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Annals of Physical and Rehabilitation Medicine 53 (2010) 319–341
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Neuromuscular and muscle-tendon system adaptations
to isotonic and isokinetic eccentric exercise
Adaptations neuromusculaires et musculo-tendineuses
à l’exercice excentrique isotonique et isocinétique
G. Guilhem, C. Cornu, A. Guével *
EA 4334, UFR STAPS, laboratoire motricité, interactions, performance, université de Nantes,
25 bis, boulevard Guy-Mollet, BP 72206, 44322 Nantes cedex 3, France
Received 12 October 2009; accepted 16 April 2010
Abstract
Objective. – To present the properties of an eccentric contraction and compare neuromuscular and muscle-tendon system adaptations induced by
isotonic and isokinetic eccentric trainings.
Synthesis. – An eccentric muscle contraction is characterized by the production of muscle force associated to a lengthening of the muscle-
tendon system. This muscle solicitation can cause micro lesions followed by a regeneration process of the muscle-tendon system. Eccentric
exercise is commonly used in functional rehabilitation for its positive effect on collagen synthesis but also for resistance training to
increase muscle strength and muscle mass in athletes. Indeed, eccentric training stimulates muscle hypertrophy, increases the fascicle
pennation angle, fascicles length and neural activation, thus inducing greater strength gains than concentric or isometric training programs.
Eccentric exercise is commonly performed either against a constant external load (isotonic) or at constant velocity (isokinetic), inducing
different mechanical constraints. These different mechanical constraints could induce structural and neural adaptive strategies specific to each
type of exercise.
Conclusion. – The literature tends to show that isotonic mode leads to a greater strength gain than isokinetic mode. This observation could be
explained by a greater neuromuscular activation after IT training. However, the specific muscle adaptations induced by each mode remain difficult
to determine due to the lack of standardized, comparative studies.
# 2010 Elsevier Masson SAS. All rights reserved.
Keywords: Lengthening contraction; Isotonic; Isokinetic; Muscle architecture; Electromyography
Résumé
Objectif. – Présenter les propriétés de la contraction excentrique et comparer les adaptations neuromusculaires et musculo-tendineuses induites
par les modes isotonique et isocinétique.
Synthèse. – Une contraction musculaire excentrique se caractérise par la production d’une force musculaire associée à un allongement du
complexe muscle-tendon. Cette sollicitation musculaire peut provoquer des microlésions auxquelles succèdent des processus de régénération des
structures musculaires et tendineuses. L’exercice excentrique est donc utilisé en rééducation fonctionnelle pour son effet positif sur la synthèse du
collagène et dans le domaine de l’entraı̂nement pour développer la force et la masse musculaire. L’entraı̂nement excentrique stimule en effet
l’hypertrophie musculaire, augmente l’angle de pennation, la longueur des fascicules musculaires et l’activation nerveuse, engendrant ainsi des
gains de force supérieurs aux entraı̂nements concentriques ou isométriques. L’exercice excentrique peut être réalisé contre une charge externe
constante (isotonique) ou à vitesse constante (isocinétique). Ces contraintes mécaniques différentes pourraient induire des adaptations structurales
et nerveuses spécifiques à chaque mode d’exercice.
* Corresponding author.
E-mail address: arnaud.guevel@univ-nantes.fr (A. Guével).
1877-0657/$ – see front matter # 2010 Elsevier Masson SAS. All rights reserved.
doi:10.1016/j.rehab.2010.04.003
320 G. Guilhem et al. / Annals of Physical and Rehabilitation Medicine 53 (2010) 319–341
Conclusion. – La littérature semble montrer que le mode isotonique induit des gains de force supérieurs au mode isocinétique, ce qui pourrait
s’expliquer par une augmentation de l’activation neuromusculaire plus importante. Toutefois, les adaptations induites par chaque modalité restent
difficiles à déterminer, en l’absence d’études comparatives standardisées.
# 2010 Elsevier Masson SAS. Tous droits réservés.
Mots clés : Contraction excentrique ; Isotonique ; Isocinétique ; Architecture musculaire ; Électromyographie
1. English version of eccentric contractions before presenting the adaptive

mechanisms induced by eccentric training on these various
1.1. Introduction factors. Finally, in a synthesis, we will present and compare
strength gains and adaptive processes, induced by the various
Eccentric contractions occur when the load torque (i.e. training protocols using IT or IK eccentric exercise, to
resistance moment) imposed on the muscle or a group of determine the specific relevance of using these two types of
muscles is greater than the muscle torque produced by all exercise modes for improving muscle qualities.
activated motor units (i.e. motor moment) [32]. The imposed
lengthening on the muscle-tendon system that goes along with 1.2. Properties of eccentric contraction
force production is given the term of ‘‘lengthening contraction’’
as opposed to concentric contractions (shortening contraction). 1.2.1. External torque
During this process, the muscle absorbs energy developed by an During an eccentric muscle contraction, the external
external load, this is why eccentric training is called ‘‘negative mechanical load exerted on the solicited muscle triggers a
work’’ [3], thus underlining its slowing-down role. The energy stretching of the elastic recoil energy components of the
absorbed can be (i) dissipated as heat, in which case the muscle muscle-tendon system, which in return, restitutes this stored
is functioning as a shock absorber or a damper (i.e. going down force-producing energy additional to the force produced by the
the stairs, slowing down at then end of a run), or it can be stored solicited muscle fibers. Consequently, this contraction mode
temporarily as elastic recoil potential energy that can requires a lesser metabolic energy cost for the same force
subsequently be recovered (i.e. locomotion, jumps with prior production compared to concentric contractions [3]. Thus, the
gained momentum) [24]. These observations brought several maximal torque developed by a muscle group during eccentric
authors to compare the muscle-tendon system behavior as a contraction is higher than the torque produced during isometric
spring that cyclically absorbs and recovers elastic recoil energy and concentric contractions (for a given angular velocity),
[105]. reaching 100 to 180% of the maximal isometric force according
During an eccentric contraction, the shock-absorber-spring to the different studies [54]. However, some research works do
properties of the muscle-tendon system contribute to the not report any significant differences between the maximal
external torque produced. The latter is higher than the one eccentric and isometric torques. There are some contradictions
developed during a concentric contraction for a given angular regarding the type of relationship between torque and
velocity [41]. The forced lengthening of the muscle associated movement velocity (‘‘torque-velocity’’) defined by using
to important tensions both cause a deterioration of the isokinetic ergometers, during eccentric contraction. Most
cytoskeleton and a local inflammation responsible for Delayed studies do agree that the maximal eccentric torque is not
Onset Muscular Soreness (DOMS) [65]. In order to protect the velocity-dependent (Fig. 1) [54]. However, the shape of
muscle-skeleton system of these damages, inhibition mechan- ‘‘torque-velocity’’ relationship seems to be related to the
isms are implemented by the nervous system. These specific individual’s sex, age or even level of strength [18].
responses of the neuromuscular system during eccentric work
brought several authors to focus on the chronic impact of 1.2.2. Muscle activation strategies
eccentric exercise in the framework of strength training [110] Eccentric contractions require unique central activation
and rehabilitation programs [103]. This type of contraction strategies by the nervous system specific to this type of action.
work can by applied to sports performance and rehabilitation In fact, electroencephalographic (EEG) measurements show a
training because it will solicit the muscle spring in a maximal higher and earlier cortical activity during submaximal and
and above-maximal manner, leading to major strength gains, maximal eccentric contraction of the forearm flexor muscles
but it will also reinforce the muscle-tendon system by [34], suggesting a different programming process in the central
stimulating collagen’s synthesis [91] and activating the genes nervous system (CNS) for eccentric contractions than
responsible for cellular development and growth [49]. concentric contractions. The electromyographic (EMG) activ-
Eccentric exercise is classically done at constant velocity ity measured on the skin surface during maximal eccentric
(isokinetic, IK) or against constant external load (isotonic, IT), muscle work is lower than the one recorded during concentric
inducing different mechanical loads depending on the type of contractions for a given angular velocity [1,58,67,130]. The
training (IK or IT), leading to specific neuromuscular and CNS seems to be incapable of recruiting in a maximal manner
muscle-tendon adaptive mechanisms. This literature review all motor units (MUs) of a muscle during an eccentric
will first explore the nervous, mechanical and structural factors contraction [42,73], this is even more visible with higher
G. Guilhem et al. / Annals of Physical and Rehabilitation Medicine 53 (2010) 319–341 321
during concentric contractions, suggesting the existence of a

pre-synaptic inhibition of the Ia afferents [26,84,111]. This
process could also explain the limited activation observed
during an eccentric contraction.
The mechanisms that highlight a specific muscle activation
during eccentric contractions [32] imply modulations in the
recruitment order, discharge rate and recruitment threshold of
MUs. Whereas some studies report that an eccentric contraction
is associated to selective activation of high-threshold MUs
[48,67,81], other studies show a recruitment order following the
size principle [38], for concentric and eccentric contractions
[23,89,120,124]. If Stotz and Bawa [124] observed the
recruitment of MUs at a high threshold during eccentric
Fig. 1. Torque evolution compared to the maximal isometric voluntary con- contractions, this was only obtained for erratic force or
traction (MVC) according to the angular velocity reported in several studies that movement profiles. These data suggest that a change in the
defined the ‘‘force–velocity’’ relationship. MVC : maximal isometric voluntary recruitment of MUs during an eccentric contraction can be
contraction; ~ Webber & Kriellaars, 1997 [127]; * Westing et al., 1998 [129];
observed at times without validating a preferential recruitment
~ Komi et al., 2000 [59]; * Dudley, 1990 [28]; ^ Colson et al., 1999 [19].
of fast MUs.
A voluntary contraction of agonist muscles is associated to
angular velocities [55]. This EMG decrease might indicate a an activation of the antagonist muscles (or co-activation) and
lower discharge rate [120] or a lower activation of the MUs can be quantified by surface EMG [33,121]. In that case the
during eccentric contractions, leading to a distribution of the external torque is the result of the torque produced by the
mechanical stress on a fewer MUs [7,82]. The unit tension agonist muscles, minus the torque generated by antagonist
generated by an active muscle fiber is then higher in eccentric muscles. The results regarding the impact of the contraction
mode, thus reducing the energy cost but also causing muscle mode on the level of co-activation are disparate according to the
damages on the recruited muscle fibers [82]. These observa- different studies. The impact could be greater during eccentric
tions have been validated by results obtained with the twitch muscle contractions (12% in eccentric versus 7% in concentric
interpolation electrical stimulation technique, showing an for plantar flexor muscles) [96] or concentric contractions (37
inhibition degree of the agonist muscle comprised between 5 to 48% in concentric versus 25 to 30% in eccentric; knee
and 30%, during eccentric contractions [6,8,28,129] versus 4 to extensor muscles) [56], whereas other studies show similar co-
21% in concentric mode [107]. Some hypotheses have been activation values regardless of neuromuscular solicitation type
suggested to explain the mechanisms responsible for this [6,99]. Such a mechanism decreases the levels of torque
limited muscle activation. The type Ib afferents from the Golgi produced and stabilizes the joint in regards to the high muscle
tendon organ (sensitive to muscle tension) can, for example, tensions induced by eccentric contractions.
have an inhibitor impact on the a motoneurons activity.
However, the impact from the Golgi tendon organ is not the 1.2.3. Behavior of muscle structures
primary mechanism able to explain this activation decrease The level of pre-activation prior to an eccentric contraction
during an eccentric contraction [96]. The efferent command can seems to have an impact on the evolution of muscles fascicles’
also be modulated by the Ia and II afferents, stemming from the length. In vitro studies on isolated muscle fibers suggest that the
neuromuscular spindles that give information on the muscle’s development of an isometric force is necessary before any
length (static sensitivity) and length variation (dynamic length change in order to ensure a complete pre-activation of
sensitivity) to the nervous system. Peripheral sensory receptors the MUs [31]. During eccentric contraction, the fascicle
exert an intense retroaction on the motoneurons a during pennation angle of the tibialis anterior is independent from
eccentric contractions; this retroaction seems to be canceled by movement angular velocity [104]. Furthermore, for high levels
the central and peripheral pre-synaptic inhibition of the Ia of angular velocity, the fascicle pennation angle measured in
afferents, as shown in studies on the Hoffmann (H) reflex in eccentric contraction is greater and the length of the fascicle is
dynamic settings [25,26]. The H-reflex is a component of the lower compared to a concentric contraction, inducing greater
stretching reflex characterized by an EMG response recorded muscular thickening in eccentric mode. Nevertheless, the
during a muscle contraction triggered by submaximal electrical mechanical behavior of the fascicles and tendons could also be
stimulation of the motor nerve. This reflex is the result of the related to the modality, intensity of the eccentric exercise as
motoneurons a activation via the Ia afferents coming from the well as the muscles and joint systems recruited [104]. However,
neuromuscular spindles. The H-reflex amplitude is linked to the all studies validate the large contribution of the tendon tissues
number of activated motoneurons and thus reflects the for the lengthening of the muscle-tendon system, estimated at
excitability of this pool of motoneurons and the synaptic 45% for the gastrocnemius medialis [16], even if we find great
transmission of Ia afferents. It represents a tool able to evaluate inter-individuals variations. The various evolutions of the
a modulation of the monosynaptic reflex activity. During fascicle length during the contraction (i.e. shortening,
submaximal eccentric contractions, this H-reflex is lower than lengthening or constant length) could be linked to the type
of exercise performed, which could have an impact on the Several authors have tried to identify if the damages induced
elastic spring recoil components and would optimize the by eccentric training affected one type of muscle fibers in
efficacy of the stretch shortening cycle. The energy restituted particular. The results of several studies, in human and animal
by the elastic spring recoil components in eccentric mode models, tend to validate that intense eccentric exercise damages
produces an additional force without any ATP consumption. especially Type II (IIb in particular) muscle fibers [36]. This can
This process, associated to a limited recruitment of MUs, leads partly be explained by the preferential recruitment of fast MUs
to a lower metabolic cost for the muscle [3]. Eccentric actions in the eccentric mode.
produce higher forces while consuming less oxygen and energy The physiological processes of cell regeneration after
than an equivalent concentric training session [88]. Energy is muscle damages are implied in muscle hypertrophy and thus
absorbed in the spring recoil properties of the muscle and are necessary to optimize training responses (Section 1.3.1.3).
tendon, thus leading to a lengthening of these structures in Eccentric muscle exercise as part of a safe muscle-strengthen-
response to the stretching. The tendon has a viscoelastic ing program remains an interesting method to improve muscle
behavior because of its nature, rich in collagen fibers (86% of strength. By affecting the motor command, it induces in return
the organic tissue mass), and the interactions existing between some adaptive processes on a structural level.
collagen and non-collagen proteins (i.e. proteoglycans). The
muscle elastic property is due to the interactions of actin and 1.3. Adaptations induced by eccentric training
myosin, the extracellular conjunctive tissue and to the elastic
structure of filaments maintaining sarcomere alignment of 1.3.1. Structural adaptation
muscle cells. This elastic complex is made of a large 1.3.1.1. Repeated bout effect. Several studies have showed
filamentous protein called titin (or connectin) and desmin, that a first session of eccentric exercise strongly limits the
both contributing to the spring property of the muscle during an DOMS perceived after the second eccentric exercise session,
eccentric contraction [66]. classically happening at a 1-week interval. This phenomenon is
The tendon’s viscoelastic properties make it more flexible called ‘‘repeated bout effect’’ (RBE) [74]. This first adaptation
than the muscle at lower stretching levels. Under these is characterized by a decrease in muscle damages, lower
conditions, tendons can absorb more energy but are less redischarge of creatine kinase (CK) and heavy myosin chains,
efficient for force transmission. At higher tension levels, they reduced loss of strength and DOMS after a second eccentric
become less flexible and stiffer, thus they are able to move exercise session, compared to the first session performed a few
heavier loads. These elements could explain that during days prior. The duration of this adaptive phenomenon varies
eccentric contractions, the muscle fascicles’ length of the from a few weeks to 6 months according to different studies.
gastrocnemius medialis remains constant when the muscle According to Chen [14], the importance of DOMS and plasma
fascicles of the vastus lateralis lengthen, because they are CK levels after the first eccentric exercise session have an
exposed to greater mechanical constraints. impact on the RBE. Thus, it is possible that an individual’s
sensitivity level to eccentric exercise conditions the importance
1.2.4. Muscle damages of this protective effect that sometimes might not be observed
The eccentric mode exerts a high so-called ‘‘above [100]. In a recent review, Lieber and Friden [65] bring up RBE’s
maximal’’ mechanical strain on the muscle-tendon system. high level of specificity, which is only valid when the two
The high tension levels that come along with the forced eccentric sessions are identical.
lengthening of this system could lead to micro lesions of the The adaptations that can potentially explain this RBE are
muscle fibers [20]. At a cellular level, these damages include a from a nervous, mechanical or cellular origin. Due to this type
disorganization of some sarcomeres and Z-line streaming of fast-setting adaptation, some authors even suggested that
disruption (i.e. ‘‘wavy’’ appearance [65]), associated to eccentric training could improve MUs synchronization and
sarcolemmal lesions, dilation of the transverse tubule system increase their activity level [74]. This process could increase
and fragmentation of the sarcoplasmic reticulum [65]. These the number of solicited muscle fibers compared to the first
damages are followed by an inflammatory response with a session, mainly by recruiting a great number of slow MUs,
gradual increase in the soreness of the involved muscles with a leading to a better mechanical stress repartition on all activated
delayed onset after exercise called Delayed Onset Muscular muscle fibers and thus limiting the risk of myofibrils damages
Soreness (DOMS). Eccentric exercise is the main contributor to [22,82]. Because of the mechanical origin of these eccentric
this pain commonly called ‘‘muscle aches’’. This pain occurs exercise-induced lesions, some authors suggested that some
12 to 48 hours after intense or unfamiliar eccentric exercises changes to the mechanical properties of the skeletal-muscle
with a peak between 24 to 72 hours before progressively system would protect the muscle against these damages.
disappearing in 5 to 7 days. DOMS are associated to a decrease Indeed, some studies did report an increase of the active or
in force production capacity ( 10 to 60% on the maximal passive stiffness of the muscle-tendon system after eccentric
voluntary isometric contraction (MVC), according to the training [98]. These adaptations were respectively attributed to
studies [74,80]), restored in about a week or so. The impact of a reorganization of the cytoskeleton proteins, in charge of
these muscle damages during a second eccentric exercise maintaining the alignment and structure of the sarcomeres (i.e.
session is lower than for the first session (i.e. protecting effect, titin, desmin) and to a quantity increase of the connective tissue
Section 1.3.1.1) [74]. in the muscle, thus allowing for a better dissipation of the
mechanical stress during eccentric mode [64]. However, some muscle for capacities is coupled to an increase in muscle mass.
studies also show that stiffer muscles are more prone to The eccentric component of the training is then essential to
damages [72]. In fact, some studies on ‘‘murine’’ models induce the surface increase of muscle fibers’ transversal section
without desmin showed that these mice were less prone to [29]. Training programs that include eccentric actions lead to
damages than healthy mice following eccentric exercise. This greater muscle mass gains than training aiming to achieve this
result was attributed to a greater muscle compliance of these goal on a concentric mode [35,58]. Higbie et al. [40] showed an
mice thus limiting the number of sarcomeres that could be increase in quadriceps cross-sectional area (CSA) after
damaged. The last hypothesis to explain the RBE is the impact 10 weeks of an eccentric training compared to a similar
of cellular factors, stating that eccentric exercise leads to an training in concentric mode. Similar results were obtained for
increase in serial sarcomeres and changes in the inflammatory elbow flexors [35]. These results recently brought researchers
response. The addition of serial sarcomeres, shown on animal to focus on the effects of eccentric training on the gene
models [79] and recently, in an indirect manner, on human expression of the solicited muscle cells. In fact, eccentric
models [13], could reduce the stretching and thus the training represents for muscle cells a specific mechanical
sarcomeres’ rupture after an eccentric exercise session. A constraint that can modify the several gene expression profiles
decrease of inflammation after repeated eccentric training has [39,57], via the mechanical signaling pathways made of
also been characterized by a reduced activation of the proteins that are sensitive to the mechanical status of muscle
monocytes and neutrophils involved in the inflammatory cell (i.e. Microtubules-Associated Proteins or MAP proteins).
process [97]. This reduced inflammatory response contributes An eccentric session triggers, 6 hours post-exercise, the
to the protective effect even if it is difficult to know if this progressive activation of genes responsible for cellular growth
adaptive strategy is the cause or the consequence of a reduced and development, involved in cellular hypertrophy processes in
number of damaged myofibrils, or even if the repeated bout humans. The expression levels of these genes are more
effect is a combination of these two processes [74]. stimulated by eccentric actions than by isometric or concentric
actions [9,15,60]. Introducing eccentric exercise during so-
1.3.1.2. Strength gains. According to several studies, called ‘‘hypertrophy’’ training phases seems to be an efficient
eccentric training seems to be the most efficient mean to mean to optimize muscle mass gain.
increase the capacities of maximal force that can be exerted in a
given movement, mainly compared to concentric training (see 1.3.1.4. Changes in muscle fiber types. The changes in fiber
Remaud et al. [107] for a detailed review). For example, types induced by eccentric training are similar to those induced
Hortobagyi et al. [43], report a strength gain of the maximal by other strength training programs, i.e. an increase of the
voluntary isometric contraction 3.5 times greater after a 6-week intermediate type muscle fibers (IIa, IIa/IIb) able to reach +12%
eccentric program of the knee extensors compared to a [43], with sometimes a decrease in the number of type IIb fibers
concentric training program of the same duration. These results [118]. However, these evolutions remain moderate even not
were validated with longer training duration (+24.6% of the statistically significant in most studies. They do not seem to be
initial strength after eccentric training versus +12.5 after responsible for the main adaptation that could explain strength
concentric training) for the elbow flexors and extensors [83]. gains observed after eccentric training.
Studies focusing on the effects of overload training (100 to
120% of the maximal repetition) during the eccentric phase 1.3.1.5. Changes in muscle architecture. Recent studies tried
unveil greater strength gains then during strength training using to identify the changes of the geometrical organization of
lighter loads [45]. Thus, it seems that the eccentric part of the muscle fibers induced by strength trainings combining
training protocol is essential to increase muscle strength. In concentric and eccentric exercises [53], eccentric trainings
fact, the contractile part of the muscle-tendon system is [12] or plyometric trainings [11]. A loaded training phase can
solicited to the maximum, thanks to the high torque levels increase the pennation angle of muscle fascicles, a process
reached in eccentric training. Strength gains obtained in regards associated to muscle hypertrophy [2]. In fact, the pennation
to the initial muscle force production capacities go from 1 to angle is modified by the increase in muscle mass, and its
116% for eccentric force, from 1 to 67% for concentric force evolution according to the interspace of aponeurosis in
and from 7 to 45% for isometric force [43,86]. If several authors hypertrophied muscles. Recently a 21.4% increase of the
remain attached to the principle of strength gains specificity pennation angle of the vastus lateralis was demonstrated after a
according to the training mode [30], it appears that eccentric 10-week eccentric training program (versus +13.3% after
training is more efficient for improving the maximal concentric concentric training) [12]. A pennation angle increase leads to a
force of a muscle group [52]. Finally, strength gains induced by decrease in the force transmitted by a muscle fiber, considered
eccentric training seem very specific to the movement velocity as an isolated fiber. However, in terms of muscle CSA, muscle
used [110]. fibers pennation leads to a physiological CSA of the muscle,
defined as the area of the muscle fibers perpendicular to the
1.3.1.3. Hypertrophy. One of the major adaptive processes longitudinal axis of each muscle fiber multiplied by the cosine
induced by eccentric training is muscle hypertrophy, represent- of the pennation angle [132], which is greater than the
ing one of the determining factors for force production anatomical muscle CSA. Since physiological CSA represents
capacities. Beyond 6 to 8 weeks of training, the improvement in the maximum number of parallel actin-myosin cross-bridges
that can be activated during a contraction, the maximal force indicator of the increase in post-eccentric training muscle
production capacity of a muscle is proportional to this compliance. This adaptation induced by eccentric training
physiological CSA and thus to the pennation angle. The suggests some applications for injury prevention. In fact, the
increase in the pennation angle leads to an increase quantity of decrease in muscle’s passive stiffness allows for transferring
parallel contractile elements, thus higher force production part of the mechanical constraint from the tendon to the muscle,
capacities [2]. reducing the risks of tendon damage. However, serial elastic
Regarding the length of muscle fascicles, if some training components (SEC) made of elastic muscle structures and the
protocols have not reported any changes for this parameter after tendon increases after eccentric training. Pousson et al. [98]
strength training [12], other protocols using overloading [116] reported an increase for this parameter after 6 weeks of
or high velocities on a plyometric mode [11] have shown an eccentric training of the elbow flexors, at 30 to 45% of the
increase in the muscle fascicles length (addition of serial maximal voluntary isometric contraction (MVC). Yet, this
sarcomeres). To our knowledge, only one study focused on the increase is not observed for higher levels of force (60 to 80% of
effect of a purely eccentric training on muscle architecture. The MVC). These results were partially validated by the work of
authors have unveiled a 3.1% length increase for the fascicles of Mahieu et al. [68] who reported a 9.5% increase for Achilles’
the vastus lateralis after eccentric training versus 6.3% for a tendon stiffness measured in active conditions after 6 weeks of
same-duration concentric training without any significant eccentric training of the plantar flexors. The increase in SEC
difference between the two training modes [12]. The authors stiffness might thus be explained by an increase in tendon
suggested that movement amplitude more than contraction stiffness. It could also be the result of an increased stiffness of
mode or movement velocity was the factor with the most the actin-myosin cross-bridges or the remodeling of the
impact on the changes in muscle fascicles’ length. The increase cytoskeletal proteins responsible for maintaining sarcomere
in muscle fascicles’ length allows also for increasing the alignment such as desmin or titin. The reorganization of these
velocity of the muscle contraction [12]. Thus, eccentric training proteins after the micro-damages induced by eccentric exercise
is an efficient process to improve not only the force production could be one of the processes responsible for the increased
capacities, but also the quality of the muscle power (i.e. result of stiffness of the elastic component of the muscle. These elastic
the torque and velocity of the muscle contractions) essential in properties’ changes could promote the restitution of mechanical
most physical and sport activities. energy, mainly during stretch shortening cycles.
Furthermore, chronic eccentric exercise is often proposed
1.3.1.6. Changes in mechanical properties. Classically, for the treatment of tendinopathies for its stimulating impact on
eccentric exercise session is followed by an increase in passive collagen synthesis, main constitutive element of the tendon and
muscle-joint stiffness [47]. The latter can be multiplied by two conjunctive tissue [91]. Several studies have also reported an
and remains elevated for 4 days after the eccentric training increase in blood flow around tendon cells after eccentric
session. This increase in passive stiffness might be explained by exercise [5,63]. These adaptive processes would justify the
the formation of residual actin-myosin cross bridges [62] relevance of eccentric exercise for strengthening tendon tissues
resulting from an increase in the calcium ions concentration at in patients suffering from tendinopathies. However some
rest in the muscle fibers, the latter being linked to the rupture of questions remain, especially regarding specific protocols (i.e.
muscle cells’ membranes. Other processes have been suggested load, movement velocity) to be used according to the
to explain this increase in passive stiffness such as edema pathology’s characteristics [103].
characterized by muscle swelling compressing the tissues and
inducing a painful resistance to passive extension thus 1.3.2. Nervous adaptations
increasing the ‘‘stiffness sensation’’ [17]. Finally, the repair During an eccentric contraction, the electric activity of a
process might lead to a permanent increase of passive stiffness muscle is lower than the one recorded during a concentric
because of the restructuring of the connective tissue as contraction for the same level of force produced (Section 1.2.2).
suggested in the work of Lapier et al. [64]. Eccentric training can increase this neuromuscular activation.
When the eccentric exercise is repeated over long periods of Hortobagyi et al. [43] report an EMG activity increase 2.6 times
time (i.e. training), the muscle-tendon stiffness determined in greater after a 6-week quadriceps eccentric training program,
passive condition decreases. Mahieu et al. [68] showed an compared to a similar concentric training program. Similar
increase in joint range of movement (ROM) and a 23% decrease results were obtained on elbow flexors [58]. This result can be
of the passive resistive torque after a 6-week eccentric training due:
program of the plantar flexors. Before that, Morgan and Allen
[79] had already reported increased serial sarcomeres in the to the low-degree of muscle activation, before training, in
muscle fibers (+11%), after 1 week of downhill running on a eccentric mode, compared to other types of contractions (i.e.
treadmill. These results are in accordance with recent studies isometric and concentric);
showing a modification in the relationship between the to an increased EMG activity in the fibers that are specifically
maximum torque produced and the joint angle with a shift recruited by eccentric exercise.
in peak torque (shift around 158) towards greater muscle
lengths right after eccentric exercise [131]. This modification in It seems that type II fibers are more likely to be recruited
muscle length capable of producing the maximal torque is an during eccentric training. We also saw that repeated eccentric
training could lead to a progressive recruitment of slow MUs in progressive increase of the movement velocity in eccentric
order to better distribute mechanical stress to protect muscles contraction and was tested on a cohort of 200 patients. The
from damages. These processes could be the cause of an results showed a complete recovery of the motor function for
increased EMG activity of the muscle trained in eccentric 44% of them. Inspired by the work Stanish, Alfredson et al. [5]
mode. The increase in MUs’ discharge frequency could also reported that the eccentric mode was an essential therapeutic
explain this gain in EMG activity. However, even if the tool and an integrant part of the results obtained with these
discharge frequency can be increased by training, only 15% of PM&R programs.
the maximal force is due to a modulation of this parameter in The physiopathology of the tendon affected, or under
large-size muscles [93]. Recently, some studies have been treatment, is not completely understood. It seems however that
started in order to better define the origin of these various during tendinopathies, the tendon cannot enter into an active
nervous adaptations through the analysis of the V-wave and H- recovery cycle [103]. Eccentric exercise could constitute an
reflex after an eccentric training phase of the plantar flexors efficient mechanical stimulus for quiescent tendon cells. Other
[27]. An increased V-wave could result in an increased number studies have shown an interruption of the blood flow around
of recruited MUs or an increased frequency of these MUs’ tendon cells during eccentric exercise [5]. This process could
discharge. This increase of the descending signals from the damage the associated micro-capillaries and nerves, thus
CNS depends also on the efficacy of the nervous transmission at reducing the pain sensation. Recently, Langberg and Kongs-
the level of the neuromuscular junction. Duclay et al. [27] gaard [63] highlighted an increase in collagen synthesis (type I
highlighted a constant increase of the V-wave during voluntary collagen) main constitutive element of tendon tissue, without
eccentric contractions, which underlines an improved excit- increasing the degrading processes, within the injured tendon
ability of motoneurons a or a reduced presynaptic inhibition of after 12 weeks of eccentric heavy resistance training. The
Ia afferents. Strength gains could also be the consequence of a authors also observed a decrease in pain perceived by the
decreased activity of the antagonist muscles (co-activation). On patients. Other studies showed an increase in the blood flow or
a nervous system level, a lesser co-activation degree could tendon volume, suggesting an improvement in the intratendi-
reduce the reciprocal inhibition level and thus facilitate the H- nous signal between tissues and interstitial area [117]. These
reflex. Nevertheless, eccentric training does not seem to have an adaptive processes justify the relevance of eccentric training in
impact on the co-activation level, the latter remaining stable tendon tissue reinforcement in affected patients. However, few
from the beginning to the end of the training period in most studies have compared these results to the effects induced by
studies [19,115]. Reciprocal inhibition cannot a priori explain concentric exercise. The first comparative studies have shown
the observed H-reflex changes [27]. However, eccentric that force peaks generated in vivo within the tendon during
training effects on the amplitude of the H-reflex are different eccentric exercise were similar than during concentric exercise
in regards to the muscle solicited (i.e. for the soleus and the [102,113]. These results show that the therapeutic benefits of
medial gastrocnemius) suggesting an organization of the eccentric training are not only due to the levels of force induced
muscle-specific reflex arcs. The origin of these different by this exercise mode. Other authors have suggested that the
nervous adaptations induced by eccentric training remains to be induced force variations would be a stimulus responsible for
defined according to the considered muscle group. tendon remodeling just like results obtained in bone tissues
using high-frequency signals [112]. These variations are
1.3.3. Role of eccentric training in rehabilitation reflected by the difficulty of the subjects to control movement
Eccentric training allows for working at high-levels of force during an eccentric contraction. Thus, if eccentric training is
with a lower metabolic cost and oxygen uptake. This exercise commonly used for treating tendinopathies, some questions
mode is then relevant for clinicians wanting to include, in their remain, mainly in regards to specific protocols (i.e. contraction
Physical Medicine and Rehabilitation (PM&R) programs, type, load, movement velocity) and how to use them according
patients that are often intolerant to intense cardiac and to the characteristics of the various pathologies [63,103].
respiratory efforts (i.e. patients with heart disorders or lung Finally, it seems that unilateral eccentric training would
pathologies). We have just brought up the protective effect of an increase muscles’ strength in the opposite limb, and this in a
eccentric training session towards muscle damages. In athletes more noticeable way than after concentric training [40,44].
getting back to a physical activity after an injury, sub-maximal Implementing eccentric contractions in rehabilitation programs
eccentric training is a relevant manner to prepare the musculo- could lead to a functional improvement in muscle groups of the
skeletal system for getting back to exercise and avoiding the opposite injured limb without it being solicited.
onset of new injuries during this re-training phase. It is also
used for therapeutic care in other pathologies such as multiple 1.4. Comparison of isotonic and isokinetic eccentric
sclerosis for its positive impact on motor command [108]. training
Initially suggested by Stanish et al. (1986), chronic eccentric
exercise is also proposed for treating tendinopathies, mainly for 1.4.1. Specificities of the two modalities of isotonic and
Achilles’ and patellar tendons. These pathologies represent a isokinetic exercise
large part of Sports Medicine [109] and are a Public Health 1.4.1.1. Isotonic eccentric mode (IT). In vivo, an eccentric IT
challenge since they also affect the general population (around solicitation opposes a muscular torque to a constant load that
2% of the active population). The Stanish program used a can overcome the force capacities production of the subject (i.e.
supra-maximal eccentric training [45]), leading to a lengthen- (Tables 1a and 1b). When we compare the mean strength
ing of the muscle-tendon system. In IT eccentric condition, a gains induced by the two eccentric modalities, we observe that
weight bar or weights, equipments classically used for strength IT eccentric training increases the force production capacities
training, most often impose the constant load. If the charge in by 1.1 1.0% per session [10,19,27,46,49–51,69,70,77,
this type of exercise is fixed, the torque exerted by the subject 85,90,92,94,98,101,119,122,123,125,126,128] for a mean
varies, mainly due to accelerations and decelerations affected to training duration of 7.5 3.4 weeks (Table 1a), against
the imposed load, and to the lever arm’s length variations 0.6 3.0%, for eccentric IK training (mean training duration
according to the apparatus [4]. This exercise mode is more an 10.6 4.9 weeks) [12,30,35,40,43,58,71,75,76,78,83,87,114,
isoinertial mechanical solicitation, but classically referred to as 115,118] (Table 1b), meaning that the gain is two times
‘‘isotonic’’ in the literature [37]. The angular velocity during greater in IT mode. Moreover, it would seem that IT
this type of movement varies and the recorded levels of solicitations, from this point of view, are more efficient than
mechanical stress are very high (up to 140% of the maximal exercises performed at constant velocity in eccentric condi-
repetition for trained subjects), mainly in the extreme joint tions. This observation has already been reported in some
angles (i.e. beginning and end of joint ROM). Thus, an IT studies [61] and in a similar evaluation study comparing IT and
eccentric contraction has a first shock-absorbing phase during IK modalities in concentric mode, bringing up average strength
which the force imposed by the load is greater than muscle gains per session from 0.5 to 1% in IT versus 0.3% in IK [106].
force, followed by a phase during which the subject controls the The differences in terms of mechanical stress (torque,
load, up to the extreme angles where load control becomes movement velocity) imposed by both modalities (IT and IK)
more difficult. This movement pattern suggests that IT according to the joint angle could also be correlated to
contraction solicits, in a preferential manner, the elastic disparities in induced strength gains. However, if the origin of
structures at these extreme angles, the muscle being more these strength gains is described for eccentric training in
solicited at the angles close to peak torque. Eccentric exercise general (Section 1.3), the specific IT and IK modalities induced
allowing for increasing muscle-joint stiffness, this action mode adaptations are still left to be determined.
could then be chosen to increase the stiffness of these elastic
structures at these joint angles. However, muscle activation is 1.4.3. Structural adaptations induced by isotonic or
not at its maximum during the entire ROM, since when isokinetic training
reaching the angle at peak torque, the production capacities of With eccentric training, skeletal muscles are solicited in a
the muscle force can sometimes exceed the resistance imposed maximal and supra-maximal way, their contractile structures
by the weights [37,46]. Eccentric IT solicitation could then adapt when the training lasts longer than 6–7 weeks [95]. Six of
have a moderate impact on nervous adaptation and on strength the 22 studies selected for this review focused on the evolutions
gain at this joint angle. of muscle mass after a phase of IT eccentric training (one study
does not report a variation in percentage and thus is not
1.4.1.2. Isokinetic mode (IK). An IK eccentric contraction is included in the mean average) and seven studies have worked
defined by the production of moment-velocity force opposed to on this parameter following IK eccentric training. IT eccentric
the movement of a lever arm mobilized at constant velocity by training induces a mean muscle hypertrophy of 3.26 3.77%
an active isokinetic dynamometer (movement isokinetics) [54]. [10,46,51,85,119,126] versus 7.26 4.07% after IK eccentric
This type of apparatus induces a maximal solicitation of the [12,30,35,40,58,114,118]. Brought down to the number of
contractile system on the overall ROM [8,59], without major sessions, the muscle hypertrophy obtained in IT eccentric
acceleration as it can be the case with IT eccentric contractions. mode is at 0.10 0.13% versus 0.29 0.17% in IK. The mean
The effects induced on muscle stiffness at the extreme angles duration of these IT protocols is 12 weeks against 10 weeks for
could not reach those obtained after IT eccentric training. IK trainings. Thus, in spite of shorter protocols’ duration, IK
However, the muscle force produced being at its maximum on eccentric training seems to induce a greater muscle hyper-
the entire joint ROM, the activity of the solicited muscles, is trophy than IT eccentric training. Furthermore, it is interesting
theoretically here also maximal, mainly at the peak torque to note that the significant gains are obtained in 12 to 20 weeks
angle, suggesting more pronounced CNS adaptations with this in IT [119] whereas an increase in muscle mass can be observed
type of training compared to IT eccentric training. This exercise in 8 weeks of IK eccentric training [35]. In the same manner,
mode is highly relevant for clinicians who want to propose the intensity required to observe muscle hypertrophy in IT
PM&R programs. Isokinetics can lead to a complete stop of the must be close to or exceed 100% of the one-repetition
movement when there is a lack of muscle force production, and maximum (1RM) if the duration does not exceed 8 to 10 weeks.
impose an adapted resistance to the individual’s capacities The fact that the exercise is at its maximal intensity during the
offering interesting safety conditions for patients under entire effort in IK eccentric training seems to promote muscle
therapeutic care [21]. mass gain. The result, however, does not seem to have an
impact on strength gains that are greater after IT eccentric
1.4.2. Strength gains training.
In order to compare the results obtained after IT and IK Regarding hypertrophy by muscle fiber types, the synthesis
eccentric trainings, strength gains were correlated to the of the different studies shows clearly a hypertrophy of type II
numbers of training sessions performed by the subjects muscle fibers in majority, regardless of the mode used (IT or
Table 1a
Strength gains induced by isotonic (IT) eccentric training.
Study’s characteristics Strength gains obtained for each velocity type test (per session)
1 1 1 1 1
Muscles Author Wks Fq Volume Intensity < 908.s 908.s < 08.s < 908.s 908.s <
(%1RM)
Knee extensors Ben-Sira et al. [10] 82 10 3 65–85 - - - +1.0 -
Housh et al. [46] 83 36 80a - +1.2 - ns ns
Johnson [49] 83 1 10 80 - +2.4 - - -
Johnson et al. [50] 63 26 120 - - - +1.0 -
Jones and Rutherford [51,52] 12 3 36 116 - - +0.2 - -
Kaminski et al. [52] 62 28 100 +2.5 +3.2 - +2.0 +2.1
Pavone and Moffat [90] 63 3 10 40–79a - - +1.3 - -
Raue et al. [101] 43 38 80 - - - +0.6 -
Smith and Rutherford [119] 20 3 3 10 100–110 - - +0.4 +0.3 +0.3
Sorichter et al. [122] 6 1–3 7 10 150 - - +0.7 - -
Spurway et al. [123] 61 36 100 +4.6 +4.3 +1.4 +2.8 +2.4
Weir et al. [128] 83 36 80a - +1.2 +0.6 +1.4 +0.6
Knee flexors Johnson et al. [50] 63 26 120 - - - +1.4 -
Mjolsnes et al. [77] 10 1–3 2–3 5–12 PC - +0.4 +0.3 - -
Elbow flexors Colson et al. [19] 73 56 100–120 +2.6 +1.6 +0.5 +0.9 +1.1
Johnson [49] 83 1 10 80 - +1.4 - - -
Martin et al. [70] 43 35 100 - - +0.4 +1.3 +0.8
Mannheimer [69] 45 25 - - +3.0 - - -
Nosaka and Newton [85] 81 1 10 79 - - +0.7 0 0
Petersen [94] 8 4–5 1 10 120–130b - - +0.2 - -
Pousson et al. [98] 62 2 4–6 90–125 - - - - -
Valour et al. [125] 73 36 57–95 - +0.7 +0.5 +1.3 -
Vikne et al. [126] 12 2–3 36 83–93 - +0.9 - +0.5 -
Johnson et al. [50] 63 26 120 - - - +3.4 -
Elbow extensors Johnson et al. [50] 63 26 120 - - - +1.6 -
Plantar flexors Duclay et al. [27] 7 2–3 66 100–120 - +1.7 +1.2 +1.6 -
Pensini et al. [92] 44 66 120 - +1.0 +1.9 +0.9 -
IK). Only three studies report a hypertrophy of type I muscle (+21.4%) and fascicles’ length (+3.1%) after a 10-week IK
fibers (from +0.62 to +0.86% per session) associated to a eccentric training program.
hypertrophy of fast-twitch muscle fibers after eccentric training
[52,87,92]. The scarce number of studies on this topic does not 1.4.4. Nervous adaptations following isotonic or isokinetic
permit us to conclude to the superiority of one training mode training
over another to trigger a hypertrophy on a preferential type of All studies focusing on nervous adaptations induced by
muscle fibers. Regarding changes in muscle fiber typology, IT eccentric training show an increase in the neuromuscular
and IK modes induce a decrease in the number of type I and IIb activation of the trained muscle groups. In this light, it is
muscle fibers with an increase in the proportion of type IIa/IIb interesting to note that muscle activation gains are higher after IT
fibers [12,43,87,101,114,118,126]. eccentric training (+1.80 1.54% per session) [19,27,92,125],
Very few studies have investigated the effect of eccentric compared to IK eccentric training (+0.43 0.17% per session)
training on mechanical properties. In fact, no study ever [40,43,58,115]. In the same manner, a study showed a significant
focused on the evolutions of the mechanical properties increase of the maximal voluntary activation (+17%) [92], a
following a phase of IK eccentric training. However, studies parameter which, to our knowledge, has not been studied on
conducted on IT mode showed an increased stiffness of the the IK mode. Finally, this same study reported a reduced
serial elastic component due to an increased collagen synthesis muscular co-activation ( 22 to 29% compared to the initial co-
[98], and a decreased muscle stiffness due to an increased activation level) after a phase of IT eccentric training [92]
fascicles’ length, characterized in passive conditions after IT whereas no significant evolution has been highlighted after IK
eccentric training [68]. Tendon stiffness tends to increase after training [12].
this type of training. However, these results still need to be Thus, IT eccentric training seems more prone to adaptation
validated and compared to the IK mode. Finally, to our of the nervous command of the solicited muscles. It does not
knowledge, only one study researched the impact on IK enable us to validate the hypothesis formulated according to
eccentric training on muscle architecture [12], which does not which IK training might be more efficient on this level due to
allow us to establish a comparison with IT eccentric training. the higher peak torque that this mode can reach (Section
The results have shown an increase of the pennation angle 1.4.1.1). However, the selected parameter (i.e. mean activation
Table 1b
Strength gains induced by isokinetic (IK) eccentric training.
Study characteristics Strength gains obtained for each velocity type test (per session)
Muscles Authors Wks Fq Volume Wf (8.s 1) < 908.s 1
908.s 1
< 08.s 1
< 908.s 1
908.s 1
<
Knee extensors Blazevich et al. [12] 10 3 4–6 6 30 - +1.3 - +0.5 -
Duncan [30] 63 1 10 120 +1.6 +1.5 - +0.2 -
Higbie et al. [40] 10 3 3 10 60 - +0.1 - +0.2 -
Hortobagyi et al. [43] 12 3 4–6 8–12 60 +3.2 +1.3 +0.1 -
Mayhew et al. [71] 43 5 10 60 - - +0.7 - -
Miller et al. [76] 20 3 1–5 6 60 - +0.7 - +0.4 -
Seger et al. [114] 10 3 4 10 90 +0.3 +1.1 +0.3 +0.2 +0.1
Seger and 10 3 4 10 90 +0.0 +0.9 - +0.0 +0.2
Thorstensson[115]
Melo et al. [75] 12 2 2–4 12 60 - +0.8 - - -
Knee flexors and extensors Nickols-Richardson 20 3 1–5 6 60 - +0.5 - +0.2 -

et al. [83]
Knee flexors Melo et al. [75] 12 2 2–4 12 60 - - - +0.5 -
Elbow extensors Farthing and 83 2–6 8 180 +0.7c - +0.9d

Chilibeck [35] 30 +0.3c - +0.7d
Komi and Buskirk [58] 74 16 - - +0.6 +0.3 +0.6 -
Paddon-Jones et al. [87] 10 3 46 180 +0.9 +1.0 +0.7 +1.0 +1.0
30 +0.4 +0.0 +0.3 +0.1 +0.2
Shepstone et al. [118] 83 1–4 10 210 +0.7 +0.2 - +1.2 +2.4
20 +0.4 +0.1 - +0.6 +0.7
Elbow flexors and extensors Nickols-Richardson 20 3 1–5 6 60 - +0.4 - +0.2 -

et al. [83]
Shoulder cuff rotators Mont et al. [78] 63 8 10 90–180 +0.8c +0.5c - +0.1e +0.5e
Strength gains obtained after isotonic eccentric training (IT, Table 1a) and isokinetic (IK, Table 1b) eccentric training for each type of velocity used : eccentric with
movement angular velocity greater than 908.s 1 (< 908.s 1), eccentric with movement angular velocity lower than 908.s 1 ( 908.s 1 <), isometric (08.s 1),
concentric with movement angular velocity lower than 908.s 1 (< 908.s 1) and concentric at movement angular velocity greater than 908.s 1 (908.s 1 <). Strength
gains are expressed in percentage of the pre-training value and brought back to the total number of sessions performed during the training period. The significant
evolutions are highlighted in bold. When several test velocities were used in each of the types, the listed value represents the mean of the obtained strength gains for all
the tested velocities.
Wks : weeks ; Fq : frequency (number of sessions per week of training); 1 RM : maximal load that the subject can lift in one repetition ; Volume : number of
series number of repetitions ; (P < 0.05).
a
Intensity defined in percentage of the maximal eccentric repetition.
b
Intensity defined in percentage of the maximal isometric voluntary contraction.
c
Mean strength gains for all test velocities in eccentric contraction.
d
Mean strength gains for all test velocities in concentric contraction.
e
Mean strength gains for internal and external rotator cuffs of the shoulder.
f
Mean angular velocity (8.s 1).
on the overall ROM, activation at peak torque) to assess the used in eccentric training, enabled to highlight the superiority
evolution of the neuromuscular activation varies according to of IT mode compared to IK mode regarding strength gains
the studies, which can influence the results obtained. These obtained. This can be explained by greater changes in the
results could explain in part greater strength gains obtained via nervous command following this type of training. Conversely,
IT eccentric training (Section 1.4.2). IK eccentric training seems to promote muscle hypertrophy.
The adaptive (structural and nervous) mechanisms still remain
1.5. Conclusion to be defined because studies on these two modes (IT, IK)
remain quite rare. The synthesis of the studies found in the
Eccentric exercise is an efficient mean to induce major literature and reported in this review allowed for identifying
structural changes on the muscle-tendon system and adapta- differences in the adaptive process induced by IT and IK
tions of the nervous central command. These adaptations allow eccentric trainings (i.e. greater muscle hypertrophy after IK
for a significant increase of the maximal force production training, greater increase in neuromuscular activation after IT
capacities, often higher to those obtained with concentric or eccentric training). However, the effects of IT and IK eccentric
isometric trainings. A comparative synthesis of the adaptive training have not, to this day, been compared in standardized
processes induced by these two solicitation modes classically settings. Such studies could better define the relevance of their
use in strength training or PM&R setting, according to the adaptations neuromusculaires et musculo-tendineuses spécifi-
targeted adaptive goals. ques. Cette revue de littérature se propose donc d’exposer
d’abord les caractéristiques de la contraction excentrique au
2. Version française niveau nerveux, mécanique et structural, avant de rendre
compte des adaptations induites par l’entraı̂nement excentrique
2.1. Introduction sur ces différents facteurs. Les gains de force et les processus
adaptatifs induits par les différents protocoles d’entraı̂nement
Une contraction musculaire est dite excentrique lorsque la ayant mobilisé l’exercice excentrique IT ou IK sont finalement
contrainte mécanique imposée (i.e. moment résistant) à un présentés et comparés dans une synthèse visant à déterminer
muscle ou à un groupe musculaire est supérieure à la force l’intérêt spécifique de l’emploi de ces deux modes d’exercice
produite par l’ensemble des unités motrices activées (i.e. dans l’amélioration des qualités musculaires.
moment moteur) [32]. L’allongement forcé du complexe
muscle-tendon qui accompagne alors la production de force est 2.2. Propriétés du régime de contraction excentrique
désigné sous le terme de lengthening contraction en opposition
au régime de contraction concentrique (shortening contrac- 2.2.1. Couple de force externe
tion). Au cours de ce processus, le muscle absorbe l’énergie Lors d’une contraction musculaire excentrique, la contrainte
développée par une charge externe, ce qui vaut au travail mécanique externe appliquée au segment mobilisé provoque
excentrique l’appellation de « travail négatif » [3], soulignant l’étirement des composantes élastiques du système musculo-
ainsi son rôle frénateur. L’énergie absorbée peut être : tendineux impliquées qui, en retour, restituent de l’énergie
productrice d’une force additionnelle à la force produite par les
dissipée sous forme de chaleur, auquel cas le muscle fibres musculaires recrutées. En conséquence, ce mode de
fonctionne tel un amortisseur (e.g. descente d’escaliers, contraction mobilise une moindre énergie métabolique pour
ralentissement en bout de course) ; une même production de force qu’en mode concentrique [3].
stockée temporairement en énergie potentielle élastique Ainsi, le couple de force maximal développé par un groupe
pouvant ensuite être restituée (e.g. locomotion, sauts avec musculaire lors d’actions excentriques est supérieur au couple
prise d’élan) [24]. de force produit en contraction isométrique et concentrique
(pour une vitesse angulaire donnée), atteignant 100 à 180 % de
Ces observations ont amené plusieurs auteurs à comparer le la force maximale isométrique selon les études [54]. Toutefois,
comportement du muscle et de ses tendons à celui d’un ressort certains travaux ne rapportent pas de différences significatives
qui emmagasine et restitue de l’énergie, de manière cyclique entre les couples de force maximaux excentrique et isomé-
[105]. trique. Il existe des divergences concernant l’allure de la
Lors d’une contraction excentrique, les composantes relation entre le couple de force et la vitesse de mouvement
élastiques du complexe musculo-tendineux contribuent ainsi (« force–vitesse ») établie à l’aide d’ergomètres isocinétiques,
au couple de force externe produit. Ce dernier est plus élevé que lors de sollicitations excentriques. La plupart des études
celui développé en concentrique pour une vitesse angulaire s’accordent néanmoins sur le fait que le couple de force
donnée [41]. L’allongement musculaire forcé associé à ces excentrique maximal ne dépend pas de la vitesse de mouvement
tensions importantes provoque une dégradation du cytosque- (Fig. 1) [54]. Toutefois, l’allure de la relation « force–vitesse »
lette et une inflammation localisée responsable de douleurs
musculaires à effet retardé communément appelées delayed
onset muscular soreness (DOMS) [65]. Afin de protéger le
système musculosquelettique de ces dommages, des méca-
nismes d’inhibition sont mis en place par le système nerveux.
Cette réponse spécifique du système neuromusculaire en
condition excentrique a amené de nombreux auteurs à
s’intéresser aux effets chroniques de l’exercice excentrique
dans le cadre de l’entraı̂nement [110] et de la rééducation [103].
Ce régime de contraction intéresse directement ces deux
champs d’application car il va permettre de solliciter l’appareil
contractile de manière maximale et supra-maximale, induisant
ainsi des gains de force importants, mais aussi de renforcer le
complexe musculo-tendineux en stimulant la synthèse de
collagène [91] et en activant les gènes du développement et de
la croissance cellulaire [49]. Fig. 1. Évolution du couple de force normalisé par rapport à la force maximale
L’exercice excentrique est classiquement réalisé à vitesse volontaire isométrique en fonction de la vitesse angulaire de mouvement selon
plusieurs études ayant caractérisé la relation « force–vitesse ». MVC : contrac-
constante (isocinétique [IK]) ou contre une charge externe tion isométrique maximale ; ~ Webber & Kriellaars, 1997 [127] ; * Westing
constante (isotonique [IT]), induisant des contraintes mécani- et al., 1998 [129] ; ~ Komi et al., 2000 [59] ; * Dudley, 1990 [28] ; ^ Colson
ques différentes selon le cas, ce qui devrait engendrer des et al., 1999 [19].
serait dépendante du sexe, de l’âge ou encore du niveau de force neuromusculaires. L’amplitude du réflexe H est liée au nombre
des sujets [18]. de motoneurones activés et reflète ainsi l’excitabilité de ce pool
motoneuronal et la transmission synaptique des afférences Ia. Il
2.2.2. Stratégies d’activation musculaire constitue un outil permettant d’évaluer une modulation de
Lors de sollicitations musculaires excentriques, le système l’activité réflexe monosynaptique. Lors de contractions sous-
nerveux met en place des stratégies d’activation centrale maximales excentriques, ce réflexe H est plus faible que lors de
spécifiques à ce mode d’action. En effet des mesures contractions concentriques, suggérant l’existence d’une inhibi-
électroencéphalographiqes indiquent une activité corticale plus tion pré-synaptique des afférences Ia [26,84,111]. Ce processus
importante et plus précoce lors de contractions excentriques peut aussi expliquer l’activation limitée qui est observée en
sous-maximales et maximales des fléchisseurs de l’avant-bras contraction excentrique.
[34], ce qui suggère une programmation des contractions Les mécanismes qui mettent en évidence une activation
excentriques différente des contractions concentriques, au musculaire spécifique lors de contractions excentriques [32]
niveau du système nerveux central. L’activité électromyogra- impliquent des modulations du recrutement et de la fréquence
phique (EMG) mesurée à la surface de la peau, lors d’actions de décharge des UMs. Tandis que certaines études rapportent
musculaires excentriques maximales, est par ailleurs inférieure qu’une contraction excentrique est associée à une activation
à celle enregistrée en concentrique, pour une vitesse angulaire sélective d’UMs à seuil de recrutement élevé [48,67,81],
donnée [1,58,67,130]. Le système nerveux central semble ainsi d’autres travaux montrent un ordre de recrutement suivant le
dans l’incapacité de recruter de façon maximale toutes les UMs principe de la taille [38], pour les contractions concentriques et
d’un muscle lors d’une contraction excentrique [42,73], et cela excentriques [23,89,120,124]. Si Stotz et Bawa [124] ont
est plus marqué à des vitesses angulaires élevées [55]. Cette observé le recrutement d’UMs à seuil de recrutement élevé lors
réduction du signal EMG pourrait être le résultat d’une de contractions excentriques, cela n’a été obtenu que pour des
diminution de la fréquence de décharge [120] ou d’un profils de force ou de mouvement variables et non contrôlés.
recrutement d’UMs moins important en excentrique, induisant Ces données suggèrent qu’une modification du recrutement en
la répartition du stress mécanique sur un plus faible nombre contraction excentrique peut être observée ponctuellement sans
d’UMs [7,82]. La tension unitaire par fibre musculaire active que cela témoigne d’un recrutement préférentiel d’UMs
est donc plus élevée en mode excentrique, ce qui participe à la rapides.
réduction du coût énergétique mais aussi à la survenue des Une contraction volontaire des muscles agonistes est
dommages musculaires au niveau des fibres sollicitées [82]. associée à une activation des muscles antagonistes (ou co-
Ces observations ont été confirmées par les résultats obtenus activation), qui peut également être quantifiée par électro-
par la technique de stimulation électrique surimposée (twitch myographie de surface [33,121]. Le couple de force externe est
interpolation), qui ont mis en évidence un degré d’inhibition alors le résultat du couple de force produit par les muscles
musculaire agoniste compris entre 5 et 30 % selon les études, agonistes, soustrait du couple de force généré par les muscles
lors de contractions excentriques [6,8,28,129] contre 4 à 21 % antagonistes. Les résultats concernant l’influence du mode de
en régime concentrique [107]. Certaines hypothèses ont été contraction sur le niveau de co-activation divergent. Celui-ci
proposées pour expliquer les mécanismes responsables de cette pourrait être plus important lors de contractions musculaires
limitation de l’activation musculaire. Les afférences Ib issues excentriques (12 % en excentrique versus 7 % en concentrique
de l’organe tendineux de Golgi (sensible à la tension pour les muscles fléchisseurs plantaires) [96] ou concentriques
musculaire) peuvent, par exemple, avoir une action inhibitrice (37 à 48 % en concentrique versus 25 à 30 % en excentrique ;
sur l’activité des motoneurones a. Néanmoins, l’influence des muscles extenseurs du genou) [56], alors que d’autres travaux
organes tendineux de Golgi ne semble pas être le mécanisme montrent des valeurs de co-activation similaires quel que soit le
majeur permettant d’expliquer cette diminution de l’activation type de sollicitation musculaire [6,99]. Un tel mécanisme
en contraction excentrique [96]. La commande efférente peut diminue les niveaux de couple de force développés et permet de
également être modulée par les afférences Ia et II, issues des stabiliser l’articulation vis-à-vis des tensions élevées induites
fuseaux neuromusculaires qui renseignent le système nerveux par les sollicitations excentriques.
sur la longueur (sensibilité statique) et les variations de
longueur (sensibilité dynamique) du muscle. Les récepteurs 2.2.3. Comportement des structures musculaires
sensoriels périphériques exercent ainsi une rétroaction accrue Le niveau de pré-activation précédant une contraction
sur les motoneurones a lors de contractions excentriques, qui musculaire excentrique aurait un effet sur l’évolution de la
semble être annulée par l’inhibition présynaptique centrale et longueur des fascicules musculaires. Des études in vitro sur des
périphérique des afférences Ia, comme le montre les travaux fibres isolées suggèrent que le développement d’une force
portant sur le réflexe de Hoffman (ou réflexe H) en conditions isométrique est nécessaire avant tout changement de longueur
dynamiques [25,26]. Le réflexe H est une composante du afin d’assurer une pré-activation complète des UMs [31]. Lors
réflexe d’étirement qui se caractérise par la réponse électro- de contractions excentriques, l’angle de pennation du tibialis
myographique enregistrée lors d’une contraction musculaire anterior est indépendant de la vitesse angulaire de mouvement
provoquée par une stimulation électrique sous-maximale du [104]. Toutefois, pour des niveaux de vitesse angulaire élevés,
nerf moteur. Ce réflexe est le résultat de l’activation des l’angle de pennation mesuré en contraction excentrique est plus
motoneurones a via les afférences Ia provenant des fuseaux important et la longueur des fascicules plus faible par rapport à
une contraction concentrique, induisant une épaisseur muscu- l’orientation linéaire de la ligne Z (i.e. aspect « distordu » [65]),
laire plus importante en excentrique. Néanmoins, le comporte- associées à des lésions au niveau du sarcolemme, des tubules
ment mécanique des fascicules et du tendon pourrait être transverses et du reticulum sarcoplasmique [65]. Ces micro-
également lié à la modalité, à l’intensité de l’exercice lésions sont succédées d’un processus inflammatoire respon-
excentrique, aux muscles et aux systèmes articulaires sable de l’apparition progressive de douleurs musculaires à
considérés [104]. L’ensemble des études confirme toutefois effet retardé ou delayed onset muscular soreness (DOMS).
l’importante contribution des tissus tendineux à l’allongement L’exercice excentrique est le principal initiateur de ces douleurs
du complexe muscle-tendon, estimée à 45 % pour le communément appelées « courbatures ». Elles surviennent
gastrocnemius medialis [16], même s’il existe de fortes dans les 12 à 48 heures qui suivent un exercice excentrique
variations inter-individuelles. Les différentes évolutions de la intense ou inhabituel et connaissent un pic entre 24 et 72 heures
longueur fasciculaire au cours de la contraction (i.e. avant de disparaı̂tre progressivement en cinq à sept jours. Les
raccourcissement, allongement ou longueur constante) seraient DOMS sont accompagnés d’une diminution des capacités de
liées au type d’exercice réalisé, qui influencerait l’étirement des production de force ( 10 à 60 % sur la force maximale
composantes élastiques et optimiserait l’efficacité du cycle volontaire isométrique, selon les études [74,80]), qui sont
étirement–détente. L’énergie alors restituée par les compo- restaurées en une semaine environ. L’impact des dommages
santes élastiques en mode excentrique produit une force musculaires lors d’une seconde session d’exercice excentrique
additionnelle sans consommation d’ATP. Ce processus, associé est cependant réduit par rapport à la première séance (i.e. effet
à un recrutement limité d’UMs, permet une économie de protecteur, Section 2.3.1.1) [74].
l’énergie métabolique totale utilisée par le muscle [3]. Les De nombreux auteurs ont cherché à savoir si les dommages
actions excentriques produisent ainsi des forces plus élevées induits par l’exercice excentrique touchaient de manière
tout en consommant moins d’oxygène et d’énergie qu’une préférentielle un type de fibre particulier. Les résultats de
session équivalente de travail concentrique [88]. Le stockage plusieurs études, obtenus chez l’homme et l’animal, tendent à
d’énergie réside dans les propriétés élastiques du tendon et du prouver que l’exercice excentrique intense engendre des lésions
muscle, qui permettent l’allongement de ces structures en principalement au niveau des fibres de type II (II b en
réponse à l’étirement. Le tendon possède un comportement particulier) [36]. Cela peut en partie être expliqué par le
viscoélastique qui résulte de sa composition, riche en collagène recrutement préférentiel d’UMs rapides en régime excentrique.
(86 % de la masse du tissu organique), et des interactions Les processus physiologiques de régénération cellulaire qui
existant entre les proteines collagéniques et non-collagéniques succèdent à ces microlésions sont impliqués dans l’hyper-
(i.e. protéoglycanes). L’élasticité du muscle est due, quant à trophie musculaire et sont donc nécessaires pour optimiser la
elle, aux interactions actine–myosine, au tissu conjonctif réponse à l’entraı̂nement (Section 2.3.1.3). La mise en place de
extracellulaire et au système élastique de filaments structurant contractions musculaires excentriques dans les programmes de
les sarcomères des cellules musculaires. Ce complexe élastique renforcement musculaire, si elle nécessite certaines précau-
est par exemple composé d’une protéine géante, la titine (aussi tions, reste donc un moyen intéressant pour améliorer les
appelée connectine) et de la desmine, qui remplissent la qualités de force musculaire et permet, tout en affectant la
fonction de ressort lorsque le muscle est soumis à une commande motrice, d’induire en retour des processus
sollicitation excentrique [66]. adaptatifs sur le plan structural.
Les propriétés viscoélastiques du tendon le rendent plus
déformable que le muscle à de faibles niveaux d’étirements. 2.3. Adaptations induites par un entraı̂nement excentrique
Dans ces conditions, les tendons peuvent absorber plus
d’énergie mais sont moins efficaces dans la transmission des 2.3.1. Adaptations structurales
forces. À des niveaux de tension plus élevés, ils deviennent 2.3.1.1. Effet de répétition de session ou « repeated bout
moins déformables et plus raides, permettant alors de déplacer effect ». De nombreuses études ont montré que la réalisation
des charges plus importantes. Ces éléments pourraient d’une première séance d’exercice excentrique a pour effet de
expliquer que lors de sollicitations excentriques, la longueur limiter fortement les DOMS perçus à la suite d’une seconde
des fascicules musculaires du gastrocnemius medialis reste session d’exercice excentrique, réalisée classiquement à une
constante alors que les fascicules musculaires du vastus semaine d’intervalle. Ce phénomène est appelé « effet de
lateralis s’allongent, car soumis à des contraintes mécaniques répétition » (repeated bout effect [74]). Cette première
plus importantes. adaptation se caractérise par une réduction des microlésions
musculaires, du relargage de créatine kinase (CK) et des
2.2.4. Dommages musculaires chaı̂nes lourdes de myosine, de la perte de force et des DOMS
Le régime excentrique permet d’imposer au système après une seconde séance d’exercice excentrique, par rapport à
musculo-tendineux une contrainte mécanique élevée ou dite la première session réalisée quelques jours plus tôt. La durée de
« supra-maximale ». Les hauts niveaux de tensions qui ce phénomène adaptatif varie de quelques semaines à six mois
accompagnent l’allongement forcé de ce complexe peuvent en fonction des études. Selon Chen et [14], l’importance des
entraı̂ner des microlésions des fibres musculaires [20]. Au DOMS et du niveau de CK plasmatique après la première
niveau cellulaire, ces dommages se caractérisent par une session d’exercice excentrique influence le degré de l’effet de
désorganisation de certains sarcomères et une perturbation de répétition. Ainsi, il est possible que le niveau de sensibilité du
sujet à l’exercice excentrique conditionne l’importance de cet notamment par rapport à un entraı̂nement concentrique (voir
effet protecteur qui peut parfois ne pas être observé [100]. Dans Remaud et al. [107] pour une revue détaillée). Par exemple,
une revue récente, Lieber et Friden [65] évoquent également le Hortobagyi et al. [43] rapportent un gain de force isométrique
haut degré de spécificité de l’effet de répétition, qui n’est volontaire maximale 3,5 fois plus important après six semaines
valable que lorsque les deux sessions excentriques sont d’entraı̂nement excentrique des extenseurs du genou en
identiques. comparaison d’un entraı̂nement concentrique de même durée.
Les adaptations pouvant potentiellement expliquer l’effet de Ces résultats ont été confirmés avec des durées d’entraı̂nement
répétition sont d’origine nerveuse, mécanique ou cellulaire. En plus longues (+24,6 % de la force initiale après entraı̂nement
raison de la rapidité de ce type d’adaptation, certains auteurs excentrique versus +12,5 % après entraı̂nement concentrique)
ont suggéré que l’exercice excentrique pouvait améliorer la pour les fléchisseurs et extenseurs du coude [83]. Les études
synchronisation des UMs et accroı̂tre leur degré d’activité [74]. s’étant intéressées aux effets d’un entraı̂nement utilisant une
Ce processus permettrait d’augmenter le nombre de fibres surcharge (100 à 120 % de la répétition maximale) lors de la
sollicitées par rapport à la première session, notamment en phase excentrique (overloading) ont mis en évidence des gains
recrutant un plus grand nombre d’UMs lentes, entraı̂nant une de force plus importants qu’avec un entraı̂nement en force
meilleure répartition du stress mécanique sur l’ensemble des utilisant des charges inférieures [45]. Ainsi, il semble que la
fibres musculaires activées et limitant ainsi les ruptures des composante excentrique d’un protocole d’entraı̂nement soit
myofibrilles [22,82]. L’origine mécanique des lésions induites essentielle pour augmenter la force musculaire. En effet,
par l’exercice excentrique a amené certains auteurs à proposer l’appareil contractile est sollicité au maximum, grâce aux hauts
que des modifications des propriétés mécaniques du système niveaux de couple de force atteints en excentrique. Les gains de
musculosquelettique protégeraient le muscle vis-à-vis des force ainsi obtenus par rapport aux capacités de production de
dommages. Certaines études ont en effet rapporté une force initiales s’échelonnent de 1 à 116 % pour la force
augmentation de la raideur active ou passive du complexe excentrique, de 1 à 67 % pour la force concentrique et de 7 à
muscle-tendon après un exercice excentrique [98]. Ces 45 % pour la force isométrique [43,86]. Si de nombreux auteurs
adaptations ont été respectivement attribuées à une réorganisa- restent attachés au principe de spécificité des gains de force en
tion des protéines du cytosquelette responsables de l’aligne- fonction du mode d’entraı̂nement [30], il apparaı̂t que
ment et de la structure des sarcomères (i.e. titine, desmine) et à l’entraı̂nement excentrique soit le moyen le plus efficace
une augmentation de la quantité de tissu conjonctif présente d’améliorer la force maximale concentrique d’un groupe
dans le muscle, permettant une meilleure dissipation de la musculaire [52]. Enfin, les gains de force induits par
contrainte mécanique en excentrique [64]. Toutefois, des études l’entraı̂nement excentrique semblent très spécifiques de la
montrent également que des muscles plus raides sont plus vitesse de mouvement utilisée [110].
susceptibles de subir des dommages [72]. En effet, des travaux
portant sur des modèles « murins » ne possédant pas de desmine 2.3.1.3. Hypertrophie. Un des processus adaptatifs impor-
ont montré que ces souris étaient moins exposées aux tants induit par l’entraı̂nement excentrique est le gain de masse
dommages que des souris saines suite à un exercice musculaire (i.e. hypertrophie), qui représente un des facteurs
excentrique. Ce résultat a été attribué à une compliance plus déterminants des capacités de production de force. Au-delà de
importante des muscles de ces souris, limitant ainsi le nombre six à huit semaines d’entraı̂nement, l’amélioration des
de sarcomères susceptibles d’être lésés. La dernière hypothèse capacités de production de force musculaire s’accompagne
avancée pour expliquer l’effet de répétition est l’influence de d’une augmentation de la masse musculaire. La composante
facteurs cellulaires, selon laquelle l’exercice excentrique excentrique d’un entraı̂nement est alors essentielle pour induire
entraı̂ne une augmentation du nombre de sarcomères en série l’augmentation de la surface de section transversale des fibres
et modifierait la réponse inflammatoire. L’addition de musculaires [29]. Les programmes incluant des actions
sarcomères en série, montrée chez l’animal [79] et récemment excentriques engendrent ainsi des gains de masse musculaire
de manière indirecte chez l’homme [13], permettrait de réduire plus importants que les entraı̂nements visant cet objectif sur le
l’étirement et donc la rupture des sarcomères après une session mode concentrique [35,58]. Higbie et al. [40] ont par exemple
excentrique. Une réduction de l’inflammation après un exercice montré une augmentation plus importante de la surface de
excentrique répété a également été caractérisée par une section transversale (CSA) du quadriceps après dix semaines
diminution de l’activation des monocytes et des neutrophiles d’entraı̂nement excentrique par rapport à un entraı̂nement
participant à la réponse inflammatoire [97]. Cette réponse équivalent réalisé en régime concentrique. Des résultats
inflammatoire réduite contribue à l’effet protecteur même s’il similaires ont été obtenus sur les fléchisseurs du coude [35].
est difficile de savoir si cette adaptation est la cause ou la Ces résultats ont récemment amené de nombreux chercheurs à
conséquence d’une diminution du nombre de myofibrilles s’intéresser aux effets de l’entraı̂nement excentrique sur
lésées, ou encore si l’effet de répétition est une combinaison de l’expression génique des cellules musculaires sollicitées. En
ces deux processus [74]. effet, l’exercice excentrique représente, pour les cellules
musculaires, une contrainte mécanique particulière qui peut
2.3.1.2. Gains de force. Selon plusieurs études, l’entraı̂ne- modifier l’expression de plusieurs gènes [39,57], via les voies
ment excentrique semble être un moyen efficace pour de mécanotransduction constituées de protéines sensibles au
augmenter les capacités de production de force maximale, statut mécanique de la cellule musculaire (e.g. microtubules-
associated proteins ou protéines MAP). Une séance excen- [12], d’autres protocoles utilisant l’overloading [116] ou des
trique déclenche, à partir de six heures post-effort, l’activation vitesses élevées sur le mode pliométrique [11] ont induit une
progressive des gènes de la croissance cellulaire et du augmentation de la longueur des fascicules musculaires
développement, impliqués dans les processus d’hypertrophie (addition de sarcomères en série). À notre connaissance, une
cellulaire chez l’homme. L’expression de ces gènes est seule étude a étudié l’effet de l’entraı̂nement purement
d’avantage stimulée par les actions excentriques que par des excentrique sur l’architecture musculaire. Les auteurs ont
actions isométriques ou concentriques [9,15,60]. L’introduction mis en évidence une augmentation de 3,1 % de la longueur des
d’exercice excentrique lors des périodes d’entraı̂nement dites fascicules du vastus lateralis après un entraı̂nement excentrique
« d’hypertrophie » semble donc être un moyen efficace pour contre 6,3 % pour un entraı̂nement concentrique de même durée
optimiser la prise de masse musculaire. sans différence significative entre les deux modes d’entraı̂ne-
ments [12]. Les auteurs ont suggéré que l’amplitude de
2.3.1.4. Modifications typologiques. Les modifications typo- mouvement, plus que le mode de contraction ou la vitesse de
logiques induites par l’entraı̂nement excentrique sont similaires mouvement, constitue le facteur ayant l’effet le plus important
à celles induites par les autres programmes d’entraı̂nement en sur les modifications de longueur des fascicules musculaires.
force, c’est-à-dire une augmentation de la part de fibres L’augmentation de la longueur des fascicules musculaires
musculaires de type intermédiaire (IIa, IIa/IIb) pouvant permet également d’accroı̂tre la vitesse de contraction
atteindre +12 % [43], avec parfois une diminution du nombre musculaire [12]. Ainsi, l’entraı̂nement excentrique constitue
de fibres de type IIb [118]. Cependant ces évolutions restent un procédé efficace pour améliorer non seulement les capacités
modérées, voire ne sont pas statistiquement significatives dans de production de force, mais aussi les qualités de puissance
la plupart des études. Elles ne semblent donc pas constituer musculaire (i.e. produit de la force et de la vitesse de
l’adaptation principale pouvant expliquer les gains de force contraction musculaires), essentielles dans la plupart des
observés après un entraı̂nement excentrique. activités physiques et sportives.
2.3.1.5. Modifications de l’architecture musculaire. Des 2.3.1.6. Modifications des propriétés mécaniques. Classique-
études récentes ont cherché à identifier les modifications de ment, la réalisation d’une séance d’exercice excentrique est
l’arrangement géométrique des fibres musculaires induites par suivie d’une augmentation de la raideur musculo-articulaire
des entraı̂nements en force combinant des actions concen- passive [47]. Celle-ci peut être multipliée par deux et reste
triques et excentriques [53], des entraı̂nements excentriques élevée pendant les quatre jours suivant l’exercice excentrique.
[12] ou des entraı̂nements pliométriques [11]. Une période Cette augmentation de la raideur passive peut potentiellement
d’entraı̂nement en force permet d’augmenter l’angle de s’expliquer par la formation de ponts actine-myosine résiduels
pennation des fascicules musculaires, processus associé à [62] résultats d’une augmentation de la concentration calcique
l’hypertrophie musculaire [2]. En effet, l’angle de pennation est au repos dans les fibres musculaires, elle-même liée à la rupture
modifié par l’augmentation du volume musculaire, et son des membranes des cellules musculaires. D’autres processus
évolution est fonction de l’espace inter-aponévroses du muscle ont été proposés pour expliquer cette augmentation de la
hypertrophié. Récemment, une augmentation de 21,4 % de raideur passive tel que l’œdème caractérisé par un gonflement
l’angle de pennation du vastus lateralis du quadriceps a été musculaire qui compresse les tissus et engendre une résistance
montrée après un entraı̂nement excentrique de dix semaines douloureuse à l’extension passive augmentant ainsi la « sensa-
(contre +13,3 % après un entraı̂nement concentrique) [12]. Une tion de raideur » [17]. Enfin, il est possible que les processus de
augmentation de l’angle de pennation engendre une diminution réparation induisent une augmentation permanente de la
de la force transmise par une fibre musculaire considérée de raideur passive en raison du remodelage du tissu conjonctif
manière isolée. En revanche, en termes de CSA musculaire, la comme suggéré par Lapier et al. [64].
pennation des fibres musculaires engendre une CSA physio- Lorsque l’exercice excentrique est répété et prolongé (i.e.
logique du muscle, qui est définie comme l’aire des fibres entraı̂nement), la raideur musculo-articulaire caractérisée en
musculaires perpendiculaires à l’axe longitudinal de chaque condition passive diminue. Mahieu et al. [68] ont mis en
fibre musculaire multipliée par le cosinus de l’angle de évidence une augmentation de l’amplitude articulaire de
pennation [132], supérieure à la CSA anatomique du muscle. mouvement et une diminution de 23 % du couple de force
Puisque la CSA physiologique représente le nombre maximal résistif passif après six semaines d’entraı̂nement excentrique
de ponts actine-myosine en parallèle qui peuvent être activés en des fléchisseurs plantaires. Antérieurement, Morgan [79] avait
contraction, la capacité de production de force maximale d’un déjà montré une augmentation du nombre de sarcomères en
muscle est proportionnelle à cette CSA physiologique et donc à série dans les fibres musculaires (+11 %), après une semaine de
l’angle de pennation. L’augmentation de l’angle de pennation course en descente sur tapis roulant. Ces résultats sont en
permet d’augmenter la quantité de matériel contractile en accord avec des études récentes mettant en évidence une
parallèle, améliorant ainsi les capacités de production de force modification de la relation entre le couple de force maximal
[2]. développé et l’angle articulaire avec un déplacement du pic de
En ce qui concerne la longueur des fascicules musculaires, si couple de force (shift d’environ 158) vers des longueurs
certains protocoles d’entraı̂nement n’ont pas montré de musculaires plus grandes immédiatement après un exercice
modification de ce paramètre après un entraı̂nement en force excentrique [131]. Cette modification de la longueur muscu-
laire permettant la production du couple de force maximal est au faible degré d’activation musculaire en excentrique avant
un indicateur de l’augmentation de la compliance musculaire entraı̂nement, en comparaison des autres modes de contrac-
post-exercice excentrique. Cette adaptation induite par tion (i.e. isométrique et concentrique) ;
l’entraı̂nement excentrique permet d’envisager des applications à une augmentation de l’activité EMG des fibres spécifique-
du point de vue de la prévention des blessures. En effet, la ment recrutées par l’exercice excentrique.
diminution de la raideur passive du muscle permet un transfert
d’une partie de la contrainte mécanique du tendon vers le Il semble que les fibres de type II soient préférentiellement
muscle, réduisant ainsi les risques de lésion tendineuse. recrutées lors d’un entraı̂nement excentrique. Nous avons vu
En revanche, la raideur de la composante élastique en série également que l’exercice excentrique répété pouvait conduire à
(CES), composée des structures musculaires élastiques et du un recrutement progressif des UMs lentes afin de mieux répartir
tendon, augmente après un entraı̂nement excentrique. Pousson le stress mécanique en vue de protéger le muscle vis-à-vis des
et al. [98] ont en effet montré une augmentation de ce paramètre dommages. Ces processus pourraient être à l’origine d’une
après six semaines d’entraı̂nement excentrique des muscles augmentation de l’activité EMG du muscle entraı̂né en régime
fléchisseurs du bras, à 30 et 45 % de la force maximale excentrique. L’augmentation de la fréquence de décharge des
volontaire isométrique (MVC). Cette augmentation n’est UMs peut également expliquer ce gain d’activité EMG.
cependant pas confirmée à des niveaux de force plus élevés Cependant, même si la fréquence de décharge peut être
(60 et 80 % de MVC). Ces résultats sont partiellement augmentée par l’entraı̂nement, seulement 15 % de la force
confirmés par les travaux de Mahieu et al. [68] qui ont maximale est due à une modulation de ce paramètre dans les
déterminé une tendance à l’augmentation de 9,5 % de la raideur muscles de taille importante [93]. Récemment, des travaux ont
du tendon d’Achille mesurée en conditions actives après six été entrepris afin de préciser l’origine de ces différentes
semaines d’entraı̂nement excentrique des fléchisseurs plan- adaptations nerveuses à travers l’analyse de l’évolution de
taires. L’augmentation de la raideur de la CES peut donc l’onde V et du réflexe H après une période d’entraı̂nement
s’expliquer par une augmentation de la raideur du tendon. Elle excentrique des fléchisseurs plantaires [27]. Une augmentation
pourrait également être le résultat d’une augmentation de la de l’onde V peut résulter d’une augmentation du nombre
raideur des ponts d’actine–myosine ou d’un remodelage des d’UMs recrutées ou d’une augmentation de la fréquence de
protéines du cytosquelette responsables de l’alignement des décharge des UMs. Cette augmentation de la commande
sarcomères, telle que la desmine ou la titine. La réorganisation centrale descendante dépend également de l’efficacité de la
de ces protéines succédant aux microlésions induites par les transmission nerveuse au niveau de la plaque motrice. Duclay
sollicitations excentriques pourrait être un processus respon- et al. [27] ont mis en évidence une augmentation constante de
sable de l’augmentation de la raideur de la composante l’onde V lors de contractions volontaires au cours de la période
élastique du muscle. Ces modifications des propriétés d’entraı̂nement excentrique, soulignant une amélioration de
élastiques peuvent ainsi favoriser la restitution d’énergie l’excitabilité des motoneurones a ou une réduction de
mécanique, notamment lors de cycles d’étirement–détente. l’inhibition présynaptique des afférences Ia. Les gains de
Par ailleurs, l’exercice excentrique chronique est souvent force pourraient également être la conséquence d’une diminu-
proposé dans le traitement des tendinopathies pour son effet tion de l’activité des muscles antagonistes (co-activation). Sur
stimulant sur la synthèse de collagène, principal élément le plan nerveux, un degré de co-activation moindre permet de
constitutif du tissu tendineux et conjonctif [91]. Plusieurs réduire le niveau d’inhibition réciproque et ainsi faciliter le
études ont également montré une augmentation du flux sanguin réflexe H. Néanmoins, l’entraı̂nement excentrique ne semble
en périphérie des cellules tendineuses après exercice excen- pas avoir d’impact sur le niveau de co-activation, celui-ci
trique [5,63]. Ces processus adaptatifs justifieraient l’intérêt de restant stable entre le début et la fin de la période
l’exercice excentrique dans le renforcement des tissus d’entraı̂nement dans la plupart des études [19,115]. L’inhibition
tendineux chez les patients souffrant de tendinopathies. réciproque ne peut donc pas a priori expliquer les modifications
Toutefois certaines questions demeurent, notamment quant du réflexe H observées [27]. Toutefois, les effets de
aux protocoles spécifiques (e.g. charge, vitesse de mouvement) l’entraı̂nement excentrique sur l’amplitude du réflexe H sont
à utiliser en fonction des caractéristiques de la pathologie [103]. différents en fonction du muscle considéré (i.e. pour le soleus et
le medial gastrocnemius), ce qui suggère une organisation des
2.3.2. Adaptations nerveuses boucles réflexes spécifiques au muscle. L’origine des diffé-
L’activité électrique d’un muscle effectuant une contraction rentes adaptations nerveuses induites par l’entraı̂nement
excentrique est inférieure à celle enregistrée lors d’une action excentrique reste donc à préciser en fonction du groupe
concentrique pour un même niveau de force produite (Section musculaire considéré.
2.2). Cette activation neuromusculaire peut être augmentée par
l’entraı̂nement excentrique. Hortobagyi et al. [43] rapportent 2.3.3. Place de l’exercice excentrique dans la rééducation
ainsi une augmentation de l’activité EMG 2,6 fois plus L’exercice excentrique permet de travailler à des niveaux de
importante après six semaines d’entraı̂nement excentrique du couple de force importants, avec un coût énergétique et une
quadriceps, par rapport à un entraı̂nement concentrique consommation en oxygène moindre. Ce mode d’exercice est
équivalent. Des résultats similaires ont été obtenus sur les donc d’intérêt pour les cliniciens qui souhaitent inclure, dans
fléchisseurs du coude [58]. Ce résultat peut être dû : leurs programmes de rééducation fonctionnelle des patients qui
présentent le plus souvent une intolérance à l’effort cardio- responsable du remodelage du tendon à l’image des résultats
respiratoire intense (i.e. patients atteints de troubles cardiaques obtenus sur les tissus osseux à l’aide de signaux haute
ou de pathologies pulmonaires). Nous avons précédemment fréquence [112]. Ces variations sont reflétées par la difficulté
évoqué l’effet protecteur que représente une séance d’exercice des sujets à contrôler le mouvement lors d’une contraction
excentrique vis-à-vis des dommages musculaires. Chez des excentrique. Ainsi, si l’exercice excentrique est couramment
sportifs reprenant l’activité physique après une blessure, utilisé dans le traitement des tendinopathies, certaines
l’entraı̂nement excentrique sous-maximales est donc un moyen questions demeurent, notamment vis-à-vis des protocoles
pertinent pour préparer le système musculosquelettique à la spécifiques (e.g. mode de contraction, charge, vitesse de
reprise de l’effort et éviter la survenue de nouvelles lésions lors mouvement) à utiliser en fonction des caractéristiques de la
de cette phase de réentraı̂nement. Il est aussi utilisé dans la prise pathologie [63,103]. Enfin, il semble que l’exercice excentrique
en charge d’autres pathologies, telles que la sclérose en unilatéral augmenterait la force des muscles du membre
plaques, pour son effet positif sur le contrôle moteur [108]. controlatéral, et cela de manière plus marquée qu’après un
Initialement suggéré par Stanish et al. (1986), l’exercice entraı̂nement concentrique [40,44]. L’introduction de contrac-
excentrique chronique est également proposé dans le traitement tions excentriques dans les programmes de rééducation permet
des tendinopathies, notamment du tendon d’Achilles et du ainsi une amélioration fonctionnelle des groupes musculaires
tendon patellaire. Ces pathologies représentent une large part du membre opposé et lésé sans avoir besoin de le solliciter.
de la médecine du sport [109] et un enjeu de santé publique
puisqu’elles affectent également la population générale 2.4. Comparaison des exercices excentriques isotonique et
(environ 2 % de la population active). Le programme de isocinétique
Stanish utilisait une augmentation progressive de la vitesse de
mouvement en contraction excentrique et a été testé sur une 2.4.1. Spécificités des modes isotonique et isocinétique
cohorte de 200 patients. Les résultats ont montré une 2.4.1.1. Mode excentrique isotonique (IT). In vivo, la
restauration complète de la fonction motrice pour 44 % d’entre sollicitation excentrique IT consiste à opposer un couple de
eux. S’inspirant des travaux de Stanish, Alfredson et al. [5] ont force musculaire à une charge constante, qui peut dépasser les
montré que la composante excentrique constituait un instru- capacités de production de force du sujet (i.e. exercice
ment thérapeutique essentiel de la réponse obtenue à la suite de excentrique supra-maximal [45]), entraı̂nant l’allongement
ces programmes de rééducation. du complexe muscle-tendon. En condition excentrique IT, la
La pathophysiologie du tendon atteint ou en cours de charge constante est le plus souvent imposée par une barre ou
traitement n’est pas complètement connue. Il apparaı̂t des poids, procédés classiquement utilisés en entraı̂nement de
cependant que lors d’une tendinopathie, le tendon ne parvient la force. Si la charge dans ce type d’exercice est fixée, le couple
pas à entrer dans un cycle actif de guérison [103]. L’exercice de force exercé par le sujet varie, notamment en raison des
excentrique pourrait constituer un stimulus mécanique efficace accélérations et décélérations qui s’appliquent à la charge
aux cellules tendineuses quiescentes. D’autres études ont imposée, et aux variations de longueur du bras de levier en
montré une interruption du flux sanguin en périphérie des fonction des appareils [4]. Cette modalité d’exercice constitue
cellules tendineuses lors d’un exercice excentrique [5]. Ce donc plutôt une sollicitation mécanique iso-inertielle, mais qui
processus pourrait endommager les microcapillaires et les nerfs est classiquement désignée sous le terme « isotonic » dans la
associés, réduisant ainsi la sensation de douleur. Récemment, littérature [37]. La vitesse angulaire de mouvement lors de ce
Langberg et Kongsgaard [63] ont mis en évidence une type de mouvement est variable et les niveaux de contrainte
augmentation de la synthèse de collagène (collagène de type mécanique enregistrés sont très élevés (jusqu’à 140 % de la
I) principal élément constitutif du tissu tendineux, sans répétition maximale pour des sujets entraı̂nés), notamment aux
augmentation des processus de dégradation, au sein du tendon angles articulaires extrêmes (i.e. début et fin d’amplitude de
lésé après 12 semaines d’exercice excentrique. Les auteurs ont mouvement). Ainsi, une contraction excentrique IT comporte
également observé une diminution de la douleur perçue par les une première phase d’amortissement au cours de laquelle la
patients. D’autres travaux ont montré une augmentation du flux force imposée par la masse dépasse la force musculaire, suivie
sanguin ou du volume du tendon, suggérant une amélioration d’une phase lors de laquelle le sujet contrôle la charge,
des échanges entre les tissus et le milieu interstitiel [117]. Ces jusqu’aux angles extrêmes où le contrôle de la charge devient
processus adaptatifs justifieraient l’intérêt de l’exercice plus difficile. Ce pattern de mouvement suggère que la
excentrique dans le renforcement des tissus tendineux chez contrainte IT sollicite de manière préférentielle les structures
les patients atteints. Toutefois, peu d’études ont comparé ces élastiques à ces angles extrêmes, le muscle étant d’avantage mis
résultats aux effets induits par l’exercice concentrique. Les à contribution aux angles proches du pic de couple de force. La
premières études comparatives ont montré que les pics de force sollicitation excentrique permettant d’augmenter la raideur
générés in vivo au sein du tendon lors d’un exercice excentrique musculo-articulaire, ce mode d’action pourrait ainsi être
étaient similaires lors d’un exercice concentrique [102,113]. privilégié en vue d’accroı̂tre la raideur au niveau des structures
Ces résultats montrent que les bénéfices thérapeutiques de élastiques à ces angles articulaires. En revanche, l’activation
l’exercice excentrique ne sont pas seulement dus aux niveaux musculaire n’est pas maximale sur l’ensemble de l’amplitude
de force induits par ce mode d’exercice. D’autres auteurs ont de mouvement, puisqu’à l’angle au pic de couple, les capacités
suggéré que les variations de force produite seraient un stimulus de production de couple de force musculaire peuvent parfois
dépasser le couple de force imposé par les masses [37,46]. La de force musculaire et impose une résistance adaptée aux
sollicitation excentrique IT pourrait donc avoir un impact capacités du sujet, ce qui offre des garanties de sécurité
modéré sur les adaptations nerveuses et sur le gain de force à cet intéressantes pour les patients pris en charge [21].
angle articulaire.
2.4.2. Gains de force
2.4.1.2. Mode isocinétique (IK). Une contraction excentrique Afin de pouvoir comparer les résultats obtenus suite à des
IK se caractérise par la production d’un couple de force entraı̂nements excentriques IT et IK, les gains de force ont été
musculaire opposé au mouvement d’un bras de levier mobilisé rapportés au nombre de séances d’entraı̂nement effectuées par les
à vitesse constante par un dynamomètre isocinétique (iso- sujets (Tableaux 1a et 1b). Lorsque l’on compare les gains de
cinétisme de mouvement) [54]. Ce type d’appareil induit une force moyens induits par les deux modes d’entraı̂nements
sollicitation maximale du système contractile sur l’ensemble de excentriques, on constate que l’entraı̂nement excentrique IT
l’amplitude de mouvement [8,59], sans accélération impor- augmente les capacités de production de force de 1,1 1,0 % par
tante, comme cela peut être le cas en excentrique IT. Les effets séance [10,19,27,46,49–51,69,70,77,85,90,92,94,98,101,119,
induits sur la raideur musculaire aux angles extrêmes pourraient 122,123,125,126,128] pour une durée moyenne d’entraı̂nement
ainsi ne pas atteindre ceux obtenus après un entraı̂nement de 7,5 3,4 semaines (Tableau 1a) contre 0,6 3,0 %, pour
excentrique IT. Cependant, la force musculaire produite étant l’entraı̂nement excentrique IK (durée moyenne d’entraı̂nement
maximale sur la totalité de l’amplitude, l’activité des muscles de 10,6 4,9 semaines) [12,30,35,40,43,58,71,75,76,78,83,87,
sollicités, est elle aussi théoriquement maximale, et notamment 114,115,118] (Tableau 1b), soit un gain près de deux fois plus
aux angles au pic de couple, suggérant des adaptations important en mode IT. Ainsi, il semblerait que les sollicitations
nerveuses plus marquées avec ce type d’entraı̂nement en IT soient, de ce point de vue, plus efficaces que les exercices
comparaison d’un entraı̂nement excentrique IT. Ce mode réalisés à vitesse constante, en condition excentrique. Ce constat
d’exercice présente par ailleurs un intérêt majeur pour les a déjà été mis en évidence par certaines études [61] et par un
cliniciens qui souhaitent proposer des programmes de réé- travail similaire de bilan comparatif des modalités IT et IK en
ducation fonctionnelle. L’isocinétisme permet en effet un arrêt régime concentrique, évoquant des gains de 0,5 à 1 % en IT
immédiat du mouvement en l’absence de production de couple contre 0,3 % par séance en moyenne en IK [106]. Les différences
Tableau 1a
Gains de force induits par l’entraı̂nement excentrique isotonique (IT).
Caractéristiques de l’étude Gains de force obtenus pour chaque gamme de vitesse de test
(par séance)
1 1 1 1 1
Muscles Auteurs Sem Fq Volume Intensité < 908.s 908.s < 08.s < 908.s 908.s <
(%1RM)
Extenseurs de la jambe Ben-Sira et al. [10] 82 10 3 65–85 - - - +1,0 -
Housh et al. [46] 83 36 80 a - +1,2 - ns ns
Johnson [49] 83 1 10 80 - +2,4 - - -
Johnson et al. [50] 63 26 120 - - - +1,0 -
Jones et Rutherford [51,52] 12 3 36 116 - - +0,2 - -
Kaminski et al. [52] 62 28 100 +2,5 +3,2 - +2,0 +2,1
Pavone et Moffat [90] 63 3 10 40–79a - - +1,3 - -
Raue et al. [101] 43 38 80 - - - +0,6 -
Smith et Rutherford [119] 20 3 3 10 100–110 - - +0,4 +0,3 +0,3
Sorichter et al. [122] 6 1–3 7 10 150 - - +0,7 - -
Spurway et al. [123] 61 36 100 +4,6 +4,3 +1,4 +2,8 +2,4
Weir et al. [128] 83 36 80 a - +1,2 +0,6 +1,4 +0,6
Fléchisseurs de la jambe Johnson et al. [50] 63 26 120 - - - +1,4 -
Mjolsnes et al. [77] 10 1–3 2–3 5–12 PC - +0,4 +0,3 - -
Fléchisseurs du bras Colson et al. [19] 73 56 100–120 +2,6 +1,6 +0,5 +0,9 +1,1
Johnson [49] 83 1 10 80 - +1,4 - - -
Martin et al. [70] 43 35 100 - - +0,4 +1,3 +0,8
Mannheimer [69] 45 25 - - +3,0 - - -
Nosaka et Newton [85] 81 1 10 79 - - +0,7 0 0
Petersen [94] 8 4–5 1 10 120–130b - - +0,2 - -
Pousson et al. [98] 62 2 4–6 90–125 - - - - -
Valour et al. [125] 73 36 57–95 - +0,7 +0,5 +1,3 -
Vikne et al. [126] 12 2–3 36 83–93 - +0,9 - +0,5 -
Johnson et al. [50] 63 26 120 - - - +3,4 -
Extenseurs de l’avant-bras Johnson et al. [50] 63 26 120 - - - +1,6 -
Fléchissuers plantaires Duclay et al. [27] 7 2–3 66 100–120 - +1,7 +1,2 +1,6 -
Pensini et al. [92] 44 66 120 - +1,0 +1,9 +0,9 -
Tableau 1b
Gains de force induits par l’entraı̂nement excentrique isocinétique (IK).
Caractéristiques de l’étude Gains de force obtenus pour chaque gamme de vitesse de test (par séance)
Muscles Auteur Sem Fq Volume Wf (8.s 1) < 908.s 1
908.s 1
< 08.s 1
< 908.s 1
908.s 1
<
Extenseurs de la jambe Blazevich et al. [12] 10 3 4–6 6 30 - +1,3 - +0,5 -
Duncan [30] 63 1 10 120 +1,6 +1,5 - +0,2 -
Higbie et al. [40] 10 3 3 10 60 - +0,1 - +0,2 -
Hortobagyi et al. [43] 12 3 4–6 8–12 60 +3,2 +1,3 +0,1 -
Mayhew et al. [71] 43 5 10 60 - - +0,7 - -
Miller et al. [76] 20 3 1–5 6 60 - +0,7 - +0,4 -
Seger et al. [114] 10 3 4 10 90 +0,3 +1,1 +0,3 +0,2 +0,1
Seger et 10 3 4 10 90 +0,0 +0,9 - +0,0 +0,2
Thorstensson [115]
Melo et al. [75] 12 2 2–4 12 60 - +0,8 - - -
Extenseurs et fléchisseurs Nickols-Richardson 20 3 1–5 6 60 - +0,5 - +0,2 -
de la jambe et al. [83]
Fléchisseurs de la jambe Melo et al. [75] 12 2 2–4 12 60 - - - +0,5 -
c d
Extenseurs de l’avant-bras Farthing et Chilibeck [35] 83 2–6 8 180 +0,7 - +0,9
30 +0,3 c - +0,7 d
Komi et Buskirk [58] 74 16 - - +0,6 +0,3 +0,6 -
Paddon-Jones et al. [87] 10 3 46 180 +0,9 +1,0 +0,7 +1,0 +1,0
30 +0,4 +0,0 +0,3 +0,1 +0,2
Shepstone et al. [118] 83 1–4 10 210 +0,7 +0,2 - +1,2 +2,4
20 +0,4 +0,1 - +0,6 +0,7
Extenseurs et fléchisseurs Nickols-Richardson 20 3 1–5 6 60 - +0,4 - +0,2 -
de l’avant-bras et al. [83]
Rotateurs de l’épaule Mont et al. [78] 63 8 10 90–180 +0,8 c +0,5c - +0,1e +0,5 e
Gains de force obtenus après entraı̂nement excentrique isotonique (IT, Tableau 1a) et isocinétique (IK, Tableau 1b) pour chaque gamme de vitesse de test utilisé :
excentrique à vitesse angulaire de mouvement supérieure à 908.s 1 (< 908.s 1), excentrique à vitesse angulaire de mouvement inférieure à 908.s 1 ( 908.s 1 <),
isométrique (08.s 1), concentrique à vitesse angulaire de mouvement inférieure à 908.s 1 (< 908.s 1) et concentrique à vitesse angulaire de mouvement supérieure à
908.s 1 (908.s 1 <). Les gains de force sont exprimés en pourcentage de la valeur pré-entraı̂nement et rapportés aux nombre total de séances effectuées durant la
période d’entraı̂nement. Les évolutions significatives sont indiquées en gras. Lorsque plusieurs vitesses de test on été utilisées dans chacune des gammes, la valeur
affichée représente la moyenne des gains de force obtenus pour l’ensemble des vitesses testées.
Sem : semaines ; Fq : fréquence (nombre de séances par semaine d’entraı̂nement) ; 1 RM : charge maximale que le sujet peut soulever une seule fois ; Volume : nombre
de séries nombre de répétitions ; ( p < 0,05).
a
Intensité fixée en pourcentage de la répétition maximale excentrique.
b
Intensité fixée en pourcentage de la force maximale isométrique.
c
Gains de force moyens pour toutes les vitesses de test en contraction excentrique.
d
Gains de force moyens pour toutes les vitesses de test en contraction concentrique.
e
Gains de force moyens pour les muscles rotateurs internes et rotateurs externes de l’épaule.
f
Vitesse angulaire moyenne.
en termes de contrainte mécanique (couple de force, vitesse de pourcentage et n’est donc pas intégrée dans les calculs moyens)
mouvement) imposée par les deux modalités (IT et IK) en et sept études ont étudié ce paramètre suite à un entraı̂nement
fonction de l’angle articulaire pourraient également être à excentrique IK. En moyenne, l’entraı̂nement excentrique IT
l’origine des disparités de gains de force induits. Cependant, si induit une hypertrophie musculaire de 3,26 3,77 %
l’origine des gains de force est décrite pour l’entraı̂nement [10,46,51,85,119,126] contre 7,26 4,07 % après entraı̂ne-
excentrique en général (Section 2.4), les adaptations spécifique- ment excentrique IK [12,30,35,40,58,114,118]. Rapportée au
ment induites par les modalités IT et IK restent à déterminer. nombre de séances, l’hypertrophie obtenue en excentrique IT
est de 0,10 0,13 % contre 0,29 0,17 % en IK. La durée
2.4.3. Adaptations structurales induites par un moyenne de ces protocoles en IT est de 12 semaines contre dix
entraı̂nement isotonique ou isocinétique semaines pour les entraı̂nements IK. Ainsi, malgré des durées
L’entraı̂nement excentrique permet de solliciter de manière de protocoles plus courtes, l’entraı̂nement excentrique IK
maximale et supra-maximale le système musculosquelettique semble induire une hypertrophie musculaire plus importante
dont les structures contractiles s’adaptent lorsque l’entraı̂ne- que l’entraı̂nement excentrique IT. Il est par ailleurs intéressant
ment se poursuit au-delà de six à sept semaines [95]. Six des de constater que les gains significatifs sont obtenus en 12 à
22 études retenues dans cette revue se sont intéressées aux 20 semaines en IT [119] alors qu’une augmentation du volume
évolutions du volume musculaire après une période d’entraı̂ne- musculaire peut être observée en huit semaines d’entraı̂nement
ment excentrique IT (une étude ne rapporte pas de variations en excentrique IK [35]. De la même manière, l’intensité nécessaire
pour observer une hypertrophie musculaire en IT doit se mode IK. Enfin, cette même étude montre une réduction de la
rapprocher ou dépasser 100 % de la répétition maximale (1RM) co-activation musculaire ( 22 à 29 % par rapport au niveau
si la durée ne dépasse pas les huit à dix semaines. Le fait que de co-activation initial) après une période d’entraı̂nement
l’intensité de l’exercice soit maximale sur la totalité de l’effort excentrique IT [92] alors qu’aucune évolution significative n’a
en excentrique IK semble donc favoriser le gain de masse été mise en évidence après un entraı̂nement IK [12].
musculaire. Ce résultat ne semble cependant pas avoir d’impact Ainsi, l’entraı̂nement excentrique IT semble plus favorable
sur les gains de force qui sont plus élevés après entraı̂nement aux adaptations de la commande nerveuse des muscles
excentrique IT. sollicités, ce qui ne permet pas de valider l’hypothèse formulée
Concernant l’hypertrophie par type de fibres musculaires, le selon laquelle l’entraı̂nement IK serait plus efficace de ce point
bilan des différentes études montre clairement une hyper- de vue en raison des pics de couple de force plus élevés que ce
trophie des fibres de type II principalement, quel que soit le mode permet d’atteindre (Section 2.4.1.1). Toutefois, le
mode utilisé (IT ou IK). Seules trois études montrent une paramètre sélectionné (e.g. activation moyenne sur l’ensemble
hypertrophie des fibres de type I (de +0,62 à +0,86 % par de l’amplitude de mouvement, activation au pic de couple de
séance) associée à une hypertrophie des fibres de type rapide force) pour évaluer l’évolution de l’activation neuromusculaire
après entrainement excentrique [52,87,92]. Le nombre peu varie en fonction des études, ce qui peut influencer les résultats
important de travaux développés sur ce sujet ne nous permet obtenus. En effet, le degré d’activation n’est pas nécessairement
pas de conclure quant à la supériorité d’un mode d’exercice par mesuré aux angles articulaires où s’exprime le pic de couple,
rapport à l’autre pour provoquer une hypertrophie typologique secteur présentant une activation musculaire plus importante en
préférentielle. En ce qui concerne les modifications typolo- IK. Ces résultats pourraient expliquer en partie les gains de
giques des fibres musculaires, les modes IT et IK induisent une force plus importants obtenus via l’entraı̂nement excentrique IT
diminution du nombre de fibres de type I et IIb avec une (Section 2.4.2).
augmentation de la proportion de fibres de type IIa/IIb
[12,43,87,101,114,118,126]. 2.5. Conclusion
Les travaux ayant investigué l’effet de l’entraı̂nement
excentrique sur les propriétés mécaniques sont très peu L’exercice excentrique est un moyen efficace d’induire des
nombreux. En effet, aucune étude ne s’est intéressée aux changements structuraux importants du complexe muscle-
évolutions des propriétés mécaniques à la suite d’une période tendon et des adaptations de la commande nerveuse. Ces
d’entraı̂nement excentrique IK. En revanche, les travaux adaptations permettent une augmentation importante des
réalisés sur le mode IT ont montré une augmentation de la capacités de production de force maximale, souvent supérieure
raideur de la composante élastique en série due à une à celle obtenue avec des entraı̂nements concentriques ou
augmentation de la synthèse de collagène [98], et une isométriques. Un bilan comparatif des processus adaptatifs
diminution de la raideur musculaire en raison d’une induits par les deux modalités de sollicitation classiquement
augmentation de la longueur des fascicules, caractérisée en utilisées en excentrique a permis de mettre en évidence une
conditions passives après un entraı̂nement excentrique IT [68]. supériorité du mode IT par rapport au mode IK en ce qui
La raideur tendineuse montre une tendance à l’augmentation concerne les gains de force obtenus. Ce constat peut s’expliquer
après ce type d’entraı̂nement. Ces résultats restent cependant à par des modifications de la commande nerveuse plus
confirmer et à comparer avec la modalité IK. Enfin, à notre importantes à la suite de ce type d’entraı̂nement. L’entraı̂ne-
connaissance, seule une étude a étudié l’effet de l’entraı̂nement ment excentrique IK semble à l’inverse plus favorable à
excentrique IK sur l’architecture musculaire [12], ce qui ne l’hypertrophie musculaire. Les mécanismes adaptatifs (struc-
nous permet pas d’établir une comparaison avec l’excentrique turaux et nerveux) restent encore à préciser car les études sur les
IT. Les résultats ont montré une augmentation de l’angle de deux modes (IT, IK) restent peu nombreuses. La synthèse des
pennation (+21,4 %) et de la longueur des fascicules (+3,1 %) travaux présents dans la littérature exposée dans cette revue a
après dix semaines d’entraı̂nement excentrique IK. permis d’identifier des différences dans les processus adaptatifs
induits par les entraı̂nements excentriques IT et IK (i.e.
2.4.4. Adaptations nerveuses suite à un entraı̂nement hypertrophie musculaire supérieure après entraı̂nement IK,
isotonique ou isocinétique augmentation de l’activation neuromusculaire plus importante
L’ensemble des travaux consacrés à l’étude des adaptations après entraı̂nement excentrique IT). Toutefois, les effets des
nerveuses induites par l’entraı̂nement excentrique montrent une entraı̂nements excentriques IT et IK n’ont à ce jour pas été
augmentation de l’activation neuromusculaire des groupes comparés en conditions standardisées. De tels travaux
musculaires entraı̂nés. Il est de ce point de vue intéressant de permettraient de statuer plus précisément sur la pertinence
noter que les gains d’activation musculaire sont plus élevés de leur utilisation dans le cadre de l’entraı̂nement ou en
après un entraı̂nement excentrique IT (+1,80 1,54 % par rééducation fonctionnelle, en fonction des objectifs adaptatifs
séance) [19,27,92,125], par rapport à un entraı̂nement visés.
excentrique IK (+0,43 0,17 % par séance) [40,43,58,115].
De la même manière, une étude montre une augmentation Conflict of interest statement
significative de l’activation maximale volontaire (+17 %) [92],
paramètre qui, à notre connaissance, n’a pas été étudié sur le The authors do not declare any conflict of interest.
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Annals of Physical and Rehabilitation Medicine 53 (2010) 319–341

Update article / Mise au point

Keywords: Lengthening contraction; Isotonic; Isokinetic; Muscle architecture; Electromyography

Mots clés : Contraction excentrique ; Isotonique ; Isocinétique ; Architecture musculaire ; Électromyographie

1. English version of eccentric contractions before presenting the adaptive

during concentric contractions, suggesting the existence of a

Knee flexors and extensors Nickols-Richardson 20 3 1–5 6 60 - +0.5 - +0.2 -

Knee flexors Melo et al. [75] 12 2 2–4 12 60 - - - +0.5 -

Elbow extensors Farthing and 83 2–6 8 180 +0.7c - +0.9d

Elbow flexors and extensors Nickols-Richardson 20 3 1–5 6 60 - +0.4 - +0.2 -

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