Effects of fatiguing constant versus alternating intensity intermittent isometric muscle actions on maximal torque and neuromuscular responses

Similar documents
Electromyographic and Mechanomyographic Time and Frequency Responses During Fatiguing, Submaximal, Isokinetic Muscle Actions of the Biceps Brachii

IMPROVEMENT OF MUSCLE STRENGTH IN REHABILITATION BY THE USE OF SURFACE ELECTROMYOGRAPHY

Effect of sex on torque, recovery, EMG, and MMG responses to fatigue

D.O.I: GEORGIOS DASTERIDIS, THEOPHILOS PILIANIDIS, NIKOLAOS MANTZOURANIS, NIKOLAOS AGGELOUSIS

Effect of cold treatment on the concentric and eccentric torque-velocity relationship of the quadriceps femoris

Effect of electrode position of low intensity neuromuscular electrical stimulation on the evoked force in the quadriceps femoris muscle

Effects of prolonged tendon vibration stimulation on eccentric and concentric maximal torque and EMGs of the knee extensors

Muscle Activation Analysis of Step and Jump Forward Lunge among Badminton Players

Sex comparisons of non-local muscle fatigue in human elbow flexors and knee extensors

POST-ACTIVATION POTENTIATION AND VERTICAL JUMP PERFORMANCE. Cody Hardwick

Systems Physiology Neuromuscular and Skeletal INFLUENCE OF ELECTRODE ORIENTATION ON ELECTROMYOGRAPHIC FATIGUE INDICES OF THE VASTUS LATERALIS

CAN TRAINING IMPROVE YOUR ABILITY TO CO-CONTRACT? Jordan Yurchevich. St. Francis Xavier University. October 9, 2006

[ research report ] 130 march 2007 volume 37 number 3 journal of orthopaedic & sports physical therapy. t Study Design: Repeated-measures experimental

Force depression in human quadriceps femoris following voluntary shortening contractions

Fatigue and Recovery from Dynamic Contractions in Men and Women Differ for Arm and Leg Muscles

EMG CHARACTERISTICS AND FIBRE COMPOSITION: STUDY ON RECTUS FEMORIS OF SPRINTERS AND LONG DISTANCE RUNNERS

Surface Electromyographic Amplitude-to-Work Ratios During Isokinetic and Isotonic Muscle Actions

Introduction ORIGINAL ARTICLE. Travis W. Beck Æ Terry J. Housh Æ Glen O. Johnson Joseph P. Weir Æ Joel T. Cramer Æ Jared W. Coburn Moh H.

Tight sportswear and physiological function

2231. The effect of accelerometer mass in mechanomyography measurements

Time course of changes in neuromuscular responses during rides to exhaustion above and below critical power

A Comparison of Plyometric Training Techniques for Improving Vertical Jump Ability and Energy Production

Biomechanics of Skeletal Muscle and the Musculoskeletal System

SITES OF FAILURE IN MUSCLE FATIGUE

ISOMETRIC SQUAT FORCE OUTPUT AND MUSCLE ACTIVITY IN STABLE AND UNSTABLE CONDITIONS

Electromyographic and kinetic comparison of a flexible and steel barbell

PERSONAL RESUME. Terry J. Housh PhD, FACSM, FNSCA. Personal Data Home: 621 West Beal Street Office: 110 Ruth Leverton Hall

Assessment of muscle contraction features by using Tensiomyography

발목관절의각도가무릎관절폄근의근활성도에미치는영향

Journal of Electromyography and Kinesiology

Acute Effects of Static and Proprioceptive Neuromuscular Facilitation Stretching on Muscle Strength and Power Output

Electromyographic Activity Recorded from an Unexercised Muscle During Maximal Isometric Exercise of the Contralateral Agonists and Antagonists

AN ELECTROMYOGRAPHIC COMPARISON BETWEEN THE SQUAT IN A DECLINE BOARD AND IN A FLAT SURFACE WITH DIFFERENT OVERLOADS

Changes in Muscle Hardness and Electromyographic Response for Quadriceps Muscle during Repetitive Maximal Isokinetic Knee Extension Exercise

Impact of Contraction Intensity and Velocity on Vastus Lateralis SEMG Power Spectrum and Amplitude

Chapter 6. Summarizing discussion

Muscle Activation Analyses for Yoga Interventions

Μέθοδοι Εμβιομηχανικών μηχ Μετρήσεων

ABDOMINAL MUSCLE ACTIVATION IN TWO TRUNK-CURL TESTS

NEUROMUSCULAR AND MORPHOLOGICAL ADAPTATIONS TO SHORT-TERM SQUAT AND DEADLIFT TRAINING IN WOMEN. Kendra Olinghouse, B.S. A Thesis

PEAK TORQUE AND IEMG ACTIVITY OF QUADRICEPS FEMORIS MUSCLE AT THREE DIFFERENT KNEE ANGLES IN

THE EFFECTS OF BIOFEEDBACK ON NEUROMUSCULAR RECRUITMENT PATTERNS DURING FATIGUING SETS OF BACK SQUAT. Daniel Arndts

Central and peripheral fatigue in sustained maximum voluntary contractions of human quadriceps muscle

COMPARISON OF FOURIER AND WAVELET TRANSFORMS ON SEMG FREQUENCY SPECTRA DURING RAPID FATIGUING MUSCULAR CONTRACTIONS

Higher Neuromuscular Manifestations of Fatigue in Dynamic than Isometric Pull-Up Tasks in Rock Climbers

Neural Recovery Rates of Knee Extensors Following a Resistance Exercise Protocol

The Contributions of Arterial Cross Sectional Area and Time Averaged Flow Velocity to Arterial Blood Flow

Intramachine and intermachine reproducibility of concentric performance: A study of the Con-Trex MJ and the Cybex Norm dynamometers

The Reliability of Four Different Methods. of Calculating Quadriceps Peak Torque Angle- Specific Torques at 30, 60, and 75

Is there inhibition during eccentric muscle contractions?

Low-frequency fatigue at maximal and submaximal muscle contractions

TMG - BMC LTD. Mirje 6 Tel: ( ) Ljubljana Fax: ( ) TENSIOMYOGRAPHY (TMG) MEASUREMENT RESULTS

Effect of Patellar Taping on Quadriceps Muscle of Knee Joint INTRODUCTION

Biomechanics of Skeletal Muscle and the Musculoskeletal System

Comparison of N-K Table Offset Angles with the Human Knee Flexor Torque Curve

In che modo le griglie bidimensionali di elettrodi ci aiutano nell interpretazione del segnale EMG di superficie?

EQA DISCUSSION QUESTIONS: INFLUENCE OF MUSCLE FIBER TYPE ON MUSCLE CONTRACTION. Influence of Muscle Fiber Type on Muscle Contraction

The Effects of Cold Water Immersion on Anaerobic Power, Dynamic Balance and Muscle Activation After a karate kumite fighting in Female Karateka


EMG analysis of shoulder muscle fatigue during resisted isometric shoulder elevation

The effect of lower limb massage on EMG and force production of the knee

The Effect of Rest Intervals on Knee-Extension Torque Production With Neuromuscular Electrical Stimulation

Analysis of EMG Signal to Evaluate Muscle Strength and Classification

JKMU. Comparison of generalized and directed co-contraction of knee joint muscles during four different movements for strengthening the quadriceps

Impact-induced soft-tissue vibrations associate with muscle activation in human landing movements: An accelerometry and EMG evaluation

Available online at Pelagia Research Library. European Journal of Experimental Biology, 2014, 4(1):

Lab Exercise 8. BIOPAC Exercise. Muscle Tissue. Muscles. What you need to be able to do on the exam after completing this lab exercise:

differentiate between the various types of muscle contractions; describe the factors that influence strength development;

The effects of poliomyelitis on motor unit behavior during repetitive muscle actions: a case report

Doctoral School on Engineering Sciences Università Politecnica delle Marche

Movement, Health & Exercise, 1(1), 39-48, 2012

Differential Fatiguability between the One and Two-Joint Muscles

Reduced muscle activity during isokinetic contractions associated with external leg compression

Electrostimulation for Sport Training

EMG Signals for Co-Activations of Major Lower Limb Muscles in Knee Joint Dynamics

Effects of KAATSU on muscular function during isometric exercise

Electromyography (EMG)

A Comparison of Measures of the Bilateral Limb Deficit During Short and Long Time Isometric Knee Extensions

The effect of the fatty tissue on EMG signal in young women

ORIGINAL ARTICLE SOLEUS MUSCLE CONTRACTION ON DRIVER S KNEE ANGLE POSTURE MORE THAN 144º FOR FOOT PRESSING AND RELEASING AN AUTOMOTIVE PEDAL

Influence of Resistance Load on Electromyography Response to Vibration Training with Sub-maximal Isometric Contractions

METHODOLOGICAL LIMITATIONS OF EMG-BASED BIO-MECHANICAL MOTION ANALYSIS. Thomas Jöllenbeck Bergische Universität - Gesamthochschule Wuppertal, Germany

Neither Stretching nor Postactivation Potentiation Affect Maximal Force and Rate of Force Production during Seven One-Minute Trials

MUSCLE SIZE AND SPECIFIC FORCE ALONG THE LENGTH OF THE QUADRICEPS IN OLDER AND YOUNG INDIVIDUALS

The effects of low-intensity resistance training with vascular restriction on leg muscle strength in older men

ABSTRACT. Associate Professor Marc A. Rogers, Department of Kinesiology. The purpose of this study was to examine the effects of nine weeks of

Resistive Eccentric Exercise: Effects of Visual

Muscle Use during Isometric muscle co-contraction compared to Abdominal Crunches and A Commercial Multi Gym Exerciser

Comparison of the relation with muscle activity (facial and leg) between Borg- and facial pictorial-rpe scale for perceived exertion

Obesity is associated with reduced joint range of motion (Park, 2010), which has been partially

Athos Wearable Technology: A Comparison Study

Evaluation of electromyographic activity and heart rate responses to isometric exercise. The role played by muscular mass and type

SINGLE- AND TWO-JOINT MOVEMENTS IN HUMANS

Looped Band Placed Around Thighs Increases EMG of Gluteal Muscles Without Hindering Performance During Squatting

Neural adaptations with sport-specific resistance training in highly skilled athletes

Evaluation of Handle Diameter in Maximum Horizontal and Vertical Torque Tasks. Yong-Ku Kong and Brian D. Lowe

ا م أ و ءا ف ج ب ه ذ يف د ب زلا ا م أ ف ي ف ث ك م يف سا نلا ع فن ي ا م ض ر لأا (17 دع ) رلا ةروس

Chapter 14 Training Muscles to Become Stronger

Evaluation of Electromyographic Frequency Domain Changes during a Three- Minute Maximal Effort Cycling Test

Strength and conditioning specialists are continually

Transcription:

J Musculoskelet Neuronal Interact 2016; 16(4):318-326 Journal of Musculoskeletal and Neuronal Interactions Original Article Effects of fatiguing constant versus alternating intensity intermittent isometric muscle actions on maximal torque and neuromuscular responses C.M. Smith, T.J. Housh, E.C. Hill, K.C. Cochrane, N.D.M. Jenkins, R.J. Schmidt, G.O. Johnson Department of Nutrition and Health Sciences Human Performance Laboratory, University of Nebraska-Lincoln, Lincoln, NE 68505, United States Abstract Objective: To determine the effects of constant versus alternating applications of torque during fatiguing, intermittent isometric muscle actions of the leg extensors on maximal voluntary isometric contraction (MVIC) torque and neuromuscular responses. Methods: Sixteen subjects performed two protocols, each consisting of 50 intermittent isometric muscle actions of the leg extensors with equal average load at a constant 60% MVIC or alternating 40 then 80% (40/80%) MVIC with a work-to-rest ratio of 6-s on and 2-s off. MVIC torque as well as electromyographic signals from the vastus lateralis (VL), vastus medialis (VM), and rectus femoris (RF) and mechanomyographic signals from the VL were recorded pretest, immediately posttest, and 5-min posttest. Results: The results indicated that there were no time-related differences between the 60% MVIC and 40/80% MVIC protocols. The MVIC torque decreased posttest (22 to 26%) and remained depressed 5-min posttest (9%). There were decreases in electromyographic frequency (14 to 19%) and mechanomyographic frequency (23 to 24%) posttest that returned to pretest levels 5-min posttest. There were no changes in electromyographic amplitude and mechanomyogrpahic amplitude. Conclusions: These findings suggested that these neuromuscular parameters did not track the fatigue-induced changes in MVIC torque after 5-min of recovery. Keywords: EMG, MMG, Fatigue, Recovery, Submaximal Introduction Fatigue has previously been defined as a gradual decrease in the force capacity of muscle or the endpoint of a sustained activity, and it can be measured as a reduction in muscle force, a change in electromyographic activity or an exhaustion of contractile function. (p.12) 1. The amplitude of the electromyographic signal represents muscle activation, and the frequency content is related to motor unit action potential conduction velocity during isometric muscle The authors have no conflict of interest. Corresponding author: Corresponding Author: Cory M. Smith, Department of Nutrition and Health Sciences, 110 Ruth Leverton Hall University of Nebraska-Lincoln Lincoln, NE 68583-0806, United States E-mail: CSmith@unl.edu Edited by: S. Warden Accepted 27 August 2016 actions 2-5. The mechanomyographic signal, however, reflects the mechanical counterpart of the motor unit electrical activity as measured by electromyographic and quantifies the low-frequency lateral oscillations of activated skeletal muscle fibers 5-7. Under some conditions, the amplitude of the mechanomyographic signal represents motor unit recruitment, and the frequency content is qualitatively related to the global firing rates of unfused, activated motor units 6,7. Therefore, simultaneous assessments of electromyographic and mechanomyographic signals have been used to examine fatigue-related neuromuscular responses and make inferences regarding motor unit activation strategies including muscle activation 2, motor unit action potential conduction velocity 2, motor unit recruitment 6,7, and global motor unit firing rate 2 from the time-dependent changes in electromyographic amplitude (root mean square), electromyographic frequency (mean power frequency), mechanomyographic amplitude, and mechanomyographic frequency, respectively. Most occupational and sporting activities involve complex motor programs that include randomly ordered levels of force production, rest intervals, number of repeated 318

contractions, durations of recovery, and modes of muscle actions, as well as the involvement of multiple muscle groups 8,9. The interactions among these factors can affect force-related outcomes during fatiguing tasks as well as the patterns of responses for electromyographic and mechanomyographic time and frequency domain parameters 10. To understand the mechanisms of fatigue as well as the time course of neuromuscular responses that describe the fatiguing process, most studies have assessed torque and neuromuscular responses while manipulating one or more of the influencing factors (i.e. force production and/or rest intervals), and controlling others (i.e. mode of muscle action and/or the muscles involved) 10-12. For example, Seghers et al. 10 reported decreases in maximal voluntary isometric contraction (MVIC) torque and electromyographic frequency as well as increases in electromyographic amplitude from the biceps brachii after a fatiguing workbout consisting of 20-min of submaximal, intermittent isometric muscle actions of the forearm flexors at 25% MVIC. In addition, Kouzaki et al. 11 reported decreases in MVIC torque as well as electromyographic amplitude, electromyographic frequency, mechanomyographic amplitude, and mechanomyographic frequency from the vastus lateralis (VL), vastus medialis (VM), and rectus femoris (RF) during 50 maximal, intermittent isometric muscle actions of the leg extensors. Bigland-Ritchie et al. 12, however, reported decreases in MVIC torque, but increases in electromyographic amplitude from the VL following 10-min of submaximal, intermittent isometric muscle actions of the leg extensors at 50% MVIC. Fowles et al. 13 also reported increases in electromyographic amplitude from the VL after 30-min of submaximal, intermittent isometric muscle actions of the leg extensors at 60% MVIC. The electromyographic amplitude reported by Fowles et al. 13, however, returned to pretest levels after 4-hr of recovery, but MVIC torque remained depressed from pretest levels. No previous studies, however, have simultaneously measured electromyographic amplitude, electromyographic frequency, mechanomyographic amplitude, and mechanomyographic frequency to identify if these neuromuscular responses track the changes in MVIC torque immediately and after 5-min of recovery from fatiguing, submaximal, intermittent isometric muscle actions of the leg extensors. Therefore, the purpose of the present study was to determine the effects of constant versus alternating applications of torque during fatiguing, intermittent isometric muscle actions of the leg extensors on MVIC torque and neuromuscular responses. Based on previous studies 13-16, we hypothesized that the alternating protocol would result in greater decreases in MVIC torque, electromyographic amplitude, mechanomyographic amplitude, electromyographic frequency, and mechanomyographic frequency than the constant protocol. In addition, we hypothesized 13-16 that MVIC torque would remain depressed for both protocols after 5-min of recovery, but the constant protocol would recover to a greater extent than the alternating protocol. Material and methods Subjects Sixteen healthy adults (11 men and 5 women, mean±sd age 22.0±2.6 yr; body mass 82.6±14.4 kg; height 177.3±7.4 cm) volunteered to participate in the investigation. The subjects regularly participated in physical activities such as resistance training, soccer, and bicycling. The study was approved by the University Institutional Review Board for Human Subjects, and all subjects completed a health history questionnaire and signed an informed consent document prior to testing. Orientation session The orientation session was used to familiarize the subjects with the testing procedures including maximal and submaximal isometric muscle actions of the leg extensors. All isometric muscle actions were performed using the dominant leg (based on kicking preference) at a knee joint angle of 120, 180 being full extension, for maximal force production on a calibrated Cybex II dynamometer. In addition, the hip was aligned with a joint angle of approximately 90. A warm-up consisting of 5-min on a cycle ergometer at a selfselected resistance was performed, as well as 5 to 10 submaximal isometric muscle actions of the leg extensors. After completion of the warm-up, 2, 6-s MVICs were performed followed by familiarization with repeated 60% MVIC, as well as alternating 40% then 80% (40/80%) MVIC. Intermittent isometric protocols A warm-up consisting of 5-min on a cycle ergometer at a self-selected resistance was performed prior to each testing session. In addition, the subjects performed 5 submaximal isometric muscle actions of the leg extensors at approximately 50% of their maximum effort, followed by 2-min of rest. After the warm-up, 2, 6-s MVICs were performed with a 2-min rest after each MVIC before performing one of the randomly ordered intermittent isometric protocols. At least 48-hrs were allowed between each of the protocols. The highest MVIC torque value on the second visit (immediately prior to the first protocol) was used to calculate the torque values at 40, 60, and 80% MVIC for both protocols. Pretest (prior to the fatiguing workbout), posttest (immediately after fatiguing workbout), and 5-min posttest (5-min after the end of the workbout) MVIC values were measured for each of the intermittent isometric protocols to track the fatigue-related changes in maximal isometric torque. The 60% MVIC protocol consisted of 50, 6-s isometric muscle actions of the leg extensors followed by 2-s of rest (6:2 work to rest ratio) at 60% MVIC (totaling 50 repetitions at 60% MVIC each separated by 2-s of rest). The 40/80% MVIC protocol consisted of 50, 6-s isometric muscle actions of the leg extensors beginning with a 6-s repetition at 40% MVIC with 2-s of rest followed by a 6-s repetition at 80% MVIC with 2-s of rest. This alternating pattern continued until 50 repetitions were performed 319

Figure 1. Depiction of the 60% and 40/80% maximal voluntary isometric contraction (MVIC) protocols. Each protocol consists of 50 intermittent isometric muscle actions of the leg extensors with equal average load. (totaling 25 repetitions at 40% MVIC and 25 repetitions at 80% MVIC each separated by 2-s of rest) (Figure 1). The 60% MVIC and 40/80% MVIC protocols performed equal, averaged load across the 50 repetitions (60% MVIC + 60% MVIC/2= 60% MVIC; and 40% MVIC + 80% MVIC/2= 60% MVIC). The subjects tracked their torque production on a computer monitor placed in front of them that displayed a real-time, digitalized torque signal overlaid onto a programmed template identifying their target torque value. The isometric template and real-time torque signal overlay used the raw voltage from the isokinetic dynamometer and was displayed using a custom written program in LabVIEW (LabVIEW version 13.0 National Instruments, Austin, TX). Electromyographic and Mechanomyographic Measurements Bipolar (20 mm inter-electrode distance) surface electrode (circular 4 mm diameter silver/silver chloride, BI- OPAC Systems, Inc., Santa Barbra, CA) arrangements were placed over the VL, VM, and RF of the dominant thigh according to the recommendations from SENIAM 17 and Barbero et al. 18. The bipolar electrode arrangement over the VL was placed 33% the distance between the lateral side of the patella and the anterior superior iliac spine, orientated 20 with respect to the reference line, and moved 5 cm laterally 17,18. The bipolar electrode arrangement over the VM was placed 20% the distance between the medial side of the patella and the anterior superior iliac spine and orientated 50 with respect to the reference line 17,18. The bipolar electrode arrangement over the RF was placed 50% of the distance between the superior side of the patella and the anterior superior iliac spine 17. The reference electrode was placed over the iliac crest. The electromyographic signals were amplified (gain: 1000x) using differential amplifiers (EMG 150 BIOPAC Systems, Inc., Santa Barbra, CA, ) and digitally bandpass filtered (fourth order Butterworth) at 10-500 Hz. The mechanomyographic signals were only detected from the VL using a miniature triaxial accelerometer (Measurement Specialties EGAS-FT 10, bandwidth 0-200 Hz, dimensions: 1.0 x 1.0 x 0.5 cm, mass 1.0 g, sensitivity 5504 mv/g) placed between the bipolar surface electrode arrangement located over the VL using double-sided adhesive tape. The locations of the bipolar electrode arrangement and accelerometer were marked with permanent black marker at the end of each testing session to ensure the same placements in subsequent sessions. Signal processing The raw electromyographic and mechanomyographic signals were digitized at 2000 Hz with a 16-bit analog-to-digital converter (Model MP150, BIOPAC Systems, Inc., Santa Barbra, CA) and stored in a personal computer for subsequent analysis. All signal processing was performed using custom programs written with LabVIEW programming software (Version 13.0, National Instruments, Austin TX). The electromyographic and mechanomyographic signals were zero-meaned and bandpass filtered (fourth-order Butterworth) at 10-500 Hz and 5-100 Hz, respectively. The electromyographic and mechanomyographic amplitude and frequency values were calculated from 2-s epochs corresponding to the middle 33% of each MVIC and normalized to the initial MVIC values. Statistical analysis All statistical analyses were performed on pooled data of men and women. Separate 2 [Protocol: 60%, 40/80%] x 3 [Muscle: VL, VM, RF] x 3 [Time: pretest, posttest, 5-min posttest] repeated measures ANOVAs were performed on the normalized electromyographic amplitude and normalized electromyographic frequency values from the MVICs. In addition, separate 2 [Protocol, 60%, 40/80%] x 3 [Time: pretest, posttest, 5-min posttest] repeated measures ANOVAs were performed on the MVIC torque, normalized mechanomyographic amplitude, and normalized mechanomyographic frequency values from the MVICs. When appropriate, Tukey post-hoc comparisons were performed. An alpha of p 0.05 was considered statistically significant for all ANOVA analyses (SPSS Version 22.0, Armonk, NY). 320

Figure 2. Maximal voluntary isometric contraction (MVIC) torque (Nm) values for the 60% MVIC and 40/80% MVIC protocols. (Note: There were no significant (p>0.05) differences between the protocols for MVIC torque. * Significantly (p 0.05) different from pretest. ** Significantly (p 0.05) different from pretest and posttest). Figure 3. Normalized marginal means for electromyographic amplitude (root mean square) collapsed across protocol and muscle. (Note: There were no significant (p>0.05) protocol or muscle-related differences or changes across time for electromyographic amplitude following 50 intermittent isometric muscle actions of the leg extensors at 60% maximal voluntary isometric contraction (MVIC) or 40/80% MVIC). Results Figure 2 includes the mean±sd and the results of the ANO- VA analyses for the MVIC torque for the 60% and 40/80% MVIC protocol at the pretest, posttest, and 5-min posttest measurements. The MVIC torque decreased from pretest to posttest and 5-min posttest for both protocols. In addition, the 5-min posttest MVIC torque values from both protocols were greater than posttest values, but did not recover to pretest values (Figure 2). The 2 x 3 ANOVA (protocol by time) for the MVIC torque values resulted in a non-significant interaction and main effect for protocol, but a significant main effect for time. Post-hoc comparisons for MVIC torque values (collapsed across protocols) indicated that pretest > posttest, pretest >5- min posttest, and 5-min posttest >posttest (Figure 2). Figure 3 and 4 include the mean±sd and the results of the ANOVA analyses for the electromyorgaphic amplitude and frequency for the 60% and 40/80% MVIC protocol at the pretest, posttest, and 5-min posttest measurements. There were no changes in electromyographic amplitude from the 321

Figure 4. Normalized marginal means for electromyographic frequency (mean power frequency) from the vastus lateralis (VL), vastus medialis (VM), and rectus femoris (RF). (Note: There were no significant (p>0.05) differences between the VL and RF following 50 intermittent isometric muscle actions of the leg extensors at 60% maximal voluntary isometric contraction (MVIC) or 40/80% MVIC. * Significantly (p 0.05) different from pretest. ** Significantly (p 0.05) different from posttest). Figure 5. Normalized marginal means for mechanomyographic amplitude (root mean square) from the vastus lateralis collapsed across protocol. (Note: There were no significant (p>0.05) protocol-related differences or changes across time for mechanomyographic amplitude following 50 intermittent isometric muscle actions of the leg extensors at 60% maximal voluntary isometric contraction (MVIC) or 40/80% MVIC). 322

Figure 6. Normalized marginal means for mechanomyographic frequency (mean power frequency) from the vastus lateralis collapsed across protocol. (Note: There were no significant (p>0.05) protocol-related differences following 50 intermittent isometric muscle actions of the leg extensors at 60% maximal voluntary isometric contraction (MVIC) or 40/80% MVIC. * Denotes significantly (p 0.05) different from pretest. ** Denotes significantly (p 0.05) different from posttest). VL, VM, and RF from pretest to posttest and 5-min posttest for either protocol (Figure 3). In addition, there were no changes in electromyographic frequency from the VM, but were decreases for the VL and RF (Figure 4). Specifically, both protocols indicated that the pretest values for electromyographic frequency from the VL and RF were lower than posttest and 5-min posttest values, however, after 5-min of recovery the electromyographic frequency was greater than posttest values (Figure 4). The 2 x 3 x 3 ANOVA (protocol by muscle by time) for the normalized electromyographic amplitude values measured during the MVIC resulted in no significant interactions or main effects. The 2 x 3 x 3 ANOVA (protocol by muscle by time) for the normalized electromyographic frequency values measured during the MVIC, however, resulted in no significant three-way interaction, but significant two-way interactions for protocol by muscle and muscle by time. Therefore, the model was decomposed in 3 separate 2 x 3 ANOVAs (protocol by time), one for each muscle (VL, VM, and RF). The follow-up 2 x 3 ANOVAs (protocol by time) for the normalized electromyographic frequency values from the VL and RF resulted in non-significant twoway interactions and main effects for protocol. There were, however, significant main effects for time for both the VL and RF muscles. Post-hoc comparisons for the normalized electromyographic frequency values indicated that for both the VL and RF muscles, pretest >posttest and 5-min posttest >posttest. The follow-up 2 x 3 ANOVA (protocol by time) for the normalized electromyographic frequency values from the VM, however, resulted in a non-significant two-way interaction, but a significant main effect for protocol (60% MVIC protocol >40/80% MVIC protocol). Figure 5 and 6 include the mean±sd and the results of the ANOVA analyses for the mechanomyographic amplitude and frequency for the 60% and 40/80% MVIC protocol at the pretest, posttest, and 5-min posttest measurements. There were no changes in the mechanomyographic amplitude from the VL from pretest to posttest and 5-min posttest for either protocol (Figure 5). Both protocols indicated that the pretest mechanomyographic frequency values from the VL were lower than posttest values, however, after 5-min of recovery the mechanomyographic frequency recovered to pretest levels (Figure 6). The 2 x 3 ANOVA (protocol by time) for the normalized mechanomyographic amplitude values measured during the MVIC resulted in no significant two-way interaction or main effects. The 2 x 3 ANOVA (protocol by time) for the normalized mechanomyographic frequency values, however, resulted in no significant interaction or main effect for protocol, but a significant main effect for time. Post-hoc comparisons indicated that pretest > posttest and 5-min posttest >posttest. Discussion In the present study, there were no differences between the 60% MVIC and 40/80% MVIC protocols for MVIC torque responses measured pretest, posttest, and 5-min posttest (Figure 2). The MVIC torque decreased by 26 and 22% im- 323

mediately following the fatiguing 60% MVIC and 40/80% MVIC protocols, respectively, and remained depressed from pretest levels by 9% for both protocols at 5-min posttest. These findings were in agreement with Fowles et al. 13 who reported 41, 28, and 26% decreases in MVIC torque immediately, 1-hr, and 4-hrs following 30-mins of intermittent isometric muscle actions of the leg extensors at 60% MVIC, respectively. Although the MVIC torque of Fowles et al. 13 did not return to pretest levels following 4-hrs of recovery, the values at 1-hr and 4-hrs following the fatiguing workbout were greater than immediately posttest. In addition, Saugen et al. 19 reported 44, 30, 27, and 21% decreases in MVIC torque, immediately, 10-min, 20-min, and 30-min following intermittent isometric muscle actions of the leg extensors to exhaustion at 40% MVIC. Thus, the findings of this investigation and previous studies 1,9,13 indicated that MVIC torque was decreased by up to 44% immediately following fatiguing, intermittent, isometric muscle actions of the leg extensors, and remained depressed by up to 26% for 4-hr. The magnitude of the fatigue-related decreases in MVIC torque following the workbouts as well as the lengths of time that MVIC torque remained depressed were likely due to the volume of work performed during the intermittent isometric protocols. In the present study, the 60% MVIC and 40/80% MVIC protocols had no effects on electromyographic amplitude from the VL, VM, and RF, or mechanomyographic amplitude from the VL immediately or 5-min posttest. Thus, varying the patterns of muscle loading (repeated 60% MVICs versus alternating 40 and 80% MVICs), while maintaining an equal average muscle loading (60% MVIC) across the 50 intermittent isometric muscle actions did not affect the amplitude responses of the electromyographic or mechanomyographic signals measured during the posttest and 5-min posttest MVICs. These findings were in partial agreement with those of Seghers et al. 10 who reported 15.2 and 15.3% decreases in MVIC torque, but no changes in electromyographic amplitude from the biceps brachii following 20-min of intermittent isometric muscle actions of the forearm flexors at 25 and 50% MVIC, respectively. Therefore the present findings, in conjunction with Seghers et al. 10 suggested that fatiguerelated decreases in MVIC torque as a result of submaximal, intermittent isometric muscle actions were not accompanied by changes in muscle activation (electromyographic amplitude) from the superficial muscles of the quadriceps femoris or the biceps brachii. Furthermore, the fatiguing, intermittent isometric protocols had no effect on motor unit recruitment as indicated by the lack of change in mechanomyographic amplitude from the VL. Unlike electromyographic amplitude, there were muscleand time-related differences for electromyographic frequency as a result of the fatiguing, intermittent isometric protocols. For the VL and RF, there were no differences between the electromyographic frequency responses for the 60% MVIC and 40/80% MVIC protocols. For both protocols, electromyographic frequency decreased from pretest to posttest (14 to 19%), but returned to pretest levels after 5-min of recovery (Figure 4). For the VM, however, there were no time-related changes in electromyographic frequency for either protocol, but the mean electromyographic frequency (collapsed across time) for the 60% MVIC protocol was greater than the 40/80% MVIC protocol. Thus, the fatiguing, intermittent isometric protocols had no time-related effect on electromyographic frequency from the VM measured during the MVICs. These results suggested muscle-specific, fatigue-related responses as indicated by decreases in electromyographic frequency for the VL and RF muscles, but not the VM. These findings were in agreement with Linssen et al. 20 who also reported no change in electromyographic frequency from the VM following fatiguing, submaximal (80% MVIC), intermittent isometric muscle actions of the leg extensors. In addition, the current findings were in partial agreement with those of Kouzaki et al. 11 who reported decreases in electromyographic frequency from the VL, VM, and RF (11 to 24%) immediately following 50 maximal intermittent isometric muscle actions of the leg extensors. The differences between the electromyographic frequency responses from the VM in the present study and that of Linssen et al. 20 versus those of Kouzaki et al. 11 may be attributable to differences in the intensity of the muscle actions (maximal versus submaximal). Kouzaki et al. 11 utilized maximal intermittent isometric muscle actions, while the present study and that of Linssen et al. 20 used submaximal (40 to 80% MVIC) muscle actions. It is also possible that the differences in electromyographic frequency responses for the VL and RF versus VM in the present study were due to muscle-specific muscle fiber type characteristics. Johnson et al. 21 reported that, on average, the VM was composed of 61.5% type I fibers, while the VL and RF were characterized by 46.9 and 42.8% type I fibers, respectively. Thus, the VM may be more fatigue-resistant than the VL and RF. Perhaps, the repeated maximal muscle actions used by Kouzaki et al. 11 were sufficiently fatiguing to cause decreases in electromyographic frequency in all three superficial muscles of the quadriceps femoris, while the submaximal muscle actions in the present study and that of Linssen et al. 20 were not. Differences in knee joint angles can also affect torque production as well as electromyographic amplitude and electromyographic frequency responses from the VL and VM 22. Therefore, it is possible that differences in the knee joint angles between the present study (120 ) and that of Kouzaki et al. 11 (90 from full extension) may have contributed to the muscle-specific electromyographic frequency responses as a result of the fatiguing protocols. Future studies should examine the effects of fatiguing, intermittent isometric protocols at various knee joint angles on the torque and neuromuscular responses of the leg extensors. In the present study, there were no differences between the 60% MVIC and 40/80% MVIC protocols for mechanomyographic frequency responses from the VL measured pretest, posttest, and 5-min posttest (Figure 6). The mechanomyographic frequency decreased by 24 and 23% immediately following the fatiguing 60% MVIC and 40/80% MVIC protocols, respectively, and then returned to pretest levels after 5-min of recovery. These findings were in agree- 324

ment with Kouzaki et al. 11 who reported 18 to 26% decreases in mechanomyographic frequency from the VL, VM, and RF immediately following 50 maximal intermittent isometric muscle actions of the leg extensors. Therefore, the present findings, in conjunction with Kouzaki et al. 11, suggested that fatigue-related decreases in MVIC torque immediately following the fatiguing, intermittent isometric muscle actions were accompanied by changes in the global motor unit firing rate (mechanomyographic frequency) from the superficial muscles of the quadriceps femoris rather than decreases in motor unit recruitment (mechanomyographic amplitude). After 5-min of recovery MVIC torque remained depressed by 9% from pretest levels, however, mechanomyographic frequency returned to pretest levels. It is plausible that MVIC torque remained depressed by 9% after 5-min of recovery due to excitation-contraction coupling failure. Thus, the decreases in global motor unit firing rate (mechanomyographic frequency) tracked the decreases in MVIC torque immediately following the fatiguing protocols, but not after 5-min of recovery. In summary, MVIC torque decreased immediately following the fatiguing, intermittent isometric muscle actions, and remained depressed after 5-min of recovery for both the 60% MVIC and 40/80% MVIC protocols. Of the neuromuscular parameters measured during the MVICs in the present study, electromyographic frequency from the VL and RF and mechanomyographic frequency from the VL tracked the fatigue-induced decreases in MVIC torque immediately following the 60% MVIC and 40/80% MVIC protocols. These neuromuscular parameters, however, returned to pretest levels following 5-min of recovery, while MVIC torque remained depressed by 9%. Furthermore, there were no changes in electromyographic amplitude from the VL, VM, and RF, or mechanomyographic amplitude from the VL for either the 60% MVIC or 40/80% MVIC protocols. Thus, there were no timerelated differences between the 60% MVIC and 40/80% MVIC protocols for the neuromuscular or torque responses and these neuromuscular parameters did not track the fatigue-induced decreases in MVIC torque immediately and/or 5-min posttest. References 1. Enoka RM, Duchateau J. Muscle fatigue: what, why and how it influences muscle function. J Physiol 2008; 586(1):11-23. 2. Basmajian JV, De Luca C. Muscles alive: their functions revealed by electromyography. 1985;278:126. 3. Linnamo V, Moritani T, Nicol C, Komi PV. Motor unit activation patterns during isometric, concentric and eccentric actions at different force levels. J Electromyog Kinesio 2003;13(1):93-101. 4. Søgaard K, Gandevia SC, Todd G, Petersen NT, Taylor JL. The effect of sustained low-intensity contractions on supraspinal fatigue in human elbow flexor muscles. J Physiol 2006;573(2):511-523. 5. Gordon G, Holbourn A. The sounds from single motor units in a contracting muscle. J Physiol 1948; 107(4):456-464. 6. Orizio C, Gobbo M, Diemont B, Esposito F, Veicsteinas A. The surface mechanomyogram as a tool to describe the influence of fatigue on biceps brachii motor unit activation strategy. Historical basis and novel evidence. Euro J Appl Physiol 2003;90(3-4):326-336. 7. Orizio C. Muscle sound: bases for the introduction of a mechanomyographic signal in muscle studies. Critic Rev Biomed Eng 1993;21(3):201-243. 8. Krustrup P, Bangsbo J. Physiological demands of topclass soccer refereeing in relation to physical capacity: effect of intense intermittent exercise training. J Sport Sci 2001;19(11):881-891. 9. Ljungberg A-S, Kilbom Å, HĀGg GM. Occupational lifting by nursing aides and warehouse workers. Ergonomic 1989;32(1):59-78. 10. Seghers J, Spaepen A. Muscle fatigue of the elbow flexor muscles during two intermittent exercise protocols with equal mean muscle loading. Clin Biomech 2004; 19(1):24-30. 11. Kouzaki M, Shinohara M, Fukunaga T. Non-uniform mechanical activity of quadriceps muscle during fatigue by repeated maximal voluntary contraction in humans. Euro J Appl Physiol 1999;80(1):9-15. 12. Bigland-Ritchie B, Cafarelli E, Vøllestad N. Fatigue of submaximal static contractions. Acta Physiol Scand 1985;556:137-148. 13. Fowles J, Green H, Tupling R, O brien S, Roy B. Human neuromuscular fatigue is associated with altered Na+- K+-ATPase activity following isometric exercise. J Appl Physiol 2002;92(4):1585-1593. 14. Yung M, Mathiassen SE, Wells RP. Variation of force amplitude and its effects on local fatigue. Euro J Appl Physiol 2012;112(11):3865-3879. 15. Yung M, Wells R. Physical variation in low-load work - physiological effects during exposure & recovery. Work 2012;41:5731-5733. 16. Yung M, Bigelow PL, Hastings DM, Wells RP. Detecting within- and between-day manifestations of neuromuscular fatigue at work: an exploratory study. Ergonomic 2014;57(10):1562-1573. 17. Hermens HJ, Freriks B, Merletti R, et al. European recommendations for surface electromyography. Roessingh Research and Development 1999;8(2):13-54. 18. Barbero M, Merletti R, Rainoldi A. Atlas of muscle innervation zones: understanding surface electromyography and its applications. Springer Science & Business Media; 2012. 19. Saugen E, Vollestad N, Gibson H, Martin P, Edwards R. Dissociation between metabolic and contractile responses during intermittent isometric exercise in man. Exper Physiol 1997;82(1):213-226. 20. Linssen WH, Stegeman DF, Joosten EM, et al. Fatigue in type I fiber predominance: a muscle force and surface EMG study on the relative role of type I and type II muscle fibers. Muscle Nerve 1991;14(9):829-837. 325

21. Johnson MA, Polgar J, Weightman D, Appleton D. Data on the distribution of fibre types in thirty-six human muscles. An autopsy study. J Neuro Sci 1973;18(1):111-129. 22. Chan AY, Lee FL, Wong PK, Wong CY, Yeung SS. Effects of knee joint angles and fatigue on the neuromuscular control of vastus medialis oblique and vastus lateralis muscle in humans. Euro J Appl Physiol 2001;84(1-2):36-41. 326