Original Research

Neuromuscular Fatigue Induced by a Mixed Martial

Art Training Protocol
Louis-Solal Giboin, and Markus Gruber
Human Performance Research Center, Sensorimotor Performance Lab, University of Konstanz, Konstanz, Germany

Abstract
Giboin, L-S and Gruber, M. Neuromuscular fatigue induced by a mixed martial art training protocol. J Strength Cond Res XX(X):
000–000, 2019—Mixed martial arts (MMA) is a full-contact sport whose popularity and professionalism are rapidly growing.
However, the specific physiological demands of this sport have been only scarcely studied so far, and especially the amount or type
of neuromuscular fatigue induced by an MMA bout remains completely unknown. We estimated neuromuscular fatigue of knee
extensors muscles during and after an MMA training protocol designed to simulate the physiological demands of MMA competition
in competitive practitioners (n 5 9) with isometric maximal voluntary force (MVF), potentiated muscle twitch at rest (Ptw), and
voluntary activation (VA). Bayesian linear mixed models showed that the training protocol induced a reduction of MVF, Ptw, and VA.
Although the largest reduction across time of VA was smaller than the largest reduction of Ptw, an effect of VA, but not of Ptw, was
found on MVF variation. The training protocol induced neuromuscular fatigue, with a larger peripheral (Ptw) than central component
(VA). However, despite the large decrease in Ptw, force production capacity was related only to VA, indicating that central control
might play an important role in the compensation of the peripheral fatigue components estimated with Ptw. This central com-
pensation can most probably prevent a too large loss of muscle force during the training protocol.
Key Words: MMA, central fatigue, peripheral fatigue, muscle fatigue, psychoneurophysiology, fight sport

Introduction to several processes, including neuromuscular fatigue. Neuro-

muscular fatigue could be defined as an exercise-induced tran-
Mixed martial arts (MMA) is a full-contact fight sport whose
sient reduction in muscle power or force, irrespective to the
concept originates from at least the classical antiquity era (e.g.,
capacity to continue or not the physical fatiguing task (9). Neu-
Pankration (11)). The currently most used set of rules has been
romuscular fatigue is composed of peripheral fatigue,
formalized in 2001 (25). Its goal consists in dominating an op-
i.e., mechanisms occurring distal to the neuromuscular junction,
ponent using strike and grappling techniques in a given space
and central fatigue, i.e., mechanisms taking place in the central
(ring or cage) and time (in general 3 3 5 minutes for non-
nervous system (10). The proportion of these different fatigue
championship fight and 5 3 5 minutes for championship fight).
components and their evolution across time throughout and after
Since the first North American fight card in 1993, MMA has
the fatiguing task seems to be task specific (6). Neuromuscular
experienced an exponential rise in popularity and pro-
fatigue and its components can be estimated with several different
fessionalism. Despite its recent popularity, little is known about
methods. Neuromuscular fatigue is most of the time estimated
the physiological demands of MMA (22). The particular nature of
with the decrease in isometric maximal voluntary force (MVF)
MMA makes it difficult to extrapolate its physiological demands
produced during isometric maximal voluntary contractions
and adaptations from studies performed on athletes from other
(MVCs) measured before and after the fatiguing task (10). In
fight sports (2). Indeed, the specific anaerobic and aerobic effort
exercise physiology, central fatigue is often estimated with the
combination due to intermittent high-intensity phases interspaced
decrease in voluntary activation (VA), i.e., the capacity to vol-
within a relatively long total effort time (8,30) and the broad skill
untarily recruit muscles. Voluntary activation can be assessed
sets that require different types of physiological adaptations are
with the twitch interpolated method that consists in stimulating
very specific to MMA (22). This gap in the knowledge may act as
electrically the motor nerve during and after the MVC (29).
a brake in regard to the optimization of the athletes’ training and
Muscle force response can be roughly estimated with the ampli-
recovery and slow down the maturing process of the sport-
tude of the potentiated muscle twitch (Ptw) elicited by maximal
specific elite performance.
motor nerve electrical stimulation at rest. A decrease in Ptw can be
It has been recently suggested that the neuromuscular charac-
considered as an indicator of decreased muscle excitability or
teristics of lower-body muscles could be a predictor of competi-
force production capability (36). These specific methods have
tive success in MMA. A high level of lower-body neuromuscular
been demonstrated to be highly reliable when used with knee
performance seems essential to display the technical repertoire of
extensor muscles in young healthy subjects (32). The amount of
athletes during competitive fights (21). However, neuromuscular
neuromuscular fatigue and the proportion of peripheral and
performance is susceptible to change during physical activity due
central fatigue, occurring during an MMA fight remain un-
Address correspondence to Louis-Solal Giboin, louis-solal.giboin@uni- known. Such information seems to be crucial for athletes and
konstanz.de. coaches as a decline of lower-body neuromuscular performance
Journal of Strength and Conditioning Research 00(00)/1–9 throughout an MMA fight due to fatigue might increase the risk
ª 2019 National Strength and Conditioning Association of defeat and injury. With such knowledge in mind, changes in

Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
 The Neuromuscular Fatigue of MMA (2019) 00:00
pacing or tactics within the fight or specific technical and physical
preparation between fights could be applied to optimize the
neuromuscular performance of lower-body muscles throughout
the whole duration of the fight and enhance probability of vic-
tory. The focus on knee extensor muscles is important not only
because of the relationship between lower body strength and
MMA competition success (21) but also because of the large ac-
cumulation of basic and applied knowledge regarding knee
extensors muscles neuromuscular fatigue.
Therefore, the aim of this study was to determine the amount
of neuromuscular fatigue in knee extensors muscles, as esti-
mated by the changes in MVF, VA, and Ptw, during and after
a MMA training protocol that simulates as best as possible
a MMA fight.
Methods
Experimental Approach to the Problem
The study was divided in 2 experimental days separated by 24
hours of rest. During the first day, we measured quadriceps
muscle isometric maximal voluntary force (MVF), potentiated
muscle twitch at rest (Ptw), and VA to investigate influence of
peripheral and central mechanisms of neuromuscular fatigue as-
sociated with a MMA training protocol. During the second day,
we measured physiological characteristics of the subjects (skin-
fold thickness, jump height, and aerobic capacity). In addition, we
assessed the sports history of subjects with a questionnaire. The
general procedure is summarized in a flowchart (Figure 1).
Subjects
Nine healthy young men participated in the study (mean 6 SD;
age: 28 6 7 years; age range: 20 to 38 years; body mass: 75.4 6 8
kg; height: 173 6 6 cm, all subjects were above 18 years old).
Before any data collection, subjects were informed of the risks and
benefits of the study and signed an institutionally approved in-
formed consent document. The study and its methods were spe-
cifically approved by the ethics committee of the University of
Konstanz and in accordance with the Declaration of Helsinki.
Subjects were included only if they were active MMA competitors
with at least one amateur or professional fight record and if they
were currently free from any lower-limb injuries. Subjects were
asked to not consume any caffeine or alcohol on the day of the
experiment and to not perform training that is not part of their
usual routine 48 hours before the experiment.
Figure 1. General procedure. Vertical arrows correspond to neuromuscular measurements which were performed
before the MMA training protocol (Pre), at the end of each round, and during the recovery period following the
training protocol. MMA 5 Mixed martial arts.
Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
Procedures
Sport History of Subjects. Individual sport history details are
displayed in Table 2. Subjects trained 7.3 6 5.1 years specifically
in MMA, 9.6 6 8.4 years in another fight sports than MMA, and
16.3 6 6.5 years in nonfight sports. At the time of the study,
subjects were training for MMA (including relevant general
physical preparation) 9.6 6 5.1 hours per week. Subjects had 2.4
6 2.5 amateur fights and 6.1 6 11.6 professional fights.
Mixed Martial Arts Training Protocol. During the training pro-
tocol, subjects interacted with a sparring partner holding mitts or
a shield during stand-up phases. Subjects were instructed to per-
form at the required intensity (low or high) and were given verbal
encouragement during the entire training protocol. Subjects did
not receive any strikes so strike-induced tissue damage would not
interact with the interpretation of neuromuscular fatigue meas-
urements. Subjects were permitted to wear their preferred training
kit (short, rash guard or not, and training or competition gloves),
but without shin-pads. Before the training protocol, each subject
completed a self-directed 10-minute warm-up based on their
previous training experience (this warm-up corresponds to warm-
up 2 in Figure 1). The protocol consisted of 3 rounds, each with
a duration of 5 minutes. Because of the neuromuscular meas-
urements performed in between, rounds 1 and 2 were separated
by 194 6 32 seconds and rounds 2 and 3 by 176 6 26 seconds.
Subjects completed the entire training protocol (including warm-
up) in approximately 30 minutes. Excluding time between
rounds, the training protocol was designed to simulate the
physiological demands of actual MMA competition. For this, the
total effort was divided in different effort types following the
observations of Miarka et al. (30), which have analyzed the effort
displayed by professional middleweight MMA fighters during the
first round (stand-up with low or high intensity, ground work
with low and high intensity, and rest sequences). Therefore, each
round consisted of upper- and lower-limb stand-up striking of
low and high intensity using focus mits, kick shields, and a heavy
bag, ground grappling and upper- and lower-limb ground strikes
of low and high intensity, and an active rest interval. Those
training activities were combined into a circuit of 75 seconds
duration, which was completed 4 times per round. Specifically,
each circuit consisted of the following: 25 seconds of standing
strikes at low intensity (;4 combinations of 2 punches and 1
kick), 10 seconds of standing strikes at high intensity (punching
and kicking with the highest intensity possible against a heavy
bag), 10 seconds of active recovery (as subjects transitioned from
 The Neuromuscular Fatigue of MMA (2019) 00:00
stand-up striking to ground grappling), 20 seconds of low-
intensity ground work with a training partner (subjects were
placed under mount, side control, or in guard with the aim to
improve their position and apply a submission hold on their
partner), and 10 seconds of high-intensity ground strikes from the
mount position (punches, elbows, and knees with the highest
intensity possible into a heavy bag placed on the mat).
Neuromuscular Measurements. Subjects started with a warm-up
consisting of 2 sets of 10 body mass squats and 2 sets of 3
countermovement jump (CMJ) (30-second rest between sets; this
warm-up corresponds to warm-up 1 in Figure 1). This warm-up
was used to ensure that the performance during the baseline
neuromuscular measurements was maximal. We taped electro-
myography (EMG) sensors (Trigno Wireless EMG system, Delsys
Inc.) on the muscle belly of the vastus lateralis (VL) and biceps
femoris (BF) according to SENIAM recommendations (www.
seniam.org). The skin underneath EMG sensors was previously
shaved, abraded, and cleaned with alcohol. Subjects sat in a cus-
tom-made chair with arm crossed on their chest and hip and trunk
and knee position maintained using noncompliant straps. The
custom-made chair had a hole in the seat to let the EMG sensor
positioned on the BF muscle go through. The knee angle was set at
110°. The right ankle strap was attached to a force transducer
(Model 9321A, Kistler). We stimulated the femoral nerve with
percutaneous electrodes (copper custom-made electrode wrapped
in water-soaked sponges). We fixed the cathode (circular, 2 cm of
diameter) over the right femoral triangle and the anode (rectan-
gular, 5 3 7 cm) over the lower part of the right gluteus maximus
muscle. The position of each electrode and EMG sensor was
marked with a waterproof pen. We stimulated the femoral nerve
with 1 ms squared electrical pulses (Stimulator DSH7, Digitimer).
The intensity of the nerve stimulation was raised until reaching
the maximal M-wave (Mmax) in the VL and until the amplitude
of the muscle twitch was not increasing anymore (average stim-
ulation intensity to reach Mmax was 56 mA). The biceps femoris
activity was monitored during femoral nerve stimulation to en-
sure that stimulation did not also increase activity in the antag-
onist muscle. The intensity was then multiplied by 1.5 and stayed
constant during the whole experiment (31). Following this pro-
cedure, subjects warmed up with incremental submaximal 5-
second isometric knee extensions until reaching around 90% of
perceived maximal effort. We used around 15–20 submaximal
contractions, depending on the force of each subject. The first
5–10 contractions were performed at very low intensity (force
output around 170 N) and were used to instruct subjects about
performing isometric contractions with a steady plateau. Then,
contraction intensity and resting time in between contractions
were gradually increased. The gradual increase in intensity be-
tween contractions was estimated according to the perceived
difficulty of each contraction by subjects. The aim was to get from
the first submaximal contractions to the last submaximal con-
traction before maximal effort in 5–10 contractions. This pro-
cedure is regularly used in our laboratory to optimize the MVF
measurement reliability while minimizing the fatigue induced by
the contractions (13,14,16). After 2 minutes of rest, we started the
VA procedure. During the VA procedure, subjects performed an
MVC of around 3 seconds, and the femoral nerve was stimulated
during the force plateau and around 1 or 2 seconds after the
muscle relaxation (see Figure 2). Subjects were encouraged with
standardized verbal encouragements just before and during the
contraction (before: “Are you ready to give your maximum!?”
and “Prepare to push as hard as possible!”; during: “Push, push,
Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
| www.nsca.com
push!”). This procedure was performed again after 2 minutes of
rest. If the MVF produced during the MVC increased by at least
5%, the procedure would have been repeated until no increase of
more than 5% in MVF was observed. However, it must be noted
that such increase did not occur for any of the subjects. The VA
procedure was repeated between rounds during the training
protocol (after round 1, 2, and 3) and after completion of the
training protocol (5, 10, and 15 minutes after the end of the VA
procedure after round 3). After the end of the training protocol, in
between neuromuscular measurements, subjects stayed on the
chair but unstrapped and had free access to water. During the
training protocol, the EMG sensors and stimulation electrodes
were removed and systematically replaced on the marks drawn on
the skin after wiping the sweat with a tissue and alcohol before
every neuromuscular measurement. To get neuromuscular
measurements as relevant as possible, we tried to position stim-
ulation and EMG electrodes in less than 2 minutes after the end of
the round. Indeed, it is known that MVF, VA, and Ptw can sub-
stantially recover in the few minutes necessary to move to the
required apparatus after the end of the fatiguing task (36). If that
was not possible, only MVF was obtained. This relatively long
delay is explained by the travel between the training protocol area
(far from any measurement devices for safety reasons) and posi-
tioning of EMG sensors and stimulation electrodes. The neuro-
muscular measurements were conducted 109 6 12 seconds, 106
6 19 seconds, and 105 6 21 seconds after the end of round 1, 2,
and 3, respectively.
Neuromuscular Measurements Analysis. EMG signals were am-
plified (3 1,000), high- and low-pass–filtered (20 6 5 and 450 6
50 Hz respectively), and sampled at 4,000 Hz. EMG and force
signals were recorded on a computer with the Signal software
(Cambridge Electronic Design) through a power 1,401 interface
(Cambridge Electronic Design). Maximal voluntary force corre-
sponded to the MVF produced during each MVC, i.e., the highest
amplitude of voluntary force. Ptw corresponded to the maximal
amplitude of the potentiated twitch at rest obtained after each
MVC. Voluntary activation was calculated using the formula VA
5 100 3 (1 2 interpolated twitch/Ptw) (29). During some trials,
the stimulation electrode could not be adequately repositioned
within time and no stimulations were elicited (2 Ptw and 3 VA
data point missing). EMGvl corresponded to the rms EMG of the
VL muscle of the epoch of 250 ms leading to the stimulation
inducing the interpolated twitch. EMGvl was normalized to the
rms of the Mmax elicited by the same stimulation (on an epoch of
20 ms starting at the beginning of the M-wave). For the Mmax
analysis, every potential was visually inspected by the main in-
vestigator and it was ensured that the epoch of 20 ms englobed the
full potential and no ongoing EMG. During some trials, the EMG
sensor could not be fixed securely within time due to the sweating
(6 data points missing). The MVF, VA, Ptw, and EMGvl of Pre
(measurement performed before the training protocol) were
obtained from the MVC that produced the highest MVF.
Physiological Characteristics Assessments. First, we estimated
body fat with skinfold thickness (Harpenden skinfold calliper, Baty
International) following ISAK standards (28). We marked and
measured 8 skinfold thickness (triceps, subscapular, biceps, iliac
crest, supraspinale, abdominal, front thigh, and medial calf) in suc-
cession and duplicated the measures. If the 2 measures differed by
more than 10%, a third one was performed. Then, subjects per-
formed 3 CMJs on a force plate (Leonardo Mechanograph) sepa-
rated by 1 minute of rest and with the instruction to jump as high as
 The Neuromuscular Fatigue of MMA (2019) 00:00

Figure 2. Measure of neuromuscular fatigue. The figure displays the force (in N)
produced by one subject before the training protocol (Pre, light line) and 5 minute
after the end of the training protocol (Recov +59, dark line). The vertical arrows
indicate the interpolated twitch.

possible with the hands on the hips (15). Finally, subjects did a ramp
test on a cycling ergometer (ergoselect200; ergoline GmbH), and gas
exchanges were monitored breath by breath (Ergostick; Geratherm
Respiratory GmbH) and analyzed with the Blue Cherry software
(Geratherm Respiratory GmbH). The test consisted of a 5-minute
warm-up at 60 W followed by a progressive increase in power
(ramp) until subject’s exhaustion or volitional termination of the
task. The ramp was followed by 1 minute of rest and 5 minutes of
cool-down consisting in cycling at 60 W. The ramp was calculated
according to the body mass of the subjects (25% of the fighter’s body
mass rounded to 15, 20, or 25 W·min21) (24). Subjects had to
maintain a constant cadence (between 70 and 90 rotations per
minute) and received strong verbal encouragements. One subject
stopped prematurely the ramp test because of discomfort (ramp test
data not analyzed). The heart rate (HR) during the training protocol
and the ramp test was measured with a chest belt (Polar V800).
However, because of system failure of the device or movement from
the belt, we could only analyze data in 7 subjects during the training
protocol and ramp test.
Physiological Characteristics Analysis. HRmax and HRmean
corresponded to the highest HR measured during each round of
the training protocol and the mean HR of each round, re-
spectively. Estimation of adiposity was made by summing the

Figure 3. Neuromuscular fatigue induced by the training protocol. This figure displays the MVF (N), the Ptw (N), and the VA (%)
before (Pre), during the training protocol (after each round), and 5, 10, and 15 min after the end of the MVF following round 3
(Recov). The dark line and points correspond to the mean values. The error bars correspond to SD. The colored lines and
points correspond to individual values. MVF 5 maximal voluntary force; VA 5 voluntary activation; Ptw 5 potentiated muscle
twitch at rest.

Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
 The Neuromuscular Fatigue of MMA (2019) 00:00 | www.nsca.com

Table 1
Posterior estimates.*†
Variable Time level Estimate SD Lower 95% CI Upper 95% CI ES
MVF (N) Intercept 697.43 38.69 620.71 774.40
Round 1 256.23 14.05 282.75 228.47 20.53
Round 2 264.74 12.66 289.76 239.73 20.57
Round 3 271.52 15.06 2101.11 241.35 20.62
Recov 5’ 298.69 19.70 2137.18 259.23 20.8
Recov 109 272.25 14.44 2101.29 244.32 20.62
Recov 159 264.87 14.53 293.61 236.09 20.58
VA (%) Intercept 94.10 2.61 88.97 99.34
Round 1 22.84 1.46 25.75 0.07 20.42
Round 2 22.41 1.22 24.82 20.06 20.35
Round 3 26.17 3.64 213.73 0.33 20.65
Recov 59 29.27 2.16 213.55 24.93 21.17
Recov 109 23.79 1.14 26.07 21.55 20.57
Recov 159 25.18 1.70 28.52 21.87 20.64
Ptw (N) Intercept 154.14 8.99 135.81 172.00
Round 1 227.72 6.73 241.15 214.36 20.88
Round 2 231.19 5.46 241.77 220.53 21.11
Round 3 251.32 12.98 277.09 225.33 21.27
Recov 59 245.34 8.59 262.11 228.87 21.3
Recov 109 229.00 5.53 239.67 217.88 21
Recov 159 226.52 5.69 237.85 215.22 20.93
EMGvl (%) Intercept 32.90 3.33 26.40 39.66
Round 1 22.15 4.12 210.12 6.15 20.1
Round 2 21.10 3.38 27.76 5.72 0.05
Round 3 21.84 3.94 29.42 5.91 20.1
Recov 59 27.49 3.28 213.85 21.04 20.92
Recov 109 21.87 3.82 29.45 5.53 20.03
Recov 159 21.62 3.18 28.01 4.57 20.15
MVF (N) Intercept 49.74 13.44 23.25 76.44
VA 0.42 0.16 0.09 0.74
Ptw 0.01 0.07 20.12 0.14
HRmean (b·min21) Intercept 160.64 8.58 143.56 177.57
Round 2 2.55 7.64 212.43 18.07
Round 3 21.35 7.80 217.09 13.70
HRmax (b·min21) Intercept 179.09 4.90 169.38 188.75
Round 2 1.47 5.51 29.77 12.70
Round 3 24.83 6.26 217.14 7.03
*CrI 5 Credible Interval; MVF 5 maximal voluntary force; VA 5 voluntary activation.
†Posterior distribution of constant estimates is described with mean, SD, and upper and lower bound of the 95% credible interval (CrI). The intercept row corresponds to the estimate at baseline level (i.e., before
the training protocol). The estimates given for round 1, 2, and 3 and at recovery time of 59, 109, and 159 must be read as the mean at time x is greater or lower by y units than the mean at baseline. ES
corresponds to effect size. The effect size is calculated against Pre values.

skinfold thickness of the 8 sites (1). Following ISAK recom-
mendations, the skinfold thickness at each site was equal to the
average of measurements if 2 measures were performed or the
median if 3 measures were performed (28). Jump height was de-
termined as the highest height estimated from the 3 CMJ (Leo-
nardo GRFP 4.3 software). V̇ O2 peak was estimated as the highest
oxygen uptake, and HRmax ramp corresponded to the highest
HR measured during the ramp test.
Statistical Analyses
R was used for statistics (3.3.0, The R Foundation for Statistical
Computing). The R package brms (5) was used to test the effect of
time on MVF, Ptw, VA, and EMGvl with the Bayesian linear
mixed model (LMM). Time was set as fixed effect (population-
level effect). The random effect structure of each model was
maximized with a varying intercept by-subject and varying slope
by-subject for the time variable (3). In addition, we calculated
within subjects effect sizes with Cohen’s dav formula, where the
mean difference is divided by the average SD and used the ap-
proximated Hedges’ correction (17). Furthermore, with variables
expressed in percent of pre, it was tested whether VA and Ptw
could explain MVF with a model including random intercept by-
subject and random slope by-subject for VA and Ptw. In addition,
the difference of HR (mean and maximal) between the rounds of
the training protocol was tested with Bayesian 1-way analysis of
variance with repeated measurement. All models were con-
structed with student family function as it allows “robust” linear
regression that is less influenced by outliers. Uninformative priors
set per default by the package were used. Four Markov chain
Monte Carlo (MCMC) with 4,000 iterations each were used for
each test (with a warm-up of 2000 iterations). The convergence of
all models was checked with the Rhat value and by visually
inspecting the MCMC (5). The fit of models’ posteriors to data
was also checked with pp_check(). Results from the models are
given with the mean estimate (95% lower credible interval, 95%
upper credible interval) of the posterior distribution. This 95%
probability interval can be interpreted as the probability that the
population parameter lies between the upper and lower bound

Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
 The Neuromuscular Fatigue of MMA (2019) 00:00

(42). In this study, we interpret that a parameter estimate is dif- HRmaxramp was equal to 182 6 2 b·min21 when including exactly
ferent from zero when zero lies outside of the 95% credible in- the same subjects for both variables (n 5 6).
terval bounds.

Physiological Characteristics of Subjects

Results
Training Protocol
Maximal voluntary force, Ptw, and VA results are displayed in
Figure 3 and posterior estimates are given in Table 1. Individual
values of physiological and training variables are given in Table 2.
Maximal voluntary isometric contraction reduced after round 1 and
did not return to baseline values during the recovery period. The
lowest MVF value is observable at recovery 59. The largest decline in
VA occurred not during the training protocol but during the recovery
period. The lowest VA value was observable at recovery 59. Ptw
declined as soon as the first round and did not fully recover at re-
covery 159. The lowest value was observable at the end of round 3,
indicating that Ptw started to recover as soon as the training protocol
was over. In addition, we can see (Figure 4 and Table 1) that EMGvl
declined only at recovery 59. For a better graphical understanding of
changes across time, MVF, Ptw, and VA were expressed in per-
centage of pre for each subject and then averaged and displayed in
Figure 5. The linear mixed model showed that VA had an effect on
MVF: For each unit increase in VA, MVF increases by 0.42
(0.09–0.74) unit. However, the estimate of the Ptw on MVF was
smaller and its 95% credible interval included zero, suggesting that
Ptw had no credible effect on MVF variation. HRmean (n 5 7, mean
6 SD) was equal to 159 6 6 b·min21 during round 1, 162 6 9
b·min21 during round 2, and 157 6 5 b·min21 during round 3.
HRmax (n 5 7) was equal to 179 6 9 b·min21 during round 1, 180
6 9 b·min21 during round 2, and 172 6 19 b·min21 during round 3.
Considering the rather large credible intervals, it could not be stated
whether the number of rounds had an effect on HRmean and
HRmax (Table 1). HRmax was equal to 184 6 7 b·min21 and
The sum of the 8 skinfolds (n 5 9) was 65.6 6 13.6 mm. The
maximal jump height during CMJ (n 5 9) was 43.8 6 4.9 cm. The
V̇ O2 peak (n 5 8) was 45.3 6 6.2 ml·kg21·min21. HRmaxramp (n
5 7) was 183 6 4 b·min21.
Discussion
This study aimed to determine the influence of a MMA training
protocol on knee extensor muscles’ neuromuscular fatigue and
inform our understanding of the mechanisms that may underpin
that fatigue in experienced MMA competitors. The main results
of this study show that MVF and Ptw were already significantly
decreased compared with baseline at the end of the first round of
the training protocol and remained below baseline during the 15
minutes of recovery period. Voluntary activation did not clearly
decrease below baseline until round 3 and remained below
baseline during the 15-minute recovery period. Although the
decrease of Ptw was larger than the decrease of VA, we observed
a significant effect of VA, but not of Ptw, on MVF changes.
Reduced knee extensor muscle MVF confirms that the MMA
training protocol used was sufficient to produce quadriceps
neuromuscular fatigue, which remained below baseline despite
a 15-minute recovery period. Our findings are supported by
a previous study that showed impaired countermovement jump
performance after participation in MMA competition (with
strikes) (12).
A reduction of Ptw was observed immediately after the end of
the first round, indicating the quick occurrence of nonvoluntary
muscle excitability reduction. The reduction of Ptw seemed to be

Table 2
Individual values of physiological and training variables.*†
MMA Other fight Training MMA training Amateur
Subjects Age Weight Height training sport training sport per week fights Pro fights CMJ
1 20 72.1 173 4 4 8 12 7 0 39
2 32 81 179.5 7.5 13.5 22 6.5 3 1 53
3 30 81.1 178.5 2 13 16 20 0 6 42
4 33 88.8 169 14 26 26 6 0 2 45
5 38 79.8 168 17 5 17 11 5 36 39
6 25 62.3 161.5 7 15 17 6 3 9 45
7 20 62 175 3 0 5 5 1 0 43
8 20 76.7 175.6 4 0 16 14 2 0 50
9 35 75 177 7 10 20 6 1 1 39
Subjects VȮ 2 peak 8 skinfolds HrRmaxramp Hrmean R1 Hrmax R1 Hrmean R2 Hrmax R2 Hrmean R3 Hrmax R3
1 49.2 66.1 183 128 177 168 185 120 145
2 50.7 63.4 177 155 176 120 163 125 146
3 na 65.6 na 142 171 144 178 150 177
4 35.3 89.9 na na na na na na na
5 47.5 55.1 180 155 167 164 175 168 177
6 48.8 54.2 186 na na na na na na
7 50.7 45.1 179 177 186 177 189 174 185
8 43.3 72.2 187 184 193 181 187 180 187
9 36.6 79.27 187 174 184 178 186 180 187
*MMA 5 mixed martial arts; CMJ 5 countermovement jump.
†Age, MMA training, other fight sport training, and training sport are given in years. Mass is given in kg. Height and CMJ are given in cm. MMA training per week is given in hours (including general physical
preparation). Amateur fights and pro fights are given in number of fights. VȮ 2 peak is given in ml·kg21·min21. Skinfold is given in mm (from 8 sites). HR is given in b·min21.

Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
 The Neuromuscular Fatigue of MMA (2019) 00:00 | www.nsca.com

Figure 4. EMGvl. Display of the rms EMG of the vl muscle normalized to the rms Mmax before (Pre),
during the training protocol (after each round), and 5, 10, and 15 minutes after the end of the training
protocol (Recov). The dark line and points correspond to the mean values. The error bars correspond to
SD. The colored points and lines correspond to individual values.

at its highest at the end of the training protocol and started to
recover already 5 minutes after. Interestingly, despite the large
reduction of Ptw occurring during the training protocol, decrea-
ses in MVF were not significantly associated with decreases in
Ptw. This phenomenon can be explained by neural mechanisms
able to counteract the effect of peripheral fatigue on voluntary
force production. An increase of neural drive can compensate the
loss of muscle force production during exercise by acting on the
motoneuron pool firing rate and recruitment (7). This suggestion
is supported by the fact that MVF changes were better explained
by the modulation of VA than Ptw. However, it must be noted
that the present EMG data show on one hand that the highest
decrease in EMG coincides with the largest decrease in VA and in
MVF, i.e., 5 minutes after the end of the training protocol, but on
the other hand that there was no increase in neural drive across
the training protocol. Nevertheless, care must be taken when

Figure 5. Neuromuscular fatigue expressed in percentage of Pre. MVF (dark circles), Ptw (teal
triangles), and VA (yellow squares) variables were expressed in percentage of Pre values for each
subject and then averaged. Error bars correspond to SD. MVF 5 maximal voluntary force; VA 5
voluntary activation; Ptw 5 potentiated muscle twitch at rest.

Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
 The Neuromuscular Fatigue of MMA (2019) 00:00
interpreting EMG with the present protocol and during MVC
because amplitude cancellation can drastically affect the signal.
Amplitude cancellation corresponds to the overlapping of posi-
tive and negative phases of motor unit potentials that cancel each
other and reduce the amplitude of the EMG signal (23).
Unlike the decreased Ptw, VA (which represents the neural ability
to activate knee extensor muscles) only clearly decreased below
baseline at the completion of the training protocol. Those findings,
i.e., a decrease of VA only close to task termination despite pre-
cocious decrease in Ptw, are supported by previous studies in-
vestigating the effect of endurance tasks on neuromuscular fatigue
(7,26,35). The failure to maintain VA and to compensate the accu-
mulating peripheral fatigue at exercise termination can be explained
by several processes. According to the “sensory tolerance limit”
model, the accumulative increase in feedforward and feedback sig-
nals from working and remote muscles and from various neural
processes during the training protocol may have reached the task-
specific threshold inducing neural drive inhibition (19). The correct
timing between neural drive inhibition and the end of the training
protocol could be explained by pacing strategies, as modeled by the
“psychobiological model of endurance performance.” This model
links performance regulation and, thereby, peripheral fatigue regu-
lation, with initial potential motivation, perception of effort,
knowledge of the task, and past experience (27,34). This interaction
between both models remains speculative, and specifically dedicated
psychoneurophysiolological studies must be conducted to support
this possible hypothesis.
It is remarkable that the largest reduction of MVF and VA did not
occur just at the end of the training protocol but 5 minutes after. At
this time point, Ptw was already recovering, which implies that the
delayed reduction of MVF cannot be completely explained by af-
ferent inhibitory feedback loops (“sensory tolerance limit” model) or
by the corollary discharge that regulates perception of effort (“psy-
chobiological model of endurance performance”). Indeed, regarding
the first model (19), a recovery of Ptw should be associated with
a reduction of the inhibitory feedback signaling from the muscles to
the central nervous system, implying a reduction of neural drive
inhibition. In this context, VA would have been expected to follow
the recovery of Ptw. In regard to the second model (27,34), the
recovery in muscle excitability implies that the same amount of force
output can be reached with a lesser neural drive. The reduction in
neural drive would lead to a reduced corollary discharge and thus
a reduced perceived effort for a given voluntary force output. In this
context, a higher MVF would have been expected. Unfortunately, in
this study, no muscle sensory afferent fiber activity and rate of per-
ceived effort were measured, which could have helped to support (or
infirm) our hypothesis. This unexpected discrepancy between the
recovery of Ptw and VA indicates that mechanisms of neuromus-
cular fatigue are yet to be fully discovered. We speculate that the
present observation could—at least partly—be explained by a shift
in the amount of self-control subjects were willing to allocate to the
task (for details, see (4,37,38)). Psychological research has shown
that such shifts can indeed affect performance in strenuous physical
tasks (16,43). In the present case, for the MMA athletes, performing
as hard as possible during the MMA training protocol would have
been valued enough to apply sufficient self-control to endure the
large accumulation of fatigue and limit neuromuscular impairments
(20). However, 5 minutes after the end of the training protocol, the
intrinsic value of producing a MVC was probably not enough to
apply the level of self-control necessary to reach previous perfor-
mance, especially in the context of still large amount of peripheral
fatigue. Therefore, we propose that the amount of VA displayed
Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
during the task (where VA compensates the occurrence of peripheral
fatigue) or after the task (where VA is more impaired despite a lower
amount of peripheral fatigue) could be under the top-down control
of the neural structures related in the application of self-control (38).
As limitations of this study, it must be noted that Ptw obtained
with a single stimulation is only a rough estimate of peripheral
fatigue. Indeed, Ptw is sensitive to postactivation potentiation,
which is specific to the fatiguing exercise performed, and to
changes in muscle stiffness (31). To better estimate peripheral
fatigue in humans, Ptw should ideally be completed with obser-
vations of Mmax amplitude changes and observation of the effect
of tetanus stimulations at different frequencies on muscle force
production (36). Similarly, the estimation of VA with motor nerve
stimulations is not without issues and is not indicative in regard to
the specific sites responsible of neural drive changes or the
mechanisms responsible for the changes in neural drive (36,40).
The present observations of neural changes during exercise
should be ideally completed by, for example, observations made
with transcranial magnetic stimulation to assess changes in cor-
tical VA or other corticospinal mechanisms (41), or even with
functional near-infrared spectroscopy to assess changes in corti-
cal sites that are known to interact with brain motor regions (16).
To clarify the present relationship between peripheral fatigue and
central compensation mechanisms, further investigation should
be performed with a larger set of methods.
Furthermore, it was difficult to recruit MMA athletes who matched
our inclusion criteria. In consequence, the sample size was small and
composed of athletes of heterogeneous level. Therefore, care must be
taken before attempting to generalize the present results to the whole
MMA athlete population. For example, the heterogeneity in training
and competition experience may have affected the neuromuscular
fatigue results differently among subjects. However, it must be noted
that the CMJ SD was relatively small, indicating a low heterogeneity
in lower-limb power among subjects. Furthermore, despite our efforts
in this regard, the time between the end of the round and the neuro-
muscular measurements remained long. Thus, the observed reduction
in VA may be underestimated (18). Yet, this underestimation does not
change the present conclusion because the largest central fatigue value
was observed 5 minutes after the end of the training protocol, which is
by far longer than the time in between the end of any round and the
following neuromuscular measurement.
We have shown that an MMA training protocol that was
designed to simulate the physiological demands of actual MMA
competition can induce considerable neuromuscular fatigue in
knee extensor muscles, as seen with the decrease in MVF, Ptw,
and VA. Despite the larger decrease in Ptw than in VA, MVF was
sensitive to VA rather than Ptw impairments. We suggest that in
MMA, neurophysiological factors are important regulators of
overall neuromuscular performance.
Practical Applications
We have shown that neuromuscular fatigue of knee extensor
muscles was occurring during the MMA training protocol. Spe-
cific training could potentially reduce the continuous decrease of
force production capability across rounds. High-intensity interval
training and task-specific training could increase the tolerance of
the central nervous system to peripheral fatigue and permit in-
creased muscular work to be performed before neuromuscular
failure (33,44). The lack of immediate neuromuscular recovery
after the end of the simulation should also be considered in
 The Neuromuscular Fatigue of MMA (2019) 00:00
competitive and training environments where repeated efforts are
required. Appropriate short-term effect recovery procedures could
be advised to reduce the deleterious effects of neuromuscular fa-
tigue on performance and accelerate neuromuscular recovery (39).
Decreased knee extensor muscles’ VA suggests that central
mechanisms contribute to the neuromuscular fatigue observed
in relation to the MMA training protocol. Therefore, proce-
dures that intend to facilitate recovery of the central nervous
system contribution to neuromuscular performance, such as
active recovery, may be advised (13).
The highest decrease in knee extensors’ neuromuscular perfor-
mance was seen following the completion of the training protocol.
Thus, including training sessions with specific competition con-
ditions (intensity, duration, specific rest between rounds, etc.) to
optimize the timing between the abrupt decrease in neuromuscular
performance and the end of a MMA competition could be
considered.
Acknowledgments
The authors thank Kristijan Milekic and Matthew Schaeberle for
their help with data collection and Fabien “Sonny” Hillairet,
a professional MMA coach, for his insights in regard to the sport
and for the discussion that led to this project. Supported by
Ausschuss für Forschungsfragen (AFF).
References
1. Ackland TR, Lohman TG, Sundgot-Borgen J, et al. Current status of body
composition assessment in sport: Review and position statement on behalf
of the ad hoc research working group on body composition health and
performance, under the auspices of the I.O.C. Medical commission. Sports
Med 42: 227–249, 2012.
2. Andreato LV, Branco BHM. Different sports, but the same physical and
physiological profiles? Sports Med 46: 1963–1965, 2016.
3. Barr DJ, Levy R, Scheepers C, Tily HJ. Random effects structure for confir-
matory hypothesis testing: Keep it maximal. J Mem Lang 68: 255–278, 2013.
4. Berkman ET, Hutcherson CA, Livingston JL, Kahn LE, Inzlicht M. Self-
control as value-based choice. Curr Dir Psychol Sci 26: 422–428, 2017.
5. Bürkner PC. brms. An R package for Bayesian multilevel models using
stan. J Stat Softw 80: 1–28, 2017.
6. Carroll TJ, Taylor JL, Gandevia SC. Recovery of central and peripheral neu-
romuscular fatigue after exercise. J Appl Physiol 122: 1068–1076, 2017.
7. Decorte N, Lafaix PA, Millet GY, Wuyam B, Verges S. Central and pe-
ripheral fatigue kinetics during exhaustive constant-load cycling. Scand J
Med Sci Sports 22: 381–391, 2012.
8. del Vecchio FB, Hirata SM, Franchini E. A review of time-motion analysis
and combat development in mixed martial arts matches at regional level
tournaments. Percept Mot Skills 112: 639–648, 2011.
9. Enoka RM, Duchateau J. Muscle fatigue: What, why and how it influences
muscle function. J Physiol 586: 11–23, 2008.
10. Gandevia SC. Spinal and supraspinal factors in human muscle fatigue.
Physiol Rev 81: 1725–1789, 2001.
11. Gardiner E. The pankration and wrestling. III. J Hellenic Stud 26: 4–22, 1906.
12. Ghoul N, Tabben M, Miarka B, et al. Mixed martial arts induces signif-
icant fatigue and muscle damage up to 24 hours post-combat. J Strength
Cond Res 33: 1570–1579, 2017.
13. Giboin LS, Amiri E, Bertschinger R, Gruber M. Active recovery affects the
recovery of the corticospinal system but not of muscle contractile prop-
erties. PLoS One 13: e0197339, 2018.
14. Giboin LS, Gruber M. Anodal and cathodal transcranial direct current
stimulation can decrease force output of knee extensors during an in-
termittent MVC fatiguing task in young healthy male participants.
J Neurosci Res 96: 1600–1609, 2018.
15. Giboin LS, Gruber M, Kramer A. Motor learning of a dynamic balance
task: Influence of lower limb power and prior balance practice. J Sci Med
Sport 22: 101–105, 2018.
16. Giboin L-S, Gruber M, Schüler J, Wolff W. Investigating performance in
a strenuous physical task from the perspective of self-control. Brain Sci 9: E317,
2019.
Copyright © 2019 National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited.
| www.nsca.com
17. Goulet-Pelletier JC, Cousineau D. A review of effect sizes and their con-
fidence intervals, Part I: The Cohen’s d family. Quantitative Methods
Psychol 14: 242–265, 2018.
18. Gruet M, Temesi J, Rupp T, et al. Dynamics of corticospinal changes during and
after high-intensity quadriceps exercise. Exp Physiol 99: 1053–1064, 2014.
19. Hureau TJ, Romer LM, Amann M. The sensory tolerance limit: A hypothetical
construct determining exercise performance? Eur J Sport Sci 18: 13–24, 2018.
20. Inzlicht M, Shenhav A, Olivola CY. The effort paradox: Effort is both
costly and valued. Trends Cogn Sci 22: 337–349, 2018.
21. James LP, Beckman EM, Kelly VG, Haff GG. The neuromuscular qualities
of higher- and lower-level mixed-martial-arts competitors. Int J Sports
Physiol Perform 12: 612–620, 2017.
22. James LP, Haff GG, Kelly VG, Beckman EM. Towards a determination of
the physiological characteristics distinguishing successful mixed martial
arts athletes: A systematic review of combat sport literature. Sports Med
46: 1525–1551, 2016.
23. Keenan KG, Farina D, Maluf KS, Merletti R, Enoka RM. Influence of
amplitude cancellation on the simulated surface electromyogram. J Appl
Physiol (1985) 98: 120–131, 2005.
24. Kramer A, Poppendieker T, Gruber M. Suitability of jumps as a form of
high-intensity interval training: Effect of rest duration on oxygen uptake,
heart rate and blood lactate. Eur J Appl Physiol 119: 1149–1156, 2019.
25. Lenetsky S, Harris N. The mixed martial arts athlete: A physiological
profile. Strength Cond J 34: 32–47, 2012.
26. Lepers R, Maffiuletti NA, Rochette L, Brugniaux J, Millet GY. Neuro-
muscular fatigue during a long-duration cycling exercise. J Appl Physiol
(1985) 92: 1487–1493, 2002.
27. Marcora SM, Staiano W. The limit to exercise tolerance in humans: Mind
over muscle? Eur J Appl Physiol 109: 763–770, 2010.
28. Marfell-Jones M, Olds T, Stewart A, Carter T. International Standards for
Anthropometric Assessment. Potchefsroom, South Africa: ISAK, 2006.
29. Merton PA. Voluntary strength and fatigue. J Physiol 123: 553–564, 1954.
30. Miarka B, Del Vecchio F, Brito C, Amtmann J. Comparisons of time-
motion analysis of mixed martial arts rounds by weight divisions. Int J
Perform Anal Sport 15: 1189–1201, 2015.
31. Millet GY, Martin V, Martin A, Vergès S. Electrical stimulation for testing
neuromuscular function: From sport to pathology. Eur J Appl Physiol
111: 2489–2500, 2011.
32. Morton JP, Atkinson G, MacLaren DP, et al. Reliability of maximal
muscle force and voluntary activation as markers of exercise-induced
muscle damage. Eur J Appl Physiol 94: 541–548, 2005.
33. O’Leary TJ, Collett J, Howells K, Morris MG. Endurance capacity and neu-
romuscular fatigue following high- vs moderate-intensity endurance training:
A randomized trial. Scand J Med Sci Sports 27: 1648–1661, 2017.
34. Pageaux B. The psychobiological model of endurance performance: An
effort-based decision-making theory to explain self-paced endurance
performance. Sports Med 44: 1319–1320, 2014.
35. Place N, Lepers R, Deley G, Millet GY. Time course of neuromuscular
alterations during a prolonged running exercise. Med Sci Sports Exerc 36:
1347–1356, 2004.
36. Place N, Yamada T, Bruton JD, Westerblad H. Muscle fatigue: From
observations in humans to underlying mechanisms studied in intact single
muscle fibres. Eur J Appl Physiol 110: 1–15, 2010.
37. Shenhav A, Botvinick MM, Cohen JD. The expected value of control: An
integrative theory of anterior cingulate cortex function. Neuron 79:
217–240, 2013.
38. Shenhav A, Musslick S, Lieder F, et al. Toward a rational and mechanistic
account of mental effort. Annu Rev Neurosci 40: 99–124, 2017.
39. Tavares F, Smith TB, Driller M. Fatigue and recovery in rugby: A review.
Sports Med 47: 1515–1530, 2017.
40. Taylor JL. Point: The interpolated twitch does/does not provide a valid measure
of the voluntary activation of muscle. J Appl Physiol (1985) 107: 354–355, 2009.
41. Todd G, Taylor JL, Gandevia SC. Measurement of voluntary activation of
fresh and fatigued human muscles using transcranial magnetic stimula-
tion. J Physiol 551: 661–671, 2003.
42. van de Schoot R, Kaplan D, Denissen J, et al. A gentle introduction to bayesian
analysis: Applications to developmental research. Child Dev 85: 842–860, 2014.
43. Wolff W, Bieleke M, Hirsch A, et al. Increase in prefrontal cortex oxy-
genation during static muscular endurance performance is modulated by
self-regulation strategies. Sci Rep 8:15756, 2018.
44. Zghal F, Cottin F, Kenoun I, et al. Improved tolerance of peripheral fatigue
by the central nervous system after endurance training. Eur J Appl Physiol
115: 1401–1415, 2015.