Scand J Med Sci Sports 2007: 17: 547–555
Printed in Singapore . All rights reserved
DOI: 10.1111/j.1600-0838.2006.00614.x
Exhaustive stretch-shortening cycle exercise: no contralateral effects
on muscle activity in maximal motor performances
S. C. Regueme*, J. Barthèlemy, C. Nicol
Department of Biology of Physical Activity, University of the Mediterranean, Marseilles, France
Corresponding author: Caroline Nicol, Department of Biology of Physical Activity, University of the Mediterranean, 163,
Avenue de Luminy, CP 910, UPRES EA 3285, Marseilles, France. Tel: 133 491 828 415, Fax: 133 491 828 409,
E-mail: caroline.nicol@univmed.fr
Accepted for publication 5 October 2006
Minor cross-over effects of unilateral muscle fatigue have
been reported after isometric exercises. The present study
re-examined this possibility after an exhaustive stretch-
shortening cycle (SSC)-type exercise. Twenty-five subjects
performed on a sledge apparatus a unilateral exhaustive
rebound exercise involving mostly the triceps surae muscle
group. Ipsilateral vs contralateral fatigue effects were
compared in uni- and bilateral tests that included a maximal
isometric voluntary contraction (MVC) and a series of 10
maximal drop jumps (DJ). These tests were carried out just
before and after (POST) the exhaustive SSC exercise, and
The general existence of ‘‘contralateral irradiation’’
of unilateral activation patterns has long been known
(e.g. Sherrington, 1906). Since then, growing evi-
dence exists that contralateral improvements in mo-
tor skill and strength rely on ‘‘cross-over effects’’ of
unilateral motor and sensory activity (e.g. Hortobá-
gyi et al., 2003). As muscle fatigue is typically
associated with motor and/or sensory changes,
cross-over effects have been examined by some stu-
dies, but mostly after maximal isometric contractions
(Zijdewind et al., 1998; Todd et al., 2003; Rattey
et al., 2005). Their results have revealed minor cross-
over effects on the voluntary activation of upper limb
muscles (Rosenbaum, 1977; Zijdewind et al., 1998;
Todd et al., 2003). According to Rattey et al. (2005),
the major implication of lower limb muscles in
locomotion and balance should favor the occurrence
of contralateral effects. Among other forms of fati-
guing exercises, natural forms of stretch-shortening
cycle (SSC)-type exercises such as running and jump-
ing (which combined eccentric and concentric muscle
actions) are typically associated with large acute and
delayed decrements in force production and volun-
*Present address: Department of Motor Function, Interactions and
Performance, University of Nantes, Nantes Atlantic Universities,
Nantes, France.
Copyright & 2007 The Authors
Journal compilation & 2007 Blackwell Munksgaard
were repeated 2 days later (D2), at the expected time of
major inflammation and pain. The exercised (fatigued) leg
analysis revealed significant declines in MVC and DJ
performances at POST and D2, the latter ones being
associated with significant decreases in voluntary muscle
activity. In contrast, no significant change was found for the
non-fatigued leg. These results do not support the existence
of cross-over effects after exhaustive SSC exercise, at least
when tested in maximal static and dynamic unilateral motor
tasks.
tary activity in maximal isometric tests (for a review,
see Nicol et al., 2006). Clear reductions in muscle
activity have also been observed in maximal SSC
testing tasks, both during the centrally programmed
preactivation phase and the subsequent braking
phase (Avela & Komi, 1998). Simultaneous reduc-
tion in the short latency stretch–reflex response (M1)
suggests that the observed EMG changes in the SSC
testing tasks are both central and reflex in nature
(Avela et al., 1999). These overall observations do
suggest existence of neural attempts of protection of
the fatigued muscle under maximal testing condi-
tions. When dealing with the hypothesis of potential
cross-over effects, the delayed recovery period (start-
ing 1–2 days post-exercise) is of particular interest.
Lasting for a few days, this slow recovery period is
usually associated with muscle pain, and attributed
to the natural time course of inflammatory and/or
remodelling processes related to the exercise-induced
muscle damage (for a review, see Nicol & Komi,
2003). Muscle inflammation and pain have repeat-
edly been shown to lead to contralateral effects in
case of chronic unilateral perturbations (e.g. Woolf,
1984; Johansson & Sojka, 1991; Lund et al., 1991;
Radhakrishnan et al., 2003). The SSC recovery
period is also very likely to result in an increased
547
Regueme et al.
effort perception for a given task that should Materials and methods
favor the occurrence of cross-over effects (Dettmers Participants
et al., 1995; Zijdewind et al., 1998). These overall Twenty-five healthy male subjects (aged 20–25 years) volun-
observations support the hypothesis of cross- teered for the experiment. The study was consistent with the
over effects of the unilateral motor, and sensory Declaration of Helsinki (2002), and was approved by the
changes associated with neural effects during the Local Committee for Human Protection in Biomedical Re-
search (visa No. 04 008). All subjects gave their informed
delayed recovery period of fatiguing SSC exercise. written consent.
Although there is no consensus on the exact neural
mechanisms that produce contralateral effects in case
of unilateral muscle inflammation and pain, small- Experimental design
diameter groups III and IV muscle afferents appear The testing part of the experimental design (Fig. 1) included
as potential candidates. This relies on the fact that maximal isometric voluntary contraction (MVC) and maximal
these afferents are for some of them nociceptive and drop jump (DJ) tests that were both performed unilaterally
(on each leg, separately) as well as on both legs (bilateral test).
mostly polymodal, being sensitive to several chemi-
cal, thermal, and mechanical parameters associated
with muscle damage (Mense, 1977; Kniffki et al.,
Practice session
1978). Additional evidence exists in the literature to - 2 weeks
Maximal Drop Jumps (DJ)
support their contribution to both ipsi- and contral- Submaximal rebounds (70%)
ateral neural effects at the spinal (Djupsjobacka
et al., 1995, Pedersen et al., 1997; Wenngren et al.,
1998) and supraspinal (Iadarola et al., 1998) levels. It Days -15 to -10 Pain scoring
is also worth noting that pain perception is usually
associated with bilateral responses of several struc-
tures of the central nervous system (for reviews, see 15 min warm'up
Verne et al., 2004; Kakigi et al., 2005).
Therefore, this study examined the possibility of PRE TESTS
2-day delayed cross-over effects of an exhaustive SSC MVC + DJ
exercise performed with one lower limb on the (Uni- & Bilaterally)
homologous contralateral voluntary muscle activity. Day 0
The fatiguing protocol consisted of series of unilat- Exhaustive SSC
eral rebounds repeated on a sledge apparatus until fatiguing exercise
exhaustion. This protocol has previously been re- POST TESTS
ported to lead to delayed sensation of muscle pain DJ + MVC
and to large decrements in maximal force and muscle (Uni- & Bilaterally)
activity of the triceps surae muscle group (Regueme Pain scoring
et al., 2005a, b). Ipsilateral vs contralateral SSC
fatigue effects were compared in two testing tasks Pain scoring
Day 1
for both uni- and bilateral testing conditions. The
first (single joint) task consisted of a commonly
used maximal isometric test of voluntary plantarflex- Pain scoring
ion. The second (multi-joint) SSC testing task in- 15 min warm'up
cluded an intermittent series of 10 maximal drop Day 2
D2 TESTS
jumps. The latter SSC testing task was chosen
MVC + DJ
to examine fatigue-induced neural adjustments
(Uni- & Bilaterally)
and more specifically those occurring during the
preactivation phase and the intervention of the
short latency M1 stretch–reflex response. When
such a stressful task is performed with both legs, Days 3 to 8 Pain scoring
two additional and opposite trends of compensatory
Fig. 1. Schematic representation of the experimental design.
contralateral strategies may be expected to occur: Fatigue was induced by an intermittent stretch-shortening
(i) an increased activity of the contralateral leg cycle (SSC) exercise, with series of 30 rebounds repeated
muscles in order to limit the risk of further injury until exhaustion. The fatigue tests included maximal iso-
of the fatigued/damaged muscles or (ii) a decreased metric voluntary contraction (MVC) and maximal drop
contralateral muscle activity in order to either jump (DJ) tests performed unilaterally (on each leg, sepa-
rately) as well as with both legs (bilateral test). These tests
reduce the inter-leg imbalance (McCabe et al., were carried out just before (PRE) and after (POST) the
1972) or to ensure maintained coordination (Rattey exhaustive SSC exercise, and were repeated 2 days later
et al., 2005). (D2).

548

These tests were carried out just before (PRE) and after
(POST) the exhaustive SSC exercise, and were repeated 2
days later (D2). Considering the well-known ‘‘protective effect
of a given eccentric bout on the subsequent ones’’ (Nosaka
et al., 2001), the practice session performed 2 weeks before the
testing protocol was limited to four to six maximal rebounds
on the sledge to determine the maximal hmax rebound height
and to the practice of a rapid isometric rise in force.
Exhaustive SSC protocol
As schematically shown in Fig. 1, a sledge ergometer (Komi et
al., 1987) inclined at 241 from the horizontal was used to
repeat until exhaustion series of 30 unilateral rebounds per-
formed in a sitting position. The target rebound height was set
at 70% of the maximal rebound (hmax) individually measured
in a previous practice session (h70 5 h0170%(hmax–h0) with h0
as the height reached with straight legs on the force plate). To
ensure that the triceps surae muscle group was involved in the
SSC exercise, no heel contact was allowed and an imposed 901
knee flexion angle had to be reached during each contact phase
on the force plate. Based on previous experience (Nicol et al.,
2003; Regueme et al., 2005a), implication of both hip and knee
extensors was limited by setting the resting seat inclination at
1201 and by instructing the subjects to let their knee flex freely
between take-off and the subsequent impact. Visual feedback
was provided to help subjects maintain the target range of
motion. To ensure that our protocol would lead each subject
to exhaustion, inter-series recovery periods of 3 min were used
to delay the progressive development of a metabolic source of
muscle fatigue and avoid having subjects stop the SSC exercise
prematurely (Nicol et al., 2003). Exhaustion was reached when
the subject could not reach the 70% set rebound height for
more than 10 successive rebounds. This protocol has been
shown to result in several hundreds of rebounds, with mostly
delayed fatigue effects (Regueme et al., 2005a, b).
Testing tasks
The ipsi- and contralateral effects of the exhaustive SSC
exercise were examined on the sledge ergometer through
MVC and DJ tests performed successively in the following
order (fatigued leg, non-fatigued leg, and with both legs).
In the MVC test, the sledge seat was bolted to the rails so
that the subjects’ knees were fully extended, with both hip and
ankle joints at 901. The test included three trials of maximal
voluntary plantar flexion performed at a 45-s interval. The
subjects were instructed to develop their maximal plantar
flexion force as fast as possible.
The DJ test included 10 DJs performed at 30-s intervals. In
each DJ, the subject was held by the experimenter before being
released from his optimal dropping height (height leading to
the maximal hmax rebound and individually predetermined
during the practice session). Once dropped, the subjects had to
rebound with maximal effort while respecting the imposed 901
knee flexion angle during the contact phase. When the subject
had reached his maximal rebound height, the sledge seat was
caught by the experimenter and replaced at the initial optimal
dropping height. Individual dropping heights were identical in
all testing sessions.
Muscle soreness
Soreness levels of the quadriceps and triceps surae muscle
groups were rated by each subject every day for 5 days
following the practice session and the fatiguing exercise at
D0. The subjects were instructed to rate their soreness on a six-
Exhaustive SSC exercise and contralateral effects
level scale – from no pain (0) to extremely sore muscle (6) –
under three testing conditions: palpation, contraction, and
passive stretch.
Data recordings and analysis
The analogue signals that were recorded simultaneously in-
cluded the normal component of the reaction force from the
sledge force plate and the EMG activity recorded from the
soleus (SOL) and the gastrocnemius medialis (GAM) muscles.
The positions of the active surface electrodes (Contrôle Graphi-
que Médical, Brie-Comte-Robert, France), which followed the
recommendations of SENIAM (1999), were marked carefully
on the skin to ensure identical positioning at D2. The EMG
signals were preamplified (
375) close to the electrodes, band-
pass filtered (8–500 Hz), and amplified (
412) in a special
measurement unit (ME3000P, MEGA, Kuoplo, Finaland). All
signals were recorded at 1 kHz sampling frequency and trans-
ferred on line to a microcomputer via an optic fiber cable.
Data analysis was performed using Matlab programs
(MathWorks, version 6.5, Release 13). MVC analysis was
based on the trial that presented the highest peak torque. This
trial was quantified through MVC performance (maximal
voluntary peak force) and mean SOL and GAM muscle
activities during the 500 ms that preceded the peak force.
The DJ trial was quantified through DJ performance (max-
imal rebound height). As illustrated in the results section of
Fig. 4, SOL and GAM muscle activities were rectified and then
integrated over the preactivation (100 ms before the impact),
the short-latency stretch–reflex response, (M1: from 40 to
70 ms after the impact), and the braking phase (from impact
to the lower sledge position) periods. To allow the analysis of
the stretch–reflex response the rectified EMG signals were
filtered using a 75 Hz low-pass Butterworth filter (second
order, Matlab toolbox). Each of the DJ parameters was
calculated separately for each DJ trial before being averaged
for the series of 10 DJ.
Statistics
The muscle soreness rating was analyzed using a two-way
ANOVA analysis [Leg (2)
Days (6: from D0 to
D5)
Conditions (3)] with repeated measures for the two
former factors. The exercise-induced changes in all variables
of the unilateral MVC and DJ tests were analyzed using a two-
way ANOVA analysis [Testing time (PRE, POST, D2)
Leg
(fatigued, non-fatigued)] with repeated measures for both
factors. For the bilateral tests, the same analysis was used
for the EMG data, whereas a one-way ANOVA analysis
(Testing time) with repeated measures was applied to the
performance ones. Newman–Keuls and HSD Tuckey post-
hoc tests were used in case of significant interactions and main
effect, respectively. As the practice session could not be used to
provide reliable reference EMG values, the MVC EMG data
of the PRE test were used instead. For this reason, the
absolute EMG data of the fatigued and non-fatigued leg could
not be compared in the PRE testing condition. In all the other
testing conditions, the SOL and GAM muscle activity values
were individually expressed as percentages of their mean
activity measured in the PRE test of MVC, and the subsequent
ANOVA analyses were performed on the normalized values.
The size of the effect of the SSC fatigue was calculated using
the omega square coefficient (o2) known as the proportion of
variance in the dependent variable, which accounted for the
independent variable (Abdi, 1987) (see equation in appendix).
For each leg, correlation tests using a general linear model
were also performed to examine the possibility of a relation-
549
Regueme et al.
ship between the fatigue-induced changes of the measured
parameters. For all statistical analyses, a P value of 0.05 (a)
was accepted as the level of significance. If a difference was not
statistically significant at the chosen a level, the b risk of an
erroneous conclusion of equivalence was chosen as a P value
lesser than or equal to 0.2.
Results
Exhaustive SSC performance
On average, the subjects performed 14  5 series of
30 rebounds, that is, 432  178 rebounds for a net
working time of 11  5 min.
Muscle soreness
None of the subjects reported any muscle soreness
during the week that followed the practice session
(P40.2). In contrast, the fatiguing protocol did
The post hoc analysis revealed significant inter-leg
difference (Po0.05–0.001) in each testing condition.
Maximal isometric (MVC) tests
The specific analysis of the unilateral MVC perfor-
mance (maximal peak force) revealed a significant
interaction between Testing time and Leg
(F2, 48 5 9.38, Po0.001), explaining 36% (o2) of the
total variance. In the bilateral testing condition, a
significant Testing time effect (F2, 48 5 4.24, Po0.05)
was found that explained 23% of the total variance.
As shown in Fig. 3(a), the post hoc analysis revealed
significant performance decreases at both POST and
UNILATERAL BILATERAL
CONDITION CONDITION

result for each of the 25 subjects in muscle pain (a) MVC Performance
40 40
within the triceps surae muscle group of the fatigued
##
leg. The two-way ANOVA analysis revealed a sig-

Delta % PRE
20 20

Delta % PRE
nificant interaction between Days and Leg
(F5, 120 5 22.6, Po0.001). As shown in Fig. 2, the 0 0
muscle pain sensation developed on the first day
− 20 − 20
post-exercise, peaked on day 2 or 3 in the palpation,
contraction and passive stretch conditions (Po0.05– − 40 − 40
0.001), and disappeared on day five. No significant *** *
difference was observed between each condition (b) Mean SOL activity
(P40.2). Nine subjects reported slight pain in the 40 40
quadriceps muscle group. No muscle pain sensation ##
Delta % PRE

20 20

Delta % PRE
was reported in any of the non-fatigued leg muscles.
0 0

Fatigue leg Non-fatigue leg − 20 − 20

Contraction Contraction
Palpation Palpation − 40 − 40
Passive stretch Passive stretch ** ***
(c) Mean GAM activity
6 Pain scoring-TS muscle group

5 40 40
Subjective scale

4
Delta % PRE

*** 20 20
Delta % PRE

***
***
3
0 0
### ***
2
### ### − 20 −20
1 ###
##
###
− 40 −40
0
D-15 D0 D1 D2 D3 D4 D5 *
to -10
Fig. 3. Maximal isometric voluntary contraction test.
Fig. 2. Pain scoring Group-averaged values (  SD) of Group-averaged values (  SD) of peak force (Performance:
subjective pain [on a six-level scale – from no pain (0) to a), mean SOL (b), and GAM (c) muscle activities in the
extremely sore muscle (6)] of the triceps surae (TS) muscle maximal isometric plantarflexion tests carried out immedi-
group under three testing conditions (palpation, contraction, ately before (PRE) and after (POST) the fatiguing exercise,
and passive stretch). The SOL and gastrocnemius medialis and repeated 2 days later (D2) ( & , fatigued leg; &, non-
(GAM) muscle pain sensations have been rated separately fatigued leg; and , bilateral data). *Po0.05, **Po 0.01,
by the subjects, and then averaged for the clarity of the ***Po0.001, significantly different from the pre-fatigue
figure. ***Po0.001, significantly different from the pre- (PRE) absolute values; ##Po0.01, ###Po0.001, significant
fatigue (PRE) absolute values; ##Po0.01, ###Po0.001, sig- inter-leg difference. SOL, soleus; GAM, gastrocnemius
nificant inter-leg difference. medialis.

550

D2 as compared with the PRE, when performed
unilaterally (upper left panel) with the fatigued leg
(PRE: 1034N; POST: 927N; D2: 821N) as well as
with both legs (PRE: 1515N; POST: 1359N; D2:
1332N) (upper right panel). No significant change
was observed in the unilateral performance of the
non-fatigued leg (PRE: 999N; POST: 1001N; D2:
942N, P40.2). This resulted in a significant inter-leg
difference in MVC performance at D2. No significant
correlation was observed between the fatigued leg
and non-fatigued leg changes (P40.2).
The EMG data analysis examined at first the
possibility of different EMG SOL and GAM muscle
activities during the PRE session between the uni-
and the bilateral testing condition. This analysis did
not reveal any significant difference (P40.2): fati-
gued SOL: 0.26 mV and 0.23 mV; non-fatigued SOL:
0.22 mV and 0.20 mV; fatigued GAM: 0.35 mV and
0.66 mV; non-fatigued GAM: 0.35 mV and 0.62 mV,
respectively. On the other hand, significant interac-
tions were found between the Testing time and Leg,
under both uni- and bilateral conditions: SOL
(F2, 48 5 4.14, o2 5 23% and F2, 48 5 5.56, o2 5 17%,
Po0.01) and GAM (F2, 48 5 7.05, o2 5 13%,
Po0.01 and F2, 48 5 4.82, o2 5 9%, Po0.05), respec-
tively. As illustrated in Fig. 3(b) and (c), the fatigued
leg was characterized by significant decreases in SOL
and GAM voluntary activities in both uni- and
bilateral testing conditions, but at D2 only. In con-
trast, no significant change was observed for the non-
fatigued leg in the unilateral testing condition
(P40.2) and the bilateral ones (Po0.2). Because of
the b risk of potential fatigue effects on the non-
fatigued leg muscle activities in the bilateral testing
condition, one-way ANOVA analyses were per-
formed on the non-fatigued leg EMG data only,
with repeated measures for the Testing time factor
(PRE, D2). When the significance of P value was set
at 0.2, the analyses revealed a significant main Test-
ing time effect for both SOL (F1, 24 5 2.01, P 5 0.16,
o2 5 1.9%) and GAM (F1, 24 5 1.95, P 5 0.17,
o2 5 1.8%).
Maximal drop jump tests
In the DJ tests, the specific analysis of the perfor-
mance (maximal rebound height) revealed a signifi-
cant interaction (Testing time
Leg) in the
unilateral testing condition (F2, 48 5 32.36,
Po0.001), and a significant main Testing time effect
in the bilateral one (F2, 48 5 37.16, Po0.001), ex-
plaining, respectively, 45% and 49% (o2) of the total
variance. In both uni- and bilateral testing condi-
tions, the EMG analysis revealed significant interac-
tions, but for the SOL EMG data only, in the
preactivation phase (F2, 48 5 5.32, Po0.01;
o2 5 26% and F2, 48 5 12.48, Po0.001; o2 5 33%),
Exhaustive SSC exercise and contralateral effects
the stretch–reflex M1 response period (F2, 48 5 3.92;
o2 5 9% and F2, 48 5 3.87; o2 5 14%, Po0.05), and
the braking phase (F2, 48 5 3.75; o2 5 17% and
F2, 48 5 3.90; o2 5 50%, Po0.05). As shown in Fig.
4, this reflected the general trend of significant EMG
decreases observed in the fatigued leg that contrasted
with an absence of significant change in the non-
fatigued one (P40.2). At POST, the DJ performance
was significantly decreased (Po0.001) when per-
formed either with the fatigued leg (  8  7%,
from 40.5 to 35.9 cm) or with both legs (  6  4%,
from 63.1 to 58 cm), but not with the non-fatigued
one (  1  6%, from 38.5 to 37.7 cm). These perfor-
mance decrements were not associated with any
significant change in SOL and GAM muscle activity
(P40.2). As shown in Fig. 4(a) and (b), significant
decreases in DJ performance were also observed at
D2 in the bilateral testing condition (  7%) and in
the unilateral test of the fatigued leg (  13%), but
not in the non-fatigued one (  1%, P40.2). In
contrast to the POST testing session, these uni- and
bilateral reductions in DJ performance were asso-
ciated with clear decreases in SOL EMG activity, and
for the fatigued leg only (P40.2 for the non-fatigued
leg SOL EMG data). Under both uni- and bilateral
testing conditions, the observed decrease in SOL
preactivation was not correlated to the subsequent
decrease in SOL stretch–reflex M1 response (P40.2).
The inter-leg analysis revealed in the unilateral
testing condition (Fig. 4(a), upper right panel) sig-
nificant differences in both DJ performance and SOL
preactivation (a). No inter-leg correlation was found
in any of the recorded changes (P40.2). In the
bilateral testing condition (Fig. 4(b), lower right
panel), significant inter-leg differences were observed
in SOL activation during the preactivation phase (a),
the M1 response (b), and the braking phase (c). No
significant inter-leg correlation was found (a:
P 5 0.18; b: P 5 0.17; c: P 5 0.10). The respective
analysis of the GAM muscle activity did not reveal
any significant fatigue-induced change, and no inter-
leg correlation (P40.2).
Discussion
The present study aimed to examine the possibility
that an exhaustive unilateral SSC exercise would lead
to both ipsi- and contralateral neural effects. As
previously observed (Regueme et al., 2005a), this
protocol did succeed in inducing limited performance
decrements at POST and large ones at D2, as
indicated by the overall o2 statistic values. The D2
testing session corresponded to the peak sensation of
muscle pain (Fig. 2), and was characterized by large
decreases in muscle activity of the exercised (fati-
gued) triceps surae muscle group in both MVC and
551
Regueme et al.
(a) Unilateral DJ Test
Fatigued Leg Performance
SOL 1600 10
Fz
EMG PRE
(N)
D2

D2 (delta % PRE)
(mV)
1200 0
0.4
−10
800

0.2 −20
400 ***
−30 ##
0.0 0
−200 0 200 400 600 800

Non-Fatigued Leg 40 SOL EMG

SOL Fz 1600
EMG (N) ##

D2 (delta % PRE)
(mV) 20
1200
0.4 a b c b c
0
a
800

0.2 −20
400

−40 **
0.0 0 ** ***
−200 0 200 400 600 800
Non-Fatigued Leg
Fatigued Leg
both Legs
(b) Bilateral DJ Test
SOL Fz 2000 Performance
PRE 10
EMG (N)
D2
(mV) 1600
D2 (delta % PRE)

0.4 0
1200

800
0.2 −10

400
***
−20
0.0 0
−200 0 200 400 600 800
SOL EMG
SOL Fz 2000 40
EMG (N) ##
(mV) 1600 #
D2 (delta % PRE)

20 ##
0.4
1200
a b c
0
800 a b c
0.2
400 −20

0.0 0 −40 ***

−200 0 200 400 600 800 *** ***

Fig. 4. Maximal drop jump test. Left panel: D2 vs PRE group mean absolute Fz and SOL EMG recordings from the fatigued
and non-fatigued legs in the unilateral (a) and bilateral (b) DJ tests. Right panel: For each testing condition, the D2 relative
changes (delta % PRE) (  SD) of the maximal rebound performance (hmax–h0), with h0, height reached with straight legs on
the force plate are presented. The D2 relative EMG changes (delta % PRE) are presented successively for the preactivation (a),
stretch–reflex M1 response (b), and the braking phase (c). ( & , fatigued leg; &, non-fatigued leg; and , bilateral data).
**Po0.01, ***Po0.001, significantly different from the pre-fatigue (PRE) normalized EMG values (% MVC PRE). #Po0.05,
##
Po0.01, ###Po0.001, significant inter-leg difference. SOL, soleus; DJ, maximal drop jump; MVC, maximal isometric
voluntary contraction.

DJ tests. These observations are typically associated
with the progressive development of inflammation
and/or regenerative muscle process, which usually
accompanies the delayed recovery phase following
exhaustive SSC exercises (Nicol & Komi, 2003).
When dealing with the possibility of contralateral
fatigue effects, it is worth noting that none of
the measured parameters of the contralateral leg
revealed any significant PRE–POST changes. As
the absence of POST fatigue effect also applies to

552

the activity of the fatigued leg muscles, the possibility
of contralateral effects will not be further investi-
gated at POST, and the discussion will concentrate
on the D2 data. This testing session is of
particular interest as it belongs to the delayed recov-
ery phase and corresponds to the peak of muscle
soreness.
At D2, large and significant neuromuscular im-
pairments were observed for the fatigued leg under
all testing conditions (Figs 3 and 4). In contrast, in
both static and dynamic testing tasks, the contral-
ateral leg did not seem to be affected when tested
unilaterally. However, potential contralateral effects
might be suspected in the bilateral testing condition
as the statistical analysis revealed P values lesser than
0.2. In the bilateral MVC test, however, the o2
coefficient indicated that the SSC effects may be
considered as negligible as they would explain less
than 2% of the total variance (Abdi, 1987). In the
bilateral DJ test, significant reductions were observed
for the fatigued leg in the SOL muscle activity during
the pre-programmed preactivation phase and the
subsequent braking phase. As evidenced by the
statistical o2 coefficient values (from 17% to 50%),
these reductions can be partly attributed to the
exhaustive SSC exercise. These observations are in
line with the findings of Avela and Komi (1998), and
support the existence of neural adjustments on a
supra-spinal level (Melvill Jones & Watt, 1971) to the
exercise-induced contractile failure. The absence of
covariance between the respective decreases in SOL
preactivation and stretch–reflex M1 response
(P40.2) might suggest the influence of additional
neural changes on a spinal level. This refers to the
fact that the present analysis of the M1 response was
based on the global EMG, which included both
central and reflex EMG components. However, de-
spite these ipsilateral neural adjustments to fatigue,
we did not observe any significant change in the
contralateral leg data. Furthermore, similar EMG
changes were observed under the unilateral and
bilateral testing conditions (Figs 3 and 4), with
associated intermediate reductions in the bilateral
MVC and DJ performances as compared with the
unilateral ones. The absence of contralateral fatigue
effects was particularly surprising in the bilateral
DJ test as several studies investigating central lower
limb coupling have showed that perturbations to
one leg invoke a coordinated response of the same
latency in both legs during stance (Dietz et al., 1989)
and locomotion (Berger et al., 1984). In the present
stressful bilateral DJ task, two opposite trends of
compensatory contralateral strategies could have
potentially occurred. In order to limit the risk of
further injury of the fatigued muscles, an increased
activity of the contralateral leg muscles was expected
prior to the impact (preactivation) and during
Exhaustive SSC exercise and contralateral effects
the subsequent braking phase. In contrast, to reduce
the inter-leg imbalance (McCabe et al., 1972) or to
ensure maintained coordination (Rattey et al., 2005),
a decreased contralateral muscle activity could
have been observed during the same time
periods. Although the exercise-induced changes in
the fatigued and non-fatigued legs were not
significantly correlated to each other, the analysis
revealed a b II type error in SOL muscle activity
(Po0.2). In this respect, the observed trend
of increased SOL preactivation in the non-fatigued
leg (Fig. 4(b)) might reveal a central attempt
to compensate for the contractile failure of the
fatigued leg.
Two major but not exclusive reasons may be
considered to explain why we did not observe any
clear contralateral neural change. Firstly, it is sug-
gested that the present SSC fatiguing exercise did not
lead to sufficient inflammation and pain to affect the
contralateral side clearly. This refers to the fact that
in animal studies, acute contralateral effects were not
examined after natural voluntary exercise, but after
local injections of either inflammatory or pain-modu-
lated substances (Djupsjobacka et al., 1995, Pedersen
et al., 1997; Wenngren et al., 1998). Although the
present MVC test revealed large decrements in plan-
tarflexion performance at D2, it should be mentioned
that this test was performed at a short muscle
length relative to the optimal 201 dorsiflexed testing
position. Owing to the expected right shift in the
muscle length–tension relationship after fatiguing
eccentric exercise (Morgan et al., 2002; McHugh &
Tetro, 2003), the observed strength loss might
have been overestimated. Also supporting the im-
portance of the present ispsilateral neuromuscular
defects at D2 (Figs 3 and 4), only half of the SSC
fatigue studies of the literature are reporting persist-
ing drops in both MVC and voluntary activity at that
time (Nicol et al., 2006). Furthermore, none of the
subjects felt any pain sensation after the practice
session whereas all of them reported still clear
pain sensation at D2 (Fig. 2). The first observation
is quite common after this type of SSC practice and
does not support the potential influence of a protec-
tive effect such as the one reported for a few weeks
after either two to six maximal eccentric actions
of the elbow extensors (Nosaka et al., 2001) or 10
eccentric actions of the knee extensors (Brown et al.,
1997). On the other hand, the exercise-induced
sensation of muscle pain at D2 could have favored
the occurrence of neural inhibition (Le Pera et al.,
2001). The occurrence of contralateral effects there-
fore remains questionable in human beings when
using natural forms of locomotion such as exhaustive
SSC exercise.
Secondly, the dynamic DJ test was used as this
task requires synchronized muscle activation to
553
Regueme et al.
optimize the rebound performance. This dynamic test
does not appear, however, to be more appropriate
than the maximal isometric test to reveal contralateral
effects. In the present DJ test, the subjects rebounded
in a well-secured sitting position along the rails of the
sledge ergometer. This did not require from the
subjects the high bipedal balance control that char-
acterizes standing and locomotion tasks, in which
contralateral compensatory changes in muscle activ-
ity have been reported to maintain balance and
stabilization of the body (Dietz & Berger, 1984). In
addition, each drop jump rebound lasted only 0.6 s.
This duration might have been, as in the isometric test
condition (3 s), too short to allow for contralateral
changes to take place. As suggested by Zijdewind and
Kernell (2001) and recently confirmed by Rattey et al.
(2005), it seems that ‘‘cross-over’’ of central fatigue is
more likely to occur in moderate but sustained con-
tractions than in maximal but brief motor tasks.
Supporting this hypothesis, a positive correlation
between the fatigued and the non-fatigued leg changes
in EMG activity has recently been reported (Regueme
et al., 2005b) at D2 when the testing task consisted of
a continuous series of 15 sub-maximal rebounds on
the sledge apparatus. In this specific test, the subjects
were instructed to reach precisely the upper and lower
rebound targets (a 70% rebound height and a 901 of
knee flexion angle during the contact phase) via visual
feedback. This precise rebound task, which required
high visual attention and effort levels for at least 15 s,
might have favored the occurrence of contralateral
neural adjustments.
To summarize, exhaustive SSC exercise leading for
the exercised leg muscles to clear pain sensation and
functional defects for a few days does not seem to
affect contralateral muscle activity when tested uni-
laterally in either maximal static or dynamic inter-
mittent tasks. However, as the absence of
contralateral effects is not so clear when tested in
tasks involving both legs, it should not be generalized
to other (upper limb) muscle groups and motor tasks.
It is suggested that ‘‘close loop motor tasks’’ (requir-
ing the use of sensory feedback) associated with
muscle pain perception might be more appropriate
to reveal contralateral neural changes.
Perspectives
According to Rattey et al. (2005), anticipatory regula-
tion exists to avoid cellular damage during fatiguing
muscular work that would reduce in a coordinated
manner the neural drive to the fatigued and non-
fatigued lower limbs. In natural forms of locomotion,
this may be considered of major importance to limit
potential risks of either falls or injuries. One of the
challenges for the future will be to determine whether
locomotor tasks requiring both anticipation strategies
and the use of sensory feedback may be more appro-
priate than the present testing tasks to lead to cross-
over fatigue effects. Furthermore, based on the recent
data of Regueme et al. (2005b), additional emphasis
should be placed on the potential influence of pain
sensation and pain anticipation.
Key words: SSC fatigue, muscle damage, pain, con-
tralateral, muscle activity.
Acknowledgements
The authors gratefully acknowledge Dr. Nancy St-Onge and
Dr. Cathy Craig for revising the English manuscript.

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