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Eur J Appl Physiol (2000) 83: 524±530 Ó Springer-Verlag 2000

ORIGINAL ARTICLE

E. Roderich Gossen á Digby G. Sale

Effect of postactivation potentiation on dynamic knee


extension performance

Accepted: 27 July 2000

Abstract Six men and four women performed, in sepa-


rate trials, maximal dynamic knee extensions with loads
Introduction
of 15%, 30%, 45% and 60% of maximal isometric knee
The force of muscle twitch contractions is increased after
extension peak torque (MVC). The dynamic extensions
a conditioning tetanus (posttetanic potentiation or PTP)
were done after postactivation potentiation (PAP) had
(O'Leary et al. 1997), a maximal voluntary contraction
been induced with a 10-s MVC, and in a control trial
(postactivation potentiation or PAP) (Hamada et al.
without PAP. PAP, measured as the increase in evoked
2000) or repeated sub-fusion stimuli (MacIntosh and
twitch torque, was 53 (4)% (SE) and 43 (3)% at the time
Kupsh 1987; MacIntosh and Willis 2000). For simplicity,
of the ®rst and second extensions with each load. PAP
all the aforementioned forms of activity-dependent pot-
failed to increase the attained peak velocity with any
entiation will be referred to as PAP. PAP is probably the
load; on the contrary, there was a trend for peak velocity
result of phosphorylation of the myosin regulatory light
to decrease in the ®rst extension, which occurred @15 s
chains via myosin light chain kinase (Sweeney et al. 1993),
after the 10-s MVC. The results suggest that fatigue
which theoretically increases the interaction between the
produced by the 10-s MVC suppressed any bene®t that
myosin cross-bridge and the thin ®lament (Levine et al.
could be derived from the induced PAP. A surface
1996; Persechini et al. 1985). In addition to increasing
electromyogram (EMG) recorded from one muscle of
peak force, PAP increases the twitch's rate of force de-
quadriceps femoris gave no indication of activation
velopment and decreases its time to peak force (Grange
failure in the ®rst knee extension; however, activation
et al. 1993; O'Leary et al. 1997; Sweeney et al. 1993).
impairment speci®c to the rate of force development
PAP also increases the force and rate of force devel-
cannot be ruled out. It is concluded that the strategy
opment of low-frequency (sub-fusion frequencies) te-
employed, namely of having knee extensions performed
tanic isometric contractions (Vandenboom et al. 1993),
soon after the 10-s MVC to maximize PAP at the time of
and under dynamic conditions (shortening contractions
performance, was unsuccessful because there had been
with various loads), again with sub-fusion frequencies,
insucient time for recovery from fatigue. It is possible
PAP increases work and power output (Grange et al.
that a longer recovery time, even at the cost of a di-
1995, 1998).
minished PAP, may have proved bene®cial.
In contrast, PAP does not increase the peak force (Po)
of high-frequency tetani; only the rate of force develop-
Key words Force±velocity relationship á Posttetanic
ment is enhanced (Vandenboom et al. 1993). Nor does
potentiation á Twitch contraction
PAP increase the maximum unresisted shortening velocity
(Vmax) of high-frequency tetani (Butler et al. 1983;
Persechini et al. 1985), or the maximum velocity of un-
resisted maximal voluntary concentric contractions
(Stuart et al. 1988). Although PAP does not increase Po or
Vmax, its positive e€ect on the isometric rate of force de-
E. R. Gossen á D. G. Sale (&) velopment raises the possibility that PAP might enhance
Department of Kinesiology, dynamic contractile performance with loads intermediate
McMaster University, Hamilton, between those representing Po and Vmax; that is, the load±
Ontario L8S 4K1, Canada
e-mail: saled@mcmaster.ca
velocity relation would be shifted upward and to the right.
Tel.: +1-905-5259140 ext. 23565 Thus, PAP could partly explain improvements in jump
Fax: +1-905-5236011 height (GuÈllich and Schmidtbleicher 1995; Young et al.
525

1998) and arm push speed (GuÈllich and Schmidtbleicher General design
1995) achieved following maximal isometric contractions
PAP and its e€ect on the load±velocity relation in the human
or weight lifting. In these studies, however, the e€ects of quadriceps muscle were studied. The experimental protocol is il-
the presence or absence of PAP was not examined. lustrated in Fig. 1. The protocol was designed so that the extent of
The purpose of the present study was to examine the PAP could be quanti®ed and monitored from evoked isometric
e€ects of PAP on dynamic knee extension performance twitch responses, and the e€ect of PAP on the load±velocity rela-
tion could be assessed by the performance of dynamic knee ex-
with imposed loads between the extremes represented by tensions (``kicks'') with various loads. An experimental session
Po and Vmax. We tested the hypothesis that PAP, by consisted of a control trial (CON), in which twitches were evoked
increasing the rate of force development (Vandenboom and two kicks were performed with a designated load, and then,
et al. 1993, 1995), would increase the acceleration and after a 15-min rest period, a PAP trial which was identical to the
thus peak velocity and power attained with these inter- CON trial, except for the inclusion of a 10-s maximal voluntary
isometric contraction (MVC) to induce PAP. Each subject repeated
mediate loads. the experimental session (CON+PAP trials) four times, one ses-
sion for each of four loads in random order (15%, 30%, 45% and
60% of the MVC peak torque determined in a previous familiar-
Methods ization session). On a given day, one session was done with one leg,
and a second session with the other leg. In two separate twitch
control experiments, one for each leg, the protocol was identical to
Subjects that of a PAP trial (Fig. 1), except that the dynamic knee exten-
sions were replaced with evoked twitches. Finally, prior to the
Ten moderately active subjects (six male, four female) between 22 CON + PAP sessions and the isometric control sessions, each
and 35 years of age volunteered for the study after giving informed, subject attended a familiarization session in which, for each leg,
written consent. To be eligible for participation, all subjects had to electrical stimulation was administered, MVC peak torque deter-
be free of lower leg injury. The study carried the approval of mined, and dynamic knee extensions practised. Altogether, each
McMaster University's Human Ethics Committee. Subjects were subject had to attend the laboratory on four occasions: familiar-
instructed to refrain from any heavy lower leg exercises or ca€eine ization session (both legs), two CON + PAP sessions (two of four
consumption 24 h prior to testing. loads per session, one load per leg), and one twitch control session
(both legs).

Fig. 1 A schematic of the control (CON) and postactivation Dynamometer


potentiation (PAP) trials that made up each test session. A trial
began with the establishment of a baseline twitch (Pre). In the CON A Biodex System 3 dynamometer (Biodex Medical Systems, Shir-
trial, this was followed by a 10-s rest period, followed, at the times ley, NY, USA) was used to test the knee extensor muscles. The
indicated, by evoked twitches and dynamic knee extensions at one of dynamometer's ``isotonic'' mode was used to set loads for the dy-
the four imposed loads. Twitches marked (*) indicate the extent to namic knee extensions (concentric contractions), which began at a
which the knee extensions themselves induced PAP. After a 15-min joint angle of 90° and continued to full extension. The dynamom-
rest period, a PAP trial was done, which was identical to the CON eter's isometric mode was used to measure the peak torque of
trial except that the 10-s rest pause after the baseline twitch was maximal voluntary isometric contractions (MVC) and the con-
replaced with a 10-s maximal voluntary isometric contraction (MVC) tractile properties of evoked isometric twitch contractions (knee
(to induce PAP). When this session (CON+PAP trials) was joint angle ®xed at 90°).
completed on one leg, another session (with the second of four The subject sat on the Biodex ``chair'', which provided thigh
dynamic loads to be tested) was done with the other leg. In a second and back support. A Velcro belt was secured over the subject's hips
visit to the laboratory, two additional sessions (one per leg) were to minimize movement in the dynamometer chair. The thigh was
conducted to test the two remaining dynamic loads. On a third visit secured with wide strap restraints and the ankle was secured to the
two additional twitch control trials (one per leg) were conducted. The leg armature by a Velcro strap. The chair position was adjusted so
twitch control trial was identical to the PAP trial, except that the that the knee's axis of rotation (tibio-femoral joint) was aligned
dynamic knee extensions were replaced with evoked twitches. See text with the dynamometer's armature axis of rotation with the knee at
for details 90° (visually estimated).
526

Stimulating and recording electrodes ratus, the time interval between the evoked twitch and dynamic
knee extensions could be no less than 10 s.
Prior to electrode attachment, skin surfaces were shaven, abraded
and swabbed with alcohol pads. Stimulation electrodes consisted of
an anode, a 6 cm ´ 4 cm lead plate coated with electrode gel Isometric control session
(Spectra 300) placed on the proximal quarter of the lateral thigh
using surgical tape (Blenderm), and a cathode, a 1.5 cm ´ 1.5 cm An isometric twitch control trial was included because of the
lead plate with 1.5-cm-thick dense foam fastened to one side, which possibility of signi®cant degradation of potentiation during the
was initially positioned immediately medial to the tendon of the period between the end of the MVC and the beginning of the ®rst
sartorius in the inguinal fold. To achieve ®nal positioning, the dynamic knee extension, and the added confounder of additional
subject was instructed to hold the electrode in place and apply time delays between switching from isometric to isotonic modes
minimal pressure on the foam backing so as to mimic the pressure on the dynamometer during each measurement interval. The
that the tape would exert on the electrode. A 250-ls square-wave twitch control trials were identical to the PAP trials except that
pulse from a Grass S11 stimulator (Grass Instruments, Quincy, isometric twitches were evoked when dynamic knee extensions
Mass., USA) was used to evoke a minimal twitch response. All would have occurred in the CON and PAP trials. This provided
twitches were evoked with the dynamometer in the o€ mode to an estimate of the amount of PAP present just as a dynamic knee
minimize signal noise. The subject was then instructed to move the extension was initiated. In addition, comparisons between
cathode medially down the inguinal fold by approximately 1 cm. twitches evoked prior to and after each contraction during the
Using the same minimal stimulation intensity, a subsequent twitch four load sessions and those of the twitch control session served
was evoked. This procedure was repeated until the electrode was to verify the consistency of potentiation within subjects across test
positioned directly over the femoral nerve, which was re¯ected by days.
the largest twitch response. The cathode was then taped into place.
The stimulus intensity was then increased until a maximal twitch
response was evoked. Measurements
To monitor agonist EMG activity, electrodes (Meditrace Ag/
AgCl ECG) were used in a monopolar con®guration. The active All isometric and dynamic analogue signals were fed directly from
electrode was placed on the vastus medialis approximately 8 cm the dynamometer head into a 12-bit A/D converter, then into a
proximal from the knee. The reference electrode was positioned on computer sampling at 2000 Hz per channel using CODAS software
the patellar ligament immediately inferior to the apex of the patella, (DATAQ electronics, Akron, Ohio, USA). Measures included
while the ground was placed over the tibia midway between the torque and armature angle. The EMG was collected using a
knee and ankle joints. The chosen electrode placement minimized bandwidth ranging from 4 to 2000 Hz (Grass P5 series ampli®er,
the e€ect of the stimulus artefact on the recording of the evoked Grass Instruments, Quincy, Mass., USA) and afterwards digitally
muscle action potential (M-wave). ®ltered during processing.
Vastus medialis is one of four heads of quadriceps femoris. It Twitch response measurements included peak twitch torque (Pt)
was chosen because M-waves could be recorded from this muscle and the amplitude of the associated muscle compound action po-
without large stimulus artefacts. While there is some evidence that tential (M-wave). Dynamic performance measures included peak
all heads of quadriceps have similar activation characteristics torque, peak power, work attained up to the point of peak power
during loaded and unloaded knee extension (Basmajian et al. (hereafter referred to as work to peak power) and peak velocity.
1972), which suggests that recording from only vastus medialis is Velocity was calculated by di€erentiating angular displacement
representative of the whole quadriceps, di€erential activation using custom residual analysis software. Velocity was then multi-
among the heads of quadriceps cannot be ruled out. plied by torque to derive power. The power±time curve was then
integrated to obtain work.
Protocol Both voluntary and evoked (M-wave) EMGs were monitored.
The voluntary EMG was intended to determine whether any per-
Familiarization session formance enhancements were in part due to increased motor unit
activation. M-wave measurements monitored any changes in the
The primary purpose of this session was to familiarize the subjects muscle compound action potential, and also served to normalize
with electrical stimulation of the knee extensors and performance voluntary EMG. Prior to analysis, all voluntary EMGs were
of maximal dynamic and isometric contractions on the dyna- highpass ®ltered (cuto€ frequency ˆ 10 Hz) with custom software
mometer. Subjects were required to perform three 10-s maximal to remove movement artefacts. To obtain a measure of activation
isometric knee extensions (MVCs) per leg. The dynamic loads for across an entire movement, an average EMG (AEMG) was cal-
subsequent test sessions were derived as a percentage of the best culated by rectifying the signal during each knee extension and
MVC peak torque achieved with each leg. Following the MVCs, averaging all data points during movement; values were then nor-
each subject was required to do a minimum of three practice malized to M-wave amplitude and expressed as a percentage of the
maximal dynamic knee extensions at each of the four loads (15, 30, latter.
45, 60% MVC).

Statistics
Control (CON) and PAP sessions
A three-factor mixed analysis of variance (ANOVA) on all dy-
Each of these sessions consisted of a CON trial (no 10-s MVC to namic measures (within, 4 loads ´ 2 conditions [CON/PAP] ´ 2
induce PAP) followed, after a 15-min rest period, by a PAP trial kicks per condition) was performed. Twitch response measure-
(10-s MVC included to induce PAP). One session was done for each ments were analysed by a three-factor ANOVA (within, 4 loads ´ 2
of four randomly assigned dynamic loads [15%, 30%, 45% and conditions ´ 4 times) to assess the degree of isometric twitch pot-
60% of the highest MVC torque obtained during the familiariza- entiation (PAP) across trials. ANOVA was also used to compare
tion (habituation) session for the leg to be tested]. The time line the twitch responses across the CON, PAP and twitch control
shown in Fig. 1 indicates when, during the course of a trial, conditions. When signi®cant main e€ects or interactions were
twitches were evoked and dynamic knee extensions performed. found with ANOVA, the Tukey post hoc procedure tested for
Since isometric twitches were used to quantify the amount of PAP di€erences among mean values. Statistical signi®cance was set at
during the dynamic actions, they were evoked as close as possible P £ 0.05. Descriptive statistics include mean and standard error of
to each dynamic knee extension. Due to constraints of the appa- the mean (SE).
527

(``kicks'') were performed. In the CON experiment, that


Results is without the 10-s MVC, it was evident that the kicks
themselves induced PAP. In the PAP trial, the e€ects of
Twitch contractile properties the 10-s MVC and kicks on PAP were additive; that is,
PAP assessed at 30 s and especially at 50 s postMVC
Baseline or ``resting'' twitch peak torque values (Pt) were was greater with than without (twitch control trial) the
determined in ten experiments: the control (CON) dy- added kicks (see Fig. 2). Although not presented, the
namic performance tests with 15, 30, 45 and 60% MVC potentiated twitches had a shortened time course and
loads, the PAP dynamic performance tests at the same increased peak rate of torque development, as shown
loads and the right and left leg twitch control experi- previously in this muscle group (Hamada et al. 2000).
ments (i.e. without dynamic performance). The mean Pt M-wave amplitude did not change signi®cantly during
values ranged from 29.1 (2.4) to 32.9 (3.3) N á m, with any of the trials (data not shown).
no signi®cant di€erences among the mean values. Of
particular note was the lack of signi®cant di€erences
between the CON and PAP trials, indicating that the 15- Dynamic knee extension performance
min rest interval between these paired experiments was
sucient for restoration of baseline (twitch) conditions; PAP did not increase the peak velocity achieved across
that is, any PAP induced during the CON trial had the four dynamic loads (Fig. 3). On the contrary, in the
subsided before the beginning of the PAP trial. Simi- PAP trial, the peak velocity achieved with kick no. 1
larly, there were no signi®cant di€erences in baseline M- tended to be depressed compared to kick no. 1 in the
wave amplitude across the ten experimental conditions. CON trial and compared to kick no. 2 in both trials.
Values ranged from 17.4 (1.3) to 19.4 (1.3) mV. All Post hoc tests to determine signi®cant di€erences among
evoked measures showed acceptable reproducibility. the mean values shown in Fig. 3 were not done because
Method errors calculated across the two test days were there were no signi®cant interactions. However, the
7.7% for Pt, 4.8% for Pt immediately following the 10-s tendency for peak velocity to be depressed in kick no. 1
MVC (i.e. PAP) and 7.1% for M-wave amplitude. The of the PAP trial resulted in two signi®cant main e€ects.
method error for the peak torque of the 10-s MVCs done There was a main e€ect for condition (P < 0.03): col-
on separate days was larger (16.3%). lapsed across kicks and loads, peak velocity was greater
Figure 2 shows the twitch responses (normalized to in CON [341.6 (10.6) °/s] than PAP [326.7 (10.6) °/s].
baseline values) throughout the course of the CON, PAP There was also a main e€ect for kick order (P < 0.02):
(both collapsed across the four loads) and twitch control collapsed across loads and conditions, peak velocity was
experiments (collapsed across right and left legs). The greater in kick no. 2 [340.2 (10.6) °/s] than kick no. 1
e€ectiveness of the 10-s MVC in inducing PAP and [328.1 (10.7) °/s].
sustaining it during the observation period is demon-
strated in the twitch control experiment; speci®cally,
PAP was present at the times when the ®rst [52.6 (3.7)%]
and second [43.2 (2.9)%] dynamic knee extensions

Fig. 2 Twitch peak torque (Pt) normalized to baseline values (PRE)


during CON and PAP trials collapsed across the four dynamic loads, Fig. 3 Load±velocity relation obtained with the ®rst and second knee
and twitch control trials collapsed across left and right legs extensions in the control (CON1, CON2) and PAP trials (PAP1,
(observations per mean for CON and PAP ˆ 40; twitch con- PAP2). Values are means (10 observations for each mean) and SE
trol ˆ 20). n Time points when dynamic knee extensions (``kicks'') (some error bars omitted for clarity). Since there were no signi®cant
occurred during CON and PAP; + greater than PRE, P < 0.01; interactions, comparisons among mean values were not made (no post
*
di€erent from all other values at same time point, P < 0.05 hoc tests)
528

The peak torque attained during the dynamic knee be representative of loads used in performance (e.g.
extensions (kicks) was also una€ected by PAP; nor did throwing, kicking, etc.).
peak torque change from kick no. 1 to 2. Only the load Our results do not agree with previous studies of the
main e€ect was signi®cant (P < 0.0001); that is, the e€ect of ``conditioning'' contractions on dynamic per-
peak torque increased in kicks against greater loads. For formance. GuÈllich and Schmidtbleicher (1995) found, in
example, in kick no. 1 of the PAP trials, peak torque a group of power event athletes, that vertical jump
increased from 67.6 (6.1) N á m at a load of 15% MVC height increased after three (presumably isometric)
to 138.9 (13.0) N á m at 60% MVC. maximal voluntary contractions (MVCs). In addition,
PAP did not enhance peak power or work to peak the movement time of an ``arm push'' test was reduced.
power (i.e. no signi®cant interactions). There were two A second study (Young et al. 1998) found, in a group of
notable main e€ects, however (Table 1). In the CON ten men with at least 1 year's experience in the barbell
condition, work to peak power was greater in kick no. squat exercise, that a preceding set of ®ve repetitions of
2 than no. 1; peak power and time to peak power the barbell squat exercise with the heaviest possible
were una€ected. In contrast, in the PAP condition, weight (i.e. 5 RM), increased the average height of a
peak power was greater in kick no. 2 than no. 1; work series of ®ve vertical jumps with a 19-kg load. There was
to peak power and time to peak power were unaf- a 4-min rest interval between the end of the set of squats
fected. and the beginning of the series of vertical jumps. There is
AEMG from the vastus medialis, expressed as a a third brief (abstract) report indicating mixed results of
percentage of M-wave amplitude, did not di€er between the e€ects of prior activity on horizontal countermove
the CON and PAP trials [CON, 4.9 (0.2)%; PAP, 4.5 jump performance (Radcli€e and Radcli€e 1996). In a
(0.3)%, P ˆ 0.26]. group of 24 male college athletes, 4 sets of 4 repetitions
in the power snatch exercise, with 75±85% of the 4 RM,
increased the distance attained in three countermove
Discussion jumps initiated within 3 min of completion of the fourth
set of the prior exercise. In contrast, 4 sets of 4 repeti-
PAP does not increase maximal tetanic isometric peak tions of the squat exercise with 75±85% 4 RM, 4 sets of
force (Po) (Vandenboom et al. 1993) or maximal un- 4 loaded jumps with 15±20% body weight added, and 4
loaded shortening velocity (Vmax) (Butler et al. 1983; sets of 4 unloaded jumps failed to improve subsequent
Grange et al. 1993; Stuart et al. 1988). On the other countermove jump performance. In a group of 11 female
hand, PAP does increase the isometric rate of force de- athletes, none of the prior activities improved perfor-
velopment (Vandenboom et al. 1993, 1995, 1997). This mance. None of these studies (GuÈllich and Schmidtb-
latter ®nding suggests that PAP might enhance perfor- leicher 1995; Radcli€e and Radcli€e 1996; Young et al.
mance (i.e. increase peak velocity and power) with loads 1998) tested for the presence of PAP following the
between the extremes represented by Po and Vmax. By conditioning activities, but it was probably present.
increasing the acceleration and hence the velocity at- The discrepancy between the present and previous
tained with a given load, the increased rate of force studies, regarding the bene®t of conditioning activities,
development induced by PAP would raise the portion of is probably related to the conditioning activity's dual
the load±velocity relation between Po and Vmax upward e€ects of inducing both PAP and fatigue (Krarup 1981).
and to the right. The present study tested the hypothesis In the present study there were two indications that the
of a PAP-induced upward and rightward shift of the 10-s MVC induced fatigue as well as PAP. First, torque
load±velocity relation. The results failed to support the declined on average by 16.3% (P < 0.001) during the
hypothesis; when a series of four loads between Po and 10-s MVC. Secondly, the peak velocity of the ®rst knee
Vmax was tested, there were no increases in peak velocity extension was depressed following the MVC. In selecting
or power during periods of twitch PAP, and thus there a strategy to take advantage of PAP, there is the di-
were no signi®cant changes in the load±velocity relation. lemma that a brief interval between the conditioning
The loads selected (15±60% MVC) were considered to contraction inducing PAP and performance maximizes

Table 1 Peak power, work attained at the time of peak power, and time to peak power of the dynamic knee extensions (``kicks''). Values
(collapsed across the four loads) are mean (SE)

Control trials PAP trials

Kick no. 1 Kick no. 2 Kick no. 1 Kick no. 2

Mean SE Mean SE Mean SE Mean SE

Peak power (W) 456.9 23.5 467.6 22.9 425.7 19.5 444.5* 20.8
Work to peak power (J) 42.7 2.7 46.4* 2.8 42.7 2.4 43.5 2.3
Time to peak power (ms) 195 7 201 6 198 7 200 9

*P < 0.05 for di€erence between kicks


529

both the PAP and fatigue e€ects. A longer recovery in- In conclusion, the present study has shown that a
terval reduces both fatigue and PAP. Compared to our 10-s MVC, known to cause a high degree of PAP, fails
recovery interval of 15 s, previous studies that showed a to enhance dynamic performance which begins @15 s
positive e€ect of conditioning contractions had longer after the MVC, when PAP is still at a high level.
recovery intervals of 3±4 min (Radcli€e and Radcli€e However, we tested a combination of type, intensity
1996; Young et al. 1998). Had we used a recovery in- and duration of conditioning activity, and recovery
terval of 4 min, PAP would probably have diminished to interval between conditioning activity and performance
@15±20% (Hamada et al. 2000; Houston and Grange that is but one of many possible strategies intended to
1990) instead of the @50% found at 15 s in the present enhance performance. The various strategies may en-
study; nevertheless, the longer recovery time may have hance, impair, or have no e€ect on performance, de-
more than compensated for the diminished PAP. pending on the interaction induced between PAP and
The fatigue caused by the 10-s MVC, which either fatigue. A series of experiments will be needed to
impaired dynamic performance or prevented it from identify the best strategy.
being enhanced by PAP, could have been the result of
central or peripheral mechanisms. In a sustained MVC, Acknowledgements John Moroz provided technical assistance. The
maximum sustainable motor unit discharge rates de- study carried the approval of McMaster University's Human
Ethics Committee, and by extension was in accordance with cur-
crease within seconds, and some high-threshold, pre- rent laws in Canada regarding research with human subjects.
sumably fast, motor units cease discharging (Grimby
et al. 1981). Since a high rate of force development
depends on high discharge rates and activation of fast
motor units (Zehr and Sale 1994), dynamic perfor- References
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paired. Further evidence implicating central of the four heads of quadriceps femoris: an electromyographic
mechanisms is that the ability to perform voluntary study. Anat Rec 172: 15±20
contractions with a high rate of force development Butler TM, Siegman MJ, Mooers SU, Barsotti RJ (1983) Myosin
deteriorates rapidly when the rate of force development light chain phosphorylation does not modulate cross-
bridge cycling rate in mouse skeletal muscle. Science 220:
of evoked contractions is not diminished (Miller et al. 1167±1169
1993). Our observed lack of change in AEMG in the Grange RW, Vandenboom R, Houston ME (1993) Physiological
vastus medialis does not indicate the presence of central signi®cance of myosin phosphorylation in skeletal muscle. Can
fatigue, but surface EMG may not be sensitive enough J Appl Physiol 18: 229±242
Grange RW, Cory CR, Vandenboom R, Houston ME (1995)
to re¯ect changes in peak motor unit ®ring rates. Fur- Myosin phosphorylation augments force±displacement and
thermore, altered activation of the other heads of force±velocity relationships of mouse fast muscle. Am J Physiol
quadriceps cannot be ruled out. 269: C713±C724
Establishing a strategy for deriving the greatest ben- Grange RW, Vandenboom R, Xeni J, Houston ME (1998) Pot-
e®t from PAP has two key parts: (1) setting the intensity entiation of in vitro concentric work in mouse fast muscle.
J Appl Physiol 84: 236±243
and duration of the conditioning activity, and (2) setting Grimby L, Hannerz J, Hedman B (1981) The fatigue and voluntary
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fatigue produced, and the second part a€ects the net tions. XVth Congress of the International Society of Biome-
e€ect of PAP decay and recovery from fatigue. Finding chanics, pp 348±349
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experiments in which the two key parts of the strategy Postactivation potentiation, ®bre type, and twitch contraction
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longer recovery interval before the ®rst knee extension, Physiol Pharmacol 68: 908±913
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cause both PAP and fatigue. In the present study, the of rabbit skeletal muscle thick ®laments. Biophys J 71:
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