2 (125250 Hz), all time points for domain 3 (62.5125 Hz), and 16 min for domains 4 (31.262.5 Hz) and 5 (15.631.2 Hz). The magnitude of increase in the intensity at 1 min post-exercise (4560 %) was largest for domains 3 and 5 (P\0.05). A signicant correlation was evident only between the magnitude of changes in the domain 5 and MNF (r = -0.56). It is concluded that DWT provides information on neuromuscular fatigue that is not detected by MNF derived from FFT. Keywords Wingate anaerobic test Neuromuscular fatigue Frequency spectrum Maximal voluntary isometric contraction Root mean square Fast Fourier transform (FFT) Mean frequency Introduction Surface electromyography (sEMG) provides information of global motor unit activation, and sEMG signals have been analysed for their changes in amplitude and frequency in the investigation of neuromuscular fatigue in various exercises (Cifrek et al. 2009; Hunter et al. 2003). A Win- gate anaerobic test (WAnT) is used to examine anaerobic capacity and also as a model of supramaximal exercise to investigate neuromuscular fatigue (Patton et al. 1985; Bogdanis et al. 1995). Previous studies (Hunter et al. 2003; Greer et al. 2006; Camata et al. 2010; Stewart et al. 2011) examined changes in sEMG frequency obtained by a fast Fourier transform (FFT) during WAnT, but controversy exists regarding the changes in mean frequency (MNF) derived from FFT. Two studies reported a signicant decrease (1015 %) in MNF over 30-s of WAnT (Hunter et al. 2003; Greer et al. 2006), but other two studies (Camata et al. 2010; Stewart et al. 2011) did not nd signicant changes in MNF during WAnT. No previous study has examined Communicated by Arnold de Haan. L. Penailillo School of Kinesiology, Ponticia Universidad Catolica de Valpara so, Valpara so, Chile L. Penailillo (&) K. Nosaka School of Exercise and Health Sciences, Edith Cowan University, 270 Joondalup Drive, Joondalup, WA 6027, Australia e-mail: l.penailillo@ecu.edu.au R. Silvestre School of Kinesiology, Universidad Mayor, Santiago, Chile 1 3 Eur J Appl Physiol (2013) 113:895904 DOI 10.1007/s00421-012-2499-1 changes in MNF after WAnT. To better understand neu- romuscular fatigue and recovery, it is important to examine post-WAnT frequency changes. It is also interesting to know how long the change (e.g. decreases) in MNF lasts after WAnT. It has been documented that further decreases in pH and ATP are seen after than during WAnT (Bogdanis et al. 1995). Thus, it is possible that further and greater changes in sEMG frequency could be observed following WAnT than during WAnT; however, this has not been investigated. Wavelet transform is a timefrequency method for sig- nal analysis, which has been proposed to be more appro- priate to analyse sEMG signals than FFT, because the wavelet transform is better than FFT for non-stationary sEMG signals (Karlsson et al. 2000), and wavelet function captures spiky signals of sEMG more efciently than FFT (Karlsson et al. 1999). The wavelet transform analysis expresses a signal as a linear combination of a particular set of functions obtained by shifting and dilating one single function called mother wavelet, and the correspondence between the resulting wavelet shape and the signal is cal- culated (Subasi and Kiymik 2010). The intensity of this coefcient is a measure of how much of the wavelet at a particular scale and time point (i.e. frequency range) is included in the signal (Subasi and Kiymik 2010). Since the mother wavelet can be moved to various locations of the signals and can be dilated (matching better with slow component of the signal) or squeezed (matching better with fast component of the signal), the wavelet transform pro- vides a mapping of the signal and decomposes the signal into several frequency domains that provide information regarding a specic portion of the frequency spectrum at a specic time (Subasi and Kiymik 2010; Letelier and Weber 2000). In the wavelet transform analyses, fast components of sEMG signals (sharp edges and steep slopes) are rep- resented by high-frequency domains and slower compo- nents of the signals (repolarisation or baseline uctuations) are represented by low-frequency domains (Letelier and Weber 2000). Discrete wavelet transform (DWT) is a type of wavelet transform and requires less computational pro- cessing time than other wavelet transform techniques such as continuous wavelet transform and wavelet packet decomposition, and provides high-quality timefrequency resolution (Constable and Thornhill 1993). Several studies (Beck et al. 2005; Constable et al. 1994; Kumar et al. 2003) used a wavelet transform for sEMG analyses in relation to neuromuscular fatigue. For example, Beck et al. (2005) compared between FFT and DWT for sEMG frequency changes over 50 maximal isokinetic concentric contractions of the elbow exors, and found that changes in MNF of FFT and the wavelet centre frequency were similar, both showing signicant decreases in fre- quency over contractions, suggesting that both FFT and wavelet analyses lead to the same physiological interpre- tations. Using a different wavelet transform, So et al. (2009) investigated changes in 11 wavelet domains during 50 maximal isokinetic concentric contractions of the knee extensors, and reported that the intensity of low-frequency domain with the centre frequency (CF) of 19.3 Hz increased in vastus lateralis muscle over 50 contractions. They speculated that the increases in the low-frequency domain were attributed to decreases in conduction velocity, which might indicate greater fatigue in fast than slow twitch muscle bres. However, no previous studies have assessed DWT changes following WAnT and the rela- tionship between changes in individual DWT domains and MNF after WAnT. The present study therefore investigated frequency changes in sEMG during maximal isometric contractions of the knee extensors following a WAnT using FFT and DWT, and assessed the relationship between the changes in MNF and DWT intensity. We hypothesised that (1) MNF would decrease and the intensity of low-frequency DWT domains would increase following WAnT, and (2) the changes in MNF would be signicantly correlated with the changes in the intensity of low-frequency DWT domains. Methods Subjects Seventeen healthy men (mean SD: age, 24.4 2.6 years; height, 173.0 5.5 cm; body weight, 72.7 8.8 kg) par- ticipated in this study. The Institutional Human Research Ethics Committee reviewed and approved the study, and all subjects gave their written informed consent before partici- pation. All subjects had not been involved in any regular training programme for at least 6 months prior to the study, and did not have any musculoskeletal or neurological dis- orders of lower limbs. They were instructed not to perform any strenuous exercise at least 48 h before the testing day, and to avoid caffeinated and alcoholic beverages for at least 8 h prior to the participation in the experiment. Protocol All subjects performed WAnT in the afternoon (14:0017:00) at least 1.5 h after lunch, and several mea- surements shown below were taken before and for 15 min following WAnT. Maximal voluntary isometric contraction (MVC) torque of the non-dominant knee extensors were measured before and 1, 3, 6, 9, 12 and 15 min following WAnT, and during which sEMG of vastus lateralis (VL) muscle was recorded. In order for the subject to perform 896 Eur J Appl Physiol (2013) 113:895904 1 3 MVC immediately after the completion of WAnT, a cycle ergometer was placed right next to a chair for the MVC measurement (explained below), and each subject was assisted by the investigator to have the MVC measures within 1 min after the WAnT. sEMG electrodes were kept on the subjects leg during WAnT, and were connected to a PowerLab system (explained below) as soon as possible. sEMG was analysed for root mean square (RMS) and MNF, and by DWT. Blood lactate concentration was assessed before each MVC torque measure. Wingate anaerobic test (WAnT) Ten minutes after the baseline measurements, the subjects performed 5 min cycling warm-up with a 2 % of their body weight load at a self-selected cadence, and two 5-s sprints were included at the third and fourth minute. 2 min after warm-up, subjects started the WAnT by cycling without a load to reach the maximal pedalling rate for 3 s on a mechanically braked ergometer (Enermax, SINEBI, Argentina) and a load of 7.5 % of their body weight (range 4.57.1 kg) was suddenly added. They were encouraged to pedal as hard and fast as possible for next 30 s. During the test, the average power output of every 5-s segment (05, 510, 1015, 1520, 2025, 2530 s) was calculated, and peak power output, mean power output over 30 s and fatigue index [(highest 5 s peak power - lowest 5 s peak power)/highest 5 s peak power] were obtained (Armstrong et al. 1983; Reiser et al. 2002). Measurements Maximal voluntary isometric contraction (MVC) torque Subjects were seated in a chair adapted for this study to keep the hip joint at 1208 exion and the knee joint at 908 exion, and knee extension torque of the non-dominant leg was measured by a load cell (MLT003/D, AdInstruments, Australia) connected to a PowerLab system ML818 (AdInstruments, Australia). The load cell was placed on the lever arm attached to the chair, and positioned 8 cm above the medial tibial malleolus. Subjects were asked to exert force against the lever arm as hard and fast as possible and verbal encouragement was given. For the measurement taken before WAnT, three 3-s MVC torque measures with 1 min rest between measures were performed, and the average of the three peak values was used for further analysis. In the measurements after WAnT (1, 3, 6, 9, 12 and 15 min post), only one MVC torque measure (3 s) was taken for each time point. Blood lactate A blood sample of 25 ll was collected from the earlobe by a capillary tube prior to each MVC measure and was analysed for blood lactate concentration using a YSI 1500 Sport Lactate Analyzer (YSI life Science, OH, USA). sEMG recording and analyses Two rectangular (20 9 30 mm) AgAgCl electrodes (Skintact, Austria) were placed on the VL of the non- dominant leg at two-third on the line from the anterior superior iliac spine to the lateral side of the patella (according to SENIAM) separated by 20 mm (Hermens et al. 2000), and the ground electrode was placed on the medial surface of tibia, after the skin was shaved and cleaned with 95 % alcohol to reduce impedance to less than 5 kX. During the 3-s MVC strength measures, sEMG signals were recorded with the PowerLab system using a Chart 5.5 software (AdInstruments, Australia) with a sampling frequency of 1,000 Hz. The sEMG signals were digitised on-line and stored in a computer for later off-line analysis with an IGOR Pro 6.0 software (WaveMetrics, Portland, USA). An epoch of the central 2 s of the raw sEMG signal during which the MVC was plateau was extracted for analyses (Merletti and Parker 2004). Using the 2-s epochs, RMS of the amplitude was calculated, and the MNF of the power spectrum density was obtained from a FFT using 2,048 points (Cifrek et al. 2009; Hostens et al. 2004). Wavelet transform processing DWT was applied for the sEMG signals of the 2-s epoch (Fig. 1) using a DWT toolbox of the IGOR Pro software. The DWT toolbox with the Daubechies (db4) mother was used to calculate the intensity of the wavelet coefcient (Beck et al. 2005) for the eight domains (Table 1). How- ever, since most of the sEMG frequency spectrum is in the range of 10250 Hz (Wakeling et al. 2001), only the middle six domains (domains 27) were focused in the present study. As shown in Fig. 1, the timefrequency matrix is shown by a grey scale for each domain (18) for the 2-s epoch, and the intensity of the wavelet coefcient is shown as white representing the highest wavelet coefcient intensity and black indicating the lowest intensity of the wavelet coefcient. The average intensity of the absolute value of the wavelet coefcient for the 2-s epoch was obtained for each domain, and percentage of change in the intensity from baseline to 1, 3, 6, 9, 12 and 15 min post- cycling was calculated. Eur J Appl Physiol (2013) 113:895904 897 1 3 Statistical analysis Changes in WAnT power output, MVC, blood lactate, RMS, MNF and average of the wavelet coefcients over time were assessed using a one-way repeated measures analysis of variance (ANOVA). When a signicant time effect was found, a Scheffes post hoc test was applied. To compare the changes in the average intensity of the DWT coefcient over time amongst six domains, a two-way repeated measures ANOVA (domain 9 time) was used, followed by post hoc pairwise comparisons using a Fishers least signicant difference (LSD) test. Relationships between the percentage change in MNF and the intensity of each DWT domain from baseline were assessed using a Pearson product moment correlation coefcient. The sig- nicance level was set at P\0.05. All statistical analyses were performed with PASW Statistics 18 software for Mac (SPSS Inc, IBM Company, USA). Data are presented in mean standard error of the means (SEM). Results WAnt All subjects completed the exercise as instructed. The peak power and the mean power output were 565.6 69.9 and 476.5 60.8 W, respectively, and the relative power per body weight was 7.8 0.6 W kg -1 for the peak and 6.6 0.4 W kg -1 for the mean power. Figure 2 shows changes in the power output normalised by body weight for 30 s. Power output decreased signicantly over time, and the fatigue index was 33.2 8.8 %. MVC torque Pre-exercise MVC torque was 324.6 146.1 Nm (range 213464 Nm), and MVC torque decreased signicantly at 3 min (23.1 14.2 %), remaining low for 15 min post- exercise without signicant changes between 3 and 15 min (Fig. 3a). Blood lactate Blood lactate concentration increased signicantly from pre (1.6 0.1 mmol l -1 ) to 1 min post-exercise (11.5 1.1 mmol l -1 ), further increased up to 6 min (15.3 0.5 mmol l -1 ) and remained the level until 15 min post- exercise (Fig. 3b). Root mean square sEMG RMS increased signicantly following the exercise by 34.4 8.3 % at 1 min, remaining the level for the rest of recovery time (Fig. 4a). MNF As shown in Fig. 4b, MNF decreased signicantly from pre (76.3 3.2 Hz) to 1 min (69.0 2.7 Hz) and 3 min Fig. 1 sEMG signal processing for DWT domain analyses. From the 3-s sEMG signal during MVC, a central window of 2 s was selected. DWT divides the signal into eight dyadic domains, and presents a timefrequency matrix that shows time (2 s) at the X-axis and the wavelet domains 18 as Y-axis. The intensity of the wavelet coefcient is shown by a grey scale at the right side of the time frequency matrix, where white indicates maximum intensity and black shows the lowest intensity. For this example, the average intensities for the domain 2, 3, 4, 5, 6 and 7 (1 and 8 domains were not analysed) are 0.20, 0.43, 0.62, 0.49, 0.32 and 0.13, respectively Table 1 Domains of the discrete wavelet transform and their frequency range Wavelet domain Frequency range (Hz) Lower frequency Higher frequency 1 250 500 2 125 250 3 62.5 125 4 31.2 62.5 5 15.6 31.2 6 7.8 15.6 7 3.9 7.8 8 1.9 3.9 898 Eur J Appl Physiol (2013) 113:895904 1 3 (70.4 2.9 Hz) post-WAnT, then returned to the pre- exercise level. DWT Figure 5 shows changes in the intensity of the wavelet coefcient from pre-exercise level for each of the six fre- quency domains following WAnT. The changes were sig- nicantly different amongst the domains, and the domains 3 and 5 showed the largest increases in the intensity of the wavelet coefcients amongst the domains. The one-way repeated measures ANOVA showed a signicant increase in the intensity of the wavelet coefcient for domains 2, 3, 4 and 5 from pre-exercise values. The intensity increased signicantly at 12 and 15 min for the domain 2, at all time points for the domain 3, and at 1, 3 and 6 min for the domains 4 and 5. Fig. 2 Changes (mean SEM) in cycling power output normalised by bodyweight every 5 s over 30-s supramaximal cycling. Fatigue index is also shown in the gure. *Signicantly (P\0.05) lower than the peak 5-s power output Fig. 3 Changes (mean SEM) in maximal voluntary isometric contraction strength (a) and blood lactate concentration (b) before (Pre) and 1, 3, 6, 9, 12 and 15 min following 30-s supramaximal cycling. *Signicantly (P\0.05) greater than the pre-value. # Sig- nicantly (P\0.05) greater than the 1 min value Fig. 4 Changes (mean SEM) in sEMG root mean square (a) and mean frequency (b) before (Pre) and 1, 3, 6, 9, 12 and 15 min following 30-s supramaximal cycling. *Signicantly (P\0.05) different than the pre-exercise value Eur J Appl Physiol (2013) 113:895904 899 1 3 Correlation between changes in MNF and intensity of DWT domains Figure 6 shows the correlation between the magnitude of changes in MNF and DWT for the domains that showed signicant changes at 1 and 3 min post-WAnT (i.e. domains 3, 4 and 5) using the two-time points of 1 and 3 min post-WAnT (n = 34: 17 subjects 9 2 time points). A signicant correlation was found (r = -0.56) for the domain 5 only (Fig. 6c). Discussion The main ndings of the present study were that (1) MNF signicantly decreased at 1 and 3 min post-WAnT from the baseline, and the DWT domains 25 signicantly increased after WAnT, exhibiting longer lasting changes (e.g. 15 min for domains 2 and 3) than MNF (13 min post-WAnT only); and (2) the relative increase in intensity of the wavelet domain 5 was signicantly correlated with the relative decrease in the MNF at 1 and 3 min post-WAnT. These results partially supported the hypotheses that MNF would decrease and the intensity of low-frequency DWT domains would increase following WAnT, and the changes in MNF would be signicantly correlated with the changes in the intensity of low-frequency DWT. The peak and mean power output, and fatigue index during WAnT (Fig. 2) and changes in blood lactate fol- lowing WAnT (Fig. 3b) were similar to those reported in previous studies (Hebestreit et al. 1996; Bogdanis et al. 1995; Beneke et al. 2007). Judging from the peak and mean power outputs, the subjects in the present study were cat- egorised as the lower 1020 % of physically active young adults according to the normative values reported by Maud and Shultz (1989). The present study was the rst to measure MVC torque of the knee extensors following WAnT. Since the fatigue index showed more than 30 % Fig. 5 The magnitude of changes (mean SEM) in wavelet coefcient intensity for the domains 27 from pre- exercise (0 %) at 1, 3, 6, 9, 12 and 15 min following supramaximal cycling. *Signicantly (P\0.05) different from pre-exercise value Fig. 6 Correlation between the magnitude of changes in mean frequency and coefcient of DWT for the domains 3 (a), 4 (b) and 5 (c) from the baseline values at 1 and 3 min following WAnT. The two-time point data (1 and 3 min post-cycling, n = 34) were used for the analyses 900 Eur J Appl Physiol (2013) 113:895904 1 3 decrease in power output, a decrease in knee extensor MVC torque immediately after WAnT was expected. However, the MVC torque did not show a signicant decrease at 1 min post-WAnT, but showed a signicant decrease from the baseline (23 %) after 3 min post-WAnT (Fig. 3b). It could be argued that repeated MVC mea- surements following WAnT would have induced additional neuromuscular fatigue and/or affected the recovery from WAnT. However, only a 3-s MVC contraction was per- formed for each time point with 2 min rest before the next contraction, thus it seems likely that the effect of the MVC measures on recovery was minimum, and they did not affect the changes in MVC and sEMG. Sahlin and Seger (1995) reported 34 % decrease in MVC immediately after cycling to exhaustion at 75 % of the VO 2max , and Giacomoni et al. (2006) reported 1315 % MVC decrease immediately after ten sets of 6-s cycling sprints. Nicolas et al. (2008) also showed 17 % MVC decrease immediately after cycling to exhaustion at 95 % of maximal power output, and Theurel and Lepers (2008) reported 712 % decrease in MVC at 1 min after 33 min cycling at 70 % of maximal aerobic power. It should be noted that the exercise duration of these studies was much longer (485 min) than that of the present study (30 s). It might be that the short duration of the WAnT delayed the decrease in MVC torque. In the present study, the blood lactate further increased from 1 to 3 min post-WAnT (Fig. 2). Bogdanis et al. (1995) showed that the intramus- cular pH was lowest at 1.5 min after WAnT, and Street et al. (2001) showed a delayed acidosis (range 0.52 min) after knee extensor fatiguing contractions. Thus, it is pos- sible that delayed peripheral metabolic changes are asso- ciated with the delayed decrease in MVC torque. It should be noted that sEMG RMS was elevated after WAnT (Fig. 4a). It might be that central fatigue was limited after WAnT, and increased cortical excitability minimised the decrease in MVC at 1 min post-WAnT. However, it can be speculated that this effect was no longer potent at the subsequent time points. Further studies are necessary to elucidate why a decrease in MVC torque was delayed after WAnT. No previous studies have reported changes in sEMG following WAnT, but some studies examined sEMG amplitude changes during WAnT and found no signicant changes (Hunter et al. 2003; Rana 2006; Camata et al. 2010; Stewart et al. 2011). For example, Stewart et al. (2011) have recently reported that the average rectied values of EMG amplitude did not change during WAnT. In contrast, the present study found 2634 % increase in RMS of sEMG during MVC measures that were performed after WAnT (Fig. 3a). Sarre and Lepers (2005) reported that RMS of VL muscle increased during cycling (from 5 to 60 min) at 65 % of maximal power output. Furthermore, Giacomoni et al. (2006) showed that RMS of VL muscle during MVC increased by 810 % from baseline at 15 min after a repeated sprint cycling exercise (ten sets of 6 s, 30 s rest between sets). Arabadzhiev et al. (2010) stated that increases in sEMG RMS indicated increases in motor unit action potential amplitude that could be related to increases in the activity of the Na ? K ? pump and intracellular action potential (IAP) length. IAP length is increased with hyperpolarisation of the muscle membrane, which is likely due to an accumulation of extracellular K ? and H ? that impairs the repolarisation of the muscle membrane (Allen et al. 2008). It can be speculated that the increases in RMS amplitude following WAnT were asso- ciated with increases of extracellular K ? and decreases in pH produced by WAnT. The sampling frequency in the present study was 1,000 Hz in agreement with the Nyquist rate stating that sampling frequency must be at least double the highest frequency components in the signal (Merletti and Parker 2004). The highest frequency components of the sEMG signals have been reported to be around 400500 Hz for isometric contractions (Ives and Wigglesworth 2003), thus the sampling frequency (1,000 Hz) seems appropriate. Although it has been reported that MNF depends on the sampling frequency such that the higher the sampling fre- quency, the greater the MNF (Sadhukhan et al. 1994), changes in the MNF shown in Fig. 4b should not be affected by the sampling frequency. Moreover, 1,000 Hz sampling frequency has been used for MNF analyses dur- ing isometric contractions in previous studies (e.g. Willems and Ponte 2012; Coorevits et al. 2008; Beck et al. 2005). Two previous studies reported a signicant decrease (1015 %) in MNF over 30 s of WAnT (Hunter et al. 2003; Greer et al. 2006). Stewart et al. (2011) found a tendency of decreases in MNF that was in parallel with decreases in power output and muscle bre conduction velocity during WAnT. Hunter et al. (2003) showed that MNF decreased by 15 % from the rst 5 s to the last 5 s during WAnT. It is important to note that in the present study, MNF was assessed during MVC measures after WAnT, not during WAnT. However, as shown in Fig. 4b, MNF decreased from pre to 1 min post-WAnT by 10 %, which was com- parable to that reported by Hunter et al. (2003). Hunter et al. (2003) stated that the decrease in MNF was probably due to fatigue of fast twitch bres and decreases in pH and phosphocreatine in those muscle bres. It is interesting that MNF was returned to the baseline value by 6 min post- WAnT in our study (Fig. 4b), when the blood lactate and presumably intracellular H ? concentration were still high (Fig. 3b). Tenan et al. (2011) have recently reported that MNF during MVC assessed intermittently over an incre- mental cycling exercise to exhaustion did not change, but blood lactate and plasma K ? increased signicantly over Eur J Appl Physiol (2013) 113:895904 901 1 3 time. They have concluded that changes in MNF are not associated with changes in blood lactate and plasma K ? . The present study results also showed the dissociation between MNF changes and blood lactate. Another possible cause of the decrease in MNF is a decrease in muscle bre conduction velocity (Kupa et al. 1995), but dissociation between the changes in MNF and muscle conduction velocity has been reported (Stewart et al. 2011; Zwarts et al. 1987). It is speculated that central factors such as changes in motor unit discharge rate and/or de-recruitment of fatigued motor units also inuence MNF changes (Hermens et al. 1992; Stewart et al. 2011). The DWT analysis showed increases in the domains 25, and the magnitude of increase in the intensity of these domains (2160 %) was similar to that of sEMG RMS (2634 %). This would suggest that the increase in the RMS was associated with the increases in the intensity of wavelet coefcients. It should be noted that the range of the frequency is not the same amongst domains, and the greater the range, the less the precision and resolution. The domains 3 and 5 (frequency range of 62.5125 and 15.631.2 Hz, respectively) showed the largest increases amongst the domains after WAnT (Fig. 5). So et al. (2009) reported that the intensity of a low-frequency domain (centre frequency = 19.3 Hz) increased largest during 50 maximal fatiguing contractions, and speculated that the large increase in the low-frequency domain was due to decreases in the ring rate and conduction velocity of fast twitch muscle bres. In the present study, domains 2 and 3 showed increases at 915 min post-WAnT, which may reect some recovery of fast twitch muscle bres. Although it is not known whether changes in the high-frequency domains 2 and 3 represent physiological changes, the results of the present study show that sEMG frequency was still affected at 15 min post-WAnT according to the DWT analysis. The DWT domain 5 showed the largest relative increase (60 % at 1 min) following WAnT amongst the six domains (Fig. 5). Furthermore, the magnitude of increase in the intensity of the domain 5 (15.631.2 Hz) was signicantly correlated with the magnitude of decrease in MNF at 13 min post-WAnT. We speculate that the decrease in MNF and the increase of domain 5, which contains a range of low frequencies, could indicate a relative decrease in muscle bre conduction velocity and/or a decrease in fast twitch muscle bre activation. It might be that the larger increases of the domain 5 compared with other domains after WAnT suggest that the compression of the frequency spectrum was shifted towards low frequencies. The chan- ges in domain 5 lasted longer (i.e. 9 min) than those of MNF (i.e. 6 min). It is possible that domain 5 could detect smaller changes in the frequency spectrum, due to its fre- quency range decomposition. Because DWT matches better the spiky shape of the sEMG signals and manages the non-stationarity in muscle contractions better than FFT (Karlsson et al. 2000), it may provide better accuracy in detecting changes in the sEMG frequency spectrum (Letelier and Weber 2000). It is plausible that since MNF is a measure of central tendency (i.e. mean, median, mode) of the whole frequency spectrum, the sensitivity of MNF analysis to detect changes in frequency spectrum is less than that of DWT. DWT appears to provide information that MNF does not provide such as the decomposition in ranges of the frequencies that allows the examination of changes at different portions of the frequency spectrum, which could increase the understanding of muscle recovery from fatiguing tasks (von Tscharner 2002; Karlsson et al. 1999). It has been documented, however, that changes in fre- quency domains do not necessarily reect a change in muscle bre recruitment or conduction velocity (Farina 2008; Farina et al. 2004). The present study assessed the sEMG during MVC measures that are often used to assess neuromuscular fatigue, and compared the relative changes in sEMG during this standardised task over 15 min fol- lowing WAnT. We speculate that the relatively larger increases in the DWT low-frequency domains most likely indicate a decrease in muscle bres conduction velocity, due to the decrease in the ring rate of fast twitch muscle bres. More studies are necessary using multi-electrode arrays and motor unit decomposition technique to investi- gate more directly changes in muscle bre recruitment and conduction velocity after WAnT, and their relationship to DWT frequency changes. In summary, both DWT and MNF frequency analyses showed a swift towards a low-frequency spectrum; how- ever, DWT domain changes lasted longer and the magni- tude of the changes in low-frequency domains were greater than those observed in MNF. Thus, it appears that DWT provides information of the frequency changes in sEMG that is not detected by FFT. It is possible that DWT, especially domain 5, could detect small changes in sEMG frequency following exercise. It seems possible that DWT has better timefrequency resolution, which could detect smaller changes in sEMG signals than MNF. It is con- cluded that DWT analysis is useful to understand neuro- muscular fatigue and recovery. References Allen DG, Lamb GD, Westerblad H (2008) Impaired calcium release during fatigue. J Appl Physiol 104:296305 Arabadzhiev TI, Dimitrov VG, Dimitrova NA, Dimitrov GV (2010) Interpretation of EMG integral or RMS and estimates of neuromuscular efciency can be misleading in fatiguing contraction. 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