Group Title: BMC Neuroscience
Title: The Representation of the verb's argument structure as disclosed by fMRI
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Title: The Representation of the verb's argument structure as disclosed by fMRI
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Creator: Assadollahi, Ramin
Meinzer, Marcus
Flaisch, Tobias
Obleser,J onas
Rockstroh, Brigitte
Publisher: BMC Neuroscience
Publication Date: 2009
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Abstract: BACKGROUND:In the composition of an event the verb's argument structure defines the number of participants and their relationships. Previous studies indicated distinct brain responses depending on how many obligatory arguments a verb takes. The present functional magnetic resonance imaging (fMRI) study served to verify the neural structures involved in the processing of German verbs with one (e.g. "snore") or three (e.g. "gives") argument structure. Within a silent reading design, verbs were presented either in isolation or with a minimal syntactic context ("snore" vs. "Peter snores").RESULTS:Reading of isolated one-argument verbs ("snore") produced stronger BOLD responses than three-argument verbs ("gives") in the inferior temporal fusiform gyrus (BA 37) of the left hemisphere, validating previous magnetoencephalographic findings. When presented in context one-argument verbs ("Peter snores") induced more pronounced activity in the inferior frontal gyrus (IFG) of the left hemisphere than three-argument verbs ("Peter gives").CONCLUSION:In line with previous studies our results corroborate the left temporal lobe as site of representation and the IFG as site of processing of verbs' argument structure.
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Research article

The representation of the verb's argument structure as disclosed by
fMRI
Ramin Assadollahi1,2, Marcus Meinzer2,3, Tobias Flaisch*2, Jonas Obleser4
and Brigitte Rockstroh2


Address: 1ExB Communication Systems GmbH, 80333 Munich, Germany, 2Department of Psychology, University of Konstanz, 78457 Konstanz,
Germany, 3University of Florida, College of Public Health Professions, Gainesville, Florida-32611, USA and 4Max Planck Institute for Human
Cognitive and Brain Sciences, 04103 Leipzig, Germany
Email: Ramin Assadollahi assadollahi@exb.de; Marcus Meinzer mmeinzer@ufl.edu; Tobias Flaisch* tobias.flaisch@uni-konstanz.de;
Jonas Obleser obleser@cbs.mpg.de; Brigitte Rockstroh brigitte.rockstroh@uni-konstanz.de
* Corresponding author


Published: 15 January 2009
BMC Neuroscience 2009, 10:3 doi: 10. 1186/1471-2202-10-3


Received: 25 September 2008
Accepted: 15 January 2009


This article is available from: http://www.biomedcentral.com/1471-2202/10/3
2009 Assadollahi et al; licensee BioMed Central Ltd.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.



Abstract
Background: In the composition of an event the verb's argument structure defines the number
of participants and their relationships. Previous studies indicated distinct brain responses depending
on how many obligatory arguments a verb takes. The present functional magnetic resonance
imaging (fMRI) study served to verify the neural structures involved in the processing of German
verbs with one (e.g. "snore") or three (e.g. "gives") argument structure. Within a silent reading
design, verbs were presented either in isolation or with a minimal syntactic context ("snore" vs.
"Peter snores").
Results: Reading of isolated one-argument verbs ("snore") produced stronger BOLD responses
than three-argument verbs ("gives") in the inferior temporal fusiform gyrus (BA 37) of the left
hemisphere, validating previous magnetoencephalographic findings. When presented in context
one-argument verbs ("Peter snores") induced more pronounced activity in the inferior frontal
gyrus (IFG) of the left hemisphere than three-argument verbs ("Peter gives").
Conclusion: In line with previous studies our results corroborate the left temporal lobe as site of
representation and the IFG as site of processing of verbs' argument structure.


Background
Most verbs describe events with one or more participants
[1]. The verb's argument structure defines the number and
relationships of participants needed for a complete event.
For instance, a sentence like "Peter gives Jim a book"
includes three participants with three thematic roles: the
agent (Peter), the recipient (Jim) and the theme (the
book; [2]). For a verb like "give" the entry in the mental
lexicon must comprise such information in addition to
phonetic and orthographic information. Neurolinguists


have always been interested whether and how this feature
of verbs is represented in the brain, the majority of studies
employing entire sentences, wh-questions (e.g. "Where
did he go?", "What did he give her?", "Why did he give her
the book?"), or sentences including syntactic or semantic
violations (e.g. [2-6]. Functional imaging studies dis-
closed the middle temporal gyrus (MTG) and the inferior
frontal gyrus (IFG, BA 45/47) of the left hemisphere [7-9]
to be involved in the processing of the verb's argument
structure. Additional activity in the left IFG (BA 44/45)


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was found, when grammatically complex sentences
requiring working memory resources [10,11] or when
argument hierarchies [3] were processed. Whenever words
were presented in the grammatically correct order in one,
and out of order in another condition, activation of the
left IFG and MTG was more pronounced to words in correct
sentences [12].

In a recent fMRI study, Thompson et al. [13] compared
nouns and verbs with increasingly more complex argu-
ment structure [one-argument (e.g. sleep) vs. two-argu-
ments (e.g., chase) vs. three arguments (put)] during a
lexical decision task. While the processing of all lexical
items activated a large bilateral network including the
occipito-temporal, superior-inferior parietal and superior
temporal areas, only verbs activated left inferior frontal
and middle temporal areas. The comparison of verbs
revealed more pronounced activity mainly in the left infe-
rior parietal cortex (IPC; angular and supramarginal
gyms) for one-argument relative to two-argument verbs.
Additional right hemispheric IPC involvement was found,
when one-argument verbs were compared to two- and
three argument verbs. Results were interpreted as reflect-
ing the integration of semantic and syntactic information
that are more pronounced for verbs that possess more
complex argument structure.

In a previous magnetoencephalographic (MEG) study
[14] the argument structure of verbs was systematically
varied between one and three, while subjects processed
the verbs in a lexical decision task. Verbs were presented
either in isolation or together with a name, thus, in a min-
imal syntactic context. Around 250 to 300 ms after stimu-
lus onset isolated one-argument verbs induced the most
pronounced activation in the left middle temporal gyms,
followed by two-argument verbs, while three argument
verbs provoke the weakest activation. Whenever the same
verbs preceded by a proper name, specifying the subject,
additional activation between 350 and 450 ms in the left
inferior frontal gyrus was larger and peaked earlier for
one-argument verbs, which require no further arguments
to form a complete sentence. This suggests that the acti-
vated areas vary depending on the linguistic context.

On this background the present study aimed at specifying
the cortical structures involved in the processing of verbs'
argument structures exploiting the spatial resolution
power of fMRI. Within a silent reading design the argu-
ment structure of verbs was varied between one and three,
and maximum activation was expected to one-argument
verbs in temporal areas of the left hemisphere. Again,
verbs were presented either in isolation or in the minimal
syntactic context of a name that filled the first position of
the argument structure. Compared to designs employing
entire sentences, this comparison should disclose,


whether the crucial information about the relationship of
participants and events was already retrieved with the verb
and its argument structure or whether a minimal context
was required. Thereby, the present study should add to the
specification of parsing and processing in the absence of
full sentences. If the context was required to activate syn-
tactic processing [3,6], this condition should activate areas
known to be involved in syntactic processing, that is, the
left inferior frontal gyrus [11]. However, the previous
MEG study [14] had suggested that already the minimal
context of a proper name filling the first argument was suf-
ficient to activate the IFG. Thus, we hypothesized that
even the minimal context of a name should automatically
start the integration of the name and the verb to a sen-
tence representation. Therefore, we expected brain
responses in the left IFG to distinguish the verb-categories
in this condition, too.

Results
Results were obtained from 20 subjects.

Basic contrasts (verbs versus fixation baseline)
The comparison of the four experimental conditions
[one-argument or three-argument verbs preceded by a
name (NI; N3); one-argument or three argument verbs
preceded by senseless letter strings (VI; V3)] with the fix-
ation baseline revealed similar activity in bilateral primary
and secondary occipital cortices and in the fusiform gyrus
(see Figure 1). The comparison of one-argument verbs
presented in isolation and in context (VI and N1) and of
three-argument verbs preceded by a name (N3) disclosed
additional activity in the middle temporal gyrus (MTG).
MTG activity in response to three-argument verbs pre-
sented in isolation was only significant for an uncorrected
cluster threshold (p. < 05). Significant activity in the infe-
rior frontal gyrus (BA 47) was restricted to one-argument
verbs preceded by a name, thus, creating a complete
phrase [Additional file 1 summarizes all contrasts].

Specific contrasts
When comparing isolated one-argument with isolated
three- argument verbs (VI > V3), more pronounced activ-
ity was found in the inferior frontal gyrus of the right hem-
isphere (BAs 44/9: peak activity x/y/z: 56/8/16; k = 44; Z
= 5.4). In the left hemisphere, isolated one-argument
verbs induced more activity than three-argument verbs the
inferior temporal fusiform gyrus (BA 37, -46/-54/-14, k =
154, Z = 4.1). No significant areas were found for the
inverse contrast (V3 > VI). When comparing one-argu-
ment verbs in context with three-argument verbs in con-
text (N1 > N3), activity was found only in one left
hemispheric cluster (k = 81) that comprised anterior supe-
rior temporal (BA 38, -48/15/-9, Z = 4.2) and inferior
frontal areas (BA 47, -53/19/-3, Z = 3.8). Again, no signif-
icant difference was found for the inverse contrast.


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Figure I
Shows the general activity pattern elicited by the four experimental conditions compared to the fixation base-
line (N 1/3 refers to one-I/three-place verbs preceded by a name; V I /3 refers to one-/three-place verbs preceded
by a senseless letter string). Only clusters surviving a threshold of p < .05 family wise error-corrected (FWE) and a cluster
extent of k>20 are reported; the voxel threshold was set to p < .001 (uncorrected). Right column = left side of the brain.




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The comparison of verbs with and without context (N1 >
VI; N3 > V3) did not disclose differences for one-argu-
ment verbs. However, three-argument verbs provoked
activation in middle and inferior temporal areas (BA 21, -
55/-3/-17, Z = 4.2) of the left hemisphere. Irrespective of
the number of arguments, verbs presented in context acti-
vated a large cluster that comprised superior/middle and
inferior temporal areas (BAs 21/38/20, -53/-5/-15, k =
231, Z = 4.7) and a cluster in the right middle temporal
gyrus (BA 21, 55/-1/-18, k = 86, Z = 3.8) relative to verbs
presented in isolation (N1/3 > Vi/3).

Discussion
Our previous MEG study [141 had suggested distinct pat-
terns of cortical activity to verbs depending on their argu-
ment structure, therefore, the present study served to
clarify the localization of involved cortical structures
exploiting the enhanced spatial resolution power of fMRI.
Confirming the previous results, verb-elicited activity var-
ied with the number of obligatory arguments in the mid-
dle temporal gyrus and the inferior frontal gyrus: one-
argument verbs "snores" led to stronger activity than
three-argument verbs ("gives") in the inferior temporal
fusiform gyrus (BA 37) of the left hemisphere. When pre-
sented in context of a proper name, one-argument verbs
("Peter snores.") induced more pronounced activity than
three-argument verbs in the inferior frontal gyrus (IFG) of
the left hemisphere. These results support (a) that syntac-
tic processing of the verb's argument structure is related to
verb itself and does not require a complete sentence and
(b) that the IFG is crucially involved in the processing of
this syntactic information.

The present fMRI study also confirmed our results of our
previous MEG-study in that less complex, one-argument
verb's elicited more pronounced activity than more com-
plex, three-argument verbs. This contrasts the intuitive
expectation, that the structurally more complex three
argument verbs would lead to stronger brain responses
compared to less complex one argument verbs. As an
explanation, it might be assumed that the brain response
may not reflect the structural complexity of the argument
structure but rather, as to what extent the verb describes a
complete event: Within the composition of words
(including verbs) into a sentence, the "completeness" of the
compositional representation of words may vary across
the parsing process, being accompanied by a sequence or
cascade of activation. Before the first word is presented,
sentence processing (or verb retrieval) is 0% complete,
and activation has not started. Upon word/verb presenta-
tion, the representation of a corresponding situation may
be activation, and retrieval of the verb describing the situ-
ation is of highest impact. While the recognition of simple
events is 50% complete (subject missing), the recognition


of complex events may be less complete (33% for two
place verbs, 25% for three place verbs; see [14]).

Stromswold et al., [11] found increased regional cerebral
blood flow (rCBF) in Broca's area (particularly in the pars
opercularis) when subjects judged the semantic plausibil-
ity of syntactically more relative to less complex sentences.
In line with this finding, name-verb-pairs with one-argu-
ment verbs in the present study were more likely to reflect
a complete event than three argument verbs. Posterior
temporal activation has been demonstrated in verb gener-
ation tasks [9,15,16]. Similarly, the argument structure of
isolated verbs was represented in the inferior temporal
fusiform gyrus (BA 37) in the present study. This is in
accordance to Bornkessel's view that this area (BAs 22/37/
39) has an enhanced sensitivity for morphological infor-
mation and the syntactic realization of the verb-based
argument hierarchy.

Relative to the processing of isolated verbs, presentation
of verbs in context of a proper name shifted the patter of
activation to Brodman's Area 47. This is in line with
higher activation in BA 47 for subject- than for object-ini-
tial sentences [4]. In the present study the name preceding
the verb might have been interpreted as the subject of the
beginning of a sentence followed by a grammatically cor-
rect inflected verb.

In their seminal studies, Petersen and colleagues [17,18]
demonstrated the association between nouns and their
related verbs: In a verb generation task, subjects articu-
lated appropriate verbs to nouns that were presented
either acoustically or visually. In a noun repetition task,
subjects repeated acoustically presented nouns or articu-
lated visually presented words. Relative to noun repeti-
tion, the verb generation task produced significantly more
activation in a left inferior prefrontal region located at or
near Brodmann area 47. This design can be compared to
the present study, in which a proper name (a noun) was
followed by a verb. One might speculate that the corre-
spondence of the verb and the subject translates into the
activity of the IFG. This again supports the significance of
BA 47 for the coordination of words reflecting the strength
of their (syntactic) connection.

Moreover, Kapur and colleagues [7] demonstrated that
nouns were better recognized in a memory task when they
were semantically processed, involving the activation of
Brodmann Area 47 (left inferior prefrontal gyrus). Kapur
et al. [ 19] conclude that "This finding suggests that the left
inferior prefrontal cortex is the anatomical region
involved in 'working with meaning', and that the activa-
tion does not reflect willed action, is not task-specific and
is not attributable to the requirements of a spoken
response."


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The present study provides strong evidence that argument
number dependent activation shifts from posterior, tem-
poral areas to anterior, frontal areas when context is
added. Rappaport and colleagues [20] assumed that a verb
activates the semantic class as an intermediate representa-
tion. Similarly, Buckner et al. [21] suggested that anterior
regions maintain or control access to higher-level seman-
tic information. We argue that this information may
resemble the dynamic representation of the sentence
which integrates the subject into the argument structure of
the verb leading to the representation of an event. Like-
wise, Wagner et al. [22] interpreted the left inferior pre-
frontal area as a semantic executive system that mediates
on-line retrieval of long-term conceptual knowledge.

Syntactic [10,11,23-27] and semantic processing has con-
sistently been found to activate left inferior frontal areas
[7,28-35]. Specifically, a region in the anterior and ventral
aspect of the inferior frontal gyrus (IFG, approximate
BA47/10) has been identified as contributing to semantic
processing, but also left middle temporal cortex [36,37]
which was also active in the present study.

Thus, it is plausible that the IFG projects back to the tem-
poral lobe to keep representations active. Such a structure
would allow for lexical items to interact when coming in
sequentially: The activation of typical fillers (subjects or
obligatory objects) is supposed to facilitate on-line lan-
guage processing. Reading is speeded up, when thematic
roles were saturated (e.g. the subject was provided) during
comprehension [38,39].

Conclusion
In summary, visually presented verbs activate their argu-
ment structure in areas assigned to the biological lexicon.
This information is used in the inferior frontal gyrus to
integrate proceeding nouns into an event representation
that is sequentially built up while new words come in.
Thus, static parts of the representation stay active in the
temporal lobe while dynamic parts may be processed in
the IFG.

Methods
Subjects
20 healthy native German speaking subjects (mean age
27.1 6.2 years, 11 females) were recruited for the study.
All subjects were right-handed as assessed with the Edin-
burgh inventory [40]. Prior to the experiment, subjects
were informed about measurement procedures and secu-
rity issues and gave written informed consent. After the
experiment, subjects received a bonus of 10 Euros. Ethical
approval was granted by the ethics committee of the Uni-
versity of Konstanz.


Experimental task and stimulus characteristics
A silent reading task was implemented during fMRI. Stim-
ulation comprised four blocked experimental conditions
[one-argument or three-argument verbs preceded by a
name (NI; N3); one-argument or three argument verbs
preceded by senseless letter strings that were matched to
the names for length (VI; V3)] and a baseline condition
(fixation cross).

Stimuli had been selected from a pilot study, in which 600
German verbs presented in third person, singular, present,
active form with different argument-structures were pre-
selected from the CELEX-database [411. Verbs were
selected to be as unambiguous as possible and having the
lowest number of different argument structures [42,431.
Ten student volunteers were asked to generate a sentence
to each verb. Non-obligatory adjuncts referring to time or
space were not considered. A verb was included into the
stimulus set, when more than 70% of the generated sen-
tences included the argument structure of the central
sense [44].

For each argument structure, 40 verbs were selected from
the pool obtained in the pilot study [141. Verbs were
selected only, if they formed a meaningful beginning of a
sentence with a name. One-argument verbs had a mean
length of 7.6 letters (SD = 1.42), three-argument verbs a
mean length of 7.6 letters (SD = 1.62). Both set were
matched for frequency (mean: 2.2 per million words,
[41]). Forty proper names were selected for the N-condi-
tion, while letter strings matched for length preceded the
verbs in the V-conditions, in order to ensure physical com-
parability of the conditions.

Each condition was repeated 8 times and consisted of 5
consecutive trials. Each verb, name or letter string was
shown only once to avoid repetition effects. The sequence
of blocks was fixed, while presentation of stimuli within
blocks was random. Two different sequences of blocks
were designed and counterbalanced across subjects. Stim-
uli were presented by a visor (VisuaStim, Resonance Tech-
nology, Inc.) in the middle of the screen. Subjects were
instructed to silently read the stimuli.

Scanning parameters
A 1.5 Tesla Philips Intera MR-System equipped with
Power Gradients Scanning was used. For functional scan-
ning, a T2*-weighted Fast-Field Echo, Echo-Planer-Imag-
ing (FFE-EPI) sequence utilizing a parallel scanning
technique (SENSE; [45]) was used. Images were acquired
in transversal orientation parallel to the AC-PC line. Each
dynamic volume consisted of 36 slices measured in inter-
leaved acquisition order with a thickness of 4.5 mm each.
In plane resolution was 2.9 x 2.9 x 3.5 mm with a squared
Field-of-View at a size of 230 mm (acquisition matrix 80


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x 80 voxels). A whole-head scan was assessed every 3 sec
(TR), the overall sequence comprising 200 continuously
acquired volumes. Eight dummy scans serving for Tl-
equilibration at the beginning of the experimental session
were discarded from data analysis.

Functional MRI post-processing was accomplished with the
Statistical Parametric Mapping (SPM2, Wellcome Depart-
ment of Cognitive Neurology, London, UK). Pre-process-
ing included correction for slice-time differences and
spatial alignment to the first volume of the image series to
adjust for head movements during the course of the exper-
iment. Subsequently, functional volumes were normal-
ized to MNI standard stereotactic space and smoothed
with a Gaussian Kernel of 8 x 8 x 9 mm full-width-at-half-
maximum (FWHM).

Pre-processed data were submitted to statistical analysis
implementing the General Linear Model (GLM). The cor-
responding design matrix comprised the 5 covariates-of-
interest representing the experimental conditions' onsets,
the duration of the different presentation epochs, and no-
interest-covariates (modelled response functions' time
and dispersion derivatives and six movement parameters
obtained during realignment). Before estimating the
modelled regressors, a high pass filter with a cut-off period
of 128 sec was applied to the data. Following estimation
of the overall model, planned contrasts-of-interest were
calculated for each subject, including the experimental
conditions (V1/V3/N1/N3) and the fixation baseline.
Moreover, specific contrasts comprised the conditions: VI
> V3 and N1 > V3 and the inverse contrasts, in order to
elucidate differential activity associated with the process-
ing of the different argument structures and the effect of
filling the argument structure. Further contrasts included
comparisons between isolated verbs and such that were
preceded by a proper name (N1 > VI and N3 > V3), verbs
with and without context (N1/3 > Vi/3).

For the group analysis a random effect model included
these contrasts of all subjects. Maximally activated voxels
within significant clusters are reported (cluster threshold
p < .05 family wise error-corrected, FWE, cluster extent
k>20; voxel threshold p < .001 uncorrected). Significant
voxels within clusters were anatomically localized with
the Talairach Demon software [46]. For graphical display
activated areas were projected onto a template of a stand-
ard MNI brain.

Authors' contributions
RA and BR proposed the general research question. RA,
MM, TF, and JO designed the experiment. MM and TF
accomplished data collection, analyzed the data and con-
tributed the methods and results sections. RA drafted the
initial version of manuscript, all authors contributed to


the discussion of the results. All authors read and
approved the final manuscript.

Additional material


Additional file 1
Basic contrasts (V1/V3/N1/N3 > Fixation). The data provided describes
details of the activity patterns for the -i... .. _n contrasts: VI > Fixation,
V3 > Fixation, NI > Fixation, N3 > Fixation
Click here for file
[http://www.biomedcentral.com/content/supplementary/1471-
2202-10-3-S1.doc]


Acknowledgements
This research was supported by the Deutsche Forschungsgemeinschaft
(SFB 471, ME 3161/2-1).

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