Sale, 2002, Postactivation Potentiation Role in Human

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ARTICLE

Postactivation Potentiation: Role in Human


Performance
Digby G. Sale
Department of Kinesiology, McMaster University, Hamilton, Ontario, Canada
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SALE, D.G. Postactivation potentiation: Role in human performance. Exerc. Sport Sci. Rev., Vol. 30, No. 3, pp. 138 –143,
2002. Postactivation potentiation (PAP) is the transient increase in muscle contractile performance after previous contractile activity.
This review describes the features and mechanism of PAP, assesses its potential role in endurance and strength/speed performance,
considers strategies for exploiting PAP, and outlines how PAP might be affected by training. Keywords: postactivation
potentiation, skeletal muscle, contractile properties, muscle endurance, muscle strength, training

INTRODUCTION (12,13). Increased sensitivity to Ca2⫹ has its greatest effect at


low myoplasmic levels of Ca2⫹, as occurs in twitch and
At any point in time, the performance of skeletal muscle low-frequency tetanic contractions; in contrast, increased
is affected by its contractile history. The most obvious effect sensitivity to Ca2⫹ has little or no effect at saturating Ca2⫹
of contractile history is fatigue, which impairs performance. levels, as occurs in high-frequency tetanic contractions.
This review is about another effect of contractile history— Thus, PAP “raises” the low- but not high-frequency portion
postactivation potentiation (PAP). In contrast to fatigue, of the force-frequency relation (1,14). In fact, the condition-
PAP serves to improve performance. The phenomenon of ing activity (e.g., 10-s MVC) can, concurrently, increase
PAP and its mechanism(s) have been studied for many years, (due to PAP) and decrease (due to fatigue) low- and high-
but its application to human performance has received less frequency force, respectively ((12); Fig. 2).
study. The purpose of the review is to consider the possible
roles of PAP in human performance and, in particular, to
discuss the hypothesis that PAP may offset fatigue in endur-
ance exercise, increase rate of force development, and, thus, CONTRACTION TYPE
improve speed and power performance.
PAP is an increase in muscle twitch and low-frequency Most studies of PAP and the force-frequency relation have
tetanic force after a “conditioning” contractile activity. In involved isometric contractions. It is important to recognize
research on PAP, the conditioning activity (or contractile that the type of muscle contraction affects both the force-
history) most commonly includes a series of evoked twitches frequency relation and the range of frequencies over which
(staircase or treppe), an evoked tetanic contraction (postte- PAP occurs. In concentric contractions, especially those at
tanic potentiation (PTP)), or a sustained maximal voluntary higher velocities, the force-frequency relation is shifted to
contraction (MVC). In this review, potentiation, no matter the right compared with isometric contractions; that is,
how induced, will be referred to as PAP. An example of PAP higher frequencies are needed to evoke a given percentage of
is shown in Figure 1. The principal mechanism of PAP is maximum force (1). In addition, PAP extends to higher
considered to be phosphorylation of myosin regulatory light frequencies in concentric versus isometric contractions (1).
chains, which renders actin-myosin interaction more sensi- Most activities involve primarily concentric (e.g., swimming,
tive to Ca2⫹ released from the sarcoplasmic reticulum rowing, cycling) or coupled eccentric-concentric contrac-
tions (e.g., running, jumping, weightlifting); therefore, PAP
may have a performance-enhancing effect beyond what
Address for correspondence: Digby G. Sale, Ph.D., Department of Kinesiology, McMaster
University, Hamilton, Ontario, L8S 4K1, Canada. (E-mail: saled@mcmaster.ca).
would be expected based on its effect on isometric contrac-
Accepted for publication: March 18, 2002. tions. The interaction between the force-frequency relation,
PAP, and contraction type is illustrated in Figure 3. Note the
0091-6631/3003/138 –143
Exercise and Sport Sciences Reviews
potentially greater role of PAP in concentric compared with
Copyright © 2002 by the American College of Sports Medicine isometric contractions.

138
motor units would be discharging at relatively low rates, the
force output of the motor units should be increased by PAP
(Fig. 2). If this happens, and if a constant force has to be
maintained, motor unit firing rates would have to decrease to
compensate for the increased force (or alternatively, some
motor units could be derecruited). In fact, motor units have
been observed to decrease their firing rates in sustained,
constant force, submaximal contractions without altered mo-
tor unit recruitment (2). A decrease in motor unit firing rate,
by reducing the number of nerve impulses and muscle action
potentials per unit time, may delay impairment of “central
drive” to motoneurons, neuromuscular transmission, muscle
action potential propagation, and excitation-contraction
coupling, all possible sites and mechanisms of fatigue. It
should be noted that, as fatigue develops in sustained sub-
maximal contractions, motor units already recruited will
eventually have to increase their firing rates to compensate
Figure 1. An example of postactivation potentiation (PAP). First, a for fatigue, and depending on the exercise intensity and the
baseline twitch is evoked in a muscle that has been at rest for some time.
Then, a conditioning contraction, such as an electrically evoked tetanic
muscle group, additional motor units will be recruited.
contraction or a maximal voluntary contraction (MVC) is done. A twitch PAP may have a special role in compensating for the
contraction evoked soon after the conditioning contraction shows the impaired excitation-contraction coupling that occurs with
increased force and shortened time course typical of PAP. For an example fatigue. Impaired excitation-contraction coupling is respon-
of actual twitch recordings, see (7). sible for low-frequency fatigue (LFF), a disproportionate loss
of low-frequency tetanic force (12). This is the exact opposite
FIBER TYPE of PAP, which is a disproportionate increase in low-frequency
tetanic force. Thus, PAP can serve to compensate for LFF.
A notable feature of PAP is that it is greater in fast, Type II Many endurance activities (e.g., running, cycling, swim-
muscle fibers because fast fibers undergo greater phosphorylation ming) consist of repeated brief concentric or eccentric-con-
of myosin regulatory light chains in response to a conditioning centric actions in which motor units discharge briefly at fairly
activity (13). Accordingly, muscles with a higher percentage of high rates; however, it should be recalled from Figure 3 that
Type II fibers (e.g., gastrocnemius vs soleus), and people with a in concentric (vs isometric) actions, the force-frequency re-
higher percentage of Type II fibers within a muscle (e.g., vastus lation is shifted to the right, extending the frequency range
lateralis, (7)), exhibit greater PAP. A person’s fiber-type distri- over which both LFF and PAP would be acting.
bution is determined primarily by genetic factors, but may also In contrast, PAP cannot compensate for so-called high-
be influenced by age and activity level. frequency fatigue, the force decline when motor units are
It might be expected that PAP offers the greatest potential
for performance enhancement in brief, maximal intensity
activities requiring maximal strength and speed (and the
product of these two, power), activities that depend on fast
fibers. But in these activities, motor unit firing rates are likely
to be at their highest, in the very range where PAP’s effect on
force is smallest or absent. Taken in this light, greater PAP in
fast fibers almost seems a “waste.” It would be better for slow
fibers to have greater PAP, because they are typically in-
volved in low-intensity activities in which motor unit firing
rates are relatively low, the range in which PAP is greatest.
However, it will be shown later that PAP has another effect
on muscle (in addition to increasing force), an effect that is
beneficial for speed and power performance, even when mo-
tor units are firing at high rates. This beneficial effect is also
most pronounced in fast fibers.

ENDURANCE PERFORMANCE
Figure 2. Effect of PAP on the isometric force-frequency relation. Force
Endurance performance typically consists of submaximal increases and then plateaus as the frequency of stimulation increases
contractions that are repeated for prolonged periods. From (solid line). After a conditioning activity, the induced PAP (dashed line)
increases low- but not high-frequency tetanic force. The conditioning
the beginning of performance the contractions themselves activity may, by causing fatigue, actually decrease high-frequency
would activate the mechanism(s) responsible for PAP (12). force, as shown. See text for references on which this schematic figure
Because, in these submaximal contractions, the recruited is based.

Volume 30 䡠 Number 3 䡠 July 2002 Postactivation Potentiation 139


within the range affected by PAP.) An athlete might also
experience this transition from initial fatigue to “feeling better”
during a training session, if LFF has persisted from the previous
training session. It would be of interest to monitor both LFF and
PAP in endurance athletes during a training program.

STRENGTH AND SPEED PERFORMANCE

Strength and speed performance typically require, in a


brief maximal effort, that all relevant motor units be re-
cruited and fire at maximum possible rates. It would appear,
with reference to Figure 3, that PAP could offer little benefit
when motor units are discharging at very high rates, because
PAP cannot increase high-frequency force. Furthermore,
PAP does not increase maximum unresisted shortening ve-
locity (for review, see (4)). Therefore, with reference to the
force (load)-velocity relation, the two extremes of this rela-
Figure 3. Effect of contraction type on the force-frequency relation
and range of frequency over which PAP extends. An isometric and a “fast”
tion, peak isometric force and maximum shortening velocity
concentric contraction condition are compared. A higher frequency is (Vmax), cannot be altered by PAP (Fig. 5). However, PAP
required to attain the plateau or peak force (solid lines) in the concentric has an additional effect; it can increase rate of force devel-
contraction. Also, PAP induced by a conditioning activity (dashed line) opment, even at very high stimulation frequencies where
extends to a higher frequency in the concentric contraction. Note that the force is not increased by PAP ((1,14); Fig. 6).
maximum isometric force is greater than the maximum concentric force,
in accordance with the force-velocity relation. Also, in this example, fa-
The effect of PAP on rate of force development raises the
tigue produced by the conditioning activity caused a decrease in high- prospect that PAP may, by increasing rate of force develop-
frequency force (see also Fig. 1 and (1)). ment and hence the acceleration attained with loads be-
tween zero (Vmax) and peak isometric force, increase the
velocity attained with these loads. Thus, PAP would shift the
firing at very high rates, because PAP cannot increase high- load (force)-velocity relation upward and rightward (making
frequency force ((12); Fig. 2 and 3). Thus, in fatiguing
exercise, low-frequency force can increase or be maintained
(by PAP) at the same time as high-frequency force is de-
creasing (Fig. 4). With reference to Figure 4, a scenario in an
endurance event can be imagined in which an athlete is
faring well while maintaining a steady submaximal pace,
because of the beneficial effect of PAP. However, if the
intensity must be increased (hill climb, strategic pace in-
crease, etc.), with the accompanying high motor unit firing
rates, the situation would be as shown by the vertical dashed
line in Figure 4. By changing from low to high motor unit
firing rates, the beneficial effect of PAP would be lost and the
full effect of fatigue would be felt.
During recovery after fatiguing endurance exercise, PAP
decays within minutes whereas LFF can persist for at least
several hours, if not days (12). The prolonged depression of
low-frequency force may partly explain the “dead” leg com-
plaint of endurance athletes during a period of high-volume
training. From rest, when the athlete begins to do “easy” daily
activities such as walking and climbing a flight of stairs, the
leg muscles feel weak and are weak in response to the low
motor unit firing normally employed to generate the required
force. Faced with the LFF force deficit, central drive to the Figure 4. Changes in low- and high-frequency force capability during
endurance exercise. As exercise proceeds, the mechanisms of PAP and
motoneurons must increase to recruit more motor units and fatigue are developing simultaneously, the former enhancing low-fre-
to increase the firing rate of those already active, to com- quency force and the latter depressing high-frequency force. If the motor
pensate for the deficit. The athlete perceives this neural units are firing at the low rates typical of the submaximal contractions in
adjustment as increased effort. Perhaps surprising to the ath- endurance exercise, the PAP benefit should prevail, at least for a time, over
lete, when the activity has been underway for a short time, it the fatigue disadvantage. If, however, the intensity of exercise must be
increased (e.g., hill climb), necessitating high-frequency firing of motor
begins to feel easier. The explanation for this is that the units, the balance between force enhancement and depression will be
activity has reactivated PAP, offsetting the effect of LFF. altered to a predominance of depression (fatigue), as represented by the
(This explanation assumes that motor unit firing rates are still vertical dashed line in the figure.

140 Exercise and Sport Sciences Reviews www.acsm-essr.org


ditioning activity may activate the PAP mechanism to a
greater extent, but it also produces greater fatigue (Fig. 7).
The second dilemma, also illustrated in Figure 7, is that the
longer the recovery period between the end of the condi-
tioning activity and the beginning of performance, the
greater the recovery from fatigue, but also the greater the
decay of the PAP mechanism.
The two dilemmas can be resolved only by trial and error.
In one study (4), the recovery period chosen was only 15 s,
so that performance would begin when PAP (assessed by the
force of twitch contraction) was still near its maximum (see
Fig. 7). However, performance (velocity attained with a
given load in concentric knee extension) was actually de-
pressed because of fatigue induced by the conditioning ac-
tivity (10-s isometric MVC). This was another example of
low-frequency (isometric) force being increased at the same
time as high-frequency (concentric) force was depressed. If a
longer recovery period had been selected, say 3 min, perfor-
mance may have been improved, provided that fatigue had
dissipated at a faster rate than PAP had decayed (Fig. 7).
Figure 5. Hypothesized effect of PAP on the load (force)-velocity rela-
tion. PAP cannot increase maximum isometric force (Po) or maximum
There is some evidence that this is what happens when
shortening velocity (Vmax), because Po and Vmax are determined with high- longer recovery periods are used (3,5,15).
frequency stimulation. In contrast, PAP can increase rate of force devel- An additional consideration is that when the performance
opment at high frequencies (see Fig. 6), an effect that may increase the is a series of contractions, the contractions themselves have
acceleration and hence velocity attained with loads intermediate between a cumulative effect in mobilizing the PAP mechanism (4,10).
the extremes of Po and Vmax. If this were to occur, the load-velocity relation
would become less concave, i.e., shifted upward and to the right.
This may partly explain the progressive increase in perfor-
mance observed in a series of jumps (5) or dynamic knee
extensions (4,10). It has also been shown that the effects of
it less concave) without changing the endpoints, as depicted a conditioning activity and repeated performance have an
in Figure 5. If this were to happen, activities like jumping, additive effect on the magnitude of PAP, at least over a few
kicking, and throwing might be improved if the muscles were repetitions of the performance (4,10). In fact, if the perfor-
in a state of PAP. There are reports of improved (5,11,15) mance consists of enough trials, the PAP induced by the
and unaffected (4,9,10) performance of this nature after a trials themselves may rival that of the conditioning activity,
conditioning activity such as isometric MVCs or a set of
repetitions with a heavy weight. The inconsistent results of
the studies could be the result of variation in the performance
to be improved, the conditioning activity, and the time
interval between the conditioning activity and the perfor-
mance. In only two studies (4,10) was the presence or ab-
sence of PAP monitored. The selection of the conditioning
activity and the interval between it and performance are part
of the strategy for exploiting PAP (see next section).
In experiments with the muscle of a small mammal (rat
gastrocnemius), PAP was shown to increase the force and power
of isovelocity concentric contractions, particularly at the higher
velocities tested (up to ~70% of Vmax). PAP did not increase
isometric rate of force development or concentric force at the
highest stimulation frequency used (1), indicating that, like
isometric force, there is a limit (albeit higher than for isometric
force) to the stimulation frequency at which PAP can have an
enhancing effect on rate of force development. Whether this
limit is reached with the motor unit firing rates attained in
human performance is presently unknown.
Figure 6. Comparison of effect of PAP on isometric force and rate of
force development of twitch and high-frequency tetanic contractions.
STRATEGIES FOR EXPLOITING POSTACTIVATION PAP increases both the rate of force development and the peak force of
POTENTIATION (PAP) twitch and low-frequency tetanic contractions (the latter not shown), but
only the rate of force development of the high-frequency tetanic contrac-
tion is increased. This latter effect may alter the shape of the load-
In exploiting PAP to enhance performance, two dilemmas velocity relation (see Fig. 5) and potentially enhance strength and
must be resolved. First, a more intense and prolonged con- speed performance.

Volume 30 䡠 Number 3 䡠 July 2002 Postactivation Potentiation 141


portion of the whole muscle, and PAP is greater in fast fibers,
PAP of the muscle as a whole would be expected to increase.
A third possible adaptation would be increased capacity for
PAP, brought about by altered myosin light chain composi-
tion (see discussion of endurance training below). As a spe-
cific example, the regular performance of jumping drills
might improve jump performance in repeated jumps as a
result of increased PAP from the adaptations outlined. Of the
few longitudinal training studies done to date (for review, see
(6)), only one showed an increase in PAP, after 12 wk of
weight training in elderly subjects (8). In the one cross-
sectional study, PAP was greater in recreational weight train-
ers than sedentary subjects (6). When an increase in PAP has
been observed, the adaptation responsible for the increase
has not been identified.
In contrast to strength training, endurance training is
likely neither to increase the ability to activate high thresh-
old fast motor units in brief maximal contractions, nor to
cause preferential hypertrophy of fast twitch fibers. Never-
theless, the one published study related to endurance training
Figure 7. Strategy for exploiting PAP to improve strength/speed per-
formance. The conditioning activity activates PAP, monitored as the
found PAP, induced by a 10-s isometric MVC, to be greater
change in twitch force, and induces fatigue, monitored as the change in in endurance athletes than sedentary control subjects. The
high-frequency tetanic force. Strength/speed performance (e.g., vertical enhanced PAP was only present in the trained muscles,
jump), also involving high-frequency motor unit firing rates, would there- pointing to a training adaptation rather than a genetic trait
fore be depressed immediately after the conditioning activity, despite the (6). In addition to the absence of adaptations that might
presence of PAP. However, if fatigue dissipates faster than PAP decays, as
illustrated, performance will transiently (optimal recovery time) exceed the
increase PAP as a result of strength training, the endurance
best performance before the conditioning activity (Pre). The optimal re- athletes likely possessed a greater than average percentage of
covery time is determined by trial and error, taking into account factors slow twitch fibers, which would promote reduced rather than
such as the performance to be enhanced, the nature of the conditioning amplified PAP.
activity, and the fiber-type composition and training status of the subjects. What adaptations could account for the increased PAP in
the endurance athletes? One adaptation would be increased
fatigue resistance, which would allow PAP to prevail over
making the latter unnecessary (10). Again, all of this has to
be sorted out by trial and error experiments.
The emphasis has been on strategies for exploiting PAP in
strength/speed performance. For endurance performance, the
optimal strategy may be simply to begin the event. PAP will
be quickly induced by the first several contractions. It might
be argued, however, that preliminary (conditioning) “warm-
up” activity, apart from other possible benefits, would allow
the athlete to benefit from PAP at the beginning of the
event. As in strength and speed performance, a balance has
to be struck between the PAP and any fatigue induced by the
conditioning activity.

EFFECT OF TRAINING ON PAP

Some adaptations to strength training could possibly in-


crease PAP induced by a conditioning activity such as a 10-s
MVC. One adaptation would be increased ability to activate,
during the conditioning activity, the high threshold, fast Figure 8. Hypothetical effect of combined strength and endurance
training on PAP, as revealed by a fatigue test consisting of a series of brief
motor units, whose muscle fibers exhibit the greatest PAP. maximal voluntary contractions (MVCs). Twitch contractions evoked in
Because the magnitude of PAP is dependent on the extent of the intervals between the MVCs monitor the interaction between two
activation in the conditioning activity, greater activation opposing effects—PAP and fatigue. After training, the greater twitch
(recruitment and increased stimulation rate) of these fibers force early in the test may reflect mainly adaptations to strength training,
would contribute to greater PAP in the whole muscle. A whereas later in the test and during recovery, adaptations to endurance
training may play an increasing role. There is some evidence in support of
second adaptation is the preferential hypertrophy of fast the effect illustrated during the test (8). During the fatigue test, MVC
versus slow muscle fibers commonly observed with strength force, unlike twitch force, would have steadily declined (see Fig. 4), but
training. If, after training, fast fibers comprise a larger pro- perhaps not as much after training (not shown in the figure).

142 Exercise and Sport Sciences Reviews www.acsm-essr.org


fatigue immediately after the conditioning activity. In fact, in PAP to improve power performance. On the other hand,
the study cited above (6), the endurance athletes suffered less increased temperature shortens twitch duration and therefore
fatigue during the 10-s MVC used to induce PAP. A second shifts the force-frequency relation to the right, an effect
possible adaptation is an increase in the content of “fast” opposite to that of PAP. Thus, PAP’s possible enhancement
myosin light chains in Type I fibers (for review, see (6)), of endurance performance would be opposed by increased
which might increase the capacity for myosin light chain muscle temperature.
phosphorylation, the principal mechanism of PAP. The sig-
nificance of this adaptation, if it occurs, is that the capacity References
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Volume 30 䡠 Number 3 䡠 July 2002 Postactivation Potentiation 143

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