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Activation of human quadriceps femoris during isometric, concentric, and

eccentric contractions

Article  in  Journal of Applied Physiology · December 2001

DOI: 10.1152/jappl.2001.91.6.2628 · Source: PubMed

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J Appl Physiol
91: 2628–2634, 2001.

Activation of human quadriceps femoris during

isometric, concentric, and eccentric contractions

NICOLAS BABAULT, MICHEL POUSSON, YVES BALLAY, AND JACQUES VAN HOECKE
Groupe Analyse du Mouvement, Unité de Formation et de Recherche Sciences et Techniques des
Activités Physiques et Sportives, Université de Bourgogne, BP 27877, 21078 Dijon Cedex, France
Received 25 January 2001; accepted in final form 14 August 2001

Babault, Nicolas, Michel Pousson, Yves Ballay, and during eccentric (13, 35, 36) or slow concentric contrac-
Jacques Van Hoecke. Activation of human quadriceps fem- tions (10, 29, 38) in humans.
oris during isometric, concentric, and eccentric contractions. The degree of voluntary activation of the motoneuron
J Appl Physiol 91: 2628–2634, 2001.—Maximal and sub- pool can be assessed by using the twitch interpolation
maximal activation level of the right knee-extensor muscle
technique (26). This technique has been frequently
group were studied during isometric and slow isokinetic
muscular contractions in eight male subjects. The activation applied to isometric contractions of different human
level was quantified by means of the twitch interpolation muscles such as the adductor pollicis (26), quadriceps
technique. A single electrical impulse was delivered, what- femoris (9, 32), elbow flexors (11, 12, 16), and plantar
ever the contraction mode, on the femoral nerve at a constant flexors (7). However, voluntary activation during dy-
50° knee flexion (0° ⫽ full extension). Concentric, eccentric namic contractions has received very little attention.
(both at 20°/s velocity), and isometric voluntary activation To our knowledge, few studies have investigated the
levels were then calculated. The mean activation levels dur- extent of activation during eccentric or concentric con-
ing maximal eccentric and maximal concentric contractions tractions (4, 5, 15, 22, 28, 36). During maximal eccen-
were 88.3 and 89.7%, respectively, and were significantly tric contractions, voluntary activation level (AL) was
lower (P ⬍ 0.05) with respect to maximal isometric contrac- generally assessed by using tetanical stimulations su-
tions (95.2%). The relationship between voluntary activation
levels and submaximal torques was linearly fitted (P ⬍ 0.01):
perimposed on the voluntary contractions. In these
comparison of slopes indicated lower activation levels during conditions, the torque recorded during the superim-
submaximal eccentric compared with isometric or concentric posed contractions was higher compared with the max-
contractions. It is concluded that reduced neural drive is imal voluntary torque alone (4, 37). This result would,
present during 20°/s maximal concentric and both maximal therefore, suggest that voluntary activation was re-
and submaximal eccentric contractions. These results indi- duced during maximal eccentric contractions. During
cate a voluntary activation dependency on both tension levels slow concentric contractions, the reduced neural drive,
and type of muscular actions in the human knee-extensor hypothesized by numerous authors (10, 13, 29, 38),
muscle group. remains, however, controversial. Indeed, voluntary ac-
twitch interpolation; isokinetic contractions; torque level; tivation assessed by the twitch interpolation technique
neural drive; knee extension during concentric contractions has not confirmed this
hypothesis. Some authors (15, 28) have concluded that
maximal concentric AL was similar to the maximal
IT IS WELL DOCUMENTED THAT voluntary torque, produced isometric level either at slow (20°/s) or at faster angu-
by human muscles, is lower during maximal concentric lar velocities (150 and 300°/s).
contractions and equal or higher, depending on train- The twitch interpolation technique requires the de-
ing status (4), during maximal eccentric contractions livery of one or more supramaximal electrical stimuli
compared with isometric contractions. This muscle be- to the appropriate motor nerve (or motor point) during
havior is explained by mechanical factors and myoelec- an attempted maximal voluntary contraction (MVC).
trical activity. However, marked differences found be- This technique assumes that the motor units that are
tween the force-velocity relationship in human and in not recruited or not discharging at their maximal fre-
isolated animal muscles, especially when eccentric or quency (incomplete activation) should yield a detect-
slow concentric contractions are performed, suggest able force increment as a consequence of the stimula-
that different activation patterns, depending on both tion of their axons. To improve the signal-to-noise ratio
contraction type and angular velocity, would affect the (24) and/or reduce the measurements variability (34),
shape of this relationship. Thus it has been postulated some studies have used multiple rather than single
that changes in torque could result from a reduction in stimuli. However, the enhanced reliability of multiple
activation, restricting maximal muscle tension, either pulses already observed (21, 24, 34) remains controver-

Address for reprint requests and other correspondence: N. The costs of publication of this article were defrayed in part by the
Babault, Groupe Analyse du Mouvement, UFR STAPS, Faculté des payment of page charges. The article must therefore be hereby
Sciences du Sport , Université de Bourgogne, BP 27877 - 21078 Dijon marked ‘‘advertisement’’ in accordance with 18 U.S.C. Section 1734
Cedex, France (E-mail: [email protected]). solely to indicate this fact.

2628 8750-7587/01 $5.00 Copyright © 2001 the American Physiological Society http://www.jap.org
 VOLUNTARY ACTIVATION AND CONTRACTION TYPE 2629

sial, as several authors have observed similar results ited every 5 s for a given stimulus intensity) were reached.
using single stimuli, doublets, or short trains of stimuli This maximal intensity was then applied under all experi-
(3, 6). Despite some variability from test to test in the mental conditions (i.e., contraction mode and torque level).
voluntary AL (2, 3, 6), Behm et al. (6) have concluded To ensure that stimulations were maximally delivered at all
times, the resultant vastus lateralis M waves were con-
that the technique, associated with a single stimulus to
trolled. No M-wave changes were measured when maximal
the motor nerve, was reliable in both the plantar flex- or submaximal isometric and dynamic contractions were
ors and quadriceps femoris. In addition, because spinal compared. During isokinetic movements, impulses were trig-
reflexes and motor axon antidromic responses are gered at a constant 50° knee-flexion angle, corresponding to
known to influence the interpolated twitch (20), single the middle of the range of motion. Although the maximal
stimuli seem preferable to minimize the distortions torque output did not occur at the same knee angle under
induced by multiple stimuli. However, single stimuli eccentric and concentric conditions, the amount of activation
superimposed during concentric contractions have was calculated at the same 50° knee flexion. According to
been used in only one study (15), and under no circum- Harridge and White (17), for angular velocities ⬍60°/s, mus-
stance has this stimulation mode been used to compare cles would have quite enough time to be as maximally acti-
vated as possible.
voluntary activation during isometric, concentric, and
Voluntary activation. Voluntary AL was calculated by ex-
eccentric contractions. pressing every increment in torque evoked during the con-
The main purpose of this study was to extend the traction (size of superimposed twitch) as a fraction of the
twitch interpolation technique (single-stimulus mode) amplitude of the response evoked by the same stimulus in the
to investigate whether the level of voluntary activation relaxed muscle (twitch size evoked at rest). The AL was then
of the human quadriceps femoris muscle was depen- quantified as the percentage AL ⫽ 关1 ⫺ 共size of superimposed
dent on the contractile conditions during both isomet- twitchⲐtwitch size evoked at rest兲兴 ⫻ 100.
ric and 20°/s isokinetic contractions (concentric and Twitch evoked at rest. The twitch evoked at rest, to be
eccentric contractions). We first determined whether compared with that superimposed during dynamic contrac-
the twitch interpolation technique could give evidence tions, was elicited while the muscle was passively shortened
or lengthened for concentric and eccentric movements, re-
of voluntary activation reduction during slow maximal
spectively. All twitches produced on relaxed muscle were
concentric and eccentric contractions. Second, we corrected by the resistive torque related to weight of the leg
tested whether voluntary activation differed among and lever arm and the passive viscoelastic forces. To find the
the different contraction modes during submaximal appropriate corrections, the resistive torque was measured
voluntary contractions. as the leg was moved passively at the same constant angular
velocity (20°/s) without any stimulation. Muscular relaxation
MATERIALS AND METHODS was checked by the absence of the vastus lateralis electro-
myographic (EMG) signal. A series of four twitches was
Subjects. Eight physical education students, with a me- evoked for each velocity (0, ⫹20, and ⫺20°/s) to obtain a
dian age of 25 yr (range: 21.3–33.1 yr), a median body mass mean evoked response. About 10 s separated each twitch.
of 78 kg (range: 71–95 kg), and a median height of 178 cm Superimposed twitch. During isokinetic contractions, mus-
(range: 171–186 cm), participated in this investigation. All cle torque was changing over the entire range of motion as a
were volunteers and gave written, informed consent to the consequence of length, velocity, lever arm changes, and de-
experiment. gree of activation. In fact, around the angle of stimulation,
Torque measurements. Measurements were made for all voluntary concentric torque decreased, whereas torque in-
subjects on the right leg extensors using a Biodex isokinetic creased during the eccentric contractions. For this reason,
dynamometer (Biodex, Shirley, NY), which allowed recording the superimposed twitch torque was calculated by subtract-
of instantaneous muscle torque at various preset constant ing the poststimulus twitch torque from the torque that
angular velocities. Subjects were seated with the trunk ver- would have occurred in the absence of stimulation for the
tical and thigh fixed at a 90° hip angle. Velcro straps were same angular position. Because the torque-time curve was
applied tightly across the thorax, pelvis, and distal thigh. linear before stimulation (Fig. 1A), the torque without stim-
Voluntary torque was measured at the angular position of ulation was estimated by linear extrapolation of the slope of
50° knee flexion (0° ⫽ full extension), during both isometric the prestimulus voluntary torque beyond the point of stimu-
and isokinetic contractions. Isokinetic contractions were per- lation. Such estimation of the interpolated twitch has already
formed at slow angular velocity (20°/s) during both concentric been used during isometric contractions (2) and concentric
and eccentric contractions. The range of motion was 60°, from contractions (15). The period used for the extrapolation was
20 to 80° knee flexion. at least 50 ms. The application of the twitch interpolation
Electrical stimulation. A single electrical stimulus was technique during maximal isometric contraction and the es-
applied percutaneously to the femoral nerve. During both timation of the superimposed eccentric twitch are presented
maximal and submaximal contractions, a ball probe cathode in Fig. 1.
(⬃10 mm diameter) was pressed in the femoral triangle, 3–5 Experimental procedure. Maximal isometric contractions
cm below the inguinal ligament. The anode (self-adhesive (5 s) were first performed at 20, 50, and 80° knee flexion to
stimulation electrode, 10 ⫻ 5 cm) was located in the gluteal determine the MVC. Two contractions were made at each
fold. Single square-wave stimulus was used with a 1-ms angular position. Second, subjects were asked to perform
duration, 400-V maximal voltage, and intensity ranging from three different tasks. 1) The first task was maximal and
50 to 180 mA (Digitimer DS7, Hertfordshire, UK). The max- submaximal isometric knee extension at 50° angular posi-
imal intensity of stimulation was set by progressively in- tion. For submaximal trials, subjects were instructed to keep
creasing the stimulus intensity until the maximal isometric a specified torque level constant (range from 20 to 100% of
twitch torque and concomitant vastus lateralis M-wave am- the MVC). The stimulation was administered ⬃3.5 s after the
plitude (determined by averaging four twitch responses elic- start of the isometric contraction. 2) The second task was
J Appl Physiol • VOL 91 • DECEMBER 2001 • www.jap.org
2630 VOLUNTARY ACTIVATION AND CONTRACTION TYPE

EMG recording. EMG was recorded by using two pairs of

silver chloride surface electrodes applied over the belly of the
vastus lateralis and biceps femoris. Low impedance of the
skin-electrode interface was obtained (impedance ⬍2 k⍀) by
abrading the skin. The interelectrode distance was 2 cm
(center to center). The reference electrode was attached to
the opposite wrist. Myoelectrical signals were amplified with
a bandwidth frequency ranging from 1.5 Hz to 2.0 kHz
(common mode rejection ratio ⫽ 90 dB; impedance ⫽ 100
M⍀; gain ⫽ 1,000). EMG as well as torque signals were
digitalized on-line at a sampling frequency of 1,000 Hz.
During isokinetic contractions, vastus lateralis activity was
evaluated and directly visualized by means of the RMS am-
plitude, calculated over subsequent 300-ms period intervals.
Vastus lateralis was chosen as it has been suggested that
surface EMG recordings from this muscle are more reliable
in predicting force output than are those from vastus medi-
alis or rectus femoris muscles (1). The antagonist coactiva-
tion (biceps femoris) was evaluated by calculating the RMS
over a 300-ms period before the superimposed stimulation.
The antagonist RMS was expressed as a fraction of its max-
imal agonist activity.
Statistical analysis. Descriptive statistics presented in RE-
SULTS or in Figs. 2 and 3 are mean values ⫾ SE. Twitches
evoked at rest, torque, AL, and the amount of coactivation
obtained during the three MVC trials were averaged for
Fig. 1. A: effect of single, superimposed electrical stimulus (up ar- statistical analysis. Because homoscedasticity and normal
row) on an eccentric contraction performed at ⬃80% maximal volun- distribution of the data were verified by means of Levene’s
tary contraction (MVC). Top trace, torque-time curve; bottom trace, and Kolmogorov-Smirnov tests, parametric statistics were
knee angle. B: estimation of the superimposed twitch response (dou-
ble arrow) during the same submaximal voluntary eccentric contrac-
used. Thus concentric, isometric, and eccentric conditions
tion. Superimposed twitch torque was calculated by subtracting the were compared by using a one-way ANOVA; subsequent least
poststimulus torque with the torque that would have occurred with- significant difference test was performed for post hoc analy-
out any stimulation (linear extrapolation of the voluntary torque sis to determine differences across velocities. Similarly, av-
preceding the stimulation) for the same angular position. Up arrow, erage resting twitch torque and average maximal voluntary
the time of delivery of the stimulation. C: superimposed twitch torque, both normalized with their maximal isometric torque,
torque after the subtracting procedure. The twitch size was calcu- were compared using a one-way ANOVA. Standard paramet-
lated as the peak torque of the reconstituted twitch. Up arrow, the
ric linear regression analysis was used to compare the degree
delivery of the stimulation; double arrow, the estimation of the
superimposed twitch response. of association between voluntary activation and voluntary
torque for all data and for each subject. Slopes of the linear
maximal and submaximal concentric knee extension; 3) the regressions for the three contraction modes (eccentric, iso-
third task was maximal and submaximal eccentric knee metric, and concentric) were tested by an ANOVA. P ⬍ 0.05
extension. Contrary to isometric contractions, during maxi- was taken as the level of statistical significance for all pro-
mal and submaximal dynamic contractions, torque could not cedures.
be maintained constant. Consequently, to be under constant
conditions, a vastus lateralis EMG activity control was made. RESULTS
At the onset of the dynamic contraction (20 or 80° knee angle
for the eccentric and concentric contractions, respectively), Maximal voluntary torque and twitch torque evoked
subjects were asked to develop a required torque level during at rest. Knee-extensor maximal voluntary or evoked
a short isometric preactivation contraction (2 s). Then they twitch torque was measured during isometric (0°/s),
were asked to modulate their level of effort during the full concentric (⫹20°/s), and eccentric (⫺20°/s) contrac-
range of motion to maintain constant the root-mean-square tions. Maximal voluntary concentric torque, measured
(RMS) level (range: ⫾5%) fixed during the preactivation
contraction. The RMS level was under visual control via a at 50° constant angular position (236.1 ⫾ 8.1 N 䡠 m),
computer monitor in front of them. Different contraction was significantly smaller (P ⬍ 0.01) than isometric and
intensities were obtained by performing preactivation con- eccentric torques, which were 310.3 ⫾ 11.6 and 313.4 ⫾
tractions ranging from 20 to 100% of the MVC. Subjects were 12.8 N 䡠 m, respectively. No significant differences were
familiarized with this procedure during a standardized observed between isometric and eccentric maximal vol-
warm-up session. Because subjects succeeded in the task untary torque values. The mean twitch values evoked
with remarkable ease, during each experiment only two to on relaxed muscles for eccentric, isometric, and concen-
three trials were excluded from the analysis. tric movements were 42.4 ⫾ 4.7, 38.5 ⫾ 4.7, and 38.3 ⫾
All maximal or submaximal contractions were conducted
with superimposed twitch stimulation (Fig. 1A). Contraction
3.8 N 䡠 m, respectively. No differences were observed
modes and levels were randomly presented. Three trials were among the three contraction modes. When twitch
made for each torque level (20, 40, 60, 70, 80, 90, and 100% of torque and maximal voluntary torque were expressed
the MVC). A 2-min rest period was permitted between each as a fraction of their respective isometric torque, rela-
three contractions to avoid the effects of fatigue. tive twitch torque was significantly greater than the
J Appl Physiol • VOL 91 • DECEMBER 2001 • www.jap.org
 VOLUNTARY ACTIVATION AND CONTRACTION TYPE 2631

Fig. 4. Torque-time profile of the superimposed twitch response

during isometric (left), concentric (center), and eccentric contractions
(right). The twitches are presented from relaxed muscle to MVCs (top
to bottom traces). Control and superimposed twitches have been
aligned so that time of stimulus (arrows) and force offset of the
superimposed twitch are coincident.

Fig. 2. Normalized torque of the right knee-extensor muscle group

during MVC or when contractions were evoked by means of a single The amount of coactivity (biceps femoris) was also
twitch (TW) during concentric (⫹20°/s) and eccentric (⫺20°/s) con- measured and averaged 25.1 ⫾ 3.0, 23.0 ⫾ 2.7, and
tractions. Torque is expressed as a fraction of the respective isomet-
ric torque. Values are means ⫾ SE. Significant differences between 26.1 ⫾ 3.5% of its agonist activity for eccentric, isomet-
TW and MVC values at identical velocity: ** P ⬍ 0.01; * P ⬍ 0.05. ric, and concentric MVC, respectively. No significant
differences were recorded among the different angular
velocities.
relative maximal voluntary torque. This was observed Submaximal voluntary torque and AL. The extent of
for both eccentric and concentric conditions (Fig. 2). activation was evaluated for different levels of effort.
AL during maximal contractions. During MVC, an As expected and whatever the contraction mode, the
electrical twitch was superimposed to measure the superimposed twitch torque decreased with increasing
amount of activation. Mean voluntary ALs obtained voluntary torque (Fig. 4). Consequently, voluntary ac-
during the three contraction modes are presented in tivation increased as a function of torque. Plots of
Fig. 3. The mean voluntary AL during maximal eccen- average AL and average voluntary torque during mus-
tric contractions (88.3 ⫾ 1.9%, range 67.1–99.5%) was cular contractions for one subject are shown in Fig. 5.
significantly lower (P ⬍ 0.01) than the maximal iso- When individual data are considered, a linear fit (P ⬍
metric AL (95.2 ⫾ 1.2%, range 85.6–99.8%). Seven of 0.001) was obtained for each contraction mode. Across
the eight subjects had a maximal eccentric AL lower the subjects, correlation coefficients ranged from 0.69
than the isometric level. AL during maximal concentric to 0.95 for eccentric contractions, 0.86 to 0.95 for iso-
contractions (89.7 ⫾ 1.4%, range 72.9–96.9%) was also metric contractions, and 0.76 to 0.96 for concentric
significantly lower compared with maximal isometric contractions. On the average, the slopes of the AL-
contractions (P ⬍ 0.05). During maximal concentric voluntary torque relationships were 0.72 ⫾ 0.44 for the
contractions, seven of the eight subjects exhibited a eccentric condition and were significantly lower than
lower AL compared with isometric contractions. No those obtained for the concentric (1.02 ⫾ 0.05) and
differences were observed between maximal eccentric isometric (0.90 ⫾ 0.05) relationships. Five of the eight
and concentric AL. For each subject, the coefficient of subjects had an eccentric slope lower than the concen-
variation across trials was calculated. Mean coeffi-
cients of variation were 4.1 ⫾ 1.1, 1.7 ⫾ 0.6, and 3.1 ⫾
0.7% for eccentric, isometric, and concentric maximal
voluntary activation, respectively.

Fig. 5. Relationship between activation level and relative torque

Fig. 3. Mean maximal voluntary activation level measured during (normalized with respect to the maximal isometric torque) for 1
maximal eccentric, isometric, and concentric contractions. Maximal subject. Slow concentric (dashed line), isometric (solid line), and slow
voluntary activation (100%) corresponds to no detectable force incre- eccentric (dotted line) relationships are best fitted by linear regres-
ment consecutive to the superimposed twitch. Values are means ⫾ sions. Equations for concentric, isometric, and eccentric contractions
SE. Activation during maximal eccentric and concentric contractions are y ⫽ 1.07x ⫺ 0.81 (r ⫽ 0.96), y ⫽ 0.94x ⫹ 5.15 (r ⫽ 0.95), and y ⫽
is significantly lower than maximal isometric activation: ** P ⬍ 0.01; 0.73x ⫹ 14.11 (r ⫽ 0.95), respectively. Eccentric slope is significantly
* P ⬍ 0.05. lower than isometric and concentric slopes (P ⬍ 0.001).

J Appl Physiol • VOL 91 • DECEMBER 2001 • www.jap.org

2632 VOLUNTARY ACTIVATION AND CONTRACTION TYPE

tric slope. Four of them had an eccentric slope lower stimulation and observed similar activation for the
than the isometric slope. No significant difference was different angular velocities. The fact that nonisokinetic
observed between isometric and concentric slopes. contractions were considered and that the authors did
not ensure constant EMG activity during the full range
DISCUSSION of motion might in part explain the differences from
our findings. Another reason for the different results
In this study, the twitch interpolation technique, could be that their study was performed on the elbow
associated with single stimuli, was adapted for appli- flexors. Indeed, in another study (28) performed on the
cation during dynamic contractions. The AL was 88.3 quadriceps muscle group under isokinetic conditions
and 89.7% for maximal eccentric and concentric con- (20 and 150°/s), results tended toward our findings. A
tractions, respectively, and 95.2% for maximal isomet- reduction of activation was effectively observed in more
ric contractions. A majority of studies using this tech- subjects at the slowest velocity than during isometric
nique during isometric MVC have concluded that or 150°/s concentric contractions.
human subjects can maximally or near maximally (vol- The conclusion supported by the twitch interpolation
untary activation ⬎95%) activate their muscles (2, 3, 7, technique during maximal voluntary efforts appears to
11, 15, 16, 26, 28, 39). Under isometric conditions, our be confirmed by the comparison between the relative
results would confirm these findings. However, as for changes in maximal voluntary torque or twitch torque
other studies, AL estimation implies some variability. evoked on resting muscles (see Fig. 2). Indeed, identi-
The range of maximal activation values obtained here cal maximal voluntary and evoked twitch torque (both
was similar to previous results obtained during isomet- normalized with respect to isometric torque) would
ric (2) and concentric contractions (15). The mean co- have revealed similar activation of the knee-extensor
efficient of variation was 1.7% for isometric MVC and muscle group by the central nervous system whatever
was identical to that obtained by Allen et al. (2). These the angular velocities. However, our results indicate
authors concluded that voluntary activation, measured that maximal voluntary torque is not dependent only
during maximal voluntary efforts, was similar from on the mechanical properties and suggest that ALs
session to session. Nevertheless, because variability would restrain the maximal voluntary concentric and
seems to increase during dynamic contractions (see eccentric torque. Thus maximal ALs depend on the
Fig. 3B in Ref. 15), the resolution of the technique types of muscle actions performed. Neural drive inhi-
would decrease slightly. Thus multiple measures are bition (incomplete motor unit activation) seems to be
required to enhance the technique reliability. active to limit high tension levels during 20°/s maximal
Results of the twitch interpolation technique indi- concentric and eccentric voluntary efforts. Initially, it
cate that, during maximal eccentric and concentric has been suggested (e.g., Ref. 4) that the neural drive
contractions, voluntary activation was lower compared reduction would contribute to the limitation of maxi-
with maximal isometric contractions. Thus application mal tension and the preservation of the musculoskele-
of single stimuli for assessment of the voluntary acti- tal integrity (patella dislocation, tendon rupturing,
vation during eccentric contractions corroborates the etc.). According to this hypothesis, maximal concentric
reduced neural drive previously hypothesized with dif- contractions should have induced higher ALs than
ferent approaches such as EMG (4, 36) or artificial those measured in our study, compared with isometric
activation evoked on relaxed muscle or superimposed ones. Nevertheless, it is possible to explain this appar-
to the voluntary contraction (4, 13, 30, 37). During ent contradiction by the fact that the “tension-limiting
concentric contractions, as already suggested without mechanism” would be mostly dependent on the type of
employment of superimposed stimulation (18, 29, 30, muscular actions (i.e., isometric or dynamic) rather
36), the same effect was also found. However, previous than on the maximal torque output. This mechanism
findings using twitch interpolation during concentric would be more efficient during maximal dynamic than
contractions (15, 28) did not confirm this result. Tetani- isometric contractions, as a higher AL was obtained
cal stimulation superimposed to maximal concentric during isometric compared with the concentric and
contraction on the knee extensors did not indicate eccentric contractions. Moreover, regarding the magni-
reduction of AL during slow concentric contractions tude of change of voluntary activation and maximal
(37). Because it has been demonstrated that training voluntary torque recorded during maximal eccentric
status did influence the reduction of neural drive (4), and concentric efforts, activation may not be the only
the differences with our results could be attributed to mechanism responsible for torque limitations. Indeed,
the population tested. Indeed, our experimental group, the similar concentric and eccentric ALs contrast with
including normally active subjects, was less accus- the considerable variations between relative voluntary
tomed to perform maximal efforts than the subjects and resting twitch torque, particularly observed during
tested by Westing et al. (37), who were, among others, concentric contractions. One must, therefore, keep in
wrestlers and weightlifters. Furthermore, the velocity mind that complementary mechanisms, dependent on
tested was threefold greater than those considered in the type of contraction, should contribute to these dis-
the present investigation and may contribute to the crepancies. For example, the resultant concentric
different results. Gandevia et al. (15) calculated max- torque is more affected (i.e., reduced) by the antagonist
imal voluntary activation during isometric and concen- coactivation than during eccentric actions (23). Indeed,
tric contractions (until 300°/s) using single electrical although antagonist EMG activity is similar whatever
J Appl Physiol • VOL 91 • DECEMBER 2001 • www.jap.org
 VOLUNTARY ACTIVATION AND CONTRACTION TYPE 2633

the angular velocity, during concentric knee exten- tractions and only maximal concentric contractions at
sions, antagonist muscles act eccentrically, thus pro- a 20°/s constant angular velocity. The pattern of vol-
ducing a greater torque in opposition to the agonist untary activation of the human knee-extensor muscle
action. Similarly, it can be postulated that maximal group would, therefore, depend on the type of muscular
voluntary concentric contractions could induce more actions performed. Neural drive inhibition can be one
viscoelastic force loss than gain obtained during eccen- possible cause of the reduced activation during maxi-
tric contractions (25). mal dynamic contractions.
During submaximal eccentric, isometric, and concen-
The authors gratefully acknowledge Nicola A. Maffiuletti for help-
tric voluntary contractions, the AL increased linearly ing with some of the experiments.
with increasing tension level. The significantly weaker
slope of the AL-torque relationship obtained during REFERENCES
eccentric contractions indicates that the voluntary ac-
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It, therefore, supports the fact that, during eccentric measurements of muscle length and voluntary activation using
contractions, less neural drive (number or discharge twitch interpolation. Muscle Nerve 18: 593–600, 1995.
3. Allen GM, McKenzie DK, and Gandevia SC. Twitch interpo-
rate of the motor unit) is needed to develop a given lation of the elbow flexor muscles at high forces. Muscle Nerve 21:
submaximal muscle tension (8). Identical findings have 318–328, 1998.
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a different technique, i.e., comparison between sub- Pousson M, and Van Hoecke J. Co-activation and tension-
maximal electrical activation and maximal and sub- regulating phenomena during isokinetic knee extension in sed-
entary and highly skilled humans. Eur J Appl Physiol 73: 149–
maximal voluntary efforts (30). Nevertheless, accord- 156, 1996.
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