The Semantics of Syntax: The Grounding of Transitive - IA de Investigación especializada en el MIT

La semántica de la sintaxis: La base de lo transitivo
and Intransitive Constructions

Wessel O. van Dam and Rutvik H. Desai

Abstracto

■ Embodied theories of language maintain that brain areas as-
sociated with perception and action are also involved in the
processing and representation of word meaning. A number of
studies have shown that sentences with action verbs elicit ac-
tivation within sensory–motor brain regions, arguing that
sentence-induced mental simulations provide a means for
grounding their lexical-semantic meaning. Constructionist the-
ories argue, sin embargo, that form–meaning correspondence is
present not only at the lexical level but also at the level of
constructions. We investigated whether sentence-induced mo-
tor resonance is present for syntactic constructions. We mea-
sured the BOLD signal while participants read sentences with

(di)transitive (caused motion) or intransitive constructions that
contained either action or abstract verbs. The results showed a
distinct neuronal signature for caused motion and intransitive
syntactic frames. Caused motion frames activated regions asso-
ciated with reaching and grasping actions, including the left
anterior intraparietal sulcus and the parietal reach region. En-
transitive frames activated lateral temporal regions commonly
associated with abstract word processing. The left pars orbitalis
showed an interaction between the syntactic frame and verb
class. These findings show that sensory–motor activation elic-
ited by sentences entails both motor resonance evoked by sin-
gle words as well as at the level of syntactic constructions. ■

INTRODUCCIÓN

Traditionally, perceptual and motor brain systems were
seen as categorically distinct from our representational
systems subserving cognition. In this symbolic view, estafa-
ceptual representations were taken to be of a symbolic
and amodal nature and lexical-semantic meaning inde-
pendent of sensory–motor systems of the brain (Kintsch,
2008; Pylyshyn, 1984; Fodor, 1983; katz & Fodor, 1963).
Opponents of such a symbolic account have raised the
problem of how symbols can be mapped to their refer-
ents in the real world (Harnad, 1990) and its failure to
provide an adequate description of the transduction pro-
cess that maps perceptual states to amodal conceptual
symbols (Barsalou, 1999). In contrast to such a symbolic
cuenta, embodied theories argue that experiential
traces stored in sensory–motor brain areas can pro-
vide the means for grounding lexical-semantic meaning
(Barsalou, 2008; Pulvermüller, 1999; Glenberg, 1997). En
la última década, a plethora of studies using various ex-
perimental techniques have directly contrasted predic-
tions from symbolic and embodied accounts. Findings
from behavioral experiments suggest that processes that
play a role in perception and action are also involved in
comprehending language about action (see Fisher &
Zwaan, 2008; Zwaan & Kaschak, 2008, for reviews). en un
similar vein, fMRI studies have shown that comprehen-
sion of action verbs, action sentences, and words denot-

University of South Carolina

ing manipulable objects frequently activate sensory–
motor brain areas (see Kiefer & Pulvermüller, 2012;
Meteyard, Cuadrado, Bahrami, & Vigliocco, 2012; Binder
& Desai, 2011, for reviews).

This work has focused almost exclusively on meanings
of nouns and verbs, either isolated or embedded in sen-
tenencias. Several studies that use sentences compared
action-related sentences with abstract sentences that have
an identical or similar sentence structure (p.ej., Desai,
Conant, Binder, Parque, & Seidenberg, 2013; Desai, Binder,
Conant, Mano, & Seidenberg, 2011; Desai, Binder,
Conant, & Seidenberg, 2009; Raposo, Moss, Stamatakis,
& tyler, 2009; Aziz-Zadeh, wilson, Rizzolatti, & Iacoboni,
2006; Tettamanti et al., 2005). Por ejemplo, Tettamanti
et al. (2005) showed that listening to sentences that de-
scribe actions (p.ej., “I bite an apple”) as compared with
similar transitive sentences with abstract content (p.ej.,
“I appreciate sincerity”) activated parts of the motor system.
These findings have been taken to show that, to convey
the meaning of a sentence as a whole, we rely on ex-
periential traces of the referent action stored in sensory–
motor brain areas. Además, recent studies have
suggested that conceptual features contribute to word
and sentence level meaning in a flexible and context-
dependent manner (Kemmerer, 2015; De la presa, Van Dijk,
Bekkering, & Rueschemeyer, 2012; Hoenig, Sim, Bochev,
Herrnberger, & Kiefer, 2008). Sin embargo, in most (if not all)
of the studies in the conceptual representation literature,
the effects of sentence structure are subtracted out by
diseño.

Revista de neurociencia cognitiva 28:5, páginas. 693–709
doi:10.1162/jocn_a_00926

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Traditionally, it is assumed that argument structures con-
vey their meaning by means of the main verb. Por ejemplo,
in “Pat gave Chris a cake,” the meaning of transfer apparent
in the overall expression is specified by the three-argument
verb “give.” That is, the three-argument verb “give” is ex-
pected to appear with phrases corresponding to the three
characters required in the act of giving: a giver, a recipient,
and something that is given. Por lo tanto, it can be argued
that the interpretation of this sentence pattern as a
whole can be determined on the basis of its main verb.
Sin embargo, in many instances, the main verb does not re-
liably determine the interpretation of a sentence. por ejemplo-
amplio, “Liza bought a book for Zach” can mean that Liza
bought a book for a third person because Zach did not
have time to buy it himself. Sin embargo, “Liza bought Zack
a book” can only mean that Liza intended to give the
book to Zach (Goldberg, 2003). Eso es, although both
sentences involve the same verb “to buy,” the interpreta-
tion of these sentences is not reliably determined by its
independent specification. The verb-centric view deals
with such cases by proposing that verbs have multiple
distinct senses because of the different linking patterns
between syntax and semantics (Levin, 1995; Pinker,
1989). The main criticism of this view is that it leads to
a proliferation of polysemous lexical entries (Goldberg,
1995).

An alternative is provided by a family of theories
known as “contructionist” approaches (p.ej., Bergen &
Chang, 2003; Goldberg, 1995, 2003; Croft, 2001; Langacker,
1999; Fillmore, kay, & O’Connor, 1988; Lakoff, 1987). Estafa-
structions are stored pairings of form and function, incluir-
ing words, idioms, or linguistic patterns that may be fully or
partially filled. A central idea in these approaches is that
constructions themselves have meaning. Eso es, form–
meaning correspondences do not only exist at the level
of verbs or single words but also at the level of construc-
ciones. Constructions are pairings of form and meaning,
and therefore, a difference in syntactic form always spells
a difference in meaning (Langacker, 1999; Wierzbicka,
1988; Haiman, 1985; bolinger, 1968). This idea that argu-
ment structure constructions themselves carry meaning is
often referred to as the “principle of no synonymy of gram-
matical forms” (bolinger, 1968).

Given that embodiment theories reject the idea that
language can convey meaning without it somehow being
grounded in real-world sensory–motor experiences, este
would suggest that semantics of constructions should be
grounded in a similar fashion. In line with this idea, recent
theorists have argued that sensory–motor activation elic-
ited by sentences might be a complex phenomenon that
entails both motor resonance evoked by single words
and word combinations, but also more general motor res-
onance at the level of constructions (Kemmerer, 2006).
Bergen and Chang (2003), in their Embodied Construction
Grammar approach, argue that syntactic constructions can
drive specific perceptual and motor simulations of a lan-
guage user. Por ejemplo, the directed motion construction

may lead to the mental simulation of an event involving an
animate mover moving along a path, even if the main verb
in the construction does not explicitly specify motion (as in
“The cat meowed down the street”). Experimental evi-
dence for the principle of no synonymy of grammatical
forms has been provided by, Por ejemplo, work showing
that comprehenders access different meanings for sen-
tences using transitive versus ditransitive constructions
(Kaschak & Glenberg, 2000). A ditransitive construction
is a construction consisting of a verb, an agent argument,
a recipient-like argument, and a theme argument (p.ej., “He
threw the ball to John”). It can be argued that, Por ejemplo,
the ditransitive (double-object) construction activates
a transfer-of-possession schema (X causes Y to have
z), whereas a prepositional dative may activate a caused-
motion schema (Goldberg, 1995).

These findings mesh with evidence from action observa-
ción, which suggests that the transitivity of an action is a
crucial factor in determining the type of simulations. El
finding that some mirror neurons of the macaque monkey
were only responsive to object-directed actions suggests
that activation of the parietal-frontal network depends on
whether an action is goal directed (Tkach, Reimer, &
Hatsopoulos, 2007; Gallese, Fadiga, Fogassi, & Rizzolatti,
1996; Rizzolatti, Fadiga, Gallese, & Fogassi, 1996).

A number of fMRI studies also suggest a neural disso-
ciation between these two types of actions. The inferior
frontal gyrus (IFG), lateral precentral gyrus, inferior pari-
etal lobule (IPL), posterior middle temporal gyrus (MTG),
and the posterior part of the superior parietal lobule have
been shown to play a role in the representation of tran-
sitive actions (Caspers, Zilles, Laird, & Eickhoff, 2010;
Sakreida, Schubotz, Wolfensteller, & por cramon, 2005;
Buccino et al., 2001). Intransitive actions, en el otro
mano, seem to rely more on posterior regions in the pa-
rietal cortex, the angular gyrus (AG), and STS regions
(Grosbras, Beaton, & Eickhoff, 2012; Lui et al., 2008). Re-
cruitment of partially different brain regions might reflect
the fact that transitive actions are object related and tend
to be more complex (p.ej., “throwing a ball”), whereas in-
transitive actions are not related to an object (p.ej., “waving
goodbye”). Additional evidence for a neural dissociation
between transitive and intransitive actions comes from
patient studies. Por ejemplo, watson, Fleet, Rothi, y
Heilman (1986) observed an isolated disturbance of tran-
sitive but not intransitive movements in bilateral apraxia
(see also Mozaz, Rothi, anderson, Crucian, & Heilman,
2002; Foundas et al., 1999).

We hypothesized that mental simulations in service of
language comprehension will be modulated by whether a
construction (or syntactic frame) is of a transitive (caused
movimiento) or intransitive nature. En el estudio actual, nosotros
investigated whether distinct patterns of sensory–motor ac-
tivation can be observed for different syntactic construc-
ciones, independent of the main verb. We also investigated
if language-induced mental simulations differ as a function
of the class of main verb used within the construction and

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whether the factors of construction and verb class interact
with each other.

Para tal fin, we measured changes in the hemodynamic
response of participants while they read transitive and in-
transitive sentences that either contained a concrete or ab-
stract main verb. We used a specific type of transitive
oraciones; transitives were defined here as a construction
with the syntactic realization (NP V NP PP). Such a transitive
construction is characterized by a transitive verb, an agent
argumento, a recipient-like argument, and a theme argu-
mento (Malchukov, Haspelmath, & Comrie, 2010). El
schematic constructional meaning of this phrase is that
of the causer argument directly causing the theme argu-
ment to move toward a recipient-like argument (X causes
Y to move to Z), when used with a concrete verb. Tenga en cuenta que
the schematic constructional meaning is closely related to
the transfer-of-possession schema (X causes Y to have Z),
with the difference that the sentence “You threw the ball to
her” does not actually entail that the recipient-like argu-
ment receives the ball (es decir., a change of possession), Alabama-
though that may typically be the case. Throughout the
manuscript, we refer to our main constructions of interest
as transitive (caused motion) and intransitive construc-
ciones. It needs to be noted that these terms are very broad
in nature and actually embrace a whole family of construc-
tions with more specific meanings. Por ejemplo, a more
specific label for the construction used in our transitive
sentences would be the “caused motion construction.”
Por lo tanto, to be more specific, we will adopt the term
“caused motion construction” from here on.

We hypothesized a main effect of verb class, with sen-
tences containing a concrete/action-related main verb eli-
citing stronger activation in the anterior IPL, primary
motor cortex, premotor cortex, posterior MTG and infe-
rior temporal gyrus (ITG), superior frontal gyrus (SFG),
and IFG. Por otro lado, for sentences containing an
abstract main verb, we expect stronger activation within
the anterior and middle STS (Desai et al., 2009, 2011,
2013). Además, given that transitive sentences are
strongly associated with a caused motion (X causes Y to
move Z) schema (and take a direct object), we expect
these sentences to elicit stronger activation in regions
within the posterior middle/inferior temporal cortex that
are involved in the processing of motion (Chen, Widick,
& Chatterjee, 2008) and within the anterior IPL, a brain
region that is involved in representing complex hand–
object interactions and tool use. Stronger activation for
intransitive sentences is hypothesized in the anterior and
middle lateral temporal lobes, given that these regions
are associated with more abstract semantics.

MÉTODOS

Participantes

Fourteen individuals participated in the study, all of whom
were right-handed and between 19 y 22 years old (m =

20.64 years old, DE = 0.93 years old; six men). All partic-
ipants had normal or corrected-to-normal vision and no
history of neurological disorders. Before the experiment,
participants were informed about the experimental pro-
cedures, signed informed consent forms, and were given
practice trials according to a protocol sanctioned by the
institutional review board of the University of South
Carolina. Participants were paid for their participation.
Every participant (excepto por uno) underwent two scans
on two different days within a 1-week period.

Estímulos

The four main experimental conditions contained 35 sen-
tences each. They were (1) concrete caused motion
(CCM; p.ej., “You threw the ball to her”; “He gave the
pizza to you”), (2) concrete intransitive (CI; p.ej., "Tú
waved at her whole family”; “He clapped after your big
concert”), (3) abstract caused motion (ACM; p.ej., "Tú
delegated the task to her”; “He communicated the news
to you”), y (4) abstract intransitive (AI; p.ej., "Tú
thought about her feelings”; “He cared about your new
program”). The caused motion sentences used a ditran-
sitive verb with a preposition. Además, 30 filler sen-
tences were also included (p.ej., “You deduced the
truth”; “He met the person”). The concrete conditions
used an action-related verb, whereas the abstract condi-
tions used verbs that did not have a strong association
with actions. Verb concreteness ratings showed no differ-
ence between the CCM and CI conditions and the ACM
and AI conditions (all ps > .5). Concreteness ratings for
the main verb used in the concrete sentences (m = 3.92)
were significantly higher than for the main verb used in
the abstract sentences (m = 2.48; pag < .001). There were also pseudoword conditions in the experiment, but we do not discuss them here and focus on the interpretable conditions. The sentences of all experimental conditions were matched on a number of psycholinguistic variables (see Table 1). Two-sample t tests ensured that all sentences were matched on number of words (all ps > .10), total
number of phonemes (all ps > .20), total number of syl-
lables (all ps > .07), total number of letters (all ps > .10),
and the number of persons mentioned in a sentence (ex-
act match). Además, 11 participants performed a be-
havioral version of our experiment in which we collected
RT data for the different sentences in a meaningfulness
judgment task. Paired-sample t tests showed that our
stimuli were not perfectly matched on RTs. Participantes
were faster to respond to ACM (m = 1356 mseg) than
to AI (m = 1506 mseg) oraciones ( pag < .01). Likewise, par- ticipants were faster to respond to CCM (M = 1404 msec) than to CI (M = 1532 msec) sentences ( p = .001). In ad- dition, participants were faster to respond to ACM (M = 1356 msec) than to CI (M = 1532 msec) sentences ( p < .001). A regressor representing the mean-centered RT for each sentence from the behavioral experiment van Dam and Desai 695 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 5 2 8 6 / 9 5 3 / 1 6 9 9 5 3 0 / 5 1 7 7 7 8 o 4 c 7 n 0 _ 7 a / _ j 0 o 0 c 9 n 2 6 _ a p _ d 0 0 b 9 y 2 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j t f / . u s e r o n 1 7 M a y 2 0 2 1 Table 1. Psycholinguistic Variables No. of words No. of phonemes No. of syllables Length Verb concreteness Noun concreteness Relative freq verb/tran Relative freq verb/intran RT Log HAL freq ACM 6.03 (0.17) 21.26 (2.77) 8.60 (1.09) 25.74 (2.60) 2.60 (0.52) 3.28 (0.83) 0.57 (0.20) 0.23 (0.15) 1356 (362) 9.84 (0.82) AI 5.91 (0.95) 22.23 (3.39) 8.71 (1.56) 27.49 (4.40) 2.38 (0.65) 3.11 (0.95) 0.31 (0.22) 0.49 (0.25) 1506 (411) 9.78 (1.50) CCM 6.26 (0.82) 20.57 (3.39) 8.03 (1.46) 26.14 (4.53) 3.82 (0.43) 4.70 (0.37) 0.46 (0.16) 0.31 (0.19) 1404 (354) 8.86 (1.10) CI 6.17 (1.18) 21.37 (3.95) 7.97 (1.93) 27.23 (5.34) 4.01 (0.48) 4.52 (0.56) 0.43 (0.16) 0.35 (0.17) 1532 (382) 9.38 (1.14) was used as an additional item-wise regressor to account for variance due to time-on-task and difficulty. In addition, we calculated the mean concreteness rat- ing of the nouns of each of the four experimental condi- tions using the Brysbaert, Warriner, and Kuperman (2013) database. In the case in which the database did not have an entry for a compound word (e.g., “dining table”), we used the concreteness rating for the head noun (i.e., “table”) of the compound word. Two-sample t tests revealed an overall difference in the mean noun concreteness ratings between the concrete sentences (M = 4.61) and the abstract sentences (M = 3.20; p < .001). No difference was observed between transitive (caused motion) and intransitive frames, neither for con- structions with a concrete verb or an abstract main verb (Table 1). A number of studies show, using single-verb stimuli, that brain activations can be affected by other character- istics of verb classes, such as whether they allow causative alternation or number of obligatory arguments. The rele- vance of these results is not clear in the present case be- cause we used sentences instead. Single verbs were used in these experiments as stimuli to examine the dominant or default response to the verb (e.g., those that require two arguments vs. three or those with and without caus- ative alternation). When the argument structure is real- ized, the salience of these defaults (beyond general difficulty effects because of low familiarity of certain us- age that is allowed but is uncommon) can be expected to reduce. We are not aware of any experiments that show similar differences between verb classes while using them in sentences. Nonetheless, we examined three such variables as a cautionary measure. First, verbs with causative alternation (those that can be used both transitively and intransitively) and those without can potentially differ. A recent study by Meltzer- Asscher, Schuchard, den Ouden, and Thompson (2012) suggests that the pattern of neural activity elicited by a verb is different for verbs, which can alternate between transitive and intransitive frames and those that cannot. We used the VALEX database to obtain information on the relative frequency with which the main verb appears in a transitive versus intransitive construction (Korhonen, Krymolowski, & Briscoe, 2006; see Table 1). Two-sample t tests indicated that caused motion and intransitive sen- tences differed in the relative frequency with which its main verb was used in a caused motion versus intransitive construction ( p < .001). That is, the main verbs in the caused motion condition occur relatively more often in caused motion constructions, whereas the main verbs in the intransitive condition occur relatively more often in intransitive constructions. Importantly, the concrete and abstract sentences did not differ in the relative fre- quency with which its main verb was used transitively versus intransitively ( p > .50).

A second factor is whether verbs used in the transitive
(caused motion) conditions obligatorily take two or three
argumentos. Some studies have indicated inferior parietal
and posterior superior temporal regions to be involved in
processing argument structure complexity, mostrando que
verbs with more obligatory arguments show a stronger
response than verbs with fewer obligatory arguments
(Thompson, Bonakdarpour, & Fix, 2010; Thompson
et al., 2007; Ben-Shachar, Hendler, Kahn, Ben-Bashat, &
Grodzinsky, 2003). We conducted an analysis on the
number of arguments a verb obligatorily takes by using
VerbNet (Kipper, Dang, & Palmer, 2000). This analysis in-
dicated that there was no difference in the number of ar-
guments a verb obligatorily takes between the CCM (m =
2.11) and ACM (m = 2.17) condiciones ( pag = .50).

Correspondingly, intransitive verbs have been subcate-
gorized as unergatives and unaccusatives. Both verb
types have the same number of arguments (es decir., a single
argumento) but differ syntactically. Unergatives and unac-
cusatives are often delineated in terms of whether the
verb takes an internal versus external argument. Such a

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distinction is, sin embargo, associated with a syntactic move-
ment analysis, which is only one of many approaches
used in linguistic studies of unaccusativity (see Roehm,
Sorace, & Bornkessel-Schlesewsky, 2012). Different ap-
proaches in this branch of linguistics tend to agree that,
in an unaccusative construction, the participant is likely
to be a patient or a nonvolitional causer of the event,
whereas in the unergative construction, some property
inherent in the argument of the verb is responsible for
bringing about the event. Some studies with healthy par-
ticipants and brain-damaged patients suggest that the
neural structures that underlie the processing of these
subtypes of intransitive verbs are dissociable (Agnew,
van de Koot, McGettigan, & Scott, 2014; Shetreet &
Friedmann, 2012; McAllister, Bachrach, Waters, Michaud,
& Caplan, 2009; Luzzatti et al., 2002). Aquí, there was no
difference in the number of unergative verbs between
the CI (norte = 5) and AI (norte = 3) condiciones ( pag = .46). Sev-
eral recent authors, sin embargo, have argued that the use of
the terms “unergative” and “unaccusative” might be
problematic, given that this distinction in intransitive
verbs cannot account for the acceptability of the use of
certain syntactic constructions (for an extensive discus-
sión, see Kuno & Takami, 2004). Además, Ha sido
argued that the labels “unergative” and “unaccusative”
are used to describe a wide variety of phenomena in dif-
ferent languages and therefore render them vague and
imprecise (dixon, 2010).

Procedimiento

The order of stimulus presentation was randomized indi-
vidually for each participant. A single trial lasted 7.4 segundo
and constituted of the presentation of a single sentence.
A variable jitter time of 0, 500, 1000, o 1500 msec was
included at the beginning of each trial to enhance the
temporal resolution of the acquired signal. After this, a
fixation cross was presented in the center of the screen.
A 400-msec blank screen followed the fixation cross; allá-
después, the sentence appeared on screen for 3000 mseg. El
time between offset of the sentence and onset of the next
trial was filled with a blank screen, to ensure that each trial
lasted exactly 7.4 segundo. Participants were instructed to read
all sentences carefully to be able to answer a surprise yes/no
question after some of the sentences (responses were
made by performing a right-hand button click with their
thumb). Thirty comprehension questions (p.ej., “Did he
clap after your big concert?") were used. This catch trial
design was used to encourage participants to read all sen-
tences for comprehension while avoiding motor execution
after every trial.

fMRI Data Acquisition

Functional images were acquired on a Siemens TRIO
3.0-T MRI system (Siemens, Erlangen, Alemania) equipado
with a 12-channel head coil. BOLD-sensitive functional

images were acquired using a single-shot gradient EPI se-
quence (echo time/repetition time = 34/1850 mseg, 34 ax-
ial slices in ascending order, slice gap = 0.60 mm, field of
view = 208 mm, flip angle = 90°, voxel size = 3.25 ×
3.25 × 3.60 mm3). High-resolution anatomical images were
acquired using a magnetization prepared rapid gradient-
echo sequence (echo time = 4.15 mseg, voxel size =
1 × 1 × 1 mm3, 192 sagittal slices, campo de visión = 256 mm).

fMRI Data Analysis

Functional data were preprocessed and analyzed with the
AFNI software package (Cox, 1996). A standardized pre-
processing pipeline involved registration of functional im-
ages to the anatomy (Saad et al., 2009). Después,
functional images were co-registered (Cox & Jesmanowicz,
1999) and projected into standard stereotaxic space
(Talairach & Tournoux, 1988). The normalized images
were smoothed with an isotropic 5-mm FWHM Gaussian
kernel, and the run mean of each voxel was scaled to
100. The ensuing preprocessed fMRI time series were ana-
lyzed on a participant-by-participant basis using an event-
related approach in the context of voxelwise multiple linear
regression with regressors for each condition (ACM, AI,
CCM, CI,) as well as the filler items and question trials con-
volved with a canonical hemodynamic response function.
Six motion parameters and the signal extracted from the
ventricles were included as noise covariates of no interest.
General linear tests were conducted to obtain the main ef-
fect of Transitivity and Concreteness and the Transitivity ×
Condition interactions.

In a random effects analysis, group maps were created
by comparing activations against a constant value of 0.
The group maps were thresholded at voxelwise p < .01 and corrected for multiple comparisons by removing clusters smaller than 800 μl to achieve a map-wise cor- rected two-tailed p < .05.1 Using the 3dClustSim pro- gram with 10,000 iterations, the cluster threshold was determined through Monte Carlo simulations that esti- mate the chance probability of spatially contiguous voxels exceeding the voxelwise p threshold. The analysis was re- stricted to a mask that excluded areas outside the brain as well as deep white matter areas and the ventricles. This mask is based on the probabilistic Desikan–Killiany atlas that contains 35 cortical areas in each hemisphere (Desikan et al., 2006) and the subcortical parcellation provided by FreeSurfer. In addition to this whole-brain analysis, two other (bilateral) ROIs were defined for a more sensitive analysis. One used the primary motor and sensory cortex (M1 and S1) as defined by the HMAT atlas (Mayka, Corcos, Leurgans, & Vaillancourt, 2006). Given its association with transitive actions and action sentence processing, we also defined a bilateral supramarginal gyrus (SMG) ROI. This ROI was defined on the basis of the maximum probability map provided with AFNI, which is based on the Destrieux atlas (Destrieux, Fischl, Dale, & Halgren, 2010). Small volume correction was applied in these ROIs to achieve a van Dam and Desai 697 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 5 2 8 6 / 9 5 3 / 1 6 9 9 5 3 0 / 5 1 7 7 7 8 o 4 c 7 n 0 _ 7 a / _ j 0 o 0 c 9 n 2 6 _ a p _ d 0 0 b 9 y 2 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j f . / t u s e r o n 1 7 M a y 2 0 2 1 corrected p < .05, determined in the same manner as above. RESULTS Concrete–Abstract Areas activated to a greater extent by the Concrete con- dition relative to the Abstract condition included the left MTG and ITG, superior frontal sulcus, inferior frontal sul- cus (IFS), and IFG (pars triangularis). The ROI analyses revealed an additional cluster in the left SMG. The Ab- stract condition relative to the Concrete condition led to stronger activation in the left superior temporal gyrus (STG), STS, fusiform gyrus, right precuneus, MTG, and AG (Figure 1, Table 2). Caused Motion (Transitive)–Intransitive Caused Motion sentences elicited greater levels of activa- tion than Intransitive sentences within the bilateral mid- dle frontal gyrus (MFG), SMG, and left AG as well as precuneus. Structures more strongly activated by the In- transitive condition relative to the Caused Motion condi- tion included bilateral STS and rectal gyrus, left occipital pole, SFG, and precentral gyrus as well as the right MTG and precuneus (Figure 2, Table 2). Overlap Some similarities as well as differences were found be- tween the Concrete > Abstract and the Caused Motion >
Intransitive contrasts (Cifra 3). Both contrasts activated
the left IFG and IFS (pars triangularis), with the former
extending ventrally and the latter extending dorsally
into MFG. A similar dorsal–ventral pattern was found
in the left IPL. Both contrasts activated the SMG, con
the Concrete > Abstract contrast extending ventrally
into the parietal operculum, whereas the Caused Motion >
Intransitive contrast extending dorsally into the anterior
intraparietal sulcus (IPS).

The Abstract > Concrete and Intransitive > Caused Mo-
tion comparisons also showed some similarities (Cifra 4),

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Cifra 1. Areas activated by the Concrete–Abstract contrast. Red-orange colors show greater activation for the Concrete condition; blue-cyan colors
show greater activation for the Abstract condition. Mean percent signal change relative to rest is shown for the CCM, CI, ACM, and AI conditions, en un
sphere of 5-mm radius around the peak voxel. L = left hemisphere; R = right hemisphere.

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Mesa 2. Activations in the Main Contrasts of Interest

Volume Max

Estructura

where the left middle and anterior STS were activated by
both contrasts.

Concrete > Abstract

3078

2268

918

675

3.61 −52 −58

0 L MTG

3.79 −40

3.15 −22

14 L IFG, IFS (pars triangularis)

44 L sup frontal s

2.95 −49 −34

32 L SMG

Abstract > Concrete

5832

2538

1377

1053

1026

810

−4.20

4 −52

26 R precuneus

−3.95 −52 −37

0 L STS

−4.09 −25 −67 −12 L occ temp g

−3.67

−3.29

61 −19 −9 R MTG

55 −55

20 R AG

−4.35 −49

13 −15 L STG

Caused Motion > Intransitive
4.53 −40

5913

26 L MFG

3294

2781

2268

1188

810

3.84 −40 −52

44 L AG, L SMG

3.58

11 R MFG

3.57 −7 −73

29 L precuneus

3.65

3.70

17 R MFG

61 −31

32 R SMG

Intransitive > Caused Motion
13986 −4.86 −52 −40

2 L STS

−3.06 −52

8 L IFG (pars triangularis)

−3.78 −34

25 −3 L IFG (pars orbitalis)

3915

1890

1674

1647

1620

1323

1188

1134

−3.93 −16 −88 −6 L occipital pole

−3.98 −43 −55

23 L STS

−4.34

52 −1 −12 R STS; R MTG

−4.62 −4

−3.61 −4

62 L SFG

32 L SFG

−3.50

−3.58

4 −52

11 R precuneus

52 −9 L rectus; R rectus

−3.99 −49 −10

44 L precentral g

The volume of the cluster (μl), peak z score, Talairach coordinates,
and the anatomical structures that the clusters overlap are shown. L =
left hemisphere; R = right hemisphere; g = gyrus; s = sulcus; sup =
superior.

CCM–ACM

The CCM condition relative to the ACM condition led to
stronger activation in the left IFG (pars opercularis),
MTG, SMG, and paracentral gyrus. Areas activated to a
greater extent by the ACM condition relative to the
CCM condition included the left fusiform gyrus, precu-
neus, posterior dorsal cingulum, right AG, and MTG
(Figure 5A, Mesa 3).

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CI–AI

Areas activated to a greater extent by the CI condition rel-
ative to the AI condition included the left middle occipital
gyrus (MOG), MTG, and SMG. Stronger activation was
observed for the AI condition relative to the CI condition
in the left MTG, STS, right MFG, and SFG (Figure 5B,
Mesa 3).

CCM–CI

Stronger activation was observed for the CCM condition
relative to the CI condition in the bilateral MFG, left IPS,
frontomarginal gyrus and sulcus, and right SMG. Areas ac-
tivated to a greater extent by the CI condition relative to
the CCM condition included the bilateral MTG, left MOG,
right rectus, and posterior ventral cingulum (Figure 5C,
Mesa 3).

ACM–AI

The ACM condition elicited stronger activation relative to
the AI condition in the left MFG, AG, and right occipito-
temporal lingual gyrus. The ROI analyses revealed an addi-
tional cluster in the left SMG. Areas activated to a greater
extent by the AI condition relative to ACM condition in-
cluded the left IFG (pars opercularis), STS, MTG, SFG,
occipital pole, and right MOG (Figure 5D, Mesa 3).

CA × TI Interaction

The left IFG (mostly in pars orbitalis, approximately BA
47) showed an interaction between Concreteness (Estafa-
crete vs. Abstracto) and Transitivity (Transitive [Caused
Motion] vs. Intransitive; Cifra 6, Mesa 3). This interac-
tion arose from a small difference between CCM and CI
conditions and a bigger difference between the abstract
condiciones (AI > ACM).

DISCUSIÓN

The current experiment explored if distinct patterns of
sensory–motor activation can be observed for different

van Dam and Desai

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Cifra 2. Areas activated by the Caused Motion–Intransitive contrast. Red-orange colors show greater activation for the Transitive/Caused Motion
condición; blue-cyan colors show greater activation for the Intransitive condition. Mean percent signal change relative to rest is shown for the CCM,
CI, ACM, and AI conditions, in a sphere of 5-mm radius around the peak voxel. L = left hemisphere; R = right hemisphere.

syntactic constructions and how these patterns relate to
those induced by verbs.

Main Effect of Verb Concreteness

En el estudio actual, constructions that contained a con-
crete verb elicited greater levels of activation in the ante-
rior part of the left IPL (aIPL) and the left inferior MTG
(ITG) and inferior temporal sulcus. The IPL has shown
to be critical for the representation of action plans and
goals and tool use (Luis, 2006; Rumiati et al., 2004)
and the performance of complex hand–object interac-
ciones (Ramayya, vidrio, & Rilling, 2010; hamilton &
Grafton, 2006). It has also been shown to be part of a tool
use network encompassing the middle/inferior temporal
and inferior frontal regions (Ramayya et al., 2010). Dam-
age to the aIPL/IPL is associated with ideomotor apraxia,
which results in impairments in skilled performance of
motor acts, difficulty imitating gestures, impairments in

Cifra 3. Areas activated by the Transitive/Caused Motion >
Intransitive (rojo) and Concrete > Abstract (azul) contrasts. Yellow
indicates the overlap between the two.

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Cifra 4. Areas activated by the
Intransitive > Transitive/Caused
Motion (rojo) and Abstract >
Concrete (azul) contrasts.
Yellow indicates the overlap
between the two.

performing appropriate actions in response to a visually
presented object, and carrying out the appropriate ac-
tion for a certain object ( Jax, Buxbaum, & Moll, 2006;
Haaland, Harrington, & Caballero, 2000; Heilman & Rothi,
1993; Heilman, Rothi, & Valenstein, 1982). Además,
sentences with action verbs elicited greater activation
within the left posterior temporal gyrus, in proximity to
the human motion area (MT+; rees, Friston, & Koch,
2000). This brain region is implicated in accessing con-
ceptual information about motion attributes and is also
associated with tool use (Gallivan & Culham, 2015; Brandi,
Wohlschläger, Sorg, & Hermsdörfer, 2014; Ramayya et al.,
2010; Saygin, McCullough, Alac, & Emmory, 2010; Kable,
Kann, wilson, Thompson-Schill, & Chatterjee, 2005). Sev-
eral studies have obtained similar effects of concreteness
within premotor and primary motor cortices (Raposo
et al., 2009; Aziz-Zadeh et al., 2006; Tettamanti et al.,
2005; Pulvermüller, Härle, & Hummel, 2000). En nuestro
estudiar, concrete sentences showed a higher BOLD signal

than abstract sentences within the left precentral gyrus,
at a reduced statistical threshold (corrected p < .09). Similarly, concrete transitive sentences showed a higher BOLD signal than abstract transitive sentences within the left precentral gyrus, at a slightly reduced statistical threshold (corrected p < .07). Activations were exclu- sively observed within the left premotor cortex. This re- sult is in line with previous findings that, during lexical decisions on manual action verbs, right-handers prefer- entially activate the left premotor cortex, whereas left- handers predominantly activate the right premotor cortex ( Willems, Hagoort, & Casasanto, 2010). In addi- tion, our activation clusters both fell within the premotor cortex, anterior to primary motor areas. Such an anterior shift of activation away from primary sensory and motor cortices has been observed across a number of studies investigating the neural correlates of action understand- ing (Chatterjee, 2010; Willems et al., 2010; Willems, Hagoort, Toni, Hagoort, & Casasanto, 2009; Wallentin, D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 5 2 8 6 / 9 5 3 / 1 6 9 9 5 3 0 / 5 1 7 7 7 8 o 4 c 7 n 0 _ 7 a / _ j 0 o 0 c 9 n 2 6 _ a p _ d 0 0 b 9 y 2 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j f . / t u s e r o n 1 7 M a y 2 0 2 1 Figure 5. Areas activated by the CCM–CI, CI-AI, and ACM–AI contrasts. Red-orange colors show greater activation for the CI, CCM and ACM conditions; blue-cyan colors show greater activation for the ACM, AI, CI, and AI conditions. L = left hemisphere; R = right hemisphere. van Dam and Desai 701 Table 3. Activations in the Remaining Contrasts Table 3. (continued ) Volume Max x y z Structure Volume Max x y z Structure CCM > ACM

3726

2781

459

3.86 −49

L IFG (pars opercularis)

3.46 −61 −52

L MTG

3.31 −55 −40

3.15 −4 −34

L SMG

L paracentral g, s

ACM > CCM

ACM > AI

2889

1377

864

432

AI > ACM

4.63 −22

3.68 −40 −49

3.45

7 −67

L MFG

L AG

0 R oc-temp lingual g

3.07 −55 −37

L SMG

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2052

1674

1647

1350

918

837

CI > AI

864

810

270

AI > CI

1755

1242

891

−3.80 −22 −67 −12

L oc-temp fusiform g

−4.54 −4 −67

L precuneus

−3.92

52 −58

35 R AG

−3.50 −4 −40

L post-dorsal cingul

−4.50

−3.59

61 −10 −15 R MTG

61 −31

0 R MTG

4.06 −46 −76

L MTG

3.18 −52 −58

L MTG

3.05 −55 −37

L SMG

−4.11 −55 −16 −12

L MTG

−4.61

29 R MFG

−3.49 −52 −37

L STS

−3.51

32 R SFG

3.52

3.28

17 R MFG

61 −40

23 R SMG

3.63 −34

3.55 −34 −49

4.36 −34

3.67 −28

L MFG

L IPS

L MFG

L frontomarginal g, s

CCM > CI

1836

1674

1647

1107

945

837

CI > CCM

1782

1323

999

945

918

−3.98

−3.93

7 −49

8 R post ventral cingul

58 −4 −9 R MTG

−3.95 −55 −1 −9

L MTG

−3.42 −31 −82 −6

L MOG

−4.38

49 −6 R rectus g

−4.74 −55

L IFG (pars opercularis)

−4.67 −55 −16 −3

L STS

−3.85 −49

7 −18

L MTG

−3.88 −4

−3.53 −34 −52

L SFG

L STS

−3.85 −25 −88 −9

L occipital pole

−4.17 −4

L SFG

−3.43

31 −85

0 R MOG

7020

6264

2943

2862

2781

2673

2160

1647

CA × TI

1026

3.70 −43

L IFG (pars orbitalis)

The volume of the cluster (μl), peak z score, Talairach coordinates, y
the anatomical structures that the clusters overlap are shown. post = pos-
terior; oc-temp = occipito-temporal; cingul = cingulate. L = left hemi-
sphere; R = right hemisphere; g = gyrus; s = sulcus; ant = anterior.

Ellegaard, Ostergaard, Ostergaard, & Roepstorff, 2005).
Several authors have proposed that such anterior activa-
tions might reflect representations that are abstracted
away from perceptually based representations and sche-
matized over the course of learning, which are neverthe-
less shared between language and perception (ver, p.ej.,
Humphreys, Newling, Jennings, & Gennari, 2013).

Constructions that contained an abstract verb elicited
greater levels of activation within lateral temporal re-
gions, most notably the left middle and anterior STS
and STG, as well as in the posterior cingulate. These re-
sults corroborate earlier findings showing that portions of
the left STS and STG play a role in processing abstract
conceptos ( Wilson-Mendenhall, Simmons, Martín, &
Barsalou, 2013; Desai et al., 2009, 2011; Wang, Conder,
Blitzer, & Shinkareva, 2010; Binder, Westbury, McKiernan,
Possing, & Medler, 2005; Wise et al., 2000). En suma-
mary, the current results of verb concreteness effects are
consistent with a number of previous studies, showing ac-
tivation of higher level sensory–motor regions for action
verb processing. These findings provide evidence for the
embodied framework insofar as they confirm that the
meaning of sentences that refer to action is represented
in a cortical network including areas that play a role in per-
ception and action.

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Main Effect of Transitivity

Of central interest was the question of whether con-
structions themselves are associated with meaning, cuando
controlling for verb meaning. In the Caused Motion–
Intransitive contrast, the effects of verbs were eliminated,
as both constructions contained similar action and abstract
verbos. The results show that sentences with caused motion
frames elicited greater levels of activation in the left aIPL,
which overlapped with the activation found because of
action verbs (Cifra 3). As discussed above, this region is
associated with action planning and control. Aunque el
same general region of the left aIPL was activated by both
action verbs and caused motion constructions, some differ-
ences were also observed. Verbs activated a more ventral
aspect of SMG, extending into the parietal operculum.
Parietal operculum is associated with secondary somato-
sensory cortex. Caused Motion constructions, en el otro
mano, activated a somewhat more dorsal section of the
SMG, extending into the anterior IPS (aIPS). The aIPS
is thought to be the human analog of monkey area AIP
(Culham & Valyear, 2006). Both of these dorsal and ventral
regions are commonly coactivated in grasping and tool use
tareas, and it is somewhat difficult to distinguish their role in
relation to control of actions. Sin embargo, the dorsal region,
and especially AIP/aIPS, is associated more strongly with
hand–object interactions, visually guided grasping, y
on-line dynamic control of movements (Reichenbach,
Thielscher, Peer, Bülthoff, & Bresciani, 2014; Rice, Tunik,
& Grafton, 2006; Tunik, Frey, & Grafton, 2005; Culham
et al., 2003; see Tunik, Rice, hamilton, & Grafton, 2007,
para una revisión).

Cifra 6. Areas showing an interaction between Concreteness
(Concrete vs. Abstracto) and Transitivity (Transitive/Caused Motion vs.
Intransitive). Mean percent signal change relative to rest is shown for
the CCM, CI, ACM, and AI conditions, in a sphere of 5-mm radius
around the peak voxel.

The left dorsal AG was also activated for caused motion
constructions relative to intransitive constructions. El
entire AG has been shown to play a role in semantic pro-
cesando (Bonner, Peelle, Cocinar, & Grossman, 2013; Binder
& Desai, 2011; Binder, Desai, Tumbas, & Conant, 2009).
Asimismo, TMS and patient studies have substantiated
the idea that the AG is involved in controlled semantic
Procesando ( Whitney, Kirk, O'Sullivan, Lambon Ralph, &
Jefferies, 2011, 2012; Corbett, Jefferies, & Lambon Ralph,
2009; Jefferies & Lambon Ralph, 2006). More specifically,
the dorsal AG has shown to play a role in action planning
and execution, especially reach-to-grasp actions. In a bi-
focal TMS study, Koch et al. (2010) examined connectiv-
ity between dorsal AG and the primary motor cortex (M1)
as well as between dorsal SMG and M1. They found that,
during early (∼50 msec) preparation of reaching and
grasping movements, AG–M1 connectivity was sensitive
to both type of grasp (whole-hand grasp) y ubicación
of the object in space, whereas SMG–M1 connectivity
was sensitive only to the type of grasp (precision grasp).
Structurally, both dorsal SMG and dorsal AG were con-
nected to M1 and to ventral premotor cortex, a través de
different bundles within the superior longitudinal
fasciculus.

Además, caused motion constructions also activated
the posterior medial aspect of the parietal lobe. Esta re-
gion, dorsal to the parieto-occipital sulcus and near the
posterior edge of the hemisphere, is termed the “parietal
reach region” (PRR) and is found in both monkeys and
humanos ( Yttri, Wang, Liu, & Snyder, 2014; Chang, Papa-
dimitriou, & Snyder, 2009; Filimon, nelson, Huang, &
Sereno, 2009; Connolly, andersen, & Goodale, 2003).
PRR plays a role in planning visually guided reaching
movimientos, possibly by computing the initial response
by taking into account the difference between the initial
hand position and gaze location.

The left MFG and IFS were also activated by both
caused motion constructions and action verbs. A dorsal–
ventral pattern, similar to the one seen in the aIPL, era
also seen in the frontal lobe, whereby the caused mo-
tion constructions activated more dorsal MFG regions,
whereas activation because of action verbs extended
more ventrally to the IFS. These frontal regions have a
wide-ranging role in executive function (p.ej., Duncan &
Owen, 2000) and can be interpreted here as reflecting
executive aspects of action processing.

En resumen, the activation of areas responsible for ac-
tion planning, alcanzando, and grasping, a saber, aIPS/aIPL,
dorsal AG, and PRR, suggests that caused motion con-
structions, regardless of the verb used, are associated
with action semantics, and this conceptual content is
grounded in regions used in action planning and control.
This is consistent with the idea that caused motion con-
structions activate a caused motion schema, cual es
grounded through areas controlling reaching and grasp-
ing actions. We did not obtain an effect of transitivity
within the premotor, primary motor, or supplementary

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motor cortex. A potential explanation is that motor reso-
nance elicited by these constructions represents relatively
abstract action plans and does not contain detailed or
specific parameters related to selection and execution of
the actions. In a study of patients with stroke, Desai,
Herter, Riccardi, Rorden, and Fridriksson (2015) presentado
that comprehension accuracy of action-related words
was selectively predicted by detailed parameters (p.ej., ini-
tial direction error or number speed maxima) of reaching
actions performed by the patients. This suggests that such
details are part of simulations at least at a single-word
nivel. Constructions, by their very nature, are general, estafa-
tain slots that can be filled by a variety of words, y son
unlikely to be grounded in specific action parameters.
Bastante, they are grounded in more general, higher-order
action plans.

Sentences with an intransitive frame elicited activations
that are similar to those seen for abstract relative to con-
crete language, both at the word and sentence levels. El
left STS plays a role in processing abstract concepts
( Wilson-Mendenhall et al., 2013; Wang y cols., 2010; Desai
et al., 2009; Sabsevitz, Medler, Seidenberg, & Binder,
2005; Noppeney & Precio, 2004; Mellet, Tzourio, Denis,
& Mazoyer, 1998). The OFC has shown to play a role in
emotion and decision-making and has been argued to
play a role in processing the affective significance of con-
cepts (Rolls & Grabenhorst, 2008; Binder et al., 2005;
Kringelbach, 2005; Bechera, Damasio, & Damasio,
2000). The IFG was activated by intransitives, mostly in-
volving pars orbitalis. An examination of the CCM–CI and
ACM–AI contrasts reveals that this activation was mostly
driven by AIs. This was also the case for most of the ac-
tivation in the left STS/STG. Given that these regions are
commonly activated for abstract relative to concrete con-
cepts, this suggests that AIs were the most abstract of the
condiciones. Abstract words occur in more diverse seman-
tic contexts (Hoffman, Rogers, & Lambon Ralph, 2011),
and their meaning is more dependent on distributional
information and integration with context (Andrews,
Vigliocco, & Vinson, 2009). The left IFG activation can
be interpreted as contributing to deriving meaning
through integration with context. The current results sug-
gest that this applies to intransitive constructions as well,
and their meaning is more abstract and constructed
through integration.

Interaction between Verb and Construction Types

To investigate if the degree to which the main verb elicits
perceptual and motor simulations is modulated by syn-
tactic structure, we calculated an interaction between
the class of the main verb (concrete vs. abstract) y
transitivity (transitive/caused motion vs. intransitive).
Our results show an interaction between these two fac-
tors in the left anterior IFG, involving the pars orbitalis
(spreading slightly into pars triangularis). An examination

of the four conditions indicates that the syntactic frame
modulates activation within the pars orbitalis of the
IFG to a greater extent for sentences with an abstract
as compared with a concrete main verb. Although both
Abstract conditions activated this region relatively more
than the Concrete conditions, the difference between
AI and ACM was greater, giving rise to this interaction.

As the meaning of a concrete noun like “rose” is less
dependent on context, a concrete verb (“throw”) would
strongly guide a specific perceptual/motor simulation
that can be expected to show a weaker modulation by
contexto. The syntactic frame is therefore expected to
have a smaller influence on these verbs. A diferencia de, ab-
stract verbs (“consider”) rely more on the surrounding
frame for their interpretation and should be modu-
lated more. This is precisely what our results show: A
stronger effect of syntactic frame for sentences with an
abstract main verb as compared with a concrete main
verb, within pars orbitalis (BA 47), which is thought to
play a role in controlled semantic retrieval ( Wagner,
Koutstaal, Maril, Schacter, & Buckner, 2000; Gabrieli, Pol-
drack, & Desmond, 1998). In a large-scale meta-analysis
(Binder et al., 2009), BA 47 was found to be sensitive to
semantics, when overall difficulty is controlled (es decir., casos
where conditions with greater executive demands are
compared with relatively easier conditions are eliminat-
ed). Our results suggest that the syntactic frame can
guide the ease with which certain semantic representa-
tions are selected, especially when selection of semantic
information is not easily achieved on the basis of the
main verb alone. Como se ha mencionado más arriba, these findings
are consistent with the proposal by Andrews and col-
ligas (2009) suggesting that concrete and abstract rep-
resentations are learned from different types of statistical
datos. Concrete representations are based on a set of per-
ceived physical properties of an object and the experien-
tial information we acquired while interacting with them.
Abstract representations, por otro lado, rely more
strongly on information of how words are distributed
across different linguistic environments. The results sug-
gest that the left pars orbitalis is especially relevant to in-
tegrating verb and construction-level meaning.

The two-level theory of verb meaning suggests that
such an interface would be provided by the template level
of the verb. This theoretical framework argues for two
separate levels of verb meaning. The composite meaning
of a verb constitutes an association of the root and tem-
plate level of a particular verb (for an overview, see Levin
& Rappaport Hovav, 2005). The uniqueness of every verb
is constituted by the root level of meaning, which is not
visible to grammar and provides subtle distinctions be-
tween each verb in a given class. In contrast to the root
level of verb meaning, the template level is visible to
grammar and provides a schematic representation that
is common to all verbs in a given class (Kemmerer,
2012; Kemmerer & Gonzalez-Castillo, 2010). Several neu-
ropsychological studies have provided evidence that

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these two levels of verb meaning are segregated in the
mente. Por ejemplo, studies with brain-damaged partici-
pants have shown a double dissociation in the impair-
ment in the discrimination of features of verb meaning
that are relevant to grammar versus features that are irrel-
evant to grammar (Kemmerer, 2000, 2003; Kemmerer &
Wright, 2002). There is some evidence for the associa-
tion of the left IFG with the schematic motor component
of template-level verb meanings. Por ejemplo, Tettamanti
and colleagues (2005) compared prototypical caused mo-
tion constructions with and without motor components,
such as “I bite an apple” and “I appreciate sincerity” (en
italiano), and found activation in the left pars opercularis
for motor sentences. Sin embargo, both verbs and nouns dif-
fered in these sentences, and it is not clear whether the
differences were due only to verb concreteness. Sobre el
other hand, Desai et al. (2011) performed a similar com-
parison (using simple caused motion sentences such as
“The thief bashed the table” and “The jury criticized the
proposal”) and did not find activation in the left IFG. En
another study, Desai et al. (2009) compared prototypical
transitive sentences (“I throw the ball”) with abstract sen-
tenencias (“I consider the risk”) and again did not find activa-
tion in the left IFG. Además, to control for the effect of
noun manipulability, they compared action sentences with
sentences with visual verbs (“I see the ball”), and no IFG
activation was found in that comparison either. De este modo, el
evidence associating left IFG with simple or prototypical
transitive constructions is mixed. A potentially relevant fac-
tor is that the posterior region of the left IFG (pars oper-
cularis/BA 44) is also involved in overt or covert speech
production and perception. If the action sentences have
longer syllable or phoneme length than the control sen-
tenencias, activation of posterior IFG can be expected. Ambos
Desai et al. (2009) and Desai et al. (2011) ensured that ac-
tion sentences were not longer (in terms of words, letters,
phonemes, or syllables) than the control abstract or visual
oraciones. Tettamanti and colleagues (2005) do not report
syllable or phoneme length of their sentences, and it is not
clear if they were matched. Studies that compare pro-
nounceable stimuli (p.ej., words or sentences) with non-
pronounceable materials (p.ej., false fonts or hash marks)
and report activation of posterior IFG can therefore be
questioned on these grounds. In our study, we did not ob-
serve activation of the pars opercularis. Es posible que
this region was activated equally by all verbs because it is
always involved in processing the argument structure and
is not seen in any contrasts. An alternative possibility is
that template-level meaning is not necessarily attached
to specific verbs, but templates or constructions them-
selves have semantics over and above those of the verb
that is filled in. Several psycholinguistic studies have pro-
vided evidence that syntactic constructions carry meanings
(Kako, 2006; Kaschak & Glenberg, 2000; Pescador, 1994,
1996). Además, a recent study by Allen, Pereira,
Botvinick, and Goldberg (2012) showed different neural
activations for the dative (p.ej., “Sally gave the book to

Joe”) and the ditransitive construction (p.ej., “Sally gave
Joe a book”), which share the same content words. Estos
results provide evidence that different grammatical con-
structions are associated with distinct patterns of neural
activación.

Finalmente, we note that several areas in various compari-
hijos, such as the left STS and MFG, showed negative ac-
tivations, relative to the baseline of gaps between the
estímulos, for all conditions. Aquí, only the differences be-
tween conditions are relevant, and not the sign, for two
razones. Primero, the sentences were separated only by
7.4 segundo, which is not long enough for the hemodynamic
response to return to baseline or even to become weak.
A “resting” state never occurs during the experiment;
there is only modulation due to reading various sen-
tenencias. De este modo, activations are not against rest but against
a baseline where general processes involved in process-
ing sentences are still active. Segundo, resting is not an
“off” or “neutral” state with respect to semantic process-
ing but is instead a very active period of rich semantic
Procesando. It has been suggested that engagement in ef-
fortful tasks reliably suppresses such task-unrelated se-
mantic processing (Binder et al., 2009). Applying this
logic leads to the prediction that, if the AG plays a role
in semantic processing than it’s expected to be deacti-
vated during nonsemantic tasks, no deactivation is
expected during semantic tasks. The finding that deacti-
vation of the left IPL (particularly the AG), which is part of
the Default Mode Network, is modulated by the semantic
nature of the task, shows that resting state and semantic
networks greatly overlap (Seghier & Precio, 2012; Seghier,
Fagan, & Precio, 2010; but see Humphreys, Hoffman,
Visser, Binney, & Lambon Ralph, 2015).

Conclusión

Our results show that the transitive/caused motion and
intransitive constructions are associated with distinct
patterns of neural activation. The caused motion con-
structions, across both action and abstract verbs, elicited
activation in areas associated with reaching and grasp-
En g. The intransitive constructions were processed simi-
larly to abstract sentences, activating lateral temporal
areas. These results provide evidence that the transitive
construction, by virtue of being associated with a caused
motion schema, elicits activations within sensory–motor
regiones del cerebro. Eso es, the syntactic construction itself
may drive the mental simulation of an event involving
the movement of an object (along a designated path).
Pars orbitalis has an important role in construction and
verb-level meaning, showing an interaction between
verb and construction type. These findings provide evi-
dence for constructionist approaches that claim that
form–meaning correspondences do not exclusively exist
at the level of verbs or lexical items but also at the level
of constructions.

van Dam and Desai

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Acknowledgment

We thank David Kemmerer and an anonymous reviewer for their
helpful suggestions. All mistakes are ours. This research was
supported by NIH/NIDCD grant R01 DC010783 (R. h. D.).

Reprint requests should be sent to Rutvik H. Desai, 220 Dis-
covery Building, 915 Greene St., University of South Carolina,
Columbia, CAROLINA DEL SUR 29208, o por correo electrónico: rutvik@sc.edu.

Nota

1. On the basis of the data from an experiment that involved
physical application of painful heat at different levels of inten-
sity, Cortejar, Krishnan, and Wagner (2014) suggested using a
voxelwise threshold of p < .001 or lower. The recommendation of such a stringent threshold is appropriate given that sensory tasks typically lead to activations that are large in extent. On the other hand, studies of higher cognitive functions often involve a comparison of conditions that are closely matched in their sen- sory properties and differ in more subtle ways. In our experi- ence, such contrasts show activations that are lower in magnitude and are much more variable between participants. To balance false positives and false negatives in group maps, thresholds in the range of p < .025 to p < .005 are more ap- propriate in these cases. This is why a large number of cognitive experiments use voxelwise thresholds in the neighborhood of p < .01, and similar values are used as defaults in some pack- ages. Although Woo et al.’s (2014) recommendation remains valuable in many cases, our choice of threshold is based on these considerations. REFERENCES Agnew, Z. 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