European Journal of Sport Science

ISSN: 1746-1391 (Print) 1536-7290 (Online) Journal homepage: http://www.tandfonline.com/loi/tejs20

Post-activation potentiation (PAP) in endurance

sports: A review

Daniel Boullosa, Sebastian del Rosso, David G. Behm & Carl Foster

To cite this article: Daniel Boullosa, Sebastian del Rosso, David G. Behm & Carl Foster (2018):
Post-activation potentiation (PAP) in endurance sports: A review, European Journal of Sport
Science, DOI: 10.1080/17461391.2018.1438519

To link to this article: https://doi.org/10.1080/17461391.2018.1438519

Published online: 01 Mar 2018.

Submit your article to this journal

View related articles

View Crossmark data

Full Terms & Conditions of access and use can be found at

http://www.tandfonline.com/action/journalInformation?journalCode=tejs20
European Journal of Sport Science, 2018
https://doi.org/10.1080/17461391.2018.1438519

REVIEW

Post-activation potentiation (PAP) in endurance sports: A review

DANIEL BOULLOSA1,2, SEBASTIAN DEL ROSSO1, DAVID G. BEHM3, &

CARL FOSTER4
1
Physical Education, Catholic University of Brasilia, Brasilia, Brazil; 2College of Healthcare Sciences, James Cook University,
QLD, Australia; 3School of Human Kinetics and Recreation, Memorial University of Newfoundland, Newfoundland, Canada
& 4Department of Exercise and Sport Science, University of Wisconsin, La Crosse, Wisconsin, United States of America

Abstract
While there is strong support of the usefulness of post-activation potentiation (PAP) phenomenon in power demanding
sports, the role that PAP could play in endurance sports has received less attention. The aim of this review is to present
evidence for a better understanding of PAP in endurance athletes; and to discuss the physiological basis and
methodological aspects necessary for better practices and designing further studies. A search for relevant articles on PAP
and endurance trained athletes was carried out using Medline and ISI Web of Knowledge databases. Twenty-two studies
were included in the review. The current evidence suggests the possible influence of PAP for performance enhancement
after appropriate conditioning activities during warm up. Evaluation of PAP responses during testing, training and
competition may be also important for athletes monitoring. There are many unresolved questions about the optimum load
parameters for benefiting from PAP in both training and competition; and the role that PAP may exert for optimal
performance while interacting with central and peripheral factors associated with muscle fatigue. Further studies should
elucidate the association between PAP responses and long-term adaptations in endurance athletes.

Keywords: Endurance, performance, testing, strength, physiology

Highlights
. Appropriate warm up exercises should be identified in every sport for performance enhacement during subsequent short
endurance events via PAP.
. It seems that endurance athletes exhibit greater PAP responses after submaximal conditioning activities as a consequence of
a better PAP/fatigue balance.
. Monitoring PAP, perceptual and muscle fatigue, during training and testing, could be very useful for a better understaning
of acute and chronic adaptations of endurance athletes.

Introduction
Post-activation potentiation (PAP) has been defined
as the phenomena by which muscular performance
characteristics are acutely enhanced as a result of
their contractile history (Tillin & Bishop, 2009).
The applicability of PAP to sport performance has
traditionally focused on power exercises, because
PAP after maximum efforts could be utilized as an
intervention that lasts several minutes after an acute
conditioning stimulus, thus increasing muscle
power output (Seitz & Haff, 2016). However, PAP
responses have also been observed in endurance
trained athletes, after maximal voluntary contractions
(MVCs) (Hamada, Sale, & Macdougall, 2000). In
addition, some recent studies have reported jump
potentiation after different endurance running exer-
cises in endurance athletes (Boullosa & Tuimil,
2009; Boullosa, Tuimil, Alegre, Iglesias, & Lusqui-
nos, 2011; Vuorimaa, Virlander, Kurkilahti, Vasan-
kari, & Hakkinen, 2006). Furthermore, Del Rosso
et al. (2016) have recently shown the possible influ-
ence of jump potentiation on pacing in a 30-km
trial. Meanwhile, scientists and practitioners in
endurance sports are often unaware of the great
potential of PAP for both training and competition
probably as a consequence of the dominant paradigm

Correspondence: Daniel A. Boullosa Physical Education, Catholic University of Brasilia, Brasilia, Brazil. E-mail: daniel.boullosa@gmail.com

© 2018 European College of Sport Science

2 D. Boullosa et al.
in PAP practices that associates maximum brief
efforts to PAP in subsequent power exercises (Seitz
& Haff, 2016).
Therefore, given the absence of a review approach-
ing this important topic, this article aims to present
the current evidence of PAP in endurance athletes;
and to discuss the physiological basis and methodo-
logical aspects of this ergogenic mechanism for
better practice during evaluation, training and com-
petition; and for designing further studies. We will
consider endurance activities as those lasting >
1 min (Chamari & Padulo, 2015). Additionally, and
following the most accepted definition of PAP
(Tillin & Bishop, 2009), we will consider as PAP
any increment in muscle performance, assessed
with different methods, after exercises of different
intensity and duration. Given the co-existence of
both muscle fatigue and potentiation after a number
of muscle contractions (Behm, Button, Barbour,
Butt, & Young, 2004; Rassier & Macintosh, 2000),
the definitions of Gandevia (2001) for ‘muscle
fatigue’ (i.e. decrease in force production capacity),
‘peripheral fatigue’ (i.e. decrease in evoked force),
and ‘central fatigue’ (i.e. decrease in maximal volun-
tary activation), should be considered onwards for
clarification.
Understanding PAP in the endurance
phenotype
Before the specific literature review, it is important
to address some physiological aspects for a better
understanding and contextualization of the evidence
of PAP in endurance athletes. Thus, the main mech-
anism for explaining PAP has been proposed to be
myosin RLC phosphorylation (Grange et al.,
1993). This transient adaptation at the molecular
level increases Ca2+ sensitivity in striated muscle,
thus enhancing force production for a given Ca2+
concentration (MacIntosh, 2010). Prior knowledge
of this mechanism was mostly obtained with single
muscles and isolated muscle fascicles or fibres,
after voluntary (i.e. PAP) or electrical stimulations
(i.e. staircase or post-tetanic potentiation) in
animal models (Vandenboom, Gittings, Smith,
Grange, & Stull, 2013). From these previous
studies, it seems that a greater potentiation could
be expected in fast-twitch fibres, at physiological
temperatures, with alkalosis, during concentric con-
tractions, and at shorter muscle lengths (MacIntosh,
2010; Vandenboom et al., 2013). It has been also
suggested, in disuse and knockout animal models,
that there is an independent mechanism to RLC
phosphorylation related to the elevation of resting
Ca2+ levels, but only after successive low-frequency
twitches (Vandenboom et al., 2013). Additionally,
the presence of adrenaline (Decostre, Gillis, &
Gailly, 2000) and estradiol concentrations (Lai,
Collins, Colson, Kararigas, & Lowe, 2016) has
been proposed to influence RLC phosphorylation
in the mouse. Meanwhile, the translational value of
this knowledge from rodents to humans and, more
specifically, to endurance athletes, should be taken
with caution. In fact, the few experiments with
humans that performed muscle biopsies, reported
similar levels of RLC phosphorylation between
fast- and slow-twitch fibres (Houston & Grange,
1991; Houston, Lingley, Stuart, & Grange, 1987;
Stuart, Lingley, Grange, & Houston, 1988), an
adaptation that may be speculated to be an evol-
utionary consequence of the physical demands of
our Homo ancestors (Boullosa, Abreu, Varela-
Sanz, & Mujika, 2013).
Other suggested mechanisms at the neuromuscular
level have been proposed after maximum or near
maximum conditioning activities mainly in power-
trained athletes. These mechanisms may include
increased recruitment of higher order motor units
(Hodgson, Docherty, & Robbins, 2005), and
changes in muscle fascicle pennation angles (Tillin
& Bishop, 2009). Moreover, the correlation reported
(r = −0.74) between the increase in twitch force after
a submaximal (75%) voluntary contraction and the
decline in discharge rate during submaximal volun-
tary contractions (10–30%) of the biceps brachii,
may suggest a link between PAP responses and a
reduced central drive for maintenance of a constant
force output (Klein, Ivanova, Rice, & Garland,
2001). However, there are recent individual studies
that do not support a contribution of an increased
motor neuron excitability through the reflex
pathway (Xenofondos et al., 2015); and voluntary
activation, corticospinal excitability, intracortical
inhibition and facilitation (Thomas, Toward, West,
Howatson, & Goodall, 2017), to PAP. Given that,
it could be concluded that RLC phosphorylation
could be considered the primary mechanism for
PAP, while other influences at the neuromuscular
level cannot be supported with the current evidence.
The use of the term PAP has been suggested to
be inappropriate without evaluation of the enhanced
response with a twitch to verify that the potentiation
occurs for the same stimulation as that used prior
to voluntary contractions (MacIntosh, 2010). It is
worth noting, however, that twitch verification is
also an indirect surrogate of the effect of RLC phos-
phorylation on muscle force production. It is
assumed that RLC phosphorylation favours an
increase in the rate of force development (RFD)
and peak tension because of the increase in the
number of cross-bridges formed (MacIntosh,
2010). This is an important consideration as an
 Post-Activation Potentiation (PAP) in Endurance Sports
increase in RFD and peak tension could be indirectly
evaluated with any voluntary exercise requiring rapid
or maximal force production. Therefore, we may
suggest the use of voluntary maximal explosive
evaluations as their relationships with PAP responses
has been extensively described (Seitz & Haff, 2016).
Furthermore, previous studies have reported the
relationship between potentiated twitch responses
and jump potentiation in different populations
(Mitchell & Sale, 2011; Nibali, Chapman, Robergs,
& Drinkwater, 2013; Requena, Sáez-Sáez de
Villareal, Gapeyava, García, & Pääsuke, 2011).
Nevertheless, caution should be also taken as jump
performances are highly variable among individuals
(Chaouachi et al., 2011), the instructions provided
to athletes could influence RFD (Sahaly, Vandewalle,
Driss, & Monod, 2001), and improvements of jump
performance after conditioning activities are not
always evident depending on the performance par-
ameter evaluated (Boullosa, Abreu, Beltrame, &
Behm, 2013; Pearson & Hussain, 2014). In fact,
not all performance improvements after conditioning
activities can be attributed to PAP. Positive acute
adaptations of warm-up activities such as elevation
of muscle temperature and increased metabolic
responses should be differentiated from potentiation
of contractile capacity per se (McGowan, Pyne,
Thompson, & Rattray, 2015). Moreover, the possible
influence of a central command during voluntary
movements should not be disregarded. Thus, assum-
ing that a true PAP effect could be solely verified with
the twitch interpolation technique, it could be
suggested the use of maximal explosive exercises as
jumps as a surrogate of PAP responses but with the
aforementioned limitations.
Any athlete could benefit from this mechanism after
any conditioning activity since the levels of muscle
fatigue were smaller than the levels of PAP. Moreover,
PAP should be maximized with low firing rates, which
occur during endurance activities, since increased sen-
sitivity to Ca2+ is maximized at low Ca2+ levels and
limited at saturated Ca2+ levels (Sale, 2002). This
phenomenon should be considered concurrently
with fatigue development with expected lower levels
of muscle fatigue at submaximal intensities (MacIn-
tosh, 2010). Given the greater fatigue resistance of
slow-twitch fibres, a better PAP/fatigue balance
would be expected with endurance trained muscles
during and after appropriate conditioning activities.
Further, differences in PAP/fatigue balance between
muscle phenotypes could be even more evident as
only slow-twitch fibres can maintain Ca2+ sensitivity
after prolonged endurance activities (i.e. 4 hrs.) as
seen in elite cyclists (Hvid et al., 2013).
Mettler and Griffin (2010) revealed a significant
positive correlation between force potentiation and
3
the force–time integral produced by the stimulation
train (r = 0.70), suggesting a dose-response relation-
ship between conditioning activity volume and the
degree of potentiation. The same authors (Mettler
& Griffin, 2012) showed that the time to achieve
maximal potentiation decreased with increased inten-
sity. Thus, force-time integral matched conditioning
contraction tests (5 s at 100% vs. 10 s at 50% vs.
20 s at 25%) did not differ in the amount of poten-
tiation produced. Of note, these previous studies
were performed in the adductor pollicies human
muscle, which is predominantly a slow-twitch
muscle (Mettler & Griffin, 2010, 2012), therefore
suggesting that, for the endurance phenotype, pro-
longed activities at submaximal intensities would
induce a greater force production capacity as a conse-
quence of a better PAP/fatigue relationship.
Given all the aforementioned evidence, it would,
therefore, be suggested that: (1) the main mechanism
behind PAP is myosin RLC phosphorylation, with
humans exhibiting similar levels between slow- and
fast-twitch fibres; (2) PAP evaluation could be per-
formed with twitch responses and indirectly with
voluntary explosive exercises although with limit-
ations; (3) for the endurance phenotype, prolonged
submaximal exercises, at least until durations that
do not promote significant levels of peripheral
fatigue, could favour a better PAP/fatigue balance.
A hypothetical model for understanding the role of
PAP in endurance activities is presented in Figure 1.
Review of literature
Search methods
A search for relevant articles on PAP in endurance
athletes and sports was carried out using Medline
and ISI Web of Knowledge databases. To be
included in this review, each article must have met
the following criteria: (1) participants should be
endurance trained athletes, (2) an endurance exercise
should be present, and (3) pre- and post-exercise
measures of muscle or motor performance.
Results
A total of 126 records were obtained from the search,
from which 86 were eliminated as duplicates or
unspecific to the topic. The remaining 40 full-text
articles were analysed for inclusion and subsequently
18 articles were eliminated as they did not meet all
inclusion criteria. Of the 22 articles included in the
literature review, two used MVC as a conditioning
activity (Hamada et al., 2000; Paasuke et al., 2007),
one used submaximal intermittent contractions
(Morana & Perrey, 2009), one compared different
4 D. Boullosa et al.
Figure 1. Hypothetical model of post-activation potentiation for endurance sports. RLC, regulatory myosin light chain, MLC-K, myosin light
chain kinase; MLC-P = myosin light chain phosphatase; AUC = area under the curve.
warm-up protocols (Skof & Strojnik, 2007), two
assessed the effects of warm-up on subsequent simu-
lated trial (Feros, Young, Rice, & Talpey, 2012; Silva
et al., 2014), 10 studied PAP using different exercise
set-ups (Boullosa et al., 2011; Boullosa & Tuimil,
2009; Garcia-Pinillos, Molina-Molina, & Latorre-
Roman, 2016; Garcia-Pinillos, Soto-Hermoso, &
Latorre-Roman, 2015; Latorre-Román, García-
Pinillos, Martínez-López, & Soto-Hermoso, 2014;
McIntyre, Mawston, & Cairns, 2012; Pageaux,
Theurel, & Lepers, 2017; Skof & Strojnik, 2006a,
2006b; Vuorimaa et al., 2006), and eight studies
looked at race simulations and competitions (Del
Rosso et al., 2016; Feros et al., 2012; Millet et al.,
2002; Millet, Martin, Maffiuletti, & Martin, 2003;
Millet, Millet, Lattier, Maffiuletti, & Candau, 2003;
Place, Lepers, Deley, & Millet, 2004; Rousanoglou
et al., 2016; Silva et al., 2014).
The most generalized output measures for asses-
sing PAP were voluntary peak torque or evoked
twitch peak torque and the countermovement jump
(CMJ). In all cases there were increases between
pre- and post-conditioning activity in the aforemen-
tioned variables. However, due to the different
research designs, the timing and degree of PAP
manifestation varied among studies. A summary of
the main characteristics for the articles included in
the literature review is provided in Table I.
Discussion
After analysis of the included studies, there are two
main issues to be considered regarding PAP in
endurance athletes: (1) the effect of PAP after
warm-up activities, and (2) the evaluation of PAP
with different methods, during testing, training, or
following endurance tests and competitions.
PAP for warm-up
PAP has been proposed as one of the most important
objectives of warm-up (McGowan et al., 2015). Fol-
lowing the traditional approach of PAP practices, the
most intuitive application of PAP is the acute
enhancement of contractile potential after warm-up
activities for training or competitive purposes.
However, very few studies in endurance sports have
demonstrated an improvement in muscle perform-
ance after different warm-up activities via PAP
Table I. Summary of the main characteristics for the articles included in the literature review.

Reference Subjects Exercise protocol Measurements Potentiation definition Main results

Boullosa and n = 12♂
Tuimil (2009) Trained distance runners
(from regional to elite,
competing at least 2
consecutive years)
Boullosa et al. n = 22♂(14) ♀(8)
(2011) Experienced endurance
athletes (8 ♀and 8
♂endurance runners,
6 ♂ triathletes)
Del Rosso et al. n = 11♂
(2016) Well trained half-marathon
runners
Maximal Run (UMTT)
Tlim (Constant running
pace at 100% MAS)
Maximal run (UMTT)
30-km self-paced multistage
trial (6 × 5-km stages)
CMJ (non-fatigued state,
CMJB)
Post-tests CMJ height (2 and
7 min)
ΔCMJ
Pre and post Ex. CMJ
Pre vs. post stages CMJ
height, stage speed
An increase in post-exercise
CMJ height
An increase in post exercise
CMJ height 20 m sprint
An increase in CMJ height
CMJB >Tlim vs. UMTT
↑CMJ at 2 (3.53%) min post Tlim
↑CMJ at 2 (12.6%) and 7 (6.76%) min post
UMTT
CMJ at 2 min post Tlim > CMJ at 2 min post
UMTT
↑CMJ 3.6% post vs. pre Ex.
↑peak power 3.4% Post vs. Pre Ex.
↓peak force 10.8% Post vs. pre Ex.
Cluster analysis (responders vs. non-responders):
↑ΔCMJ (4.9%), vs. Pre Ex. in responders
↑Δpeak power (5.8%) vs. pre Ex. in responders
↓ΔCM vertical path (9.7%) vs. pre Ex. in non-
responders
↓peak force (29.9%) vs. Pre Ex. in non-
responders
↔Speed at 5, 10, 15 and 20 km
↓Speed at 25, 30 km vs. 5, 10, 15 and 20 km
↑CMJ at 5, 10, 15, 20, 25 and 30 km vs. Pre

Post-Activation Potentiation (PAP) in Endurance Sports

Feros et al. n = 10♂(9) ♀(1) (PW) Self-selected warm-up
(2012) Elite level rowers + 5 × 5 s isometric
conditioning contraction
with 15 s recovery
between sets + 1000 m
rowing ergometer time
trial
(NW) Self-selected warm-
up + 1000 m rowing
ergometer time trial
Garcia-Pinillos n = 30♂ 4 × 3 × 400 m runs, 85–
et al. (2015) Experienced recreational 100% of MAS; 1 min
long-distance runners passive recovery between
(6 years of training and runs and 3 minutes
competition) between sets
↑CMJ at 25 km vs. 20 km
↑ΔCM at 25 km vs. 20 km
MPO, mean SR, and split Acute enhancement in > MPO at 0–500 m in PW vs. NW
time for every 100 m strength or power output >SR at 0–500 m in PW vs. NW
<500 m time split in PW vs. NW
<SR at 0–1000 m in PW vs. NW
Pre vs. Post Sets: Time for An increase in CMJ ↑CMJ height (∼5%). Rest vs. Set 1, 3 and 4
400 m (T400), CMJ height, performance (height or ↑Peak force (∼6–10%), Rest vs. Set 2, 3 and 4
peak force, peak power, mechanical parameters) ↑Peak power (∼8–13%), Rest vs. Set 2, 3 and 4
eccentric work, and Cluster Analysis (Responders):
concentric work. ↑CMJ (∼8–13%), Rest vs. Set 1, 2, 3 and 4
↑Peak force (∼10–13%), Rest vs. Set 2, 3 and 4
↑Peak power (∼13–17%), Rest vs. Set 2, 3 and 4
Responders vs Non-Responders
↑ΔCMJ (13.89%) in Responders vs. ↔ in Non-
Responders

(Continued)

5
 6
Table I. Continued.

D. Boullosa et al.
Reference Subjects Exercise protocol Measurements Potentiation definition Main results

Garcia-Pinillos n = 33 Maximal run (Léger Pre and post test CMJ height An increase in CMJ height ↑CMJ vs. Pre test
et al. (2016) Recreationally trained multistage test) and kinematic variables Cluster analysis (responders and non-
endurance runners (20♂ (video analysis) responders)
and 13♀) ↑CMJ in responders (∼8%) vs. Pre test
↑angular position of the ankle in non-responders
↔In any other kinematic variable
Hamada et al. n = 40 ♂ 10-s MVC of the elbow MVC post 5 min of exercise. % change in peak twitch ↑ PT in all groups (triathletes and runners >
(2000) 10 triathletes, 10 distance extensor and ankle PT, TPT, HRT. torque post-MVC active controls and sedentary)
runners, 10 active control plantar-flexor muscles ↓ TPT in all groups
subjects, 10 sedentary ↓ HRT in all groups
Significant
negative correlations between PAP and pre-MVC
twitch TPT and HRT
Latorre-Román n = 16♂ 4 × 3 × 400 m runs, 85– Pre vs. Post Sets: Time for An increase in CMJ ↑CMJ height (∼ 2–6%)
et al. (2014) Experienced sub-elite long- 100% of MAS; 1 min 400 m (T400), CMJ height, performance (height or ↔Peak velocity vs. Pre
distance runners passive recovery between peak velocity, flight time, mechanical parameters) ↑Peak Force (∼8–15%) vs. Pre
(six years of training and runs and 3 min between peak force, peak power, ↑Peak Power (∼10–13%) vs. Pre
competition) sets eccentric work and
concentric work
McIntyre et al. n = 10♂ Prolonged submaximal cycle Time to exhaustion, PPO, Increase in peak isokinetic Six cyclists completed two exercise and four
(2012) ten competitive or test to exhaustion while Peak isometric MVC, torque completed four to six stages
recreationally active (4 × 20 min at 70% isokinetic torque, DJ At stage 2 or 3 ↑PPO (∼140% vs. initial) in 5 of
cyclists VOpeak) + 30 s all-out height, ground contact and the 10 cyclists
sprint at the 17th min of reactivity coefficient (all At stage 2 ↓PPO (∼ 56–95% vs. initial) in the
each bout measured throughout the other 5 cyclists peak isometric
trial) ↓MVC torque (14%) at exhaustion.
At exhaustion:
↓PPO (81 ± 25% vs. Pretest)
↓Peak isometric torque (MVC) of quadriceps (86
± 11% vs. pretest)
↓peak concentric torque (quadriceps) (83 ± 10%
vs. pretest)
↓peak concentric torque (hamstrings) (93 ± 7%
vs. Pretest)
↓DJ height 92% vs. Pretest
↔Ground-contact time
Millet et al. n = 9♂ 65 km ultra-marathon Non fatigued (one week ND ↑PT, Average RFD, MRFDt, and MRFRt of PF
(2002) Trained triathletes and (altitude 2500 m) before) vs. Fatigued (∼ and KE vs. Non-Fatigued condition
endurance runners 2 min after the race), PT, ↓CT, HRT and MRFRt of PF and KE vs. non-
CT, MRFDt, HRT, fatigued condition
average RFD, MRFRt, and ↔M-Wave duration and amplitude except for
M-Wave (amplitude and muscle soleus (↑ M-Wave amplitude vs. non-
duration) of plantar flexors fatigued condition)
(PF) and knee extensor
(KE) muscles
Millet, Martin n = 11♂ 140 km road race Non fatigued vs. fatigued (Pre An increase in peak twitch ↓MVC (∼9%) vs. Pre
et al. (2003) Trained cyclists (regional vs. Post 15–30 min) trial, tension ↔VA vs. Pre
level) MVC, VA, Peak twitch ↑Peak twitch tension (∼12%) vs. Pre
tension, CT, Average RFD, ↑RFD (∼11%) vs. Pre
MRFDt, MRFRt ↔CT, MRFDt, MRFRt
Millet, Millet n = 11♂ Ski Skating Marathon Non-fatigued (2 days before) The post-tetanic ↓MVC (8.4%)
et al. (2003) Trained cross-country (∼159.7 ± 17.9 min) vs. Fatigued (5 min after potentiation (PTP) was ↑PT, MRFDt, HRT and MRFRt
skiers the race), MVC plus PT, calculated as Pt of tw2 ↔ P020, P080 and MRFD80
CT, MRFDt, HRT, divided by Pt of tw1 ↑MRFD20
average RFD, MRFRt ↑ P020/P080 ratio
(From evoked twitch), ↓PTP
P020, P080, MRFD80 and ↓RMS (∼30%) and M-wave amplitude
MRFD20 (from evoked ↔ M-Wave duration
tetanus) and M-wave (All vs. Non-fatigued condition)
(amplitude and duration)
plus RMS of KE muscles.
Morana and n = 15♂ Intermittent submaximal Pre and Post Ex. Increase in PT during the At 1 min of exercise, ↑PT (53%) for
Perrey (2009) (8 endurance athletes; exercise: 10-minute, 5 s × PT, TPT, HRT, VA, MRFD submaximal exercise POW vs. (52%) for END
distance 5 s cycles, of submaximal and MRFR protocol ↑PT in END still increased at 1.5 min
runners and triathletes, knee extensor (PT was monitored (56%) and remained high until the end of
training 7 h·wk−1 during contractions (50% of throughout the exercise exercise
the previous six months, MVC) protocol) At 5.5 min of exercise ↓PT in POW (30%)
END) In END ↓TPT throughout

Post-Activation Potentiation (PAP) in Endurance Sports

(7 power-trained athletes;
i.e. professional rugby
players
And weightlifters, POW)
Paasuke et al. n = 36♀
(2007) PW (sprinters, jumpers and
volleyball players), ET
(long-distance runners
and cross-country skiers),
UT (students, no history of
regular participation in
any sport)
10 s conditioning MVC of
the KE muscles
Pre vs. Post 2 s, 1, 3, 5, 10 and
15 min of recovery: MVC,
PT, CT, HRT, MRFD,
MRFR
In POW, ↓TPT only between 1 and 4 min of Ex.
↓MVC > in POW (36.2%) vs. END (15.3%)
At post Ex. ↑PT 28.2% for END vs. ↓PT 22.7%
for POW
↑MRFD (63.3%) and ↑MRFR (32.2%) for END
Enhancement of the ↓PT in PW (27%), ET (18%) and UT (21%),
electrically evoked twitch (PW > ET)
contraction peak torque ↑Twitch PT (51%) for PW, (44%) for ET and
after a brief MVC (30%) for UT, at 2 s vs. Pre Ex.
PAP for PT was still present for ET at 1 min
(13%), and for PW (14%) and UT (13%) at
5 min of recovery
↑PT at 2 s, 1, 3 and 5 min recovery >PW vs. ET
↑MRFD (125%) for PW, (79%) for UT and
(51%) for ET at 2 s vs. Pre Ex.
PAP for MRFD was still present for ET at 1 min
(24%) and for UT and PW at 3 min (26%) and
5 min (25%) of recovery
↑MRFD at 2 s, 1, 3, 5 and 10 min >PW vs. ET
↑MRFR (124%) for PW, (95%) for UT and
(76%) for ET at 2 s vs. Pre Ex.
PAP for MRFR was still present for ET at 2 s post
Ex., for UT at
3 min (26%) and for PW (12%) at 10 min

7
↔CT and HRT

(Continued)
 8
D. Boullosa et al.
Table I. Continued.

Reference Subjects Exercise protocol Measurements Potentiation definition Main results

Pageaux et al. n = 9♂
(2017) Endurance Athletes
(Triathlon or duathlon from
regional to national level)
Three conditions: Post Warm-Up (baseline) vs. An increase in TTW and ↓MVC Torque at Pre Run and Post Run vs.
Warm Up = 5 min at 33% of Pre run (after 1-h cycling / RFD of the KE Baseline (only in condition “A”)
MAP + 5 min 50% of uphill walking) ↓MVC Torque at Post Run vs. Pre Run (in
MAP vs. Post-15 min of rest (in the condition “C”)
A. 1-h cycling + 10 km 10 km only condition) vs. Post ↑MVC Torque at Post Run vs. Pre Run (in
running time trial 10 km running, MVC, TTW, conditions “A”)
B. 1-h uphill walking (at DT, tRFD, MMA, M-Wave >MVC Torque in conditions “C” and “D” vs.
the same HR than the condition “B” at Pre run
1-h cycling exercise) + ↓TTW at Pre run vs. Baseline (only in condition
10 km running time “A”)
trial (at the same ↑TTW at Pre run vs. Baseline (in condition “B”);
running velocity as at Post run vs. Baseline (in conditions “A” and
condition “A”) “B”) and at Post Run vs. Pre Run (in condition
C. 10 km running time “A”)
trial (at the same >TTW at Pre Run in conditions “B” and “C” vs.
running velocity as condition “A”
condition “A”) ↓DT at Pre Run vs. Baseline (in condition “A”)
and at Post Run vs. Baseline (in condition “C”)
↑DT at Post Run vs. Pre Run (in condition “A”)
>DT at pre run in conditions “B” and “C” vs.
condition “A”
↑tRFD at pre run vs. baseline (in condition “B”),
at post run vs. baseline (in conditions “A”, “B”
and “C”) and at post run vs. pre run (in
condition “A”)
>tRFD at pre run in conditions “B” and “C” vs.
condition “A”
>tRFD at post run in condition “A” vs. condition
“B”
↔M-wave (between time points and conditions).
↓MMA at pre and post run vs. baseline in
conditions “A” and “B”
Place et al. n=9
(2004) Well-trained triathletes and
endurance runners
Rousanoglou n = 27,
et al. (2016) Recreationally trained ultra-
marathon runners
Silva et al. (2014) n = 11♂
Trained cyclists
(specialist in speed
cycling trials with an
average training
background of 4 years)
Skof and Strojnik n=7 ♂
(2006a) Well-trained middle- and
long-distance runners
Treadmill running Pre vs. intra-trial (1 to 5 h) vs.
(300 min at 55% MAS) Post-Trial (30 min) MVC,
RMS, RMS/RMSM, VA,
PT, CT, HRT, MRFDt,
MRFRt, doublet maximal
torque (DMT), P020, P080,
M-Wave amplitude and
duration
23 km-Mountain race Pre-Race (Pre Warm-Up) and
(Altitude = 554 m, initial post race (1 and 5 min)
5.2 km uphill, followed by CMJ (mechanical variables)
10.5 km of alternating
downhill and uphill trails
and a final downhill of
8 km)
Leg press, 4 sets of 5RM Pre vs. Mid vs. Post 20 km
Time Trial
Power output, pedal cadence,
cycling economy.
PAP vs. Non-PAP
comparisons
5 × 300 m, interval runs at Pre and Post (50–60 s, 3, 10,
95% of 400 m speed (77% 20, 30, 40, 60, and
Smax) + 1 min recovery 120 min) TTW; CT;
between the runs EMD; HRT; TF20; TF100
An increase in doublet ↓MVC at 4 h (26 ± 23%) and 5 h (28 ± 27%) vs.
torque Pre
↓RMS/RMSM at 4 h (50 ± 21%), 5 h (45 ± 27%)
and Post-30 min (50 ± 30%) vs. Pre
↔VA during the first 3 h
↓VA at 4 h (12 ± 17%), 5 h (16 ± 21%) and Post-
30 min (21 ± 28%) vs. Pre
↑PT at 5 h (18 ± 18%) vs. pre-trial
↑DMT at 2 h (12 ± 5%), 4 h (15 ± 15%) and 5 h
(14 ± 14%) vs. pre-trial
↔CT, HRT, MRFDt and MRFRt in any time.
↓M-Wave amplitude at 4 h (33 ± 21%) and 5 h
(34 ± 21%) vs. pre
↑M-wave duration at Post-30 min (20 ± 25%) vs.
Pre
↔P20, P80 or P20/P80 at any time
ND ↓CMJ height at Pots 5 vs. Pre (∼8%)
↔TECC and TCON displacements and durations
(both relative and absolute)
↓Anterior-posterior force during CMJ in TECC
(Pre vs. Post 5) and in TCON (Pre vs. Post 1
and Pre vs. Post 5)
↓TCON Power (Post 1 vs. Post 5)
↓Vertical Force at Post 1 vs. Pre in TECC and at
Post-Activation Potentiation (PAP) in Endurance Sports
Post 5 vs. Pre in TECC and TCON
Earlier vertical force peak at Post 5 vs. Pre in
TECC
Later vertical force peak at Post 1 vs. Pre in TCON
ND ↑Cycling economy vs. control
↓20 km TT (6.1%)
↔Power output or pedal cadence
An increase in twitch torque ↓ TTW (28 ± 3.7%) vs. Pre Ex.
↓ EMD (5.1 ± 0.6%) vs. Pre Ex.
↓ CT (7.6 ± 0.4%) vs. Pre Ex.
↔ HRT
↓ TF20
↓ TF100 7.5 ± 2.3% vs. Pre Ex.
↔ peak MVC torque and LA of quadriceps
during MVC
At Post 10 min ↑ TTW 51 ± 8% (13.5% vs. Pre
Ex.).
↑ TTW was maintained for 40 min after the
exercise.
(Continued)
9
 10
D. Boullosa et al.
Table I. Continued.

Reference Subjects Exercise protocol Measurements Potentiation definition Main results

Skof and Strojnik n=7 ♂ 6 km Continuous running at
(2006b) Well-trained middle- and VOBLA (∼ 56% of Smax)
long-distance runners on an athletic track.
Skof and Strojnik n = 7♂ Two warm-up protocols:
(2007) Well-trained middle- WU1: 10-min continuous
distance runners (On run at 80% VLT, 5-min
average they competed for stretching, 6 × 50 m
9 ± 3 years) bounding exercises (2 ×
skipping, 2 × hopping,
2 × strides), and 5 × 80 m
acceleration runs
WU2: 10-min continuous
run at 80% VLT and 5-
min stretching
Vuorimaa et al. n = 22 ♂ MR = Maximal run until
(2006) Long-distance runners (> 5 exhaustion
years of experience) TR = 40-min tempo run at
80% vVO2max
IR = 40-min intermittent
run (2 min × 2 min) at
100% vVO2max
Pre (after the warm up) vs.
Post
1, 3, 10, 20, 30, 40, 60, and
120 min of recovery TTW,
EMD, CT, HRT, TF20,
TF100, TMVC, AL
Pre vs. post Ex. TTW, EMD,
CT, HRT, torque during
maximum isometric knee
extension torque (TMVC),
AL, torque during MVC
with added electrical
stimulation (TMVC + ES)
Pre vs. Post Ex.
CMJ height (post 5–15 s)
Mechanical power (½ Squat,
35% 1RM, 10 reps, 3 s of
rec. between reps.)
and EMGrms
An increase in Twitch ↓TTW (5.1%) Post 1–3 min vs. Pre Ex.
torque (Post-tetanic ↓EMD (6.1%) Post 1–3 min vs. Pre Ex.
potentiation) ↓CT (8.0%) Post 1–3 min vs. Pre Ex.
↔HRT
↓TF20 (20.3%) Post 1–3 min vs. Pre Ex.
↔TF100
At Post 10 min ↑ TTW (10%) vs. Pre Ex.
↑ TTW (5–14%) was maintained for 120 min
after the exercise.
An increase in torque of ↑TTW (∼4) in WU1 vs. Pre
voluntary and electrically ↓CT (∼9%) in WU1 vs. Pre
stimulated muscle ↔EMD and HRT with both warm-up protocols
contraction ↑TMCV (∼11%) in WU1 vs. Pre and > WU2
↑AL (∼8) in WU1 vs. Pre and > WU2
ND ↓EMGrms vs. Pre Ex. in all three conditions
(−7.5% MR vs. −17.1 TR vs. −12.2% IR, NS
between trials)
↑ Mechanical power (2.3% MR vs. 4.1% TR vs.
3.3% IR, NS between trials)
↑CMJ vs. Pre Ex. in all three conditions (8.9%
MR vs. 14.5% TR vs. 10.7% IR; NS between
trials)

Notes: NS, non-significant; ↑, significant increase; ↓, significant reduction; ↔, no change; ND, not defined; PAP, post-activation potentiation; KE, knee extensors; EMG, electromyography;
EMGrms, EMG root mean-square; ES, electrical stimulation; MVC, maximal voluntary contraction; PT, peak torque, TPT, time to peak torque; HRT, half-relaxation time; TTW, twitch peak
torque; CT, contraction time; EMD, elecromechanical delay; TF20, torque during 20-Hz ES; TF100, torque during 100-Hz ES; AL, Activation level; VA, voluntary activation; MMA, maximal
muscle activation; t RFD, twitch rate of force development; MRFD, twitch maximal rate of torque development; MRFR, twitch maximal rate of torque relaxation; DT, Doublet Torque; HRmax,
maximum heart rate; VOBLA, speed at anaerobic threshold; MAP, maximum aerobic power; MAS, maximum aerobic speed; Smax, maximal speed, 1RM, one repetition maximum; UMTT,
Université of Montreal Track Test, CM, centre of mass; MPO, mean power output; PPO , peak power output; SR, stroke rate; TECC, Eccentric Phase of CMJ, TCON, Concentric Phase of CMJ
 Post-Activation Potentiation (PAP) in Endurance Sports
(Feros et al., 2012; Silva et al., 2014; Skof & Strojnik,
2007).
In a study with well-trained middle-distance
runners (Skof & Strojnik, 2007), it was suggested
that incorporating bouncing and sprinting exercises
into a standardized warm-up would enhance neuro-
muscular efficiency via PAP, although the authors
did not exclude an increase in muscle activation via
improved central input and enhanced excitation of
the α-motor neuron. While the results of this study
(Skof & Strojnik, 2007) are promising, it presents a
number of limitations including the absence of a
control condition, and no verification whether differ-
ences in neuromuscular function after the two differ-
ent warm-up protocols would effectively influence
running performance as only neuromuscular func-
tion after the warm-up protocols were evaluated.
Subsequently, Feros et al. (2012) reported with
national level rowers that the addition of 5 × 5 s iso-
metric contractions to individualized warm-up rou-
tines significantly improved mean power and
performance time over the first 500 m of a 1000 m
rowing ergometer time trial. However, performance
time did not significantly improve over the whole
trial while stroke rate was significantly increased.
Although not significant, the effect size in perform-
ance improvement (ES = 0.21, small) along with
the observation of a high variability in individual
responses, may suggest that a more controlled study
with a standardized warm-up would provide a signifi-
cant improvement in performance time. Thus, it is
unknown if the individualized warm up plus the iso-
metric exercises did effectively promote PAP in the
musculature of all athletes or if it induced more
fatigue than PAP in some of them.
Silva et al. (2014) found a ∼6% time reduction in a
20-km cycling time trial after adding 4 sets of 5RM
leg press to a 5 min submaximal cycling warm-up.
This improvement in performance was associated
with an improved cycling economy. While the PAP
effect did not affect pacing strategy, a trend (p
= .06) for a higher mean power output during the
first 10% of the trial was observed. A noticeable limit-
ation of this study is that the control condition
included a very short warm-up, which was likely
insufficient for an optimum activation of the aerobic
metabolism.
The application of PAP responses after specific
warm-up protocols for endurance performance has
been an under-explored area. However, caution
should be taken given that the PAP effect of con-
ditioning activities during warm-up lasts < 12 min
(Seitz & Haff, 2016). Therefore, its positive influence
could be expected only during the first minutes of the
subsequent exercise, with potentially limited applica-
bility in competitive settings (Docherty & Hodgson,
11
2007). This acute effect would be more important
in efforts of short duration as the duration of PAP
represents a significant percentage during such brief
events. PAP eliciting warm-up strategies may also
have a potential for performance enhancement in
events longer than sprints (Kilduff et al., 2011)
when added to PAP elicited by muscle activity itself
during endurance competition, considering that any
little advantage in elite sport could be decisive for
success. Further studies should elucidate in every
sport, the mode, duration and intensity of different
exercises and their combinations for the enhance-
ment of muscle force production while avoiding the
deleterious effect of fatigue of different origin
(Tomaras & MacIntosh, 2011). Given that previous
studies have used brief high-intensity exercises to
induce PAP, it is still to be resolved if a more specific
stimuli for endurance athletes (e.g. continuous
specific exercise below the lactate threshold over >
20 min), or a combination of both stimuli would
induce a greater acute PAP effect than a standard
warm up. Importantly, it would be recommended
to differentiate with appropriate experimental
designs the influence of PAP from other warm-up
objectives as muscle temperature elevation and meta-
bolic activation (McGowan et al., 2015). This issue
could be solved with the use of a standardized
warm-up with known exercises for athletes, while
controlling both muscle temperature and cardiome-
tabolic responses. In addition, while a recent study
(Stoter et al., 2016) has suggested that an intended
fast start does not induce an improvement in
short endurance performances, it could be speculated
that the increments in power output via PAP could
be advantageous as they are not consciously regulated
and, therefore, would not consume any cognitive
resources that could influence on perceived exertion.
PAP during testing, training and
competitions
Enhancement of contractile activity during and after
testing, training and competition has been described
in the literature. Previous studies have observed sim-
ultaneously the presence of potentiation and muscle
fatigue of different origin in single muscles or whole
body measures. Collectively, these previous findings
may be suggesting that exercise load and athletes’
training background could be the main modulators
of the PAP/fatigue relationship.
The current evidence would confirm that endur-
ance athletes exhibit greater PAP responses after pro-
longed submaximal conditioning activities, contrary
to power athletes who benefit more from brief
maximal or near maximal conditioning activities, fol-
lowing the principle of training specificity. For
12 D. Boullosa et al.
instance, Morana and Perrey (2009) showed a pro-
longed prevalence of potentiation over muscle
fatigue by means of percutaneous stimulation in
endurance trained (n = 8) compared to power-
trained athletes (n = 7) during 10 min of submaximal
(50% of MVC) intermittent contractions of knee
extensors. Interestingly, in this study (Morana &
Perrey, 2009), the maximal rate of torque develop-
ment during evoked contractions in the endurance
group at the end of the conditioning activity increased
(from 966 ± 245 to 1580 ± 632 Nm·s−1) until levels
similar to those observed in the power group at the
start of the conditioning activity (1544 ±
352 Nm·s−1). Furthermore, the peak torque of the
endurance group was significantly elevated during
the whole protocol (from 30.0 ± 5.9 to 38.3 ±
11.0 N·m), while the power group was significantly
depressed from the second half of the protocol until
the end (from 43.4 ± 9.5 to 31.1 ± 10.4 N·m). On
the other hand, another study reported greater PAP
after 10 s MVCs of knee extensors in power-trained
compared to endurance-trained female athletes who
also exhibited a more rapid decline in PAP after
MVCs (Paasuke et al., 2007). Meanwhile, the
classic study of Hamada et al. (2000) showed that
endurance athletes exhibited PAP in trained
muscles after 10 s MVCs compared to active and
sedentary controls. Thus, although it seems that
endurance athletes could benefit from both
maximal and submaximal contractions in trained
muscles, it seems that they could benefit more from
specific submaximal contractions when testing PAP
responses with the twitch interpolation technique.
The results of previous findings in laboratory con-
ditions with different neuromuscular evaluations
(Millet et al., 2002; Millet, Martin et al., 2003;
Millet, Millet et al., 2003; Skof & Strojnik, 2006a,
2006b; Vuorimaa et al., 2006) have confirmed an
enhanced contractile capacity in endurance athletes
after endurance exercises of different duration. Pre-
viously, Millet et al. (2002) reported a severely
depressed maximal voluntary force capacity and a
potentiated twitch mechanical response after an
ultra-marathon. Subsequently, the only study with
cyclists (Millet, Millet et al., 2003) evaluated the
twitch responses after a prolonged 140-km race and
reported an enhancement of the rate of twitch force
development despite a decrease in MVC, although
some caution should be taken given the low number
(n = 4) of participants evaluated. Similarly, Millet,
Martin et al. (2003) showed greater peak mechanical
response during electrically evoked single twitches,
faster RFD, and shorter contraction time in the fati-
gued state after a ski skating marathon. More
recently, (Vuorimaa et al., 2006) showed a jump
potentiation, after constant pace running of 40 min
at 80% of velocity associated with maximum oxygen
consumption (vVO2max) (∼14.5%), that was greater
compared to 40 min of 2 min intervals at vVO2max
interspersed with 2 min of recovery (∼10.7%), and
after a treadmill incremental test (∼8.9%). In
another study with endurance runners, Skof and
Strojnik (2006b) observed concurrent evidence of
fatigue and potentiation after a non-exhausting 6-
km run at anaerobic threshold. In this study (Skof
& Strojnik, 2006b), muscle activation level, (i.e. %
change in knee extension torque assessed with and
without a superimposed twitch of 100 Hz, during a
MVC) was similar before and after the run, while
maximum twitch torque was enhanced 10 min after
the run and remained elevated over 60 min of recov-
ery. In another study, the same authors (Skof & Stroj-
nik, 2006a) reported that, 10 min after 5 × 300 m
intervals at ∼24 km·h−1 (i.e. 5% lower than the best
400 m run), the twitch torque exceeded the pre-
workload value by 11% (p < .01) and that the poten-
tiation lasted 40 min, despite evidence of low-fre-
quency fatigue (i.e. a more pronounced reduction
in torque at low-frequency stimulation vs. high fre-
quency stimulation) and a high blood lactate concen-
tration. Collectively, this evidence would suggest that
higher exercise intensities could induce a worse
potentiation/fatigue balance than prolonged endur-
ance activities, with metabolic fatigue symptoms (e.g.
elevated lactate) being not related to PAP responses.
Other recent studies in the field (Boullosa et al.,
2011; Boullosa & Tuimil, 2009; Garcia-Pinillos
et al., 2015, 2016) have confirmed the acute effect
of different endurance running exercises on jump
potentiation of endurance athletes. For instance, it
has been previously reported a greater (∼12.7 vs.
∼3.5%) and more prolonged (7 min vs. 2 min)
jump potentiation after the Université de Montréal
Track Test (UMTT) (∼30 min) compared to a
time limit at vVO2max test (∼5 min) (Boullosa &
Tuimil, 2009). Subsequently, another study (Boul-
losa et al., 2011) with runners and triathletes revealed
jump potentiation, 2 min after the UMTT. Athletes
with lower peak force loss during the eccentric
phase of the CMJ exhibited a greater peak power in
the concentric phase and subsequently a greater
jump potentiation after exhaustion. Moreover, the
increment in peak power was related to the increment
in sprint velocity after exhaustion (Boullosa et al.,
2011) in those athletes who exhibited more jump
potentiation. The jump potentiation after incremen-
tal tests and interval training sessions were also con-
firmed by Garcia-Pinillos et al. (2016) who
observed jump potentiation after an incremental
shuttle run test, and over an extended interval train-
ing session (4 × 3 × 400 m) (Garcia-Pinillos et al.,
2015). Interestingly, the recorded jump potentiation
 Post-Activation Potentiation (PAP) in Endurance Sports
at the end of the interval training session showed a
correlation with the enhancement in handgrip
strength in the responders group (Garcia-Pinillos
et al., 2015), maybe suggesting a possible central
origin of this improvement in contractile capacity.
Overall, these studies in the field do confirm how
potentiation responses could be monitored with
simple exercises as jumps for a better understanding
of the effect of different training and evaluation exer-
cises on acute neuromuscular adaptations of endur-
ance runners. However, it is still unknown if the
jump potentiation does effectively correspond to
PAP responses assessed with the twitch interpolation
technique. In addition, it would be necessary to
understand how these responses could vary, not
only after different exercises, but also during different
moments of the season when examining the appropri-
ate PAP/fatigue responses during training sessions to
illustrate the best chronic adaptations.
There are five studies (Del Rosso et al., 2016;
McIntyre et al., 2012; Pageaux et al., 2017; Place
et al., 2004; Rousanoglou et al., 2016) evaluating
the neuromuscular function during and after endur-
ance multistage trials that, in some cases, observed
PAP or jump potentiation during the course of the
endurance activity. Place et al. (2004) were the first
to describe a depression of maximal voluntary force
generating capability in the final stages of a submax-
imal (i.e. 55% of maximum aerobic speed) 5-h
running exercise, but with a simultaneous peak
twitch potentiation. More recently, McIntyre et al.
(2012) evaluated cycling peak power output, drop
jump performance, and isokinetic strength during
and after 20-min stages at 70% of VO2peak until
exhaustion. They found that peak power output
during cycling was potentiated in some athletes at
the end of the first stages, and that the decline of
this measure after exhaustion was strongly correlated
to the change in drop jump height (McIntyre et al.,
2012). Interestingly, a jump potentiation was
observed over a 30-km trial in half-marathon
runners (i.e. after every 5-km stage) with its
maximum at 25 km when the athletes exhibited the
greatest rise in rating of perceived exertion (RPE)
and decline in speed (Del Rosso et al., 2016). The
results of this study (Del Rosso et al., 2016) may
suggest that PAP may be a mechanism by which the
neuromuscular system counteracts the deleterious
effect of fatigue of different origins. In another
recent study (Rousanoglou et al., 2016), it was
found that jump height was maintained immediately
after a mountain half-marathon race but was sub-
sequently reduced at 5 min of recovery. The
authors of this study observed several changes in
the timing kinetics during jumping, and suggested
that these responses could be attributable to the
13
existence of concurrent PAP and muscle fatigue,
probably because of the greater eccentric demands
of the last downhill part of the race. More recently,
Pageaux et al. (2017) evaluated the influence of per-
forming different exercise modes during 1 h on a
10 km running race in a group of triathletes, and
found that both running and uphill walking induced
PAP whereas cycling did not. Thus, these studies
(Del Rosso et al., 2016; McIntyre et al., 2012;
Pageaux et al., 2017; Place et al., 2004; Rousanoglou
et al., 2016) would confirm that PAP could be
observed in competitive settings, with multistage
trials demonstrating to be a valid paradigm to evaluate
PAP in relation to endurance performance. However,
studies analysing the kinetics of PAP and fatigue of
different origins during continuous prolonged endur-
ance exercises are still lacking (Millet & Lepers, 2004).
From the current evidence, it is clear that PAP
responses can be assessed during and after intermittent
and continuous endurance exercises for training moni-
toring in different settings. In practical terms, simple
measures such as jump height could be very useful to
monitor PAP responses and their interaction with
fatigue of different origin, although the sensitivity of
different instruments and the selection of performance
measures should be taken into account (Boullosa et al.,
2011; Boullosa, Abreu, Beltrame, et al., 2013). Studies
verifying PAP during jumping with twitch verification
are still lacking in endurance athletes, with squat
jump being a potentially valid test in this context
(Nibali et al., 2013). Additionally, well controlled
studies are needed to identify the optimal intensity
and duration of work and relief intervals in intermittent
exercises for the best PAP/fatigue balance when
looking for the best chronic adaptations. Further
studies should also verify the influence of PAP during
and after different modes of exercise (e.g. cycling vs.
running) and their combinations, given the variable
PAP responses during different muscular actions
(MacIntosh, 2010). Thus, while jumping could be
very useful for evaluation of endurance runners, it
could be recommended that specific ergometers be
used for evaluation of PAP in sports with other neuro-
muscular demands (e.g. skiers, skaters). Recording of
different neuromuscular, metabolic, and perceptual
measures, could be very useful for better understand-
ing the true significance of PAP responses in endur-
ance performance.
Future perspectives
Determination of the best warm-up routines for max-
imizing the PAP effects on endurance efforts of differ-
ent duration is warranted in every sport. Further
studies should elaborate on the role of PAP for per-
formance and pacing in endurance events of different
14 D. Boullosa et al.
duration. In this context, it should be also verified if
the capacity of force preservation at exhaustion
(Marcora & Staiano, 2010; Morales-Alamo et al.,
2015) may be influenced by PAP. Given that twitch
force potentiation can be accompanied by a decline
in motor units discharge rates (Klein et al., 2001), a
phenomenon that could be speculated to induce a
greater efficiency during endurance activities, future
studies should also evaluate how efficiency could be
influenced by PAP. In this regard, attention should
be taken to consider a number of methodological
aspects, given the contradictory results observed
when examining running economy changes after
ultra-marathons (Vernillo, Millet, & Millet, 2017).
As only longitudinal studies for the evaluation of
endurance training on PAP were performed with
non-athletes in laboratory settings (Mettler &
Griffin, 2012, 2016), chronic studies with athletes
are urgently needed for verifying the best training
stimuli that improve the PAP/fatigue balance. Given
that RLC kinase polymorphisms have recently
demonstrated to influence muscle damage and
power production in runners after a marathon (Del
Coso et al., 2016), new venues could be open with
examination of genetic variants that influence both
kinase and phosphatase activities.
Conclusions
The current review suggests the existence of PAP
mechanism during and after endurance performances
in endurance athletes. It is still unknown what the
exact role that this acute adaptation could play
during training and competitions. While different exer-
cises in the warm-up may enhance muscle power pro-
duction during subsequent short endurance activities
via PAP, it would seem that the submaximal endurance
activity itself could be more important in longer events
with PAP maybe counteracting the negative conse-
quences of fatigue of different origin, possibly enhan-
cing muscular efficiency and pacing. However, this
later hypothesis should be appropriately tested. There
is an urgent need for chronic studies for verifying the
effectiveness of different training interventions in
both PAP responses and endurance performance.
Monitoring PAP, muscle and perceptual fatigue
during training sessions and simulated competitions
could be very useful for a better understanding of
acute and chronic adaptations of endurance athletes.
Acknowledgement
The authors would like to thank the editor and
reviewers involved during the peer review process
for their outstanding contributions.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
Daniel A. Boullosa is supported by CNPq with a productivity
research grant (Grant number 305131/2015-0).
References
Behm, D. G., Button, D. C., Barbour, G., Butt, J. C., & Young,
W. B. (2004). Conflicting effects of fatigue and potentiation
on voluntary force. Journal of Strength and Conditioning
Research, 18(2), 365–372. doi:10.1519/R-12982.1
Boullosa, D. A., Abreu, L., Beltrame, L. G., & Behm, D. G.
(2013). The acute effect of different half squat set configur-
ations on jump potentiation. Journal of Strength and
Conditioning Research, 27(8), 2059–2066. doi:10.1519/JSC.
0b013e31827ddf15
Boullosa, D. A., Abreu, L., Varela-Sanz, A., & Mujika, I. (2013).
Do Olympic athletes train as in the Paleolithic era? Sports
Medicine, 43(10), 909–917. doi:10.1007/s40279-013-0086-1
Boullosa, D. A., & Tuimil, J. L. (2009). Postactivation potentiation
in distance runners after two different field running protocols.
Journal of Strength and Conditioning Research, 23(5), 1560–
1565. doi:10.1519/JSC.0b013e3181a3ce61
Boullosa, D. A., Tuimil, J. L., Alegre, L. M., Iglesias, E., &
Lusquinos, F. (2011). Concurrent fatigue and potentiation in
endurance athletes. International Journal of Sports Physiology
and Performance, 6(1), 82–93.
Chamari, K., & Padulo, J. (2015). ‘Aerobic’ and ‘Anaerobic’ terms
used in exercise physiology: A critical terminology reflection.
Sports Medicine – Open, 1(1), 1–4. doi:10.1186/s40798-015-
0012-1
Chaouachi, A., Poulos, N., Abed, F., Turki, O., Brughelli, M.,
Chamari, K., Drinkwater, E. J., & Behm, D. G. (2011).
Volume, intensity, and timing of muscle power potentiation
are variable. Applied Physiology, Nutrition, and Metabolism, 36
(5), 736–747. doi:10.1139/h11-079
Decostre, V., Gillis, J. M., & Gailly, P. (2000). Effect of adrena-
line on the post-tetanic potentiation in mouse skeletal muscle.
Journal of Muscle Research and Cell Motility, 21(3), 247–254.
Del Coso, J., Valero, M., Lara, B., Salinero, J. J., Gallo-Salazar, C.,
Areces, F. & Weber, C. R. (2016). Myosin light chain kinase
(MLCK) gene influences exercise induced muscle damage
during a competitive marathon. PLoS One, 11(8), e0160053.
doi:10.1371/journal.pone.0160053
Del Rosso, S., Barros, E., Tonello, L., Oliveira-Silva, I., Behm, D.
G., Foster, C., … Piacentini, M. F. (2016). Can pacing be
regulated by post-activation potentiation? Insights from a self-
paced 30 km trial in half-marathon runners. PLoS One, 11(3),
e0150679. doi:10.1371/journal.pone.0150679
Docherty, D., & Hodgson, M. J. (2007). The application of post-
activation potentiation to elite sport. International Journal of
Sports Physiology and Performance, 2(4), 439–444.
Feros, S. A., Young, W. B., Rice, A. J., & Talpey, S. W. (2012).
The effect of including a series of isometric conditioning con-
tractions to the rowing warm-up on 1000-m rowing ergometer
time trial performance. Journal of Strength and Conditioning
Research, 26(12), 3326–3334. doi:10.1519/JSC.
0b013e3182495025
Gandevia, S. A. (2001). Spinal and supraspinal factors in human
muscle fatigue. Journal of Applied Physiology, 81(4), 1725–1789.
 Post-Activation Potentiation (PAP) in Endurance Sports
Garcia-Pinillos, F., Molina-Molina, A., & Latorre-Roman, P. A.
(2016). Impact of an incremental running test on jumping kin-
ematics in endurance runners: Can jumping kinematic explain
the post-activation potentiation phenomenon? Sports
Biomechanics, 15(2), 103–115. doi:10.1080/14763141.2016.
1158860
Garcia-Pinillos, F., Soto-Hermoso, V. M., & Latorre-Roman, P.
A. (2015). Acute effects of extended interval training on coun-
termovement jump and handgrip strength performance in
endurance athletes: Postactivation potentiation. Journal of
Strength and Conditioning Research, 29(1), 11–21. doi:10.1519/
JSC.0000000000000591
Grange, R. W., Vandenboom, R., & Houston, M. E. (1993).
Physiological significance of myosin phosphorylation in skeletal
muscle. Canadian Journal of Applied Physiology, 18(3), 229–242.
Hamada, T., Sale, D. G., & Macdougall, J. D. (2000).
Postactivation potentiation in endurance-trained male athletes.
Medicine & Science in Sports & Exercise, 32(2), 403–411.
Hodgson, M., Docherty, D., & Robbins, D. (2005). Post-acti-
vation potentiation: Underlying physiology and implications
for motor performance. Sports Medicine, 35(7), 585–595.
Houston, M. E., & Grange, R. W. (1991). Torque potentiation and
myosin light-chain phosphorylation in human muscle following
a fatiguing contraction. Canadian Journal of Physiology and
Pharmacology, 69(2), 269–273.
Houston, M. E., Lingley, M. D., Stuart, D. S., & Grange, R. W.
(1987). Myosin light chain phosphorylation in intact human
muscle. FEBS Letters, 219(2), 469–471.
Hvid, L. G., Gejl, K., Bech, R. D., Nygaard, T., Jensen, K.,
Frandsen, U., & Ortenblad, N. (2013). Transient impairments
in single muscle fibre contractile function after prolonged
cycling in elite endurance athletes. Acta Physiologica, 208(3),
265–273. doi:10.1111/apha.12095
Kilduff, L. P., Cunningham, D. J., Owen, N. J., West, D. J.,
Bracken, R. M., & Cook, C. J. (2011). Effect of postactivation
potentiation on swimming starts in international sprint swim-
mers. Journal of Strength and Conditioning Research, 25(9),
2418–2423. doi:10.1519/JSC.0b013e318201bf7a
Klein, C. S., Ivanova, T. D., Rice, C. L., & Garland, S. J. (2001).
Motor unit discharge rate following twitch potentiation in
human triceps brachii muscle. Neuroscience Letters, 316(3),
153–156.
Lai, S., Collins, B. C., Colson, B. A., Kararigas, G., & Lowe, D. A.
(2016). Estradiol modulates myosin regulatory light chain phos-
phorylation and contractility in skeletal muscle of female mice.
American Journal of Physiology-Endocrinology and Metabolism,
310(9), E724–733. doi:10.1152/ajpendo.00439.2015
Latorre-Román, PÁ, García-Pinillos, F., Martínez-López, E. J., &
Soto-Hermoso, V. M. (2014). Concurrent fatigue and postacti-
vation potentiation during extended interval training in long-
distance runners. Motriz: Revista de Educação Física, 20, 423–
430.
MacIntosh, B. R. (2010). Cellular and whole muscle studies of
activity dependent potentiation. Advances in Experimental
Medicine and Biology, 682, 315–342. doi:10.1007/978-1-4419-
6366-6_18
Marcora, S. M., & Staiano, W. (2010). The limit to exercise toler-
ance in humans: Mind over muscle? European Journal of Applied
Physiology, 109(4), 763–770. doi.org/10.1007/s00421-010-
1418-6
McGowan, C. J., Pyne, D. B., Thompson, K. G., & Rattray, B.
(2015). Warm-up strategies for sport and exercise:
Mechanisms and applications. Sports Medicine, 45(11), 1523–
1546. doi:10.1007/s40279-015-0376-x
McIntyre, J. P., Mawston, G. A., & Cairns, S. P. (2012). Changes
of whole-body power, muscle function, and jump performance
15
with prolonged cycling to exhaustion. International Journal of
Sports Physiology and Performance, 7(4), 332–339.
Mettler, J. A., & Griffin, L. (2010). What are the stimulation par-
ameters that affect the extent of twitch force potentiation in the
adductor pollicis muscle? European Journal of Applied Physiology,
110(6), 1235–1242. doi:10.1007/s00421-010-1625-1
Mettler, J. A., & Griffin, L. (2012). Postactivation potentiation and
muscular endurance training. Muscle & Nerve, 45(3), 416–425.
doi:10.1002/mus.22313
Mettler, J. A., & Griffin, L. (2016). Muscular endurance training
and motor unit firing patterns during fatigue. Experimental
Brain Research, 234(1), 267–276. doi:10.1007/s00221-015-
4455-x
Millet, G. Y., & Lepers, R. (2004). Alterations in neuromuscular
function after prolonged running, cycling, and skiing exercises.
Sports Medicine, 34(2), 105–116. doi.org/10.2165/00007256-
200434020-00004
Millet, G. Y., Lepers, R., Maffiuletti, N. A., Babault, N., Martin,
V., & Lattier, G. (2002). Alterations in neuromuscular function
after an ultramarathon. Journal of Applied Physiology, 92(2),
486–492. doi:10.1152/japplphysiol.00122.2001
Millet, G. Y., Martin, V., Maffiuletti, N. A., & Martin, A. (2003).
Neuromuscular fatigue after a ski skating marathon. Canadian
Journal of Applied Physiology Journal, 28(3), 234–245.
Millet, G. Y., Millet, G. P., Lattier, G., Maffiuletti, N. A., &
Candau, R. (2003). Alteration of neuromuscular function
after a prolonged road cycling race. International Journal of
Sports Medicine, 24(3), 190–194. doi:10.1055/s-2003-39088
Mitchell, C. J., & Sale, D. G. (2011). Enhancement of jump per-
formance after a 5-RM squat is associated with postactivation
potentiation. European Journal of Applied Physiology, 111(8),
1957–1963. doi:10.1007/s00421-010-1823-x
Morales-Alamo, D., Losa-Reyna, J., Torres-Peralta, R., Martin-
Rincon, M., Perez-Varela, M., Curtelin, D., … Calbet,
J. A. L. (2015). What limits performance during whole-body
incremental exercise to exhaustion in humans? The Journal of
Physiology, 593, 4631–4648. doi:10.1113/JP270487
Morana, C., & Perrey, S. (2009). Time course of postactivation
potentiation during intermittent submaximal fatiguing contrac-
tions in endurance- and power-trained athletes. Journal of
Strength and Conditioning Research, 23(5), 1456–1464. doi:10.
1519/JSC.0b013e3181a518f1
Nibali, M. L., Chapman, D. W., Robergs, R. A., & Drinkwater, E.
J. (2013). Validation of jump squats as a practical measure of
post-activation potentiation. Applied Physiology, Nutrition, and
Metabolism, 38(3), 306–313. doi:10.1139/apnm-2012-0277
Paasuke, M., Saapar, L., Ereline, J., Gapeyeva, H., Requena, B., &
Oopik, V. (2007). Postactivation potentiation of knee extensor
muscles in power- and endurance-trained, and untrained
women. European Journal of Applied Physiology, 101(5), 577–
585. doi:10.1007/s00421-007-0532-6
Pageaux, B., Theurel, J., & Lepers, R. (2017). Cycling vs uphill
walking: Impact on locomotor muscle fatigue and running exer-
cise. International Journal of Sports Physiology and Performance, 1–
22. doi:10.1123/ijspp.2016-0564
Pearson, S. J., & Hussain, S. R. (2014). Lack of association
between postactivation potentiation and subsequent jump per-
formance. European Journal of Sport Science, 14(5), 418–425.
doi:10.1080/17461391.2013.837511
Place, N., Lepers, R., Deley, G., & Millet, G. Y. (2004). Time
course of neuromuscular alterations during a prolonged
running exercise. Medicine & Science in Sports & Exercise, 36
(8), 1347–1356.
Rassier, D. E., & Macintosh, B. R. (2000). Coexistence of poten-
tiation and fatigue in skeletal muscle. Brazilian Journal of
Medical and Biological Research, 33(5), 499–508.
16 D. Boullosa et al.
Requena, B., Sáez-Sáez de Villareal, E., Gapeyava, H., García, I.,
& Pääsuke, M. (2011). Relationship between postactivation
potentiation of knee muscles, sprinting and vertical jumping
performance in professional soccer players. Journal of Strength
and Conditioning Research, 25(2), 367–373. doi:10.1519/JSC.
0b013e3181be31aa
Rousanoglou, E. N., Noutsos, K., Pappas, A., Bogdanis, G.,
Vagenas, G., Bayios, I. A., & Boudolos, K. D. (2016).
Alterations of vertical jump mechanics after a half-marathon
mountain running race. Journal of Sports Science and Medicine,
15(2), 277–286.
Sahaly, R., Vandewalle, H., Driss, T., & Monod, H. (2001).
Maximal voluntary force and rate of force development in
humans-importance of instruction. European Journal of Applied
Physiology, 85(3-4), 345–350. doi.org/10.1007/s004210100451
Sale, D. G. (2002). Postactivation potentiation: Role in human
performance. Exercise and Sport Sciences Reviews, 30(3), 138–143.
Seitz, L. B., & Haff, G. G. (2016). Factors modulating post-acti-
vation potentiation of jump, sprint, throw, and upper-body bal-
listic performances: A systematic review with meta-analysis.
Sports Medicine, 46(2), 231–240. doi:10.1007/s40279-015-
0415-7
Silva, R. A., Silva-Junior, F. L., Pinheiro, F. A., Souza, P. F.,
Boullosa, D. A., & Pires, F. O. (2014). Acute prior heavy
strength exercise bouts improve the 20-km cycling time trial
performance. Journal of Strength and Conditioning Research, 28
(9), 2513–2520. doi:10.1519/JSC.0000000000000442
Skof, B., & Strojnik, V. (2006a). Neuro-muscular fatigue and
recovery dynamics following anaerobic interval workload.
International Journal of Sports Medicine, 27(3), 220–225.
doi:10.1055/s-2005-865632
Skof, B., & Strojnik, V. (2006b). Neuromuscular fatigue and
recovery dynamics following prolonged continuous run at
anaerobic threshold. British Journal of Sports Medicine, 40(3),
219–222. discussion 219-222. doi:10.1136/bjsm.2005.020966
Skof, B., & Strojnik, V. (2007). The effect of two warm-up proto-
cols on some biomechanical parameters of the neuromuscular
system of middle distance runners. The Journal of Strength and
Conditioning Research, 21(2), 394–399. doi:10.1519/R-18055.1
Stoter, I. K., MacIntosh, B. R., Fletcher, J. R., Pootz, S.,
Zijdewind, I., & Hettinga, F. J. (2016). Pacing strategy,
muscle fatigue, and technique in 1500-m speed-skating and
cycling time trials. International Journal of Sports Physiology and
Performance, 11(3), 337–343. doi:10.1123/ijspp.2014-0603
Stuart, D. S., Lingley, M. D., Grange, R. W., & Houston, M. E.
(1988). Myosin light chain phosphorylation and contractile per-
formance of human skeletal muscle. Canadian Journal of
Physiology and Pharmacology, 66(1), 49–54.
Thomas, K., Toward, A., West, D. J., Howatson, G., & Goodall,
S. (2017). Heavy-resistance exercise-induced increases in
jump performance are not explained by changes in neuromus-
cular function. Scandinavian Journal of Medicine & Science in
Sports, 27(1), 35–44. doi:10.1111/sms.12626
Tillin, N. A., & Bishop, D. (2009). Factors modulating post-acti-
vation potentiation and its effect on performance of subsequent
explosive activities. Sports Medicine, 39(2), 147–166. doi:10.
2165/00007256-200939020-00004
Tomaras, E. K., & MacIntosh, B. R. (2011). Less is more:
Standard warm-up causes fatigue and less warm-up permits
greater cycling power output. Journal of Applied Physiology,
111(1), 228–235. doi:10.1152/japplphysiol.00253.2011
Vandenboom, R., Gittings, W., Smith, I. C., Grange, R. W., &
Stull, J. T. (2013). Myosin phosphorylation and force poten-
tiation in skeletal muscle: Evidence from animal models.
Journal of Muscle Research and Cell Motility, 34(5-6), 317–332.
doi:10.1007/s10974-013-9363-8
Vernillo, G., Millet, G. P., & Millet, G. Y. (2017). Does the
running economy really increase after ultra-marathons?
Frontiers in Physiology, 8, 783. doi:10.3389/fphys.2017.00783
Vuorimaa, T., Virlander, R., Kurkilahti, P., Vasankari, T., &
Hakkinen, K. (2006). Acute changes in muscle activation and
leg extension performance after different running exercises in
elite long distance runners. European Journal of Applied
Physiology, 96(3), 282–291. doi:10.1007/s00421-005-0054-z
Xenofondos, A., Patikas, D., Koceja, D. M., Behdad, T., Bassa,
E., Kellis, E., & Kotzamanidis, C. (2015). Post-activation
potentiation: The neural effects of post-activation depression.
Muscle & Nerve, 52(2), 252–259. doi:10.1002/mus.24533