Theoretical Background and Practical Applications of the Physiological Mechanism of Post-Activation Potentiation

Abstract

Post-activation potentiation (PAP) is a mechanism that has gained interest in research during the past years. This is due to the fact that several studies use this physiological mechanism as a tool to improve athletic performance. This review will describe the basic principles of PAP and how exercise can influence the existence and magnitude of PAP. As definition, PAP is the phenomenon of increased force or twitch output that occurs after a high-intensity voluntary or evoked contraction. Although PAP is assumed to occur within the muscle due to the increased phosphorylation of the myosin light chains, several other factors can influence its appearance. The fact that fatigue counteracts the level of PAP, muscular as well as neural properties of the neuromuscular system contribute to the final outcome, despite the fact that PAP is mainly a mechanism that is attributed to changes in muscular level. During the past 5 years, our research group has examined in depth central and peripheral mechanisms that influence performance under the co-existence of PAP and fatigue. Numerous techniques combining the application of electromyography, electrical stimulation and force recording are briefly presented and discussed with emphasis on the practical aspect of interpretation on athletic performance. Using such techniques, some of the recent findings of our research group are presented, giving an insight in the PAP effect immediately after a contraction or after series of contractions and the effectiveness of long-term combined training programs.

Keywords: Post-activation potentiation, physiology, isometric, training

Introduction

Post-activation potentiation (PAP) is a topic of increased research interest during the past years and

more recently has been more intensively investigated among the athletic research community as well.

The phenomenon is rather complex and currently there is a significant body of literature that targets to

explain its underlying mechanisms and to determine more precisely the range of applications that might

have in human performance. Therefore, the purpose of this study is to analyse the basic principles of

PAP and its impact during exercise by analysing the factors that contribute to the PAP effect and

eventually determine performance.

1.1. Definitions

Earlier studies have shown that after repetitive stimulation over a muscle the produced twitch torque

increases for a short period of time after the end of the repetitive stimulation (Garnett, O'Donovan,

Stephens, & Taylor, 1979). In this case the repetitive stimulation represents the,

which in later studies has be replaced by an intense voluntary contraction with similar effects on twitch

torque (Vandervoort, Quinlan, & McComas, 1983). The phenomenon of an increased production in

twitch torque after an intense voluntary contraction is called or PAP,

whereas when the conditioning stimulus is induced by electrical stimulation of high frequency rate the

phenomenon is called (O'Leary, Hope, & Sale, 1997). The current study will

cover issues regarding the PAP.

1.2. Possible mechanisms

Numerous authors attribute the phenomenon of PAP to the phosphorylation of the myosin

regulatory light chains, in the presence of active myosin light chain kinase (Pääsuke, Ereline, &

Gapeyeva, 1996; Rassier & MacIntosh, 2000; Sale, 2004). This phosphorylation increases the

sensitivity of the released from the sarcoplasmic reticulum Ca2+ with the troponin-C (MacIntosh,

2003). This increases the level of cross-bridge formation for a given amount of Ca2+ release (Hodgson,

Docherty, & Robbins, 2005; Sale, 2004). Furthermore, when Ca2+ bind to calmodulin, the myosin light

chain kinase is activated, and the phosphorylation of the regulatory myosin light chains is catalysed

(Manning & Stull, 1982). As a result, more ATP is phosphorylated, improving the formation rate of

actin-myosin cross-bridge. Hence, the power output per cross-bridge is increased which might have a

positive effect during explosive activities (Houston, Lingley, Stuart, & Grange, 1987).

1.3. Factors influencing PAP

The amount of PAP under different circumstances is affected by multiple factors (Tillin & Bishop,

2009; Xenofondos et al., 2010). Numerous conditioning stimuli have been tested for PAP in the past,

varying for type of contraction and intensity, number of sets and repetitions. According to these studies

there is consensus that the contraction should have maximum or near maximum intensity and that

isometric contractions may induce higher PAP than dynamic ones (Rixon, Lamont, & Bemben, 2007).

Furthermore, it should be mentioned that there is an accumulative PAP effect when repetitive

conditioning stimuli are applied (Gossen & Sale, 2000; Tillin & Bishop, 2009). Furthermore, fast

twitch (type II) muscle fibers contain more myosin light chain kinase and less or its antagonist myosin

light chain phosphate than slow twitch (type I) muscle fibers (Moore & Stull, 1984). Therefore,

muscles that contain higher percentage of type II fibers are prone to higher PAP (Hamada, Sale,

MacDougall, & Tarnopolsky, 2000; Xenofondos, Patikas, Koceja et al., 2014). Another factor that

could affect PAP is the muscle architecture that might change after the conditioning stimulus. It has

been reported 3-6 minutes after a maximal voluntary contraction the pennation angle decreased

(Mahlfeld, Franke, & Awiszus, 2004), which works in favour for force production. However, a more

recent study did not verify this finding (Reardon et al., 2014).

As described in 1.2, PAP is a phenomenon that takes place on muscular level, and therefore it would

be expected that events within the muscle milieu should be responsible for its expression. However, the

conditioning stimulus does not exclusively result in the PAP effect, but induces fatigue as well (Sale,

2002). Fatigue and PAP act antagonistically and therefore, the measured muscle performance is the net

result of the effect of both PAP and fatigue (Rassier & MacIntosh, 2000). For this reason, it could be

argued that the final performance after a conditioning stimulus could be subjected to the effect of both

peripheral-muscular and central-neural factors.

Research methods and results

There are numerous methods that are used from many research groups to investigate the function of

the neuromuscular system. Many of them have been applied in the investigation of PAP as well and

two of them, one for peripheral and one for central aspects, are shortly presented below.

2.1. Twitch torque

When a muscle is electrically stimulated the produced force that is transmitted to the tendons, bones

and joints results in a measureable torque, the so-called twitch torque, which is independent of the

volition and skill of the subject. Twitch torque properties are a milestone for the evaluation of PAP.

The electrical stimulation can be applied over the muscle or the nerve that innervates the muscle or

muscle group of interest. From the twitch torque numerous parameters can be evaluated, which

describe several contractile properties of the examined muscles. Peak twitch torque is the maximum

torque that the muscle can produce with a single stimulus and is related to the capacity of the muscle to

produce force, which is a product of the muscle fibre diameter (Bowden & Goyer, 1960), the integrity

of excitation-contraction coupling (Ingalls, Warren, Williams, Ward, & Armstrong, 1998), the activity

of the myosin ATPase (Drachman & Johnston, 1973), and effectiveness of neuromuscular transmission

along the neuromuscular junction (Fuglevand, Zackoaski, Huey, & Enoka, 1993). Time to peak torque

and the rate of torque development (mean, maximum or over predefined intervals) are parameters that give information about the intracellular Ca2+ release kinetics (Harridge et al., 1996) and may be

influenced by the elastic properties of the muscle-tendon unit as well (Winter & Brookes, 1991).

After a conditioning stimulus, studies have shown 9.2%-133.3% increase in peak twitch torque

(Baudry & Duchateau, 2007; Folland, Wakamatsu, & Fimland, 2008; Gilbert & Lees, 2005; Hamada et

al., 2000; O’Leary et al., 1997; Xenofondos, Patikas, Koceja et al., 2014) and 40%-135% increase in

the rate of torque development (Froyd, Beltrami, Jensen, & Noakes, 2013; Gago, Arndt, Tarassova, &

Ekblom, 2014; Gago, Marques, Marinhio, & Ekblom, 2014; Shima, Rice, Ota, & Yabe, 2006).

Moreover, in most of the studies time to peak along and half relaxation time decreased (Alway,

Hughson, Green, Patla, & Frank, 1987; Hamada et al., 2000; Klein, Ivanova, Rice, & Garland, 2001;

Krarup, 1977; O’Leary et al., 1997; Vandenboom & Houston, 1996). These changes in twitch torque

indicate changes in Ca2+ kinetics, i.e. the rate of release from the sarcoplasmic reticulum and the

sensitivity to bind with troponin-C, as well as the rate of Ca2+ removal from the sarcoplasm (Hamada et

al., 2000; O’Leary et al., 1997).

2.2. Hoffmann reflex (H-reflex)

The H-reflex is a technique used to examine the motoneurone excitability via the Ia afferent reflex

pathway (Zehr, 2002). It can be affected by changes in the excitability of the motoneurone membrane

(post-synaptic), and the excitability of the Ia afferent terminals (pre-synaptic). Therefore, changes in

the H-reflex amplitude are explained by post-synaptic and pre-synaptic mechanisms, which are

governed by central-descending commands and peripheral-sensory input (Schieppati, 1987). The H-

reflex is evoked by low-intensity electrical stimulation over the nerve, which selectively stimulates the

Ia afferent axons, which in-turn recruit the homonymous α-motoneurones. The muscle action potential

which is recorded with EMG electrodes over the muscle represents the H-reflex, and the most common

measure of it is the peak-to-peak amplitude (Zehr, 2002).

Potentiation of the H-reflex has been suggested as a possible mechanism contributing to PAP, since

there are indications that, despite the small sample size, H-reflex is increased in the medial

gastrocnemius muscle 5-13 minutes after a conditioning stimulus (Güillich & Schmidtbleicher, 1996).

Nonetheless, according to other studies the contribution of such neural contribution on PAP appears to

be conflicting (Folland et al., 2008; Trimble & Harp, 1998; Xenofondos, Patikas, & Kotzamanidis,

2014). According to the recent findings of our research group, it seems that H-reflex decreases

immediately after the end of the conditioning contraction due to post-activation depression of the Ia

afferents and recovers to the pre-conditioning values with no further potentiation within 1-3 minutes

(Xenofondos, Patikas, Koceja et al., 2014).

PAP and human performance

The PAP effect on human performance has gained interest during the past years. Previous studies

have used conditioning contractions targeting to enhance subsequent performance, especially in

athletes. More particularly, they demonstrate an increase in performance for movements that demand

power after several different conditioning stimuli such as a maximum isometric voluntary contraction

(Tillin & Bishop, 2009), high intensity resistance training bouts (Tillin & Bishop, 2009), plyometrics

(Tobin & Delahunt, 2014) or whole body vibration (Avelar et al., 2014). The positive effect of PAP

was observed mainly in rate of force development, jump height and sprint time, but not in strength

(Tillin & Bishop, 2009).

Although these results have been attributed to PAP and a recent study showed that in some extend

PAP in twitch torque is correlated with performance enhancement (Nibali, Chapman, Robergs, &

Drinkwater, 2013), it is questionable whether this could be the case. For instance, the gain in

performance does not appear at the same time interval after conditioning stimulus as PAP appears

when obtained by twitch torque measures. More specifically, twitch torque properties reach their

maximal values directly after or within the first minute after the end of the conditioning stimulus, and

thereafter the PAP effect gradually decreases (Hamada et al., 2000). On the contrary, in many cases

performance enhancement is not observed immediately after conditioning stimulus but after several

minutes (up to 10 minutes, with optimum at ~5 minutes), when actually PAP effect is basically absent

(Batista et al., 2007; Tillin & Bishop, 2009).

During and after a high intensity resistance training protocol fatigue and PAP coexist and compete

out each other (Tillin & Bishop, 2009) and therefore a diminished PAP effect would be expected.

Untrained subjects demonstrated that between sets, jump height occasionally increased and then

decreased (Smilios, Pilianidis, Sotiropoulos, Antonakis, & Tokmakidis, 2005), while sprint time was

not affected (Tsimahidis et al., 2010). In trained athletes, however, sprint performance between sets

increased, underlying the importance of training background (Tsimachidis, Patikas, Galazoulas, Bassa,

& Kotzamanidis, 2013). Regarding the potentiating effect after high intensity resistance training there

are some studies that showed no effect in performance (Chatzopoulos et al., 2007) or increased

performance (Gourgoulis, Aggelousis, Kasimatis, Mavromatis, & Garas, 2003), which indicates that

further research in this field is recommended to be conclusive.

One plausible explanation for the discrepancy between PAP and performance is that voluntary

execution of a movement is influenced by many more parameters than twitch torque. For instance

although children and adults demonstrated similar PAP levels in terms of twitch torque (Pääsuke,

2000), children are not able to improve their jumping performance as adults do after a conditioning

stimulus (Arabatzi et al., 2014). This indicates that deterioration in technique could mask the PAP

effect, especially immediately after the conditioning stimulus when the fatigue effect is more

prominent.

Conclusions

PAP is a complex phenomenon that is affected by events occurring on muscular level, but since it

can be opposed to some extent by fatigue it may be influenced by neural factors as well. Despite the

large number of studies, further research in a more targeted and controlled manner is required to

explain more precisely relationship between PAP and human performance. Regarding performance, the

maximal augmentation in athletic performance attributed to PAP is expected to occur when a

conditioning stimulus or a series of conditioning stimuli have an optimal duration, intensity and rest

interval that maximize the PAP effect and limit as much as possible fatigue, taking into account their

accumulative effect that exists.

References

• Alway, S. E., Hughson, R. L., Green, H. J., Patla, A. E., & Frank, J. S. (1987). Twitch potentiation after

• fatiguing exercise in man. European Journal of Applied Physiology, 56, 461-466.

• Arabatzi, F., Patikas, D., Zafeiridis, A., Giavroudis, K., Kannas, T., Gourgoulis, V., & Kotzamanidis, C. (2014). The post-activation potentiation effect on squat jump performance: Age and sex effect. Pediatric Exercise Science, 26, 187-194.

• Avelar, N. C., Salvador, F. S., Ribeiro, V. G., Vianna, D. M., Costa, S. J., Gripp, F., Coimbra, C. C., & Lacerda, A. C. (2014). Whole body vibration and post-activation potentiation: A study with repeated measures. International Journal of Sports Medicine, 35, 651-657.

• Batista, M. A. B., Ugrinowitsch, C., Roschel, H., Lotufo, R., Ricard, M. D., & Tricoli, C. A. A. (2007). Intermittent exercise as a conidtioning activity to induce postactivation potentiation. Journal of Strength and Conditioning Research, 21, 837-840.

• Baudry, S., & Duchateau, J. (2007). Postactivation potentiation in human muscle: Effect on the rate of torque development of tetanic and voluntary isometric contractions. Journal of Applied Physiology, 102, 1394-1401.

• Bowden, D. H., & Goyer, R. A. (1960). The size of muscle fibers in infants and children. Archives of Pathology & Laboratory Medicine, 60, 188-189.

• Chatzopoulos, D. E., Michailidis, C. J., Giannakos, A. K., Alexiou, K. C., Patikas, D. A., Antonopoulos, C.

• B., & Kotzamanidis, C. M. (2007). Postactivation potentiation effects after heavy resistance exercise on running speed. Journal of Strength and Conditioning Research, 21, 1278-1281.

• Drachman, D. B., & Johnston, D. M. (1973). Development of a mammalian fast muscle: dynamic and biochemical properties correlated. Journal of Physiology, 234, 29-42.

• Folland, J. P., Wakamatsu, T., & Fimland, M. S. (2008). The influence of maximal isometric activity on twitch and H-reflex potentiation, and quadriceps femoris performance. European Journal of Applied Physiology, 104, 739-748.

• Froyd, C., Beltrami, F. G., Jensen, J., & Noakes, T. D. (2013). Potentiation increases peak twitch torque by enhancing rates of torque development and relaxation. Journal of Human Kinetics, 38, 83-94.

• Fuglevand, A. J., Zackoaski, M. M., Huey, K. A., & Enoka, R. M. (1993). Impairment of neuromuscular propagation during human fatiguing contractions at submaximal forces. Journal of Physiology (London), 460, 549-572.

• Güillich, A., & Schmidtbleicher, D. (1996). MVC-induced short-term potentiation of explosive force. New Studies in Athletics, 11, 67–81.

• Gago, P., Arndt, A., Tarassova, O., & Ekblom, M. M. (2014). Post activation potentiation can be induced without impairing tendon stiffness. European Journal of Applied Physiology, 114, 2299-2308.

• Gago, P., Marques, M. C., Marinhio, D. A., & Ekblom, M. M. (2014). Passive muscle length changes affect twitch potentiation in power athletes. Medicine and Science in Sports and Exercise, 46, 1334-1342.

• Garnett, R. A. F., O'Donovan, M. J., Stephens, J. A., & Taylor, A. (1979). Motor unit organisation of human medial gastrocnemius. Journal of Physiology (London), 287, 33-43.

• Gilbert, G., & Lees, A. (2005). Changes in the force development characteristics of muscle following repeated maximum force and power exercise. Ergonomics, 48, 1576-1584.

• Gossen, E. R., & Sale, D. G. (2000). Effect of postactivation potentiation on dynamic knee extension performance. European Journal of Applied Physiology, 83, 524-530.

• Gourgoulis, V., Aggelousis, N., Kasimatis, P., Mavromatis, G., & Garas, A. (2003). Effect of a submaximal Half-Squats warm-up program on vertical jumping ability. Journal of Strength and Conditioning Research, 17, 342-344.

• Hamada, T., Sale, D. G., MacDougall, J. D., & Tarnopolsky, M. A. (2000). Postactivation potentiation, fiber type, and twitch contraction time in human knee extensor muscles. Journal of Applied Physiology, 88, 2131-2137.

• Harridge, S., Bottinelli, R., Canepari, M., Pellegrino, M. A., Reggiani, C., Esbjörnsson, M., & Saltin, B.

• (1996). Whole-muscle and single-fibre contractile properties and myosin heavy chain isoforms in humans. Pflügers Archives, 432, 913-920.

• Hodgson, M., Docherty, D., & Robbins, D. (2005). Post-activation potentiation: Underlying physiology and implications for motor performance. Sports Medicine, 35, 585-595.

• Houston, M. E., Lingley, M. D., Stuart, D. S., & Grange, R. W. (1987). Myosin light chain phosphorylation in intact human muscle. FEBS Letters, 219, 469-471.

• Ingalls, C. P., Warren, G. L., Williams, J. H., Ward, C. W., & Armstrong, R. B. (1998). E-C coupling failure in mouse EDL muscle after in vivo eccentric contractions. Journal of Applied Physiology, 85, 58-67.

• 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, 153-156.

• Krarup, C. (1977). Electrical and mechanical responses in the platysma and in the adductor pollicis muscle: in normal subjects. Journal of Neurology, Neurosurgery and Psychiatry, 40, 234-240.

• MacIntosh, B. R. (2003). Role of calcium sensitivity modulation in skeletal muscle performance. News Physiol Sci, 18, 222-225.

• Mahlfeld, K., Franke, J., & Awiszus, F. (2004). Postcontraction changes of muscle architecture in human quadriceps muscle. Muscle and Nerve, 29, 597-600.

• Manning, D. R., & Stull, J. T. (1982). Myosin light chain phosphorylation-dephosphorylation in mammalian skeletal muscle. American Journal of Physiology, 242, C234-241.

• Moore, R. L., & Stull, J. T. (1984). Myosin light chain phosphorylation in fast and slow skeletal muscles in situ. American Journal of Physiology, 275, C462-C471.

• 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, 306-313.

• O'Leary, D. D., Hope, K., & Sale, D. G. (1997). Posttetanic potentiation of human dorsiflexors. Journal of Applied Physiology, 83, 2131-2138.

• Pääsuke, M. (2000). Twitch contraction properties of plantar flexor muscles in pre- and post-pubertal boys and men. European Journal of Applied Physiology, 82, 459-464.

• Pääsuke, M., Ereline, J., & Gapeyeva, H. (1996). Twitch potentiation capacity of plantar-flexor muscles in endurance and power athletes. Biology of Sport, 15, 171-178.

• Rassier, D. E., & MacIntosh, B. R. (2000). Coexistence of potentiation and fatigue in skeletal muscle.

• Brazilian Journal of Medical and Biological Research, 33, 499-508.

• Reardon, D., Hoffman, J. R., Mangine, G. T., Wells, A. J., Gonzalez, A. M., Jajtner, A. R., Townsend, J. R., McCormack, W. P., Stout, J. R., Fragala, M. S., & Fukuda, D. H. (2014). Do changes in muscle architecture affect post-activation potentiation? Journal of Sports Science and Medicine, 13, 483-492. Rixon, K. P., Lamont, H. S., & Bemben, M. G. (2007). Influence of type of muscle contraction, gender, and lifting experience on postactivation potentiation performance. Journal of Strength and Conditioning Research, 21, 500-505.

• Sale, D. G. (2002). Postactivation potentiation: Role in human performance. Exercise and Sport Sciences Reviews, 30, 138-143.

• Sale, D. G. (2004). Postactivation potentiation: Role in performance. British Journal of Sports Medicine, 38, 386-387.

• Schieppati, M. (1987). The Hoffman Reflex: A means of assessing spinal reflex excitability and its descending control in man. Progress in Neurobiology, 28, 345-376.

• Shima, N., Rice, C. L., Ota, Y., & Yabe, K. (2006). The effect of postactivation potentiation on the mechanomyogram. European Journal of Applied Physiology, 96, 17-23.

• Smilios, I., Pilianidis, T., Sotiropoulos, K., Antonakis, M., & Tokmakidis, S. P. (2005). Short-term effects of selected exercise and load in contrast training on vertical jump performance. Journal of Strength and Conditioning Research, 19, 135-139.

• Tillin, N. A., & Bishop, D. (2009). Factors modulating post-activation potentiation and its effect on performance of subsequent explosive activities. Sports Medicine, 39, 147-166.

• Tobin, D. M., & Delahunt, E. (2014). The acute effect of a plyometric stimulus on jump performance in professional rugby players. Journal of Strength and Conditioning Research, 28, 367-372.

• Trimble, M. H., & Harp, S. S. (1998). Postexercise potentiation of the H-reflex in humans. Medicine and Science in Sports and Exercise, 30, 933-941.

• Tsimachidis, C., Patikas, D., Galazoulas, C., Bassa, E., & Kotzamanidis, C. (2013). The post-activation potentiation effect on sprint performance after combined resistance/sprint training in junior basketball players. Journal of Sports Sciences, 31, 1117-1124.

• Tsimahidis, K., Galazoulas, C., Skoufas, D., Papaiakovou, G., Bassa, E., Patikas, D., & Kotzamanidis, C.

• (2010). The effect of sprinting after each set of heavy resistance training on the running speed and jumping performance of young basketball players. Journal of Strength and Conditioning Research, 24, 2102-2108.

• Vandenboom, R., & Houston, M. E. (1996). Phosphorylation of myosin and twitch potentiation in fatigued skeletal muscle. Canadian Journal of Physiological Pharmacology, 74, 1315-1321.

• Vandervoort, A. A., Quinlan, J., & McComas, A. J. (1983). Twitch potentiation after voluntary contraction. Experimental Neurology, 81, 141-152.

• Winter, E. M., & Brookes, F. B. (1991). Electromechanical response times and muscle elasticity in men and women. European Journal of Applied Physiology, 63, 124-128.

• Xenofondos, A., Laparidis, K., Kyranoudis, A., Galazoulas, C., Bassa, E., & Kotzamanidis, C. (2010). Post-activation potentiation: Factors affecting it and the effect on performance. Journal of Physical Education and Sport, 28, 32-38.

• Xenofondos, A., Patikas, D., Koceja, D. M., Behdad, T., Bassa, E., Kellis, E., & Kotzamanidis, C. (2014). Post-activation potentiation: The neural effects of post-activation depression. Muscle & Nerve.

• Xenofondos, A., Patikas, D., & Kotzamanidis, C. (2014). On the mechanisms of post-activation potentiation: the contribution of neural factors Journal of Physical Education and Sport, 14, 134-137.

• Zehr, E. P. (2002). Considerations for use of the Hoffmann reflex in exercise studies. European Journal of Applied Physiology, 86, 455-468.