ORIGINAL ARTICLE

Motor imagery practice may compensate for the slowdownof sensorimotor processes induced by short-term upper-limbimmobilization

Aurore Meugnot • Nounagnon Frutueux Agbangla •

Yves Almecija • Lucette Toussaint

Received: 10 April 2014 / Accepted: 24 May 2014

� Springer-Verlag Berlin Heidelberg 2014

Abstract Recently, it has been demonstrated that senso-

rimotor representations are quickly updated following a

brief period of limb non-use. The present study examined

the potential of motor imagery practice (MIP) and inves-

tigated the role of motor imagery instructions (kinesthetic

vs. visual imagery) to counteract the functional impairment

induced by sensorimotor restriction. The participants were

divided into four groups. Three groups wore a splint on

their left hand for 24 h. Prior to the splint removal, two of

the three groups performed 15 min of MIP, with kines-

thetic or visual modalities (KinMIP and VisMIP groups,

respectively). The third group did not practice motor

imagery (NoMIP group). Immediately after the splint

removal, the participants were assessed using a hand lat-

erality task known for evaluating sensorimotor processes.

A fourth group served as the control (i.e., without immo-

bilization and MIP). The main results showed slower left-

hand response times for the immobilized NoMIP group

compared with the controls. Importantly, faster response

times for the left-hand stimuli appeared for the KinMIP

groups only compared with the NoMIP group. No differ-

ence between the four groups was observed for the right-

hand stimuli. Overall, these results highlighted the soma-

totopic effect of limb non-use on the efficiency of senso-

rimotor processes. Importantly, the slowdown of the

sensorimotor processes induced by 24 h of sensorimotor

deprivation may be counteracted by a kinesthetic MIP,

whereas no beneficial effect appeared with visual imagery.

We discuss the importance of imagery modalities for sen-

sorimotor reactivation.

Introduction

Motor imagery (MI) is ‘‘a dynamic state during which the

representation of a given motor act is internally rehearsed

within working memory without any overt motor output’’

(Decety & Grezes, 1999). Psychophysical and brain

imaging studies have demonstrated that movement simu-

lation and movement execution involve similar sensori-

motor representations. Behavioral experiments have shown

that motor imagery preserved the spatiotemporal charac-

teristics of the actual actions (Cerritelli, Maruff, Wilson, &

Currie, 2000; Decety & Jeannerod, 1995) and obeyed the

same joint limits (Petit, Pegna, Mayer, & Hauert, 2003) or

inertial and gravitational constraints (Papaxanthis, Pozzo,

Skoura, & Schiappati, 2002). Neuroimaging studies have

revealed an overlap between the brain regions activated

during MI and the actual movement (Gerardin et al., 2000;

Guillot et al., 2009; Hanakawa, Dimyan, & Hallett,

2008; Solodkin, Hlustik, Chen, & Small, 2004). Another

similarity emerged from studies which recorded autonomic

nervous system activity, with similar increases in ventila-

tion and systolic blood pressure (Wang & Morgan, 1992)

or in electrodermal and thermovascular responses (Guillot

& Collet, 2005) during simulation and the actual execution

of movements.

Overall, these findings allowed to us to consider MI as

the cognitive level of actions (Jeannerod, 2001), and MI

tasks appeared as a relevant means to highlight the central

functioning of the sensorimotor system. MI tasks are often

used in clinical studies, with patients suffering from

somatic motor disorders, to provide information on the link

A. Meugnot � N. F. Agbangla � Y. Almecija � L. Toussaint (&)

Universite de Poitiers, Centre de Recherches sur la Cognition et

l’Apprentissage, CeRCA, CNRS/UMR 7295, 5 rue Theodore

Lefebvre, 86000, Poitiers, France

e-mail: lucette.toussaint@univ-poitiers.fr

URL: http://cerca.labo.univ-poitiers.fr

123

Psychological Research

DOI 10.1007/s00426-014-0577-1

between a cortical structure and its functional outcome. In

studies with apraxic patients, Sirigu and collaborators

highlighted the contribution of the parietal cortex in the

generation of action representations (Sirigu, Daprati, Pra-

dat-Diehl, Franck, & Jeannerod, 1999; Sirigu et al., 1996).

Other experiments examined the function of the basal

ganglia in both inner and behavioral states of action by

comparing the performance of parkinsonian patients and

healthy participants in actual and imagined sequential fin-

ger movements (Dominey, Decety, Brousolle, Chazot, &

Jeannerod, 1994). In other cases, MI studies were used to

better understand complex pathologies, such as schizo-

phrenia (de Vignemont et al., 2006), focal hand dystonia

(Fiorio, Tinazzi, & Aglioti, 2006), anosognosia for hemi-

plegia (Jenkinson, Edelstyn, & Ellis, 2009), developmental

coordination disorder in children (Wilson et al., 2004) and

Asperger syndrome (Conson et al., 2013).

Recently, MI tasks were also used in studies of func-

tional brain plasticity that explored the effects of peripheral

perturbations on sensorimotor representations, such as limb

amputation (Curtze, Otten, & Postema, 2010; Malouin

et al., 2009; Nico, Daprati, Rigal, Parsons, & Sirigu, 2004),

chronic deafferentation (ter Horst, Cole, van Lier, &

Steenbergen, 2012), congenitally absent limb (Funk &

Brugger, 2008) and visual loss and disuse (Malouin et al.,

2009). Overall, these studies have shown that kinesthetic

afferent signals are crucial to maintain efficient sensori-

motor representation and implicitly use internal motor

representation. Importantly, ter Horst et al. (2012) revealed

that long-term loss of kinesthetic afferents may be com-

pensated by visual information. In order to compensate for

the chronic loss of proprioceptive afferents, the deaffe-

rented patient used visual information to construct a rep-

resentation of his body, highlighting the multimodal nature

of body representation. More interested by transient

peripheral perturbations, some authors examined the effect

of regional anesthesia (Silva et al., 2011) or short-term

limb non-use (Meugnot, Almecija, & Toussaint, 2014;

Toussaint & Meugnot, 2013) on sensorimotor representa-

tion. Toussaint and collaborators (Meugnot et al., 2014;

Toussaint & Meugnot, 2013) specifically explored the

effect of short-term hand immobilization (24 or 48 h delay)

on motor simulation processes in healthy participants. The

authors clearly demonstrated that motor imagery processes

(i.e., mental rotation of body stimuli) were affected by left-

or right-hand immobilization, whereas visual imagery

processes were not affected (i.e., mental rotation of non-

body stimuli). For the mental rotation of body stimuli, the

immobilization-induced effects have been demonstrated by

the slower response times for the stimuli that depicted the

immobilized hand. Toussaint and collaborators suggested

that the limb non-use slowed down the corresponding

sensorimotor processes. This quick update of sensorimotor

representations induced by the decrease of input/output

signals could be the origin of motor performance distur-

bances reported in other experiments on movement accu-

racy (Huber et al., 2006; Moisello et al., 2008) and

movement kinematics (Bassolino, Bove, Jacono, Fadiga, &

Pozzo, 2012).

Motor imagery practice (MIP), ‘‘the act of repeating the

imagined movements several times with the intention of

learning a new ability or perfecting a known skill’’ (Jack-

son et al., 2001), has been investigated in both motor

learning and rehabilitation domains. The benefits of MIP in

improving or recovering motor skills have been widely

highlighted in sports (Driskell, Copper, & Moran, 1994;

Feltz & Landers, 1983; Guillot & Collet, 2008), motor

learning (Gentili, Han, Schweighofer, & Papaxanthis,

2010; Jackson, Lafleur, Malouin, Richards, & Doyon,

2003; Toussaint & Blandin, 2010) and rehabilitation pro-

grams (Braun et al., 2013; Dunsky, Dickstein, Marcovitz,

Levi, & Deutsch, 2008; Malouin, Jackson, & Richards,

2013; Malouin, Richards, Durand, & Doyon, 2008). Exe-

cution facilitation following MIP was attributed to the

rehearsal of the corresponding brain structures and the

solicitation of the same sensorimotor representations that

are used in physical practice (Decety & Grezes, 1999;

Jeannerod, 2001). This suggestion is particularly supported

by neurophysiological data that showed MIP produced a

similar reorganization of the motor cortex compared with

physical practice (Jackson et al., 2003; Lacourse, Turner,

Randolph-Orr, Schandler, & Cohen, 2004; Pascual-Leone

et al., 1995). Therefore, MIP appears to be an effective tool

for the maintenance of sensorimotor schemes, especially

when physical execution is not possible.

The benefit of MIP following short-term reduction of

proprioceptive information is actually not known. In this

study, we specifically investigated whether MIP could be

useful to counteract the slowdown of the sensorimotor

processes induced by the decrease in the input/output sig-

nal processing during 24 h of left-arm immobilization.

Moreover, we investigated the importance of the imagery

instructions on the sensorimotor reactivation by focusing

on the visual vs. kinesthetic motor imagery. Motor imagery

based on the visual modality (i.e., visual motor imagery)

requires the self-visualization of the movement, and indi-

viduals are instructed to focus on the spatial components of

the action (i.e., visualization of space, size, amplitude or

form of the movements). Motor imagery based on the

kinesthetic modality (i.e., kinesthetic motor imagery)

requires an individual to ‘‘feel’’ the movement, to mentally

perceive the muscle contractions, including the force and

the effort perceived during the movement and to focus on

the temporal component of the movements (rhythm, speed,

duration). Neural data have demonstrated that kinesthetic

motor imagery had a greater association with the

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123

sensorimotor system compared with visual motor imagery,

which was predominately activated by the visual associa-

tion cortex (Guillot et al., 2009; Ruby & Decety, 2003;

Solodkin et al., 2004; Stinear, Byblow, Steyvers, Levin, &

Swinnen, 2006). Therefore, the authors recommended the

use of a kinesthetic MIP rather than a visual MIP for

reactivating motor function within rehabilitation programs

(Jackson et al., 2001; Mulder, 2007; Stinear et al., 2006).

However, until now, no study has investigated whether the

motor imagery instructions have specific effects in the

functional rehabilitation domain (Braun et al., 2013).

Moreover, behavioral studies in sport or laboratory situa-

tions currently remain controversial. Many studies have

emphasized the benefits associated with kinesthetic MIP in

skill learning, but differences existed depending on the task

constraints (Fery, 2003; Guillot, Collet, & Dittmar, 2004;

Hardy & Callow, 1999; White and Hardy, 1995). Recently,

Toussaint and Blandin (2010) demonstrated the impact of a

previously stored sensory-specific movement representa-

tion (visual vs. kinesthetic) on the efficiency of MIP, which

favored either visual or kinesthetic motor imagery

instructions. The authors reported the dominance of visual

information over kinesthetic information when participants

executed hand movements with visual feedback. In that

specific case, a subsequent visual MIP promoted motor

learning, whereas no benefit of a kinesthetic MIP was

observed. In contrast, when participants experienced the

task without vision (the eyes closed), their performance

increased after a subsequent kinesthetic MIP, whereas no

benefit of a visual MIP appeared. Consequently, in the

present experiment, the sensory-specific condition of the

task experienced before MIP was accurately controlled to

test the potential of kinesthetic and/or visual MIP to

counteract the functional impairment induced by 1 day of

sensorimotor restriction.

In the present experiment, to examine whether MIP may

reactivate the sensorimotor processes that have been slo-

wed down by immobilization and whether kinesthetic or

visual imagery may be favored, four groups of participants

were used: a control group and three left-hand immobili-

zation groups. Two of the three immobilized groups were

subjected to a MIP with either visual (VisMIP group) or

kinesthetic imagery (KinMIP group) during the immobili-

zation (immediately prior to the splint removal), whereas

the third group did not practice motor imagery (NoMIP

group). All participants were required to judge whether a

hand figure displayed at the center of the screen was a left

or a right hand. As emphasized in the literature, we

expected slower response times for the left (immobilized)

hand identification in the immobilized NoMIP group

compared with the control group. The VisMIP and KinMIP

groups should allow us to examine the effect of motor

imagery instructions on sensorimotor reactivation. In

agreement with neuroimaging studies, if the use of bodily

information is crucial to reactivate the sensorimotor sys-

tem, a kinesthetic imagery practice (KinMIP group) should

further compensate for the slowdown of the sensorimotor

processes compared with a visual imagery practice

(VisMIP group), especially for stimuli that depict the

immobilized hand. In contrast, regarding the multimodal

nature of action representation, we may argue that senso-

rimotor memories might recover vividness by working on

their visual component. In this case, the VisMIP group

should improve its ability to identify the hand immobilized

stimuli similar to the KinMIP group.

Another aim of the present experiment was to examine

whether the effect of sensorimotor deprivation varied

according to the accuracy requirement of the simulated

movements. A previous study conducted by Gentilucci,

Benuzzi, Bertolani, Daprati, and Gangitano (2000) showed

that the type of grip influenced the recognition of hand

laterality, with the response times increasing with the

increase in precision grip (small sphere vs. large sphere).

Moreover, they observed similar grip-precision effects for

both the simulated and the actual grasping movements. The

authors suggested that to recognize hand laterality, the

participants used a motor hand representation that retrieved

the main aspect of the motor act. Therefore, in the present

experiment, we presented the participants with pictures of

the hand grasping a small or a large sphere to determine

whether the effect of sensorimotor deprivation varied

according to the precision of the simulated movements.

Methods

Participants

Fifty-two university students (18–26 years, mean

age = 19.7 years, 27 males) completed the experiment. All

participants were self-reported right-handers and non

ambidextrous. The participants were divided into four

groups of 13 participants: a control group (mean

age = 20 years, 9 males) and three left-hand immobiliza-

tion groups. Two of these 3 groups performed a MIP with

either visual (VisMIP, mean age = 20 years, 6 males) or

kinesthetic imagery instructions (KinMIP, mean

age = 19.5 years, 6 males) during the immobilization,

whereas the third group did not practice motor imagery

(NoMIP, mean age = 19 years, 6 males). The participants

were healthy, had normal or corrected-to-normal vision and

no history of motor or neurological disorders. The local

ethics committee approved the study protocol. All partici-

pants provided their written informed consent prior to their

inclusion in this study. Before testing, they were naıve to

the aims of the experiment.

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123

Materials and tasks

The participants were seated *60 cm in front of a com-

puter screen, with their hands resting palm down on their

thighs. All participants were asked to identify, as accu-

rately and as quickly as possible, whether a hand figure

displayed at the center of the computer screen corre-

sponded to a right or a left hand (Fig. 1). Each trial began

when a fixation cross was displayed on the center of the

screen for 500 ms. Then, a hand figure was presented and

remained visible until the participant provided his/her

response. The participants provided a verbal response

(‘‘left’’/‘‘right’’). The E-Prime 2.0 software package (Psy-

chology Software Tools Inc., Pittsburgh, USA) was used to

present the stimuli and record the participants’ response

times via a microphone connected to the computer. For

each trial, the experimenter wrote the participants’

responses on a data sheet.

The stimuli consisted of pictures of right or left hands

and forearms (see Gentilucci et al., 2000 for similar

material). They were presented on a rectangular

(12 9 9.44 cm) dark background. They were horizontally

located and could be presented in two directions. The fin-

gers were located either on the right (right-oriented hand)

or on the left (left-oriented hand). The hands held a sphere

that was one of two sizes (‘‘small’’ or ‘‘large’’). The

diameters of the small and large spheres were 2 and 14 cm,

respectively (see Fig. 1). Eight stimuli (2 hands 9 2

grips 9 2 directions) were used in the present experiment.

Procedure

We used a rigid splint (model DONJOY ‘‘Comfort Digit’’,

DJO, Surrey, UK) to immobilize the left wrist and three

fingers (index, middle and ring fingers) of the participants

for the three immobilization groups for 24 h. Their

immobilized arm was placed in a sling to ensure that the

participants kept their hand at rest as much as possible

during the day of immobilization. The participants were

instructed to never remove the splint and to keep the left-

hand at rest during the immobilization period. Back in our

lab, none of the participants reported having removed the

splint when we asked them. The fourth group served as the

control (i.e., without immobilization).

Before the splint removal, two of the three immobilized

groups participated in a motor imagery session for 15 min,

whereas the third immobilized group did not practice motor

imagery (NoMIP). The first two groups differed with

respect to the imagery instructions.1 The participants who

practiced visual imagery (VisMIP) had to imagine them-

selves producing specific hand and fingers movements,

which focused on visual information. The participants who

practiced kinesthetic imagery (KinMIP) were asked to feel

themselves producing specific hand and finger movements,

which focused on bodily information. MIP was divided

into three series of 5 min. In each series, the participants

repeated four motor imagery exercises with the immobi-

lized (left) hand, after having physically executed it with

the non-immobilized (right) hand (with vision for the

VisMIP group, without vision for the KinMIP group; see

Toussaint & Blandin, 2010 for the importance of the sen-

sory-specific experience prior to motor imagery). In the

first exercise, the participants simulated finger tapping

movements (i.e., alternatively touched the thumb starting

with the index, middle, ring and little fingers and then in

the opposite direction). The second and third exercises

consisted of adduction/abduction and flexion/extension

movements of the wrist, respectively. In the fourth exer-

cise, they imagined successively clenching and unclen-

ching their hand. In each exercise, the participants

simulated *25 hand and finger movement repetitions with

the immobilized hand. The participants kept their eyes

closed during each mental exercise. The experimenter

ensured that the participants kept their eyes closed during

each mental exercise, by watching them continuously. The

participants were instructed to say ‘‘top’’ when they started

and finished a repetition. Chronometry of the imagined

movement was used to gauge the engagement in the MIP

training and to control that all participants practiced the

motor imagery for *15 min.

The hand identification task was performed immediately

after the splint removal for the immobilized groups. For all

participants (control, NoMIP, VisMIP and KinMIP

groups), the task was divided into two phases. During the

first familiarization phase, the participants were shown 8

randomly presented trials (2 hands 9 2 grips 9 2 direc-

tions). No time constraint was imposed during the famil-

iarization phase. During the second experimental phase, the

participants were shown 6 blocks of 32 trials (i.e., 192 trials

Left-oriented hand Right-oriented hand

Small sphere

Large sphere

Fig. 1 Illustration of the right-hand stimuli used in the present

experiment. Mirror images were used for the left-hand stimuli

1 Note that visual imagery is less motor than kinesthetic imagery.

Psychological Research

123

per participant) presented in a random order, with a break

(1 min maximum) after the second and fourth blocks.

At the beginning of the experiment and before splinting

for the immobilized groups, all participants completed the

Vividness of Movement Imagery Questionnaire (VMIQ;

Isaac, Marks, & Russell, 1986). The VMIQ consists of 24

items relevant to both the visual imagery of movement

(i.e., movement is simulated from an external, third-person

perspective) and the imagery of kinesthetic sensations (i.e.,

movement is simulated from an internal, first-person per-

spective). For each imagery perspective, the measures of

imagery vividness were assessed on a 5-point Likert-type

scale. High scores (120 maximum) represented a low viv-

idness of movement imagery, with the reverse being true

for low scores (24 minimum). The visual and kinesthetic

scores for each group are shown in Table 1. Analyses of

variance (ANOVA) performed on the imagery scores with

group (control, NoMIP, KinMIP and VisMIP) as the

between-subjects factor did not reveal significant differ-

ences between the participants for the visual imagery

scores [F(3,48) = 0.288, p = 0.834] or the kinesthetic

imagery scores [F(3,48) = 0.288, p = 0.834].

Data analysis

For the hand recognition task, the accuracy and response

times were recorded and analyzed. Only data from the

correct responses were used to analyze the response times.

We first computed the mean response times (ms) and the

mean accuracy scores (%) for each participant. Then, we

examined the Pearson’s correlation between the response

times and the accuracy scores for each group. We did not

find significant correlations in any group (control:

r = 0.16, t(12) = 0.59, p = 0.57; NoMIP: r = 0.20,

t(12) = 0.73, p = 0.48; KinMIP: r = -0.25, t(12) = 0.90,

p = 0.38; VisMIP: r = -0.26, t(12) = 0.94, p = 0.37),

indicating the absence of a speed–accuracy trade-off.

Finally, ANOVA tests were performed on both the accu-

racy (%) and the response times (ms) with the group

(control, NoMIP, VisMIP and KinMIP) as the between-

subjects factor and the hand (left vs. right), the grip (small

sphere vs. large sphere) and the hand orientation (left-ori-

ented vs. right-oriented hand) as the within-subjects fac-

tors. Post-hoc comparisons were carried out by means of a

Newman–Keuls test. Alpha was set at 0.05 for all analyses.

Results

The ANOVA performed on the accuracy scores (i.e., the

percentage of the correct responses) yielded no main

effects [group, F(3,48) = 1.12; hand, F(1,48) = 0.14;

hand orientation, F(1,48) = 0.48; grip, F(1,48) = 0.01,

ps [ 0.34] and no significant interactions (see Table 2)

[group 9 hand, F(3,48) = 0.86; group 9 hand orientation,

F(3,48) = 0.28; group 9 grip, F(3,48) = 0.47; hand 9

hand orientation, F(1,48) = 1.69; hand 9 grip, F(1,48) =

2.70; hand orientation 9 grip, F(1,48) = 1.77; group 9

hand 9 hand orientation, F(3,48) = 0.53; group 9

hand 9 grip, F(3,48) = 1.62; group 9 hand orientation 9

grip, F(3,48) = 0.60; hand 9 hand orientation 9 grip,

F(1,48) = 0.67; group 9 hand 9 hand orientation 9 grip,

F(3,48) = 0.57; ps [ 0.11]. Regardless of the group, the

task was succeeded in 95 % (±7 %) of the cases.

The ANOVA performed on the response times showed

significant main effects for the group [F(3,48) = 3.14,

p = 0.034, g2p = 0.16] and the hand [F(1,48) = 81.85,

p = 0.000, g2p = 0.63]. No main effect was observed for

hand orientation [F(1,48) = 0.29, p = 0.59] and grip

[F(1,48) = 0.46, p = 0.51]. We also found three signifi-

cant interactions: group x hand [F(3,48) = 4.74,

Table 1 Mean (standard deviation) scores that measured the visual

and kinesthetic imagery capacities for each group

Control NoMIP KinMIP VisMIP

Visual

imagery

scores

50.8 (18.3) 48.0 (13.1) 53.2 (15.2) 48.6 (12.4)

Kinesthetic

imagery

score

49.8 (17.0) 51.6 (14.9) 49.8 (20.1) 50.8 (16.7)

The Vividness of Movement Imagery Questionnaire (or VMIQ; Isaac

et al., 1986) was used in the present experiment

Table 2 Mean (standard

deviation) accuracy scores for

each group (control, NoMIP,

KinMIP and VisMIP) as a

function the hand (left vs. right),

the hand stimuli orientation

(left-oriented vs. right-oriented)

and the grip (large sphere vs.

small sphere)

Left-hand Right-hand

Left-oriented Right-oriented Left-oriented Right-oriented

Small Large Small Large Small Large Small Large

Control 95 (7) 94 (6) 96 (6) 96 (6) 96 (8) 96 (5) 93 (6) 94 (8)

NoMIP 96 (5) 95 (9) 97 (5) 95 (5) 96 (5) 97 (4) 96 (5) 96 (5)

KinMIP 94 (7) 92 (9) 96 (5) 93 (6) 93 (9) 92 (8) 91 (7) 95 (6)

VisMIP 94 (9) 94 (8) 94 (6) 96 (5) 96 (8) 94 (8) 94 (6) 96 (5)

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123

p = 0.006, g2p = 0.23], hand 9 grip [F(1,48) = 7.99,

p = 0.007, g2p = 0.14] and hand 9 orientation [F(1,48)

= 8.52, p = 0.006, g2p = 0.15]. No other significant

interaction appeared [group 9 hand orientation, F(3,48)

= 0.28; group 9 grip, F(3,48) = 0.99; hand orienta-

tion 9 grip, F(1,48) = 2.74; group 9 hand 9 hand ori-

entation, F(3,48) = 0.77; group 9 hand x grip, F(3,48)

= 0.07; group 9 hand orientation 9 grip, F(3,48) = 0.27;

hand 9 hand orientation 9 grip, F(1,48) = 0.85; group 9

hand 9 hand orientation 9 grip, F(3,48) = 0.56; ps [0.11].

The group 9 hand interaction showed (Fig. 2) that the

response times were significantly slower for the left

(immobilized) hand stimuli identification for the NoMIP

group compared with the control (p \ 0.002) and the

KinMIP groups (p \ 0.008), with no difference between

the two latter groups (p = 0.58). In contrast, the response

times did not significantly vary between the VisMIP group

and the other groups (ps [ 0.13). For the right (non-

immobilized) hand stimuli, the response times did not vary

between the groups (ps [ 44). For each group, response

times were significantly slower for the left hand than for

the right hand (ps \ 0.02).

The hand 9 grip interaction showed (Fig. 3a) that the

response times were significantly faster for the large grip

compared with the small grip (p \ 0.02) for the right-hand

stimuli, whereas the response times did not vary with the

grip for the left-hand stimuli (p = 0.13). For both the small

and large grips, the response times were slower for the left

hand compared with the right hand (ps \ 0.001). The hand

x hand orientation interaction revealed (Fig. 3b) that the

response times for the left hand were significantly slower

for the right-oriented compared with the left-oriented hand

(p \ 0.01), whereas they tended to be faster for the left-

oriented compared with the right-oriented hand for the

right hand (p = 0.07).

Discussion

Over the past decade, some studies reported that limb non-

use for 10 or 12 h induced cortical changes (Avanzino,

Bassolino, Pozzo, & Bove, 2011; Avanzino et al., 2013;

Huber et al., 2006) and affected motor performance

(Bassolino et al., 2012; Huber et al., 2006; Moisello et al.,

2008). The authors suggested that the behavioral alterations

were due to a dysfunction of the proprioceptive signals and

a modification of the sensorimotor memories that corre-

sponded to the immobilized limb. These suggestions were

in agreement with the functional plasticity of the sensori-

motor representations induced by short-term upper-limb

immobilization (Meugnot et al., 2014; Toussaint & Me-

ugnot, 2013) or upper-limb anesthesia (Silva et al., 2011) in

healthy participants. The authors reported that the motor

imagery processes were slowed down by the upper-limb

non-use during a brief period of sensorimotor deprivation.

The present study was a continuation of these studies. It

was specifically designed to examine the potential of MIP

to counteract the functional impairment induced by short-

term upper-limb immobilization (during 24 h). Moreover,

the role of the motor imagery instructions (visual vs.

Control group

VisMIP group

KinMIP group

NoMIP group

Res

pons

eti

me

(ms)

900

800

700

Left-hand Right-hand

1000

***1100

Fig. 2 Response time (ms) as a function of the hand (left vs. right)

and the group (control, NoMIP, KinMIP and VisMIP). Error bars

indicate the standard error of the mean

Res

pons

eti

me

(ms)

900

850

800

Left-hand Right-hand

950

Large sphere

Small sphere

*

**

Res

pons

eti

me

(ms)

900

850

800

Left-hand Right-hand

950

Left-oriented hand

Right-oriented hand

*

**

A BFig. 3 a Response time (ms) as

a function of the hand (left vs.

right) and the grip (large sphere

vs. small sphere). b Response

time (ms) as a function of the

hand (left vs. right) and the hand

stimuli orientation (left-oriented

vs. right-oriented). Error bars

indicate the standard error of the

mean

Psychological Research

123

kinesthetic) in reactivating the sensorimotor processes

(assessed using a motor imagery task) was of particular

interest in this experiment.

Short-term upper-limb immobilization

and the slowdown of the sensorimotor processes

As previously observed for the hand mental rotation tasks

(Meugnot et al., 2014; Toussaint & Meugnot, 2013), the

results of the present experiment showed that the immo-

bilization-induced effects became widespread in the hand-

grasping object recognition. One day of sensorimotor

restriction for the NoMIP group induced a slowdown of the

hand-stimuli recognition (i.e., slower response times) that

corresponded to the immobilized hand only compared with

the performance of the participants who did not undergo

the immobilization procedure (the control group). The

immobilization-induced effects provided evidence for the

sensorimotor representations involvement in solving the

hand-grasping object task and confirmed the embodied

nature of the motor imagery processes (Meugnot et al.,

2014). The involvement of the sensorimotor representa-

tions when identifying hand laterality was confirmed in the

present experiment by the slower identification processes

for the left (non-dominant) hand compared with the right

hand in the four groups (see also Gentilucci, Daprati, &

Gantigano, 1998; Ionta & Blanke, 2009; Ni Choisdealbha,

Brady, & Maguiness, 2011; Parsons, 1987, 1994, for sim-

ilar results). The greater expertise of the dominant hand

(vs. the non-dominant hand), which is associated with a

finer sensorimotor representation, is at the origin of the

difference in the response times between the hands for both

actual and simulated movements.

Did the immobilization-induced effects vary as a function

of the accuracy requirement of the simulated movements

(i.e., small vs. large sphere grasping)? In the present exper-

iment, the results indicated that left-hand non-use (i.e., the

non-dominant hand) was not the appropriate way to answer

this question. Contrary to Gentilucci et al. (2000), who

reported an increase in the response times for the small

sphere compared with the large sphere for both left and right

hands, we observed a grip-precision effect for the right hand

only. In our experiment, grasping small spheres was more

time-consuming compared with grasping large spheres for

the dominant hand only. This lower engagement of the

sensorimotor processes when identifying the left-hand

stimuli was also suggested by the influence of the spatial

orientation of the hand pictures on the screen. The effect of

the spatial incompatibility, which was more important for the

left-hand stimuli, suggested its greater dependence on the

visual context. The overall results for the right- and left-hand

differences indicate that the use of a motor imagery strategy

to solve the hand laterality task was stronger in identifying

the dominant hand compared with the non-dominant hand.

Therefore, these unexpected results show that immobilizing

the left non-dominant hand of the participants was not the

right solution to determine whether the effects of sensori-

motor deprivation varied according to the precision of the

simulated movements. Note, however, that the sensorimotor

processes remained involved in the identification of the left-

hand stimuli. As reported at the beginning of this section,

24 h of left-hand immobilization effectively led to the

slowdown of the left-hand movement simulation, without

affecting the motor imagery processes for the right-hand.

Motor imagery practice and the reactivation

of the sensorimotor processes

The major aim of the present experiment was to examine

the potential role of MIP in reactivating the sensorimotor

processes that have been slowed down by short-term upper-

limb immobilization. Importantly, we questioned the

influence of the motor imagery instructions (visual vs.

kinesthetic) in counteracting the deleterious effect of 24 h

of sensorimotor deprivation (i.e., left-hand immobilization).

The results of the present experiment showed that the

response times were faster for the left-hand identification

for the participants who practiced 15 min of kinesthetic

motor imagery (KinMIP group) compared with the par-

ticipants who did not practice imagery (NoMIP group).

Moreover, although imagery abilities were similar for the

KinMIP and VisMIP groups (i.e., ‘‘clear and vivid’’ or

‘‘fairly clear and vivid’’, as determined by the VMIQ

scores), a visual imagery practice did not improve the

performance of the immobilized participants, whereas a

kinesthetic imagery practice did improve the performance.

In accordance with neuroimaging (Guillot et al., 2009;

Solodkin et al., 2004) and behavioral studies (Fery, 2003;

Hardy & Callow, 1999; Toussaint & Blandin, 2010), this

finding supports a neuronal and functional distinction

between kinesthetic and visual imagery, with kinesthetic

imagery having a greater association with the sensorimotor

system compared with visual imagery, which would pre-

dominantly activate the visual association cortex (Guillot

et al., 2009). These results clearly indicated that a kines-

thetic MIP was particularly effective in reactivating the

sensorimotor representations of the immobilized hand,

thereby canceling the slowdown of the motor imagery

processes induced by sensorimotor deprivation. As previ-

ously suggested (Moisello et al., 2008; Toussaint & Me-

ugnot, 2013) and in line with the theory of internal models

(Wolpert & Flanagan, 2001), it is likely that the acute

decrease in the sensory and motor flow into the CNS during

the 24 h of limb non-use disrupted the sensorimotor

memories, thereby affecting the ability of the forward

processes to accurately form the movement predictions.

Psychological Research

123

Indeed, the central representation theory (Jeannerod, 2001)

states that motor imagery relies on the same cognitive

processes as motor execution. During simulation, the for-

ward models would provide sensory and motor information

similar to that of the physical execution to mentally predict

the movement (i.e., in the laterality task, the future states

and sensory consequences of hand matching the stimulus

on the computer screen). During the period of limb non-

use, the processing of the input/output in the contralateral

area was greatly reduced, and the sensorimotor represen-

tations were no longer updated. Thus, the left-hand inner

memories would be less vivid and less easily retrieved by

the forward models after immobilization. However, during

a kinesthetic MIP, proprioceptive and motor processing

would be progressively reactivated to simulate left-hand

movements. The motor planning routines would progres-

sively recover their initial level of functioning.

The present experiment showed that kinesthetic MIP is

particularly effective in reactivating the motor imagery pro-

cesses following short-term limb non-use. However, no clear

conclusion can be drawn with regard to visual MIP. The

results did not reveal a significant difference in hand recog-

nition between the VisMIP and KinMIP groups, as well as

between the VisMIP and NoMIP groups. These ambiguous

results could be due to several possibilities. First, we cannot

exclude the possibility that participants who performed a

visual MIP still relayed on residual proprioceptive informa-

tion. Indeed, before practicing Visual MIP, participants exe-

cuted right hand movements with both visual and

proprioceptive information. However, the residual of propri-

oceptive information could contribute to sensorimotor reac-

tivation without being sufficient enough to induce a significant

improvement compared with the KinMIP group. Second, the

benefit of visual MIP might necessitate more times to reacti-

vate the sensorimotor system following a brief period of

sensorimotor deprivation compared with kinesthetic MIP.

Visual motor imagery is known to predominately activate the

visual association cortex, whereas kinesthetic motor imagery

has stronger association with the sensorimotor system (Guillot

et al., 2009). It may be possible that the lesser association of

visual motor imagery with the sensorimotor system, combined

with the slowdown of the sensorimotor processes due to

immobilization, decreases the potential benefit of visual

imagery. Whether a longer visual MIP will effectively help to

reactivate the sensorimotor system following a relatively short

period of limb immobilization is a relevant question that must

be clarified in the future.

Conclusion

Overall, the results of the present experiment showed for

the first time that a kinesthetic MIP was efficient in

counteracting the deleterious effect of short-term upper-

limb immobilization on motor imagery processes. When

movement is not possible, the kinesthetic imagery rehearsal

would allow an individual to maintain the recursive pro-

cessing of sensory and motor inflow into the system and

prevent the disruption of sensorimotor memories. To our

knowledge, only one study has investigated the potential of

kinesthetic imagery in modulating the brain changes fol-

lowing short-term limb non-use (Bassolino, Campanella,

Bove, Pozzo, & Fadiga, 2013). The authors used trans-

cranial magnetic stimulation to assess the effect of short-

term immobilization (10 h) on the excitability of the pri-

mary motor cortex contralateral to the limb immobilization

(cM1). Moreover, they examined the effect of a kinesthetic

MIP during the period of sensorimotor restriction to pre-

vent the cortical depression. Surprisingly, they observed no

compensatory effect and concluded that motor imagery

was not effective in modulating brain plasticity during

inactivity. In fact, the contribution of the cM1 during motor

imagery remains controversial, as neuroimaging studies did

not systematically report the activation of the cM1 during

motor imagery (De Vries & Mulder, 2007; Guillot, Di

Rienzo, MacIntyre, Moran, & Collet, 2012). In contrast,

there is substantial evidence that kinesthetic motor imagery

involves many others brain structures (i.e., the ventral and

dorsal parts of the premotor cortex, the supplementary

motor area, the inferior parietal lobule; Gerardin et al.,

2000; Guillot et al., 2009; Munzert, Lorey, & Zentgraf,

2009; Solodkin et al., 2004). Therefore, the lack of benefit

of kinesthetic imagery practice on the cM1 excitability

does not necessarily indicate that MIP is not a reliable tool

for reactivating the sensorimotor system during limb non-

use. In fact, the results of the present experiment support

the use of kinesthetic MIP.

The present study supports the potential use of a kin-

esthetic MIP in a rehabilitation program as a means of

promoting the recovery of motor function, especially when

physical practice is not possible. In fact, the potential of MI

in rehabilitation is now well accepted, even though no

definite conclusions on the effects of mental practice can be

drawn at this time because the evidence base is relatively

small (Braun et al., 2013; Malouin et al., 2013). In the

specific case of limb disuse, to our knowledge, only two

studies have examined the potential of mental practice on

functional recovery (Crews & Kamen, 2006; Stenekes,

Geertzen, Nicolai, De Jong, & Mulder, 2009). However, in

these studies, the positive effect of motor imagery on motor

performance following at least 1 week of immobilization

was rather weak or non-significant. Further research on the

use of kinesthetic motor imagery in injury rehabilitation

(i.e., with immobilized patients after the effects of ortho-

pedic injury, such as sprain, fracture or surgical interven-

tion) would be of interest. One cannot exclude the

Psychological Research

123

possibility that a kinesthetic MIP is effective only when

sensorimotor deprivation is transient. As reported in studies

examining the effect of long-term loss of kinesthetic

afferents on the sensorimotor representation (ter Horst

et al., 2012), the availability of kinesthetic afferent signals

is crucial to incorporate bodily information into the

imagery processes. Therefore, when patients are immobi-

lized longer because of injury, a visual MIP could be more

effective to construct a body representation than a kines-

thetic MIP. Studies in healthy individuals have shown that

motor imagery use in addition to physical practice allows a

reduction in the amount of physical practice required to

attain a given level of motor performance (Allami, Pau-

lignan, Brovelli, & Boussaoud, 2008; Pascual-Leone et al.,

1995). The challenge over the coming decades will be to

ensure the field is readily amenable to integrating MIP into

the current standard of care used in clinical practice.

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