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www.frontiersin.org October 2010 | Volume 1 | Article 150 | 1
Original research article
published: 04 October 2010
doi: 10.3389/fpsyg.2010.00150
Context effects in embodied lexical-semantic processing
Wessel O. van Dam*, Shirley-Ann Rueschemeyer, Oliver Lindemann and Harold Bekkering
Donders Institute for Brain, Cognition and Behavior, Radboud University Nijmegen, Netherlands
The embodied view of language comprehension proposes that the meaning of words is
grounded in perception and action rather than represented in abstract amodal symbols. Support
for embodied theories of language processing comes from behavioral studies showing that
understanding a sentence about an action can modulate congruent and incongruent physical
responses, suggesting motor involvement during comprehension of sentences referring to
bodily movement. Additionally, several neuroimaging studies have provided evidence that
comprehending single words denoting manipulable objects elicits specific responses in
the neural motor system. An interesting question that remains is whether action semantic
knowledge is directly activated as motor simulations in the brain, or rather modulated by the
semantic context in which action words are encountered. In the current paper we investigated
the nature of conceptual representations using a go/no-go lexical decision task. Specifically,
target words were either presented in a semantic context that emphasized dominant action
features (features related to the functional use of an object) or non-dominant action features. The
response latencies in a lexical decision task reveal that participants were faster to respond to
words denoting objects for which the functional use was congruent with the prepared movement.
This facilitation effect, however, was only apparent when the semantic context emphasized
corresponding motor properties. These findings suggest that conceptual processing is a context-
dependent process that incorporates motor-related knowledge in a flexible manner.
Keywords: embodiment, semantics, action, conceptual flexibility
Edited by:
Anna M Borghi, University of Bologna,
Italy
Reviewed by:
Veronique Boulenger, Centre National
de la Recherche Scientifique, France
Rutvik Desai, Medical College of
Wisconsin, USA
*Correspondence:
Wessel O. van Dam, P.O. Box 9104,
6500 HE Nijmegen, Netherlands.
e-mail: w.vandam@donders.ru.nl
Taylor (2006) have shown that motor resonance is activated on-line
during sentence comprehension. For example, while reading the
sentence The marathon runner opened the water bottle, evidence
for motor resonance is seen in conjunction with presentation of
the verb opened. Furthermore motor resonance is elicited by words
which unambiguously specify kinematic properties of actions, even
if these words are not verbs (Taylor et al., 2008). For example, in the
text He looked at the pie and turned the oven dial. The baking time
needed to be shorter/longer evidence for motor resonance is seen
in conjunction with the sentence final adjective (which specifies
how the oven dial should be turned) rather than in conjunction
with the verb turned.
In addition to words and sentences describing active events,
isolated words denoting manipulable objects have also been shown
to interact with motor behaviors and to activate neural motor
areas. For example, Glover et al. (2004) show that words denot-
ing objects that afford particular actions are sufficient to activate
motor representations. Participants in their study were primed
with the names of objects of different sizes (e.g., apple, grape) and
required to reach out and grasp a wooden block. Interestingly, the
maximum grip aperture during the reach (i.e., maximum dis-
tance between forefinger and thumb) was influenced by the size
of the object denoted by the prime word, despite the fact that
this information was irrelevant for executing the grasping task.
Similarly, Rueschemeyer et al. (2010a) had participants perform
a lexical decision task to words denoting objects typically brought
toward or away from the body for functional use (e.g., cup or key,
respectively). The authors found that responses to words were
facilitated if the required response action was congruent to the
IntroductIon
According to an embodied view of language comprehension, lan-
guage concepts are grounded in motor and perceptual systems
(Glenberg, 1997; Barsalou, 1999; Pulvermueller, 1999; Barsalou
et al., 2003). Lexical-semantic representations are postulated to
rely on sensori-motor brain areas and to reflect real-world experi-
ence with words’ referents. For example an object such as a ham-
mer is experienced by most people visually (i.e., we know what
hammers look like) and motorically (i.e., we know what it feels
like to wield a hammer). Thus, embodied language theories postu-
late that upon encountering the word hammer, experiential traces
stored in modality specific (e.g., visual and motor) brain areas are
activated. Importantly, these activations are seen to contribute to
the lexical-semantic meaning of the word form hammer. This view
differs fundamentally from symbolic (disembodied) accounts, in
which conceptual representations are symbolic and amodal, and
lexical-semantic meaning independent of real-world experience
(Kintsch, 2008).
Converging evidence for the idea that sensory-motor brain areas
are involved in language comprehension comes from behavioral,
electrophysiological, and neuroimaging studies. Glenberg and
Kaschak (2002) present the action-sentence compatibility effect
(ACE), in which reading a sentence that implies an action toward
or away from the body (e.g., open the drawer/close the drawer)
facilitates a congruent action (i.e., moving a hand toward or away
from the body). The authors argue that responses are facilitated
because comprehending language about action recruits the same
neural resources as required for action execution; thus compre-
hending the sentence primes a congruent motor act. Zwaan and
Frontiers in Psychology | Cognition October 2010 | Volume 1 | Article 150 | 2
van Dam et al. Conceptual flexibility
studies provide evidence that motor responses rather than being
automatic and invariable, depend on the context (i.e., sentence
context in the case of Raposo et al., 2009, and morphological con-
text in the case of Rueschemeyer et al., 2007) in which action words
are encountered. That is, a crucial factor for observing activity in
motor and premotor regions during action word processing seems
to be that the context emphasizes motor properties (suggesting
that representations are flexible).
The failure of some studies to report motor activation for action-
related words embedded in various contexts already speaks against
a strictly automatic interpretation, and suggests that motor activa-
tions may be called on in a flexible manner during word processing.
In the two studies cited above, motor information is not useful
in comprehending the given language utterances (i.e., idiomatic
phrases or morphologically complex words), and indeed the results
indicate that in these cases the motor system is not reliably activated.
However, even in cases in which motor information is helpful in
processing semantic content, a certain degree of flexibility on the
level of specific motor programs might be expected. For example,
the motor programs associated with the word cup are quite differ-
ent in the sentences She filled her cup at the tap/She drank from the
cup. In the first case, the actor in the sentence brings the cup away
from her body and toward the tap, in the second she brings the
cup toward her mouth. Previous research has shown that words
referring to manipulable objects activate motor areas (Chao and
Martin, 2000; Saccuman et al., 2006; Rueschemeyer et al., 2010a),
but since most objects can be used in multiple ways, does the motor
contribution to lexical-semantic comprehension also vary depend-
ing on the language context in which a word is presented?
In the current study, we investigated precisely this issue: within a
given modality (i.e., action) we investigated whether a word always
activates a specific motor program, or whether the motor program
activated depends on the context in which a word is presented.
Previously we (Rueschemeyer et al., 2010a) showed that compre-
hension of words denoting manipulable objects (e.g., cup, ham-
mer) are facilitated by the prior planning of an action congruent
to the prototypical use of the object. In other words, participants
were faster to respond to the word cup when they had planned an
action toward their body (the most common use of the cup being
a vessel to drink from) than when they had planned an action away
from their body; the opposite pattern was true for words denoting
objects typically brought away from the body (e.g., hammer). We
thus show a strong congruency effect between the typical action
associated with a word’s referent and processing of the individual
word. The target words in this previous study were presented in
isolation, i.e., without any explicit language context. In the current
study we investigated whether embedding the same words in a
lexical context suggesting a non-prototypical (but not unfamil-
iar) use of the objects would influence the observed congruency
effects. To this end object words were always preceded by a prime
word providing a context that emphasized either the dominant
action feature (thirst – cup) or a non-dominant action feature
(sink – cup). As previous studies have shown that the retrieval of
a particular conceptual feature depends on the context in which
a word is encountered, we hypothesized that the effect of move-
ment preparation on word processing would interact with the
action typically performed on the word’s referent (i.e., responses
to cup were faster if a movement was made toward the body rather
than away from the body). This indicates that very specific infor-
mation about how an object is manipulated is activated during
lexical retrieval.
Evidence in favor of embodied theories of language also comes
from recent neuroimaging studies. Specifically, the comprehen-
sion of action verbs (Kemmerer et al., 2008), action sentences
(Desai et al., 2010), and words denoting manipulable objects
(Chao and Martin, 2000; Saccuman et al., 2006; Rueschemeyer
et al., 2010b) all reliably activate the cerebral motor system.
Furthermore, embodied lexical-semantic representations activate
the neural motor system in a somatotopic manner (Hauk et al.,
2004; Tettamanti et al., 2005; Aziz-Zadeh et al., 2006; but see also
de Zubicaray et al., 2010). Findings from the studies reviewed
above provide strong evidence for functional links between the
neural motor system and lexical-semantic processing of words
that entail a motor component.
It has been suggested that embodied lexical-semantic repre-
sentations are fast, automatic and invariant. Pulvermueller et al.
(2000) demonstrated that category-specific activation can be
observed as early as about 200 ms after word onset (see also Hauk
and Pulvermueller, 2004), and occurs irrespective of attention to
action words (Shtyrov et al., 2004; Pulvermueller et al., 2005).
Pulvermueller proposes that strong functional links between lan-
guage and motor systems have developed as a consequence of the
fact that actions and their referents often co-occur near-simulta-
neously. Specifically, the action and the word co-occur frequently,
and thereby, neural populations recruited for processing a word
form and those involved in processing the referent body movement
frequently fire together and become strongly linked (Pulvermueller,
1999, 2001). Due to the strong within-assembly connections that
link language and action representations, action word recognition
will thereby automatically and invariably trigger activation in spe-
cific action-related networks.
However, in contrast to what would be expected if embodied
representations are indeed automatic and invariant, several stud-
ies have failed to find motor-related activity for words with an
action-semantic component (Rueschemeyer et al., 2007; Raposo
et al., 2009). Raposo et al. (2009) showed in a recent fMRI study
that action verbs in isolation (e.g., kick) or in literal sentences
(e.g., kick the ball) elicit a response in motor/premotor cortices.
Action verbs in an idiomatic context (e.g., kick the bucket), how-
ever, did not elicit such activations. These findings strongly chal-
lenge the automaticity of motor-related activity for action words
and rather suggests that the activation of meaning attributes of
words is a flexible and contextually dependent process (but see
also Boulenger et al., 2009). In a similar vein, Rueschemeyer et al.
(2007) showed that processing of morphologically complex verbs
built on motor stems showed no differences in involvement of
the motor system when compared with processing complex verbs
with abstract stems. For example, the difference between the verb
begreifen (to comprehend) vs. bedenken (to think). In the first case
the morphologically complex verb is a prefixed form of the sim-
ple motor verb greifen (to grasp), whereas in the latter case it is
a prefixed form of the abstract verb denken (to think). All these
www.frontiersin.org October 2010 | Volume 1 | Article 150 | 3
van Dam et al. Conceptual flexibility
semantic context in which a word is encountered. Specifically, in
trials in which the semantic context emphasizes the dominant
motor properties of a concept, we expect faster onset latencies to
words denoting objects for which the functional use is congruent
with the prepared movement in directionality. Furthermore, we
expect that this facilitation effect is not present if the semantic
context emphasizes motor properties of a concept that are not
related to the object’s functional use.
ExpErImEnt
mEthod
Participants
Thirty-five subjects participated; the average age was 21.3 years.
All subjects were students at the Radboud University Nijmegen
and participated either for money or course credit. No subject was
aware of the purpose of the experiment.
Stimuli
Words were presented in white Arial fonts on a black background.
The average word length was 10 letters. For a viewing distance
of 100 cm, the stimuli subtended on average a visual angle of
0.57° × 2.86°. A total of 120 letter string stimuli were created for
the experiment (stimuli with English translations can be seen in
Table 1). Eighty were real Dutch words denoting familiar objects
and comprised the critical experimental stimuli. The remaining 40
stimuli were Dutch pseudowords (i.e., phonotactically and ortho-
graphically legal letter strings with no meaning in Dutch). Target
stimuli belonged to one of two experimental conditions: (1) words
denoting objects for which the functional use is associated with a
movement toward the body (e.g., telephone, photo camera), and
(2) words denoting objects for which the functional use is associ-
ated with a movement away from the body (e.g., hammer, pencil).
Target stimuli were presented in two contexts: (1) target words were
preceded by a word that emphasized the action feature related to
the functional use (e.g., conversation-telephone, nail-hammer), and
(2) target words were preceded by a word that emphasized an action
feature not related to the functional use (e.g., adapter-telephone,
tool belt-hammer).
To test that stimuli were truly matched with regards to important
psycholinguistic variables, a questionnaire was administered to 15
native Dutch speakers who did not participate in the behavioral
experiment (see Table 2 for results). In this questionnaire, par-
ticipants were asked to rate words on a 7-point scale with respect
to (1) the imageability of the noun (1 = very difficult to imagine
the referent noun, 7 = very easy to imagine the referent noun),
(2) whether the noun denoted an object that you typically bring
toward or away from the body (−3 = toward the body, +3 = away
from the body).
The results of the questionnaire showed that nouns were matched
across the two conditions with respect to imageability (body:
M = 6.82, SE = 0.024; world: M = 6.76, SE = 0.029), t(1,78) = 1.46,
p > 0.1. For object nouns, participants agreed that body words
referred to objects that you typically bring toward the body, world
words referred to objects you typically bring away from the body
(body: M = −1.13, SE = 0.049; world: M = 2.33, SE = 0.057), both
means significantly differed from 0 as indicated by one-sample
Table 1 | Dutch words associated with a movement toward the body
(body words) and a movement away from the body (world words).
English translations are printed in italics.
Body words World words
Haarband Hair ribbon/Hairband Zwabber Swab/Mob
La Drawer Spaarpot Money box
Sjaal Scarf Spade Spade/Shovel
Microfoon Microphone Zaag Saw
Loep Magnifying glass Vaas Vase
Hoed Hat Naald Needle
Nagelvijl Nail file Kaars Candle
Pleister Band-Aid Plant Plant
Fluit Flute Flesopener Bottle opener
Bril (pair of) Spectacles Deegroller Rolling pin
Wijnglas Wine glass Koekenpan Frying pin
Mok Mug Voetbal Football
Make-up Make up Theepot Teapot
Zakdoek Handkerchief Speld Pin
Lolly Lollipop Stempel Stamp
Halssnoer Necklace Sleutel Key
Helm Helmet Lamp Lamp
Telefoon Telephone Schep Scoop/Shovel
Shampoo Shampoo Knikker Marble
Armband Bracelet Bijl Axe
Tondeuse (pair of) Clippers Baksteen Brick
Mondharmonica Mouth organ Fakkel Torch
Want Toothbrush Bel Bell
Tandenborstel Mitten Hamer Hammer
Handdoek Towel Computer Computer
Ring Ring Hengel Fishing rod
Trompet Trumpet Pen Pen
Schoen Shoe Boor Drill
Lippenstift Lipstick Gloeilamp Lightbulb
Lepel Spoon Trommel Drum
Oorbel Earring Kapstok Coat rack
Borstel Brush Verfpot Pot/Tin of paint
Verrekijker Binoculars Vergiet Stainer/Colander
Stropdas Tie/Necktie Karaf Decanter/Carafe
Gordel Belt Garde Whisk
Vork Fork Paraplu Umbrella
Parfum Perfume Mes Knife
Horloge Watch Potlood Pencil
Jas Jacket Ventilator Fan
Fototoestel Photo camera Dobbelsteen Dice
t-tests (all p-values <0.001). In order to obtain an objective measure
of word frequency, we calculated the mean lemma frequency per
million for each condition using the lexical database (Baayen et al.,
1993). This gave a mean of 567 (SE = 128.3) for the body words
and 487 (SE = 119.3) for the world words. An independent sample
t-test indicated words were matched on frequency, t(1,78) = 1.48,
p > 0.1. Additionally, independent sample t-tests indicated that
nouns were matched with regard to length (body: M = 6.8, world:
M = 6.3), t(1,78) = 0.90, p > 0.2.
Frontiers in Psychology | Cognition October 2010 | Volume 1 | Article 150 | 4
van Dam et al. Conceptual flexibility
Data analysis
We measured the latencies to recognize that a presented Dutch
word was lexically valid defined as the time difference between
word onset and release of the start button. Average RTs can be seen
in Figure 2. Additionally, we recorded movement times (MTs; i.e.,
the time from releasing the start button until depressing the target
button). Trials with extreme RTs or MTs (2 × STD ± mean) were
treated as outliers and excluded from further analysis. This led to
an exclusion of 9.12% of the data. The significance criterion for all
analyses was set to α = 0.05.
rEsults
The error rates in the lexical decision task were on average lower than
2% and therefore not further analyzed. RTs were averaged for each
participant in each condition (see Table 3 for means) and submitted
to a two-way repeated measures analysis of variance (ANOVA) with
factors Action Congruency (congruent vs. incongruent) and Context
(focus on dominant vs. non-dominant action feature). The main effect
for Action Congruency and Context did not reach significance, both
p-values >0.2. Importantly, in line with our hypothesis, the ANOVA
revealed a significant interaction F(1,34) = 5.76, p < 0.05,
ηp
20145=.,
indicating that the RT differences between action congruent and
action incongruent words were modulated by the different context
conditions. To further explore this interaction, we calculated post hoc
paired sample t-tests. When the context emphasized the dominant
action feature, words congruent with the prepared action were proc-
essed faster than incongruent words, t(34) = −2.21, p < 0.05. However,
if the context focused on non-dominated action features, the mean
RTs to action congruent and action incongruent words did not differ
statistically, t(34) < 1, and the RT pattern reversed descriptively.
Movement times were averaged for each participant in each
condition and submitted to a two-way repeated measures ANOVA.
The main effect for Action Congruency and Context did not reach
significance, both p-values >0.05. Furthermore, the ANOVA did not
reveal a significant interaction (p > 0.5).
Procedure and design
Participants were seated comfortably in front of a computer moni-
tor, and responded by means of a key press (i.e., by pressing either
a key that was located nearer or further from the body). To start a
trial participants had to press the start button of a response device
(located in the middle of the response device). Subsequently, they
received a cue (i.e., the letters A or B) that signaled them to prepare
a movement (either toward or away from the body) which they
only executed if the second word was lexically valid. Participants
were instructed to read both words carefully. The advent of the
two words was signaled by a fixation cross (appearing centrally
for 500 ms). The first word was presented for 1000 ms. The second
word appeared 1000 ms after the offset of the first word, calling
for a speeded response to the identity of the word (i.e., a response
in the case that the second word was a real word in Dutch). The
second word remained visible until participants responded, or for
a maximum of 2000 ms (see Figure 1 for an illustration of a Go
and NoGo trial). That is, the first presented word was not clearly
associated with a specific direction or action, the differences
we report are relative differences between reaction times (RTs)
acquired in response to the second presented word. The experi-
ment comprised 160 critical trials composed by 20 replications
of the factorial combination of two movement cues, two word
types, and two contexts. That is, the direction of the participant’s
response and the motor program generally associated with the
word’s referent either corresponded or not (action congruent
vs. action incongruent) and the word was either presented in a
language context highlighting the functional use or a less typical
use of the object (focus on dominant vs. non-dominant action
feature). The order in which word pairs were presented was coun-
terbalanced over participants. That is, half of the participants
first saw a target word (e.g., telephone) preceded by a word that
emphasized the dominant action feature (conversation) and then
the same target word preceded by a word that emphasized a non-
dominant action feature (adapter) and vice versa.
FIGURE 1 | Illustration of a Go and NoGo trial.
www.frontiersin.org October 2010 | Volume 1 | Article 150 | 5
van Dam et al. Conceptual flexibility
flexible and contextually dependent process, and not a process
of an automatic and invariant simulation of a specific motor
program. In other words, semantic features are flexibly recruited
with respect to the type of properties emphasized by the semantic
context (see also Hoenig et al., 2008; Masson et al., 2008; Raposo
et al., 2009).
The congruency effect in the present study extends the results of
our previous research on motor involvement in language processing
(Rueschemeyer et al., 2010a) and demonstrates the important role
of contextual information in the embodied processing of language.
We show that preparing an action congruent to the typical, func-
tional use of an object facilitates processing of the word denoting
the object. We suggest that words presented in isolation rely on a
default representation, which highlights the typical functional use
of the object. In our current results we extend this finding to show
(not surprisingly) that words presented in a context highlighting
the denoted object’s functional use are also facilitated by preparing
a congruent action.
Interestingly, the action congruency did not affect the word
processing time if the semantic context emphasized action fea-
tures not belonging to the core of the concept. There are two
alternative explanations for this finding: first, it is possible that
motor semantic features are not activated at all in a non- dominant
action context. Alternatively, it is possible that dominant concep-
tual features are co-activated along with non-dominant features.
In other words, cup in the context of sink activates both motor
programs associated with moving a cup toward the sink and
motor programs associated with bringing the cup toward the
mouth. This activation of two opposite motor programs might
have resulted in a null effect for trials in which properties unre-
lated to the functional use of the object are emphasized. In this
scenario, the priming effect of movement preparation on lexi-
cal access is canceled out, because motor codes underlying two
opposing movements are activated simultaneously. This explana-
tion is in line with the findings of Hoenig et al. (2008) in which
they show that activation in regions coding non-dominant object
features increased if semantic context encouraged participants
to focus on the non-dominant feature. However, brain regions
coding information about dominant features were co-activated,
even when not directly probed by the task. The authors argue that
dominant object features are co-activated with non-dominant
object features through collateral support (see also Farah and
McClelland, 1991).
The reliable congruency effect for trials in which there was a
contextual focus on action properties related to the functional use
of the object, bolsters the claim that in processing of a word with
an action-semantic component we activate information stored in
modality specific sensory-motor systems (Hoenig et al., 2008). The
present study, however, cannot determine whether motor system
involvement is a fundamental necessity or a consequence of word
comprehension. One might argue that the reason for observing
a congruency effect between prepared movement direction and
associated movement direction is because the person voluntarily
images the functional use of the referent object, after the meaning
of the object word is already understood. The design of our current
study prevents us from ruling out this possibility. For a better under-
standing of the exact nature of the interaction between context
dIscussIon
The present study clearly demonstrates contextual effects on
embodied word processing, evidenced by a reliable Action
Congruency × Context interaction effect. Specifically, a congru-
ency effect (i.e., faster word recognition times) is observed for
trials in which the direction of the participant’s response and
the motor program generally associated with the word’s refer-
ent correspond, but only if the word is presented in a language
context highlighting the functional use of the object (e.g., thirst
– cup). In a language context highlighting a less typical use of
the object (e.g., sink – cup), the congruency effect disappears.
These findings suggest that activation of modality specific (in
this case motor) information during word comprehension is a
FIGURE 2 | Average reaction times (RTs) for words, as a function of the
congruency between the cue and associated movement direction, and
the contextual focus (focus on dominant action feature vs. non-dominant
action feature).
Table 3 | Average performance rates (PR), reaction times (RTs), and
movement times (MTs) with standard errors for congruent and
incongruent trials in both the dominant focus and non-dominant focus
condition.
PR (SE) RT (SE) MT (SE)
DOMINANT FOCUS
Congruent 98.4 (0.26) 543 (14.73) 276 (16.05)
Incongruent 98.3 (0.37) 553 (16.21) 272 (15.55)
NON-DOMINANT FOCUS
Congruent 98.6 (0.35) 551 (15.54) 272 (15.51)
Incongruent 98.3 (0.37) 548 (15.84) 269 (15.69)
Table 2 | Mean ratings of the pre-tests.
Body words World words
Length 6.8 6.3
Lemma frequency 567 487
per million (CELEX)
Imageability 6.82 6.76
Action association −1.33 2.33
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van Dam et al. Conceptual flexibility
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Conflict of Interest Statement: The
authors declare that the research was
conducted in the absence of any com-
mercial or financial relationships that
could be construed as a potential conflict
of interest.
Received: 16 June 2010; paper pending pub-
lished: 03 July 2010; accepted: 21 August
2010; published online: 04 October 2010.
Citation: van Dam WO, Rueschemeyer
S-A, Lindemann O and Bekkering H
(2010) Context effects in embodied lexical-
semantic processing. Front. Psychology
1:150. doi: 10.3389/fpsyg.2010.00150
This article was submitted to Frontiers
in Cognition, a specialty of Frontiers in
Psychology.
Copyright © 2010 van Dam, Rueschemeyer,
Lindemann and Bekkering. This is an open-
access article subject to an exclusive license
agreement between the authors and the
Frontiers Research Foundation, which
permits unrestricted use, distribution, and
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original authors and source are credited.
and embodied word processing, we need to further investigate the
temporal dynamics of the observed contextual effects on embodied
word processing.
conclusIon
The current study demonstrates that the interaction between
lexical-semantic processing and movement preparation processes
was modulated by the context in which these words were encoun-
tered. Together the data suggest that context plays a fundamental
role in sensory-motor activations during language processing. That
is, the activation of specific motor properties in language compre-
hension is flexible and context-dependent.
AcknowlEdgmEnts
The authors thank P. de Water, and G. van Oijen for technical
support. The study was supported by NWO-Veni grant nr (016-
094-053) to Shirley-Ann Rueschemeyer and NWO-VICI grant nr
(453–05–001) to Harold Bekkering.
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