ARTICLE IN PRESS
Brain and Language xxx (2005) xxx–xxx
www.elsevier.com/locate/b&l
Large-scale neural network for sentence processing
Ayanna Cooke a, Murray Grossman a,¤, Christian DeVita a, Julio Gonzalez-Atavales a,
Peachie Moore a, Willis Chen a, James Gee b, John Detre a,b
a
b
Department of Neurology, University of Pennsylvania School of Medicine, Philadelphia, PA, USA
Department of Radiology, University of Pennsylvania School of Medicine, Philadelphia, PA, USA
Accepted 11 July 2005
Abstract
Our model of sentence comprehension includes at least grammatical processes important for structure-building, and executive
resources such as working memory that support these grammatical processes. We hypothesized that a core network of brain regions
supports grammatical processes, and that additional brain regions are activated depending on the working memory demands associated with processing a particular grammatical feature. We used functional magnetic resonance imaging (fMRI) to test this hypothesis by comparing cortical activation patterns during coherence judgments of sentences with three diVerent syntactic features. We
found activation of the ventral portion of left inferior frontal cortex during judgments of violations of each grammatical feature.
Increased recruitment of the dorsal portion of left inferior frontal cortex was seen during judgments of violations of speciWc grammatical features that appear to involve a more prominent working memory component. Left posterolateral temporal cortex and
anterior cingulate were also implicated in judging some of the grammatical features. Our observations are consistent with a largescale neural network for sentence processing that includes a core set of regions for detecting and repairing several diVerent kinds of
grammatical features, and additional regions that appear to participate depending on the working memory demands associated with
processing a particular grammatical feature.
 2005 Elsevier Inc. All rights reserved.
Keywords: fMRI; Grammar; Working memory; Frontal; Temporal
1. Introduction
Early studies of sentence comprehension hypothesized a cognitive architecture focusing on a grammatical
structure-building component of sentence processing
(Chomsky, 1981; Fodor, 1983). More recent work also
implicates a component involving executive resources.
The latter component includes elements that facilitate
sentence processing, such as selective attention to speciWc sentence features, and the temporary storage and
manipulation of phrasal information kept active in
working memory (WM) during the processing of long-
*
Corresponding author. Fax: +1 215 349 8464.
E-mail address: [email protected] (M. Grossman).
0093-934X/$ - see front matter  2005 Elsevier Inc. All rights reserved.
doi:10.1016/j.bandl.2005.07.072
distance syntactic dependencies in a sentence (Berwick &
Weinberg, 1984; Caplan & Waters, 1999; Gibson, 1998;
Just & Carpenter, 1992; Marcus, 1980; Zurif, Swinney,
Prather, WingWeld, & Brownell, 1995). In the present
study, we used fMRI to learn about the neuroanatomic
basis for the WM component of sentence processing
through judgments of diVerent types of grammatical features in a sentence.
Our model of sentence processing includes two major
components. Grammatical structure-building involves
the elements of sentence processing that specify grammatical relationships in meaningful, multi-word utterances. While it is beyond the scope of the present study
to determine the extent to which sentence-level processes
depend on information represented in single words or on
an independent set of rules specifying grammatical
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A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
movement, we assume that a modality-neutral component governs allowable rule-based movement and inXectional morphology sequences in an utterance, and that
these help interpret relationships between a sentence’s
meaningful units. Even though the constituents of a sentence may emerge over time in a linear and sequential
fashion, the relationships between words that best reXect
the meaning of the resultant sentence are often characterized in terms of an embedded or hierarchical structure
that does not necessarily correspond to the linear manner in which its words may occur. There are many
advantages to this Xexible relationship between surface
linearity and non-linear meaningful associations in the
structure-building system. For example, this arrangement supports phenomena such as communicating two
diVerent messages with the same words but a slight
change in word order. The cost associated with this Xexibility, however, is that the structure-building system can
be quite resource-demanding. The second major component of sentence processing thus involves executive
resources. While the details remain to be worked out,
several resources appear to contribute to sentence processing. The detection of subtle morphology inXections
or changes in word order may require selective attention,
for example, and since utterances are not necessarily
structured in a linear manner, WM may help maintain
crucial components of a sentence in an active state until
the correct grammatical relationships can be established
during the course of sentence processing. Our assumption is that not all resources are needed to support all of
the structure-building processes of all sentences, and that
any one sentence may require diVerent combinations of
resources to interpret its grammatical structure. The
present study focuses on WM.
One source of evidence for the claim that both structure-building and WM components contribute to
sentence processing comes from electrocortical eventrelated potential (ERP) studies describing diVerent time
courses for the activation of each component. An early
left anterior negativity (LAN) at about 300–500 ms following stimulus onset is associated with a structurebuilding component (Friederici, 1995; Hahne &
Friederici, 2002; Mecklinger, Schriefers, Steinhauer, &
Friederici, 1995), while resource-demanding components
such as WM are related to sustained LAN (Fiebach,
Schlesewsky, & Friederici, 2002) or a late positivity
(called the P600) at about 500–1200 ms (Friederici, 1998;
Hagoort, Brown, & Groothusen, 1993; Kaan, 2000;
Osterhout & Holcomb, 1992; Rosler, Putz, Friederici, &
Hahne, 1993). In a series of elegant studies, Friederici
and her co-workers have shown that the early LAN and
P600 are recruited variably, depending on the speciWc
grammatical features of a passive-voice sentence (Friederici, 1995, 2002). Morphosyntactic information such as
the suYx of a past-tense participle appears to have minimal WM demands and thus is associated only with the
early LAN, while other grammatical features such as the
transitivity of a passive verb seem to be more resourcedemanding and correspondingly elicit a more prominent
P600.
The neuroanatomic distribution of grammatical
structure-building and WM resources during sentence
comprehension has been the subject of some debate.
Many studies of aphasic patients following stroke have
associated grammatical comprehension diYculty with
insult to left inferior frontal cortex (IFC) (Alexander &
Benson, 1991; Tramo, Baynes, & Volpe, 1988; Zurif,
1996), although exceptions have been reported (Caplan,
Hildebrandt, & Makris, 1996; Dick, Bates, Wulfeck,
Aydelott Utman, & Dronkers, 2001; Grodzinsky, 2000).
The precise anatomical basis for grammatical comprehension has been diYcult to establish in stroke patients
because of limitations associated with mapping vascular
insult determined by the cerebrovascular system onto a
cortical function that does not correspond to a particular perfusion bed. Functional neuroimaging studies of
healthy adults over the past decade have begun to circumvent this limitation. This work has shown activation
of IFC and posterolateral temporal cortex (PLTC) in the
left hemisphere during a wide variety of tasks probing
grammatical aspects of sentences (Caplan, Alpert,
Waters, & Olivieri, 2000; Constable et al., 1998; Cooke
et al., 2002; Dapretto & Bookheimer, 1999; Friederici,
Meyer, & von Cramon, 2000a; Friederici, Opitz, & von
Cramon, 2000b; Kang, Constable, Gore, & Avrutin,
1999; Kuperberg et al., 2003; Luke, Liu, Wai, Wan, &
Tan, 2002; Moro et al., 2001; Ni et al., 2000; Roder,
Stock, Neville, Bien, & Rossler, 2002). For example, one
recent study demonstrated the speciWc role of left IFC
and PLTC in structure-building by comparing sentences
requiring syntactic movement with sentences not requiring syntactic movement (Ben-Shachar, Palti, & Grodzinsky, 2004). Although there are subtle diVerences in the
location of peak neuroanatomic activation in fMRI
studies of healthy subjects depending on factors such as
the visual or auditory modality of sentence presentation
(Caplan, Alpert, & Waters, 1999; Friederici et al., 2000a;
Michael, Keller, Carpenter, & Just, 2001), the use of
comprehension or expression (Indefrey, Hagoort, Herzog, Seitz, & Brown, 2001), and the age of the subjects
(Caplan, Waters, & Alpert, 2003; Grossman et al., 2002),
and although some work has implicated other brain
regions in grammatical processing such as left anterior
superior temporal cortex (Humphries, Willard, Buchsbaum, & Hickok, 2001), the overall pattern of results is
relatively consistent in emphasizing the contribution of
peri-Sylvian frontal and temporal regions of the left
hemisphere during sentence processing.
There is also reason to believe that these brain regions
may support WM during sentence processing. Studies of
patients with WM limitations thus implicate frontal and
temporal-parietal brain regions in supporting resource-
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related components of a large-scale neural network for
sentence processing (Grossman, 1999; Grossman &
White-Devine, 1998; Grossman, Rhee, & Antiquena,
2003; Kempler, Almor, Tyler, Andersen, & MacDonald,
1998; Martin & Romani, 1994; Waters & Caplan, 1997;
Waters, Rochon, & Caplan, 1998). Other patient work
has begun to examine whether a form of WM is dedicated to sentence processing, or whether WM of all sorts
depends on a common pool of available resources (Martin & Romani, 1994; Martin & Freedman, 2001). fMRI
studies also demonstrate activation of frontal and temporal-parietal areas following manipulations of WM
during sentence processing. Manipulations have
included varying the number of words retained in WM
while processing long-distance syntactic dependencies in
sentences (Carpenter, Just, Keller, Eddy, & Thulborn,
1999; Cooke et al., 2002; Fiebach, Schlesewsky, Lohmann, von Cramon, & Friederici, 2005; Keller, Just, Carpenter, & Thulborn, 1998; Keller, Carpenter, & Just,
2001), varying the numbers of propositions in a sentence
(Caplan, Alpert, & Waters, 1998), varying concurrent
articulation that occupies a subvocal rehearsal component of WM (Caplan et al., 2000), and varying the frequency of content words used in a sentence (Keller et al.,
2001). Some work emphasizes the role that left IFC
plays in supporting a form of WM that is dedicated to
language processing (Caplan et al., 2000; Cooke et al.,
2002; Fiebach et al., 2005; Grossman et al., 2002),
although a similar distribution of left IFC activation has
been reported in studies of WM for letters and other
non-linguistic verbal material (Chein, Fissell, Jacobs, &
Fiez, 2002; Smith, Jonides, Marshuetz, & Koeppe, 1998;
Smith & Jonides, 1999). There is also some evidence that
PLTC supports WM during sentence processing
(Caplan et al., 1998; Cooke et al., 2002; Just et al., 2001;
Keller et al., 2001; Michael et al., 2001). Yet other work
has associated additional brain regions with other executive resources. There is the apparent role of anterior cingulate in sustaining selective attention to detect
conXicting choices (Corbetta & Shulman, 2002), for
example, and together with dorsolateral prefrontal cortex, this system modulates planning and decision-making through selective inhibitory control by monitoring
errors and regulating response contingencies in a topdown manner (Badre & Wagner, 2004; Barch et al.,
2001; Botvinick, Braver, Barch, Carter, & Cohen, 2001;
Braver, Barch, Gray, Molfese, & Snyder, 2001; Carter
et al., 1998, 2000; Kerns, Cohen, Stenger, & Carter, 2004;
MacDonald, Cohen, Stenger, & Carter, 2000).
In the present study, we sought to test a dynamic
model of sentence processing. The model involves the
Xexible implementation of grammatical structure-building and resource-related components of a large-scale
neural network for sentence processing. There are several sources of evidence consistent with this view. Recent
fMRI studies of healthy young adults showed diVerent
3
patterns of recruitment depending on the syntactic or
thematic nature of the sentence probe (Kuperberg et al.,
2003; Newman, Just, Keller, Roth, & Carpenter, 2003).
Another study showed diVerent distributions of activation within left IFC depending on whether subjects were
judging the gender or grammatical subcategory properties of a word (Heim, Opitz, & Friederici, 2003). While it
is often diYcult to dissociate WM resources from structure-building in studies such as these since WM support
for sentence processing is so pervasive, it is nevertheless
clear that varying levels of WM are recruited depending
on the demands of a sentence (Gibson, 1998). Other
fMRI work thus has shown diVerent patterns of left IFC
recruitment depending on the strategic placement of
additional words in a sentence to stress WM during sentence processing (Cooke et al., 2002; Grossman et al.,
2002). Based on the ERP work of Friederici (1995, 2002),
the present study focused on the neuroanatomic substrate of three diVerent kinds of grammatical features in
a sentence, and the WM resources contributing to these.
We hypothesized that the neuroanatomic constituents of
the sentence processing network vary depending on the
speciWc nature of the sentence material used to challenge
comprehension. We used a grammatical acceptability
task, where subjects judge the coherence of sentences
that occasionally contain a grammatical violation. This
technique has been used productively to study the neural
basis for processing diVerent aspects of a sentence,
although it should be noted that detecting a grammatical
violation also may involve repair operations. We speciWcally hypothesized that a grammatical feature of a sentence may vary in its relative reliance on structurebuilding and WM resources, and this would be reXected
in the anatomic distribution of cortical activation during
coherence judgments of sentences containing violations
of various grammatical features.
2. Methods
2.1. Subjects
All subjects were young, healthy, right-handed, native
English speakers from the University of Pennsylvania
community who were paid for their participation and
completed an informed consent procedure approved by
the Institutional Review Board at the University of
Pennsylvania. A semi-structured interview conWrmed
that the subjects were medically healthy, and without
psychiatric disorders such as depression and anxiety.
None of the subjects were taking centrally acting medications. Twenty-one healthy young adults [11 female, 10
male; age range 17–27 years; mean (§SD) age D 23
(§3.0) years; mean (§SD) years of education D 16 (§2.5)
years] participated in Stimulus Development 1, and 17
young healthy adults [12 female, 5 male; age range 17–29
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years; mean (§SD) age D 22 (§3.5) years; mean (§SD)
education D 15 (§2.6) years] participated in Stimulus
Development 2. Fifteen young healthy adults [5 female,
10 male; age range 18–30 years; mean (§SD) age D 22.4
(§3.3) years; mean (§SD) education D 15.5 (§2.2) years]
participated in the fMRI study.
2.2. Materials
We used two techniques to develop our sentence
material, including an “on-line” study that focused on
the structure-building component of sentence processing, and an oV-line study that manipulated WM resource
demands during sentence processing.
2.2.1. Stimulus Development 1: On-line study of
grammatical features—Methods
We evaluated three types of grammatical features in
this study, based on ERP work demonstrating distinct
neuroanatomic and time course proWles of neural activity.
We focused on violations of a passive sentence in the present progressive of the sort “The manager is being confronted by the crazy company president.” One feature we
examined is the inXectional form of the past participle
with the dental suYx. For this type of stimulus, the morpheme -ed was omitted from some of the stimulus sentences that subjects heard (e.g., “The wig is being create
for the famous blonde actress”). This manipulation
appears to elicit only an early LAN, suggesting that it is
associated primarily with a rapid, structure-building component of sentence processing (Friederici, 1995; Hahne &
Friederici, 2002; Mecklinger et al., 1995). A second feature
we assessed is verb transitivity, that is, those verbs that
take a direct object. We violated the restriction associated
with transitivity by using a verb that does not take a direct
object in a present progressive construction (e.g., “The
toddler is being slept in the day care center”). This manipulation does not elicit a prominent early LAN, but instead
is strongly associated with a late positivity (Friederici,
1998). This late ERP signal appears to be related more
closely to eVort-demanding resources such as WM that
emerge over a slower time course during sentence processing (Friederici, 1998; Hagoort et al., 1993; Kaan, 2000;
Osterhout & Holcomb, 1992). A third type of feature we
assessed in this study focuses on the major grammatical
category of a word. A violation of this category feature
involved substituting a noun related in meaning to the
semantic content of the sentence for a verb (e.g., “The
dance is being rehearsal prior to the debut performance”).
This feature appears to be associated with equally prominent early LAN and late ERP responses (Rosler et al.,
1993). It is important to emphasize that these stimuli were
not intended to test a speciWc aspect of grammar, but
rather were selected because of suggestions in the literature that associate these features with diVerent levels of
resource demand during sentence processing.
We administered 192 randomly ordered sentences
involving manipulation of the “inXectional morphology,” “grammatical category,” or “transitivity” features
of a sentence (see Appendix A). All sentences were 13
words long, modeled after ERP studies by Friederici
(1995). We also administered 192 Wller items: half of
these contained no error, and the remainder contained a
diVerent kind of violation. A total of 384 sentences thus
was presented, with subjects exposed to 16 of each type
of feature violation of interest.
We presented these sentences using an on-line “word
detection” paradigm. The “word detection” technique
minimizes task-related resource demands since subjects
are not making explicit judgments of sentence grammaticality. A target word was presented and subjects were
asked to detect the word’s occurrence in the sentence
that followed (Marslen-Wilson & Tyler, 1980). Unbeknownst to subjects, half of each type of sentence contained a feature violation and half were control items
with the corresponding correct structure. By manipulating the placement of the target word relative to the violation and by monitoring the latency to respond to the
occurrence of the target word, it is possible to infer
whether subjects are sensitive to the grammatical feature. The correct and incorrect sentences thus were
equally divided across three conditions. One-third of the
sentences of each type of feature included a target word
immediately after the feature violation (“near”), and
one-third located the target word at a point seven words
after the feature violation in the sentence (“distant”).1 In
the “near” condition, the target word is strategically
placed immediately after the feature of interest so that its
occurrence overlaps with the envelope of mental activity
associated with the structure-building component of sentence processing. Processing of an adjacent grammatical
feature that is incorrect prolongs the latency to detect
the target word in comparison to word detection immediately following a correct grammatical feature. In the
“distant” condition, the target word is several words
downstream from the grammatical feature of interest.
Rapid, structure-building components of sentence processing should have been completed by the time that the
target word is encountered, so there should be little
diVerence in the latency to detect a word following a violation compared to a correct grammatical feature. If
resource-related demands slow the processing needed for
a particular type of stimulus, however, on-going processing may delay detection of the target word following a
violation of this kind in the “distant” condition. We varied the target word form class to ensure that target
words were not predictable based on the sentence context, and we conWrmed this with a cloze procedure in
1
The remaining third of the sentences had a target word located at
an intermediate point four words after the target stimuli. For the sake
of clarity, we did not consider these further.
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2.2.2. Stimulus Development 1: On-line study of
grammatical features—Results and discussion
Fig. 1 illustrates the results of response latencies for
target word detection. Fig. 1A shows the absolute
response latencies to target words embedded in correct
and incorrect sentences at a “near” or a “distant” location, averaged across the three grammatical features.
Latencies to respond following an anomalous item are
longer than following a coherent item in the “near” location [t (19) D 2.03, p < .05] but not the “distant” location
[t (19) D 0.61, ns].
2
Unpublished studies in our laboratory have shown no diVerence in
performance between a condition where the sentence is interrupted following a response compared to a condition where the full sentence is
heard.
3
We could not probe a sentence containing a violation because the
violation is likely to have rendered the sentence diYcult to interpret.
A
620
600
5
*
580
correct
violation
560
540
520
500
480
near
distant
Position of target word
B
50
40
*
30
DifferenceScore (msec)
another group of healthy young subjects. We ensured
that nouns in the phrase structure violation were only
nouns or were much more frequent (>7 times) as nouns
than as verbs, and that past participles occurred more
frequently (>7 times) as verbs than as nouns, according
to form-class sensitive frequency counts (Francis &
Kucera, 1982).
The sentences were digitized at 16 bits using SoundEdit v2.0 software involving the same female voice, and
sentences were presented on a Macintosh G3 laptop
computer. We used PsyScope (Cohen, MacWhinney,
Flatt, & Provost, 1993) to present the experiment. Subjects were informed that a word would be heard, followed (1000 ms later) by a beep, and then (1000 ms later)
a sentence was heard that contains this word. Subjects
pressed the space bar of the computer with their right
hand as soon as they detected the word in the sentence.
Detection of the target interrupted the sentence,2 and
initiated the onset of the next item after 1500 ms. Premature responses (responses prior to the occurrence of the
target word in the sentence—<.5%) were not recorded
by PsyScope, and a small number (<.5%) of extremely
long latency responses were eliminated. All remaining
latencies to detect a target word were analyzed after a 2
SD Wlter (based on each subject’s mean response latency)
eliminated outliers. A brief practice test was administered prior to the experiment, including both correct sentences and sentences with violations, to familiarize the
subjects with the task. Subjects were not informed of the
speciWc nature of the anomalies they were about to hear.
Throughout each block, the content of 6.7% of correct
sentences was probed to ensure that subjects were
attending to stimuli.3 Sentences were distributed over
four randomly ordered and counterbalanced blocks so
that a correct sentence was in one block and the version
of the same sentence with an error was in a diVerent
block.
Latency (msec)
A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
*
20
10
near
distant
0
-10
-20
-30
-40
morph
categ
trans
Violation Type
Fig. 1. On-line processing of grammatical features. (A) Overall sensitivity to grammatical violations illustrated with absolute latencies to
detect a target word following a correct grammatical structure or following a grammatical feature violation. The target word is placed
immediately following the feature (“near”) or seven words following
the feature (“distant”). (B) DiVerence scores (judgments of violations
minus judgments of correct stimuli) in “near” and “distant” positions
relative to each of the three grammatical features we studied. *SigniWcant diVerence between conditions at least at p < .05 level, according to
t tests. Morph D inXectional morphology feature; categ D grammatical
category feature; trans D transitivity feature.
Fig. 1B illustrates “diVerence scores,” or the discrepancy equal to latencies for target word detection in
incorrect sentences minus control sentences, for each
type of feature, associated with “near” and “distant” target words. The diVerence score is greater in the “near”
position than in the “distant” position for inXectional
morphology [t (19) D 2.47, p < .03] and grammatical category [t (19) D 2.69, p < .02] violations, emphasizing the
relatively prominent structure-building component associated with these stimuli, corresponding to the early
LAN (Friederici, 1995). However, a non-signiWcant bias
toward a greater diVerence score in the “distant” position for transitivity violations suggests that this feature
may involve a more prominent WM component. We
also did not Wnd a signiWcant diVerence between “near”
and “distant” diVerence scores for transitivity items,
consistent with the relatively WM-dependent nature of
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A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
this feature. These Wndings are consistent with a prominent structure-building component in inXectional morphology and grammatical category types of stimuli. In
the next study, we assessed the WM demands associated
with processing grammatical features in sentences more
directly.
2.2.3. Stimulus Development 2: WM demands during
sentence processing—Methods
The basic task presented the same sentences as above
(i.e., containing an inXectional morphology, grammatical
category, or transitivity feature—see Appendix A) to
subjects auditorily through headphones. Half of each
type of sentence was correct and half contained a violation, as above. The subjects made a grammaticality judgment: they were asked to state “no” into a microphone
as soon as an error was encountered. A voice-activated
relay triggered the latency measurement and then the
onset of the next stimulus following a warning 1500 ms
later. Latencies were calculated in the same manner as
above, following the elimination of 15.8% of the sentences that were judged incorrectly. Premature responses
were eliminated (responses that occurred at a point that
preceded the occurrence of the target word in the sentence—<.5%), and a small number of very long latencies
(<.5%) were eliminated. A 2 SD Wlter (based on each
subject’s mean response latency) was then applied to the
data. A practice test was administered to familiarize subjects with the task, and to adjust microphone gain.
In this context, we examined WM resources through
two approaches. First, we stressed WM by manipulating
the location of three additional words relative to the two
contributing elements of the inXectional morphology,
grammatical category, and transitivity features (Table 1).
Sentences were modiWed such that elements were separated by three adjectives or adverbs (“long” distance) in
half of the sentences to stress WM, and were adjacent
(“short” distance) in the other half. These sentence stimuli were distributed in a systematic manner (a control
version of a sentence was in a diVerent block from the
violation version of the same sentence) over four counterbalanced blocks of 96 sentences that were administered in a random order.
In the second WM-stressing manipulation, this auditory grammaticality judgment task was presented alone
and then during the simultaneous presentation of a
visual secondary task. Visually presented secondary
tasks consisted of blocked lexical decisions for simple
letter detection (“indicate whether or not the presented
word contains the letter ‘a”’), grammatical class categorization (“indicate whether or not the presented word is
a noun”), and semantic categorization (“indicate
whether or not the presented word is an animal”). Half
of the words in each series were targets meeting the speciWed criterion and half were foils. Unique words were
used as targets and foils in each secondary task. To
account for individual diVerences in WM capacity, the
words were presented serially at a rate such that subjects
were 60–80% correct in their responses when each task
was presented alone, as determined during a pre-experimental trial of each secondary task. This also served as
practice for performing the dual task. All lexical stimuli
were matched for frequency and familiarity across
blocks, and noun category membership required a word
to have a frequency rating seven times greater for a noun
than a verb if the word was both a noun and a verb. Subjects were instructed to press one of two computer keys
with the index Wnger of either hand to indicate their
response. Instructions were presented prior to each task
performed in isolation, and prior to each secondary task.
2.2.4. Stimulus Development 2: WM demands during
sentence processing—Results and discussion
The results of sentence grammaticality judgments
performed alone and during a WM-demanding secondary condition are summarized in Fig. 2. Panel A shows
that concurrent performance of a secondary task signiWcantly decreases accuracy judging grammatical category
[t (16) D 2.88, p < .02] and transitivity [t (16) D 4.40,
p < .001] types of violations relative to judging the
acceptability of sentences with these features during presentation without a secondary task. There was no diVerence between the three diVerent secondary tasks,
suggesting that the use of the “noun” target in the secondary task did not preferentially contribute to slowing
during the grammatical category judgments. By compar-
Table 1
Examples of stimulus materials and behavioral measuresa
Violation type
Sentence example
InXectional morphology
The test is being vaguely yet slowly explain/explained*
to excited new college students
The dance is being not too seriously rehearsal/rehearsed*
prior to the debut performance.
The woman is being brazenly and lewdly winked/harassed*
by the building workers outside.
Grammatical category
Transitivity
Accuracy mean
(§SD) % correct
Latency mean
(§SD) ms
92 (2.2)
989 (68)
95 (1.7)
914 (78)
79 (2.4)
1301 (86)
a
All sentences have the same overall structure and vary only in the speciWc nature of the feature violation. Sentences in Experiment 1 do not contain (italicized) Wller material placed between agreeing elements. Accuracy and latency data are from Experiment 3. Latency reported for correctly
answered items with a violation, timed from the end of the violating word, indicated by an asterisk.
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A 100
95
% correct
90
85
*
*
80
75
alone
dual task
70
65
60
55
50
morph
categ
trans
Violation judgment type
% correct
*
*
features involve diVering structure-building and WMdemanding components: inXectional morphology stimuli involving the past tense morpheme appear to involve
a relatively prominent structure-building component
with few WM demands; transitivity stimuli concerned
with the direct object of a passive verb apparently
involve relatively prominent WM demands compared to
its structure-building component; while grammatical
category stimuli concerned with a noun/verb substitution involve both structure-building and WM-demanding components.
2.3. Imaging materials and procedure
B 100
95
90
85
80
75
70
65
60
55
50
7
short
long
morph
categ
trans
Violation judgment type
Fig. 2. Grammatical sensitivity under working memory-demanding
conditions. (A) Relative decrement in grammaticality judgments under
the dual task condition compared to grammaticality judgments performed alone. (B) Relative decrement in grammaticality judgments
when agreeing elements are separated by Wller material (“long” condition) compared to judgments when agreeing elements are not separated by Wller material (“short” condition). *SigniWcant diVerence
between conditions at least at p < .05 level, according to t tests.
Morph D inXectional morphology; categ D grammatical category;
trans D transitivity.
ison, simultaneous performance of a secondary task does
not have a noticeable eVect on the subjects’ grammaticality judgments of inXectional morphology violations.
This is consistent with the claim that WM resources contribute to grammatical category and transitivity types of
items, and that processing an inXectional morphology
type of stimulus requires fewer WM resources. Fig. 2B
shows that grammaticality judgments of sentences are
more diYcult when Wller material is embedded between
the components contributing to grammatical category
[t (16) D 5.57, p < .001] and transitivity [t (16) D 4.70,
p < .001] features, compared to sentences with the Wller
material not interrupting the components of these grammatical features. In contrast, the inclusion of Wller material has less of an eVect on judgments of sentences with
inXectional morphology stimuli [t (16) D 1.96, ns].
The results from these two stimulus development
experiments, together with ERP work (Friederici, 1995,
1998; Hahne & Friederici, 2002; Mecklinger et al., 1995;
Rosler et al., 1993), suggest that the three grammatical
While in the bore of an MRI scanner, subjects made
acceptability judgments of written sentences that are
correct or anomalous, presented one word at a time at a
rate of 2 Hz. Sentences were identical to those described
above, except that they were presented in a word-byword written format. Because of time limitations, we
were able to study only stimuli with a “long” distance
between critical elements of a phrase, as illustrated in
Table 1 (see Appendix A). Event-related design allowed
for randomization of sentence presentation order for the
24 sentences in each of 5 balanced runs. This resulted in
a total of 80 sentences with an error (20 violations of
each type of sentence) and 40 correct sentences across
the entire experiment.4 The proportion of correct-toincorrect stimuli was 1:2 because of scanner time limitations. Subjects responded with a hand-held response
box, and were instructed to press the left button with the
left thumb as soon as possible if the sentence contained
an error. Subject response interrupted sentence presentation. Subjects pressed the right button with the right
thumb at a sentence-terminal prompt if no error was
encountered. A response was immediately followed by a
blank screen until the onset of the next sentence stimulus. We decided not to use passive reading in this experiment because of the importance of determining whether
subjects correctly judged sentence grammaticality. Sentence onsets were separated by 20 s to allow the hemodynamic response for the stimulus sentence to return to
baseline. The time from the subjects’ response to the
beginning of the next sentence ranged from 12–16 s,
depending on when the subject responded. Psyscope
software (Cohen et al., 1993) on a Macintosh G3 laptop
controlled projection of the written words of each sentence onto the central portion of a screen projected to
the magnet bore by a system of mirrors. Accuracy and
latency data were collected during scans by the software
program. Latency data were recorded from the onset of
the violation in a sentence. Incorrect judgment trials
4
A fourth type of violation was a Wller involving a stimulus that was
not of interest. These items were excluded from analyses.
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A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
were removed from subsequent imaging analysis so
accuracy was equated across conditions.
2.4. Imaging procedure
The experiment was carried out at 1.5 T on a GE
Signa MRI Scanner using a standard clinical quadrature
radiofrequency head coil and a prototype gradient system for ultrafast imaging. T1 weighted images were
acquired for anatomic localization, followed by BOLDsensitive gradient echoplanar images in 18 £ 5 mm transaxial slices with TR D 2 s, TE D 50 ms, 3.75 £ 3.75 £ 5 mm
voxels. A separate acquisition lasting 1–2 min was
needed for phase maps to correct for distortion in echoplanar images (Alsop, 1995). Initial data processing was
carried out with Interactive Data Language (Research
Systems) on a Sun Ultra 60 workstation. Raw image
data were reconstructed using a 2D FFT with a distortion correction to reduce artifact due to magnetic Weld
inhomogeneities.
Data were analyzed with SPM99, operating on a
MatLab platform, developed by the Wellcome Department of Cognitive Neurology (Frackowiak, Friston,
Frith, Dolan, & Mazziotta, 1997). A random eVects
procedure allowed us to Wlter out data corresponding
to incorrect responses. BrieXy, the images in each subject’s time series were registered to the initial image in
the series. The images were then aligned to a standard
coordinate system (Talairach & Tournaux, 1988). We
used an 8 mm Gaussian kernel for spatial smoothing,
and low-pass temporal Wltering was implemented to
control auto-correlation with a Wrst-order auto-regressive method. ERP studies (Friederici, 1995, 1998;
Hahne & Friederici, 2002; Mecklinger et al., 1995; Rosler et al., 1993) and fMRI studies (Kang et al., 1999; Ni
et al., 2000) call attention to early and late time courses
during which grammatical processing is active, so we
examined two hemodynamic response functions that
correspond to the 2-s window when the grammatical
structure occurs in the stimulus sentence and the 2-s
window immediately following the grammatical structure (referred to below as “early” and “late” time windows). Data thus were modeled with hemodynamic
response functions that peaked at 4 s (“early”) and 6 s
(“late”) after the occurrence of the violation in the sentence. The remainder of the sentence was modeled as a
variable of no interest.
We also performed a region of interest (ROI) analysis of activation in ventral and dorsal portions of left
IFC. Based on the anatomic work of Amunts et al.
(1999), we divided inferior frontal cortex into two
regions, a dorsal portion consisting of Brodmann area
44 and the dorsal-posterior half of Brodmann area 45,
and a ventral portion consisting of Brodmann area 47
and the ventral-anterior half of Brodmann area 45.
These regions were identiWed on transaxial slices of a
digitized version of the Talairach and Tournaux (1988)
template atlas, and then transferred in a user-independent manner to the normalized brain volumes of each
subject. We then identiWed the number of suprathreshold (p < .001 uncorrected for multiple comparisons)
voxels in the left dorsal IFC and left ventral IFC
regions during early and late time windows for processing each of the grammatical structures.
We examined the coordinated activation patterns
within the network of brain regions recruited during this
study using stepwise multiple linear regression analyses,
where the dependent variable was each brain region [left
ventral IFC (vIFC), left dorsal IFC (dIFC), left PLTC,
or left anterior cingulate], and the independent variables
were the remaining three brain regions within the same
temporal window or an earlier temporal window. The
direction of the connectivity was established by the independent and dependent status of the variables entered
into the regression analyses. For example, prediction of
left vIFC activation by other brain regions was determined by a regression analysis that modeled left vIFC as
the dependent variable; prediction of the contribution of
left vIFC activation to left PLTC activation was determined by a regression analysis where left vIFC was modeled as an independent variable. We decided not to
examine right hemisphere regions in this analysis
because of their inconsistent and unreliable activation in
the literature. For the purpose of this analysis, activation
in each region was based on the percent of increased
activation compared to baseline at each SPM-identiWed
peak (using the coordinates speciWed in Table 2) for each
structure during each temporal window (including the
2 s epoch prior to the early time window).5 Activation in
each region included the peak voxel and extended in X,
Y, and Z axes by one voxel to form an average of seven
adjacent voxels. These analyses were not implemented in
SPM, but used software developed in our laboratory by
Dr. Gonzalez.
3. Results
Table 1 summarizes the behavioral measures collected during fMRI scanning. Subjects judged transitivity sentences less accurately than inXectional
morphology sentences [t (14) D 4.77, p < .001] and grammatical category sentences [t (14) D 5.44, p < .001]. They
were also slower at judging transitivity sentences than
inXectional morphology sentences [t (14) D 5.10, p < .001]
and grammatical category sentences [t (14) D 8.32,
p < .001]. This suggests behaviorally that transitivity
5
For regions without a statistically signiWcant peak in Table 2, we
identiWed the peak activation within a speciWc region-of-interest as follows: vIFC, BA 47 and anterior BA 45; dIFC, BA 44 and posterior BA
45; PLTC, BA 21; cingulate, BA 24.
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9
Table 2
Coordinates of peak activation and associated Brodmann areas for each grammatical feature during early and late time windowsa
Time course
InXectional morphology
Early
Coordinates
X
Y
Z
¡60
¡60
4
8
¡52
16
24
8
48
Z score
Region (Brodmann areas)
4.50
5.00
4.80
Left inferior frontal cortex (47, 45, 44, 6)
Left posterolateral temporal cortex (21, 37)
Bilateral anterior cingulate (24, 32)
Late
Grammatical category
Early
Late
Transitivity
Early
Late
No activated regions
¡44
¡60
¡12
48
36
12
¡52
8
8
¡64
24
4
52
32
44
5.02
5.73
5.00
5.26
5.03
Left inferior frontal (47, 45, 44, 6)
Left posterolateral temporal (21, 37)
Bilateral anterior cingulate (24, 32)
Right inferior frontal (44, 6)
Right parietal (7)
¡48
¡44
12
¡52
0
0
4.03
4.60
Left inferior frontal (47, 45, 44, 6)
Left posterolateral temporal (21)
¡52
4
48
0
8
8
40
52
40
4.78
4.75
4.57
Left inferior frontal (47, 45, 44, 6)
Bilateral anterior cingulate (24, 32)
Right inferior frontal (44, 6)
¡48
8
12
4.42
Left inferior frontal (47, 45, 44, 6)
a
All regions signiWcant at least at p < .05 level following correction for multiple comparisons, except the left inferior frontal region during the
grammatical category—late condition which is signiWcant at p D .15 (corrected), and minimal extent >20 adjacent voxels.
Fig. 3. Activation associated with each grammatical feature in early and late time windows. (A) InXectional morphology stimuli: early window (there
was no signiWcant activation for the inXectional morphology stimuli during the late time window). (B) Grammatical category: early window. (C)
Grammatical category: late window. (D) Transitivity: early window. (E) Transitivity: late window.
sentences have relatively greater WM demands compared to inXectional morphology and grammatical category sentences.
Fig. 3 illustrates the fMRI activation patterns associated with judgments of each grammatical violation in
the early and late time windows. The anatomic distribu-
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A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
tion of peak activations and the extents of activation
corresponding to these images are summarized in Table 2.
Fig. 3A shows that judgments of inXectional morphology sentences elicited left IFC and left PLTC activation
during the early time window, but no suprathreshold
activity during the late window. Activation restricted to
the early time window is consistent with the prominent
structure-building component that occurs relatively
early in the time course of processing this grammatical
feature, and with the central contribution of left IFC and
PLTC areas during judgments of grammatical features
involving a prominent structure-building component.
Fig. 3B and Table 2 show that judgments of grammatical
category sentences elicited early activity in left IFC and
left PLTC as well as right IFC and right parietal cortex.
Fig. 3C shows that some activation associated with
grammatical category judgments persisted into the late
time window, unlike activation associated with judgments of inXectional morphology sentences. Late
recruitment for grammatical category stimuli was
restricted to left IFC and left PLTC. Fig. 3D and Table 2
show that judgments of transitivity sentences elicited
early activation of bilateral IFC. Like grammatical category judgments but unlike inXectional morphology judgments, Fig. 3E shows that transitivity judgments
sustained recruitment into the late time window. Late
recruitment for the transitivity stimulus involved left
IFC. These Wndings suggest partially overlapping and
partially distinct anatomic patterns of activation during
grammaticality judgments that diVer in the relative
prominence of their structure-building and WMdemanding components.
We investigated diVerences in the anatomic distributions of activation associated with judgments of grammatical category and transitivity compared to
inXectional morphology stimuli to learn about the neural basis for WM resources during sentence processing.
We report contrasts only in the early time window
because we did not Wnd signiWcant activation associated
with all three grammatical stimuli during the late time
window. These Wndings are illustrated in Fig. 4, and the
Fig. 4. Contrasts demonstrating relatively greater recruitment for
grammatical category and transitivity than inXectional morphology
during early time window. (A) Greater activation during grammatical
category judgments compared to inXectional morphology judgments.
(B) Greater activation during transitivity judgments relative to inXectional morphology judgments.
Table 3
Coordinates of signiWcant activations that are greater for judgments of
grammatical category and transitivity than inXectional morphology
featuresa
Coordinates
Z score
Region (Brodmann areas)
X
Y
Z
Grammatical category > inXectional morphology
¡56 ¡48
4
3.71
Left posterolateral temporal cortex
(21)
Transitivity > inXectional morphology
¡48 8
24
3.02
Left dorsal inferior frontal (44, 6)
a
All contrasts signiWcant at the p < .001 level uncorrected for multiple comparisons.
loci of peak activation are summarized in Table 3.
Fig. 4A illustrates the direct subtraction of the inXectional morphology condition associated with a prominent
structure-building
component
from
the
grammatical category condition that also has a WMrelated component. Judgments of grammatical category
acceptability appeared to be associated with greater left
PLTC activation than inXectional morphology stimuli.
Fig. 4B shows the direct subtraction of the inXectional
morphology condition from the transitivity condition
that has a prominent WM-demanding component. Judgments of transitivity acceptability were associated with
greater left IFC activation compared to judgments of
inXectional morphology stimuli. Left IFC and left PLTC
thus appear to be associated with a relatively prominent
WM component of some grammatical features, and thus
may support in part a WM-related component of grammatical processing in sentences.
We observed activation of both vIFC and dIFC during grammaticality judgments of these features (Figs.
3A, B, and D), but visual inspection of the contrast of
transitivity and inXectional morphology features
(Fig. 4B) suggested a relatively dorsal locus of activation
within left IFC. This is important because of anatomical
work suggesting diVerent cytoarchitectonic and connectivity patterns in these regions (Amunts et al., 1999). In
the analyses described below, we examined the role of
left dIFC in WM-dependent aspects of sentence processing, and left vIFC in the structure-building component
of sentence processing.
Fig. 5 illustrates the percent of signiWcantly activated
voxels in the ventral and dorsal IFC ROIs. Repeated-measures ANOVAs in each temporal window, with a grammatical feature (3) £ region-of-interest (2) design, showed
signiWcant interaction eVects in both early [F(2,28) D 4.95;
p < .02] and late [F(2,28) D 3.42; p < .05] time windows.
Transitivity judgments were associated with greater activation in left dIFC than left vIFC in the early time window
[t(14) D 3.89, p < .005], consistent with the prominent WMdemanding component of this feature. Moreover, left
dIFC was recruited more for the transitivity judgments
than the inXectional morphology judgments during both
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early [t(14) D 3.61, p < .003] and late [t(14) D 2.02, p < .06]
time windows. InXectional morphology judgments were
associated with the least dIFC activation. Grammatical
category judgments recruited left dIFC at a level that falls
between inXectional morphology and transitivity features.
Fig. 5 also shows that left vIFC activity did not signiWcantly diVer across the three types of stimuli, although activation tended to be greater in the early time window. The
relatively invariant left vIFC activation in the early time
window is consistent with the constant contribution of a
grammatical component to all three grammatical features.
However, these Wndings suggest that transitivity sentences,
and to a lesser extent the grammatical category sentences,
diVer from inXectional morphology sentences in that the
former activate the dorsal portion of left IFC. This may be
due to the WM demands associated these relative to the
inXectional morphology stimuli.
Neuroanatomic studies have demonstrated reciprocal
projections between vIFC, dIFC, PLTC, and anterior
cingulate regions in the left hemisphere that are likely to
mediate activation within a sentence processing network
(Pandya & Kupers, 1969). We examined the coordinated
activation patterns within this network using stepwise
multiple linear regression analyses. The dependent variable in this series of analyses was each brain region (left
vIFC, left dIFC, left PLTC, or left anterior cingulate),
and the independent variables were the remaining three
brain regions within the same temporal window or an
earlier temporal window. Fig. 6 shows the results of
these regression analyses. We display only the results of
regression models that are signiWcant at a p value that
exceeds the .01 level. Arrows project from independent
regions to dependent regions according to the results of
the regression analyses. The arrows thus illustrate how
neural activity in one region predicts activity in another
region of the network.
30
% activated voxels/ROI
25
20
11
Fig. 6A summarizes the functional network associated with inXectional morphology judgments. Regressions for the inXectional morphology judgments were
signiWcant only for the early time window. In this early
time window, we found reciprocal functional projections
between core language components in left vIFC and left
PLTC. Left dIFC did not contribute to the functional
neuroanatomy of inXectional morphology judgments.
We also observed a projection within the early time window from anterior cingulate to left PLTC. Anterior cingulate may help attend to the unstressed grammatical
morpheme. These observations suggest a crucial role for
left vIFC and left PLTC in the structure-building component of inXectional morphology judgments.
Fig. 6C summarizes the functional network associated with judgments of the transitivity feature. We found
a functional neuroanatomic projection from left dIFC to
left PLTC that spans early and late time frames. This
suggests that the WM-related processes supported by
left dIFC interact with the peri-Sylvian sentence processing network during transitivity judgments. We also
observed a functional projection from left vIFC to the
anterior cingulate region. Left dIFC may keep elements
of the grammatical structure active in WM during the
course of sentence processing, while the anterior cingulate may help focus attention on speciWc components of
the sentence relevant to the structure.
Regression analyses associated with judgments of the
grammatical category feature are summarized in Fig. 6B.
We found functional projections from left vIFC to left
PLTC and to left dIFC. The left vIFC projection to left
PLTC was not restricted to the early time window, as it
was for the inXectional morphology judgments, but was
evident across early and late time windows. Similarly, the
left vIFC projection to left dIFC spanned two time windows. Anterior cingulate also projected to left PLTC
across two time windows. Resembling ERP observations,
these observations suggest that left PLTC may play several roles during sentence comprehension, depending on
the time course of activation: an early role in grammatical
structure-building, as in the inXectional morphology sentences, and a later role in processing grammatical features
that also involve a WM-related component.
15
4. Discussion
10
5
early
0
morph
categ
VENTRAL
trans
late
morph
categ
DORSAL
trans
Fig. 5. Percent of voxels signiWcantly activated in regions-of-interest
for ventral and dorsal portions of left inferior frontal cortex during
grammatical
judgments.
Morph D inXectional
morphology;
categ D grammatical category; trans D transitivity.
Converging lines of evidence suggest that structurebuilding and WM-related processing components both
contribute to sentence comprehension (Berwick & Weinberg, 1984; Caplan et al., 2000; Caplan & Waters, 1999;
Cooke et al., 2002, 2003; Fiebach et al., 2005; Friederici,
1995, 2002; Grossman et al., 2002; Just & Carpenter,
1992; Just, Carpenter, Keller, Eddy, & Thulborn, 1996;
Marcus, 1980). In this study, we assessed the neuroanatomic basis for these two components of sentence pro-
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A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
Fig. 6. Schematic representation of regression analyses showing functional connectivity among brain regions activated during grammatical judgments. (A) InXectional morphology; (B) grammatical category; (C) transitivity. Arrows project from independent to dependent variables according
to regression analyses computed for each region at early and late time windows. Arrow thicknesses reXect signiWcant r values derived from correlation analyses, as shown in arrow heads. White arrows represent activity within the “early” window (4 s post-anomaly), black arrows represent activity originating before the early window predicting activity in the late window, and the transparent arrow represents activity before the early window
predicting activity in the early window. V D vIFC, D D dIFC, T D PLTC, and C D anterior cingulate (medial structure).
cessing by examining the distribution of cortical
activation related to three diVerent grammatical features. The results associated left vIFC and PLTC activation with a grammatical feature that appears to have a
prominent structure-building component with modest
WM demands—inXectional morphology; in addition,
left dIFC, right IFC, and parietal cortex were associated
with grammatical features that have a more prominent
WM-related component—transitivity and grammatical
category. These Wndings are consistent with the hypothesis that WM-related components of a large-scale neural
network are systematically recruited to support the processing of speciWc grammatical features during sentence
comprehension.
Grammaticality judgment tasks have been used to
examine the neural basis for sentence processing in several previous functional neuroimaging studies (Kang
et al., 1999; Kuperberg et al., 2000, 2003; Moro et al.,
2001; Newman, Pancheva, Ozawa, Neville, & Ullman,
2001; Newman et al., 2003; Ni et al., 2000). Although this
kind of challenge involves a meta-cognitive judgment,
the activations seen in studies involving grammaticality
judgments largely parallel the results of work involving
passive sentence listening with an oddball eVect (Ni
et al., 2000) and direct probes of sentence comprehension (Caplan et al., 2000; Cooke et al., 2002; Indefrey
et al., 2001). The same neuroanatomic substrate involved
in processing grammatically coherent sentences thus
may be recruited to detect and repair grammatical violations in these sentences, although direct comparisons of
these diVerent approaches remain to be performed to
rule out the contribution of repair mechanisms mediated
by other lexical, semantic, or pragmatic processes.
Regardless of the speciWc nature of the technique, these
studies underline the importance of left IFC during the
structure-building component of sentence processing.
However, close inspection reveals some inconsistencies
in the speciWc neuroanatomic distribution of activation
within left IFC that contributes to grammatical processing. Ni et al. (2000) recruited BA 47, 45, and 44 for grammaticality judgments of short sentences and for an
oddball eVect involving syntactically anomalous brief
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sentences. Just et al. (1996) and Keller et al. (2001) also
showed activation in regions of interest that appear to
correspond to BA 47, 45, and 44 during probes for comprehension of sentences containing an object-relative
center-embedded clause. Kang et al. (1999) found
recruitment only of BA 45 and 44 for grammaticality
judgments of two-word phrases. Stromswold, Caplan,
Alpert, and Rauch (1996) and Caplan et al. (1998) also
showed activation in BA 45 and 44 during anomaly
judgments of object-relative sentences with a centerembedded clause compared to subject-relative sentences
with a clause in a sentence-terminal location.
Others have associated left IFC activation during sentence processing with WM. However, the precise nature
of WM associated with IFC activation is unclear. A
large body of work has demonstrated dIFC activation in
BA 6/44 during WM tasks involving letters and other
verbal non-linguistic materials, suggesting that this
region supports a non-speciWc form of WM for verbal
materials (Awh et al., 1996; Cohen et al., 1997; Paulesu,
Frith, & Frackowiak, 1993; Schumacher et al., 1996).
One such study found deactivation of BA 47 (Jonides
et al., 1997), emphasizing the dissociation of dIFC from
vIFC during a verbal WM challenge that did not involve
sentence structure-building. By comparison, a recent
study associated dIFC activation with syntactic working
memory (Fiebach et al., 2005). This work assessed German wh-questions with an object-extracted or a subjectextracted clause, where additional material had been
strategically positioned between syntactically dependent
features in some of these sentences. The investigators
showed left IFC activation in BA 44/45 during directly
comparisons of object-extracted sentences with a long
distance between syntactically dependent features, and
structurally similar sentences with a short distance
between syntactically dependent features.
IFC may not be anatomically homogeneous, but
instead there appear to be diVerent histologic features
and connectivity patterns in dorsal and ventral portions
of IFC (Amunts et al., 1999). This raises the possibility
of a functional dissociation between a relatively ventral
locus of activation within left IFC compared to a more
dorsal locus, and that both structure-building and WM
functions can be accommodated by IFC. In sentences
constructed to vary the grammatical structure of a sentence independently of its WM demands, young adults
and healthy seniors recruited BA 47, 45, and 44 during
comprehension of sentences with an object-relative center-embedded clause that have a long linkage between
the head noun and the trace in the subordinate clause
(Cooke et al., 2002; Grossman et al., 2002). Direct comparison of healthy seniors with age-related WM limitations yet matched for sentence comprehension accuracy
with young adults revealed signiWcantly more prominent
activation of the dorsal portion of left IFC in seniors.
This implicates left dIFC in the support of WM during
13
sentence comprehension. Since left vIFC activation was
otherwise equivalent across these age-deWned groups
matched in their sentence comprehension accuracy, we
concluded that left vIFC contributes to the structurebuilding component of sentence processing that does not
appear to change with age. Additional support for this
approach came from a study of patients with frontotemporal dementia. Patients with poor comprehension of
grammatically demanding sentences due to Progressive
Non-Xuent Aphasia had diYculty activating left vIFC,
while non-aphasic frontotemporal dementia patients
with executive and working memory limitations had
diYculty activating left dIFC during sentence processing
(Cooke et al., 2003).
Other studies associate diVerent neuroanatomic mechanisms with WM during sentence processing. Some work
described up-regulation of the entire left peri-Sylvian sentence processing network during WM challenges, at times
including IFC and PLTC homologues in the right hemisphere (Just et al., 2001; Keller et al., 2001; Michael et al.,
2001). Yet other studies have implicated left PLTC
(Caplan et al., 1998; Grossman et al., 2002) or right
PLTC (Cooke et al., 2002; Grossman et al., 2002). Fiebach et al. (2005) also showed bilateral PLTC activation
during their critical contrast assessing WM. Finally, it is
important to note that IFC has been associated with
other aspects of language, including phonologic judgments (Poldrack et al., 1999; Zatorre, Meyer, Gjedde, &
Evans, 1996) and selection and retrieval from semantic
memory (Thompson-Schill, D’Esposito, Aguirre, &
Farah, 1997; Wagner, Pare-Blagoev, Clark, & Poldrack,
2001). Other frontal brain regions have been reported to
play a role in a variety of executive resources as well.
Anterior cingulate has been associated with selective
attention (Corbetta & friederici, 2002) and with regulating responses in a top-down manner during decisionmaking (Badre & Wagner, 2004; Barch et al., 2001; Botvinick et al., 2001; Braver et al., 2001; Carter et al., 1998,
2000; Kerns et al., 2004; MacDonald et al., 2000), for
example, and it may be necessary to recruit anterior cingulate to support these executive resources during the
course of processing certain sentence material. In this
context, we consider each of the feature judgments below.
4.1. InXectional morphology
In the present study, we sought to examine the neural
basis of sentence processing with fMRI by comparing
several grammatical features. The behavioral studies used
to develop our sentence materials are consistent with
ERP work showing that these grammatical features
implicate several diVerent processes (Friederici, 1995,
1998; Hahne & Friederici 2002; Mecklinger et al., 1995;
Rosler et al., 1993). Past-tense morphology appears to
depend on a prominent structure-building component,
for example, although this feature apparently involves a
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A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
modest WM component. We observed signiWcant left
IFC activation during grammaticality judgments of pasttense morphology, suggesting that left IFC is related to
processing the structure-building component of this
grammatical feature. Moreover, this activation tended to
occur in a temporal window that is relatively early in the
time course of sentence processing. While this is consistent with ERP studies of structure-building that demonstrate an early LAN during grammaticality judgments, it
is important to point out that fMRI has poor temporal
resolution, and inferences based on the time course of
fMRI activation must be made cautiously. This early
activation is not due to the ability to detect a violation
earlier during the course of a sentence containing an
inXectional morphology feature compared to the other
features, as shown in Table 1. ROI and regression analyses emphasized the prominent role of the ventral portion
of left IFC in inXectional morphology judgments, and the
early time window when this occurred. Although SPM
contrasts did not reveal statistically signiWcant activation
in the late time window associated with the inXectional
morphology feature, we do not intend to imply that there
is no left IFC activity during the late time window associated with this type of stimulus since the ROI analyses
showed some activated voxels in IFC in the late time window. The ROI analysis also showed that inXectional morphology judgments are associated with the least
prominent activation of the dorsal portion of left IFC.
Taken together, the behavioral and imaging Wndings
appear to associate the process of grammatical structurebuilding closely with left vIFC.
There has been considerable debate about the cognitive basis for past tense formation in regular and irregular verbs (McClelland & Patterson, 2002; Pinker &
Ullman, 2002). The present study cannot address this
issue directly since we investigated only past tense formation for regular verbs. Nevertheless, noun–verb structures with modest resource demands such as number
agreement (Newman et al., 2003) and gender agreement
(Heim et al., 2003) also have been shown to activate left
IFC. Other studies have pointed out the contribution of
left IFC to other aspects of language such as phonologic
processing (Poldrack et al., 1999; Pugh et al., 1996;
Zatorre et al., 1996). We cannot rule out that subjects
were reading the sentences to themselves subvocally as
they were presented, although this would not explain the
selective contribution of left vIFC during the processing
of sentences containing inXectional morphology stimuli
seen in the ROI and regression analyses. With these
caveats in mind, it appears that left vIFC contributes to
inXectional morphology judgments that depend more on
rapid structure-building than on WM resources.
It is also important to point out that other left hemisphere regions such as left PLTC were activated during
judgments of inXectional morphology. Sentence processing may be disturbed following left PLTC disease in
aphasic stroke patients (Berndt, Mitchum, & Haendiges,
1996; Caplan et al., 1996, Caplan, Waters, & Hildebrandt, 1997; Grodzinsky, 2000). However, the precise
role of left PLTC activation during sentence processing
is unclear. While left PLTC activation during sentence
processing has been related to the structure-building
process (Caplan et al., 2001; Friederici et al., 2000b; Just
et al., 1996; Stowe et al., 1998), studies comparing syntactic and thematic role features during sentence processing have associated left PLTC with the latter (Kang
et al., 1999; Kuperberg et al., 2000, 2003; Newman et al.,
2003; Ni et al., 2000). This is consistent with the frequent
activation of this area during studies of single word
meaning that may also contribute to the appreciation of
thematic role (Joseph, 2001; Price, 1998). Others have
related left PLTC activation in part to WM during sentence processing (Caplan et al., 1998; Just et al., 2001;
Keller et al., 2001; Michael et al., 2001). This account
seems less likely, given the evidence we acquired during
stimulus development that inXectional morphology
judgments involve minimal WM demands. Regardless of
the speciWc role that left PLTC plays in sentence processing, regression analyses in the present study showed an
intimate reciprocal link between left ventral IFC and left
PLTC early during the course of processing the inXectional morphology feature.
Regression analyses suggested that activation in cingulate cortex projects into this left peri-Sylvian system.
Previous fMRI studies of sentence processing also have
shown cingulate activation (Caplan et al., 1998, 1999,
2000). Disease involving cingulate cortex has been associated with disturbed sentence processing in brain-damaged patients (Alexander & Schmitt, 1980; Freedman,
Alexander, & Naeser, 1984), providing converging evidence for the contribution of this area to a large-scale
neural network for language. However, agrammatic
speech does not appear to be a property of patients with
insult to this cingulate region. According to non-linguistic studies (Corbetta, Miezin, Dobmeyer, Shulman, &
Petersen, 1991; Corbetta & Shulman, 2002), it is possible
to speculate that the cingulate may help support selective
attention to detecting an unstressed grammatical morpheme. This would be consistent with other cognitive
and imaging work implicating poor selective attention in
the sentence processing limitations of patients with Parkinson’s disease (Grossman et al., 2003; Lee, Grossman,
Morris, Stern, & Hurtig, 2003). Given the absence of
anterior cingulate recruitment during contrasts of activation patterns for diVerent grammatical features, we
cannot rule out that anterior cingulate supports a taskrelated resource involved in deciding between two possible responses (Botvinick, Nystrom, Fissell, Carter, &
Cohen, 1999; Botvinick et al., 2001; Carter et al., 1998).
Additional work is needed to determine more precisely
the role of anterior cingulate cortex in sentence comprehension.
ARTICLE IN PRESS
A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
4.2. Transitivity
We observed activation of left vIFC during judgments
of the transitivity feature as well. Regression analyses
showed that left vIFC is linked to anterior cingulate, an
area that may help selectively attend to the relevant features of a sentence involving transitivity. When the pattern of activation for inXectional morphology was
directly contrasted with the activation proWle for verb
transitivity, moreover, we found signiWcantly greater left
dIFC activation during transitivity judgments. Our preliminary behavioral assessments suggested that verb transitivity involves a more prominent WM component.
Together with the results of published ERP studies
(Friederici, 1998; Hagoort, et al., 1993; Kaan, 2000;
Osterhout & Holcomb, 1992; Rosler et al., 1993), this is
consistent with the hypothesis associating a greater WM
component with dIFC during processing of some grammatical features. ROI analyses also emphasized the relatively dorsal distribution of left IFC activation for verb
transitivity judgments. SPM and ROI regression analyses
showed that left dIFC activation during processing of
transitivity extends into a later time window as well, associating left dIFC recruitment with WM.
Examinations of the speciWc distribution of recruitment
within left IFC have been performed by several investigators. Some contrasts have attempted to elucidate the locus
of activation for diVerent tasks in left IFC, such as phonologic judgments compared to semantic memory judgments
(Poldrack et al., 1999). Other work has contrasted judgments of a grammatical feature relative to a thematic role
feature (Kang et al., 1999; Kuperberg et al., 2000, 2003;
Newman et al., 2003; Ni et al., 2000). Stimuli showing prolonged WM-related activation in an ERP study of sentence processing (Fiebach et al., 2002) were used to
demonstrate a dorsal locus of activation within left IFC
associated with WM for these materials (Fiebach et al.,
2005). While Fiebach et al. (2005) also identiWed two loci
of activation within IFC, the functional signiWcance of this
dissociation was not clear in their study. Recruitment of
the dorsal portion of left IFC is consistent with studies
associating left dIFC with verbal WM (Chein et al., 2002;
Smith & Jonides, 1999; Smith, Marshuetz, & Geva, 2002).
Left dIFC activation during judgments of transitivity compared to morphology appears to be related to the relatively
prominent WM component of the transitivity items in the
present study, but the precise nature of this WM-associated activation requires additional investigation.
We also observed right IFC activation for grammatical structures involving transitivity, resembling other
fMRI work assessing grammatical comprehension
(Fiebach et al., 2005; Friederici et al., 2000a; Just et al.,
1996; Luke et al., 2002; Moro et al., 2001). While some
of these investigators have suggested that right IFC
supports WM demands during sentence processing
(Just et al., 1996), right IFC activation is not observed
15
in other studies assessing WM during sentence comprehension (Cooke et al., 2002). There is little converging
evidence from studies of stroke to suggest that right
IFC insult plays a role in WM during sentence processing, although right hemisphere insult may interfere
with discourse features of comprehension, prosody,
and two-term reasoning problems (Gardner, Brownell,
Wapner, & Michelow, 1983; Goel, Gold, Kapur, &
Houle, 1998; Goel, Buchel, Frith, & Dolan, 2000; Gorelick & Ross, 1987; Hough, 1990; Joanette & Goulet,
1990; Martin & McDonald, 2003; Siegal & Varley,
2002). Right IFC also has been implicated in recovery
of language functioning following left IFC stroke,
although this is not a consistent observation (Basso,
Gardelli, Grassi, & Mariotti, 1989; Heiss, Kessler,
Karbe, Fink, & Pawlik, 1993; Heiss, Kessler, Thiel,
Ghaemi, & Karbe, 1999; Rosen et al., 2000). Some
work in healthy aging proposes that contralateral
homologues are activated during cognitive challenges
to supplement the neural network recruited in your
adults (Cabeza, Anderson, Locantore, & McIntosh,
2002), although this was not seen in a study of sentence
processing in healthy seniors (Grossman et al., 2002).
Additional work is needed to establish the basis for
right IFC activation during sentence comprehension.
Regardless of the precise nature of WM associated
with dIFC recruitment, regression analyses linked left
dIFC to left PLTC activity in a late time window. This
may be consistent with studies emphasizing a role for left
PLTC in supporting WM during sentence processing
(Caplan et al., 1998; Cooke et al., 2002; Grossman et al.,
2002; Just et al., 2001; Keller et al., 2001; Michael et al.,
2001), although a direct contrast of transitivity minus
inXectional morphology did not reveal left PLTC activation. It is also possible that two adjacent PLTC regions
contribute independently to structure-building and WM.
Inspection of some studies suggests a more dorsal locus of
activation associated with WM compared to structurebuilding during sentence processing (Grossman et al.,
2002). However, it may be diYcult to dissociate these two
adjacent regions anatomically given the limited spatial resolution of fMRI. Other work has suggested that a single
brain region may play multiple roles during sentence processing, where the region may have increased activity
under one condition but suppressed activity under
another condition (Kuperberg et al., 2003). From this perspective, left PLTC may contribute to grammatical structure-building in an early time window, but may help
support lexical–semantic or WM-related processes in a
later time window. Additional work is needed to evaluate
the role of left PLTC in sentence processing.
4.3. Grammatical category
Judgments of sentences involving a word’s grammatical category showed left vIFC and left PLTC
ARTICLE IN PRESS
16
A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
activation. This is consistent with our observations of
other stimuli, and emphasizes the central role of these
brain areas in judging many kinds of grammatical features. Stimulus development studies demonstrated that
stimuli involving grammatical category appear to
entail a WM-demanding component, although at a
level that appears to lie between inXectional morphology and transitivity types of features. Activation of
right IFC and right parietal cortex was also seen on
fMRI during judgments of grammatical category.
Recruitment of these areas has been seen in other fMRI
studies of sentence processing (Friederici et al., 2000a;
Just et al., 1996; Luke et al., 2002; Moro et al., 2001).
Both right IFC (Just et al., 1996) and right PLTC
(Cooke et al., 2002) activation have been associated
with a WM component of sentence processing. One
possibility is that grammatical category features
involve a diVerent kind of WM compared to transitivity features.
These Wndings emphasize the dynamic nature of the
large-scale neural network underlying sentence comprehension. Left vIFC appears to play a constant role
in all three grammatical judgments. Regression analyses also implicate left PLTC in judgments of all three
features. We believe that the ubiquitous involvement
of these brain regions during the processing of all
three grammatical structures emphasizes their central
role in grammatical structure-building during sentence comprehension. Importantly, the core vIFC and
PLTC areas of the left hemisphere appear to interact
selectively with other brain regions such as left dIFC,
anterior cingulate, right dIFC, and right parietal cortex, depending on the speciWc nature of the grammatical feature. This variable pattern of activation
suggests that some brain regions are not a necessary
component of a sentence processing network. Instead,
areas such as dIFC and anterior cingulate appear to
contribute as needed during the course of sentence
processing to support speciWc, WM-related demands
of a particular grammatical feature. Observations
such as these emphasize that the large-scale neural
network for sentence processing adapts to the challenges it confronts during the course of
comprehension.
Acknowledgments
This research was supported in part by National
Institutes of Health Grants DC00237, AG17586,
NS35867, and AG15116. Portions of this work were presented at the CUNY Sentence Processing Conference,
Philadelphia, PA, March 2001, and the meeting of the
Cognitive Neuroscience Society, New York City, New
York, March 2001.
Appendix A
InXectional morphology-short-correct
1 The manager is daily quite visibly being confronted by the crazy company president.
2 The glass is skillfully and gingerly being selected because of its
romantic value.
3 The computer is eagerly but earnestly being replaced by a fancy new
machine.
4 The biker is by next week being imprisoned for drunken and disorderly behavior.
5 The gymnast is not too fairly being evaluated in the Xoor exercise
Wnals.
6 The lamb is gently and gracefully being bathed by the sick little girl.
7 The zoo is all too frequently being walked through by groups of children.
8 The jack is preferably quite quickly being installed by the capable
telephone workman.
9 The ballerina is seemingly quite often being stalked by the insanely
jealous fan.
10 The wig is expertly and neatly being created for the famous blonde
actress.
11 The test is vaguely yet slowly being explained to excited new college
students.
12 The family is against all reason being evicted by the sleazy realty
agents.
13 The Xoor is slowly but steadily being polished by us this very evening.
14 The boy is sternly and eternally being scolded by the impatient angry
parents.
15 The crew is really only mildly being entertained because the movie
has subtitles.
16 The mail is shockingly but eYciently being delivered even during the
winter holidays.
17 The concert is thankfully only brieXy being postponed until the conductor’s return tomorrow.
18 The soldier is regularly and thoroughly being examined by the serious
drill sergeant.
19 The maid is duly and cunningly being watched because of her bad
attitude.
20 The plumbing is hopefully by tomorrow being Wxed by my landlord
and him.
21 The basement is hastily and ruthlessly being destroyed due to the rain
storm.
22 The oYce is being quietly and cautiously observed during the faithful
employee’s exhibit.
23 The book is carefully and closely being picked through by the curious
students.
24 The lesson is usually not completely being prepared in time for the
class.
InXectional morphology-short-violation
25 The zoo is all too frequently being walk through by groups of children.
26 The jack is preferably quite quickly being install by the capable telephone workman.
27 The ballerina is seemingly quite often being stalk by the insanely jealous fan.
28 The wig is expertly and neatly being create for the famous blonde
actress.
29 The test is vaguely yet slowly being explain to excited new college students.
30 The family is against all reason being evict by the sleazy realty agents.
31 The manager is daily quite visibly being confront by the crazy company president.
32 The glass is skillfully and gingerly being select because of its romantic
value.
ARTICLE IN PRESS
A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
The computer is eagerly but earnestly being replace by a fancy new
machine.
The biker is by next week being imprison for drunken and disorderly
behavior.
The gymnast is not too fairly being evaluate in the Xoor exercise
Wnals.
The lamb is gently and gracefully being bathe by the sick little girl.
The maid is duly and cunningly being watch because of her bad attitude.
The plumbing is hopefully by tomorrow being Wx by my landlord and
him.
The basement is hastily and ruthlessly being destroy due to the rain
storm.
The oYce is being quietly and cautiously observe during the faithful
employee’s exhibit.
The book is carefully and closely being pick through by the curious
students.
The lesson is usually not completely being prepare in time for the
class.
The Xoor is slowly but steadily being polish by us this very evening.
The boy is sternly and eternally being scold by the impatient angry
parents.
The crew is really only mildly being entertain because the movie has
subtitles.
The mail is shockingly but eYciently being deliver even during the
winter holidays.
The concert is thankfully only brieXy being postpone until the conductor’s return tomorrow.
The soldier is regularly and thoroughly being examine by the serious
drill sergeant.
InXectional morphology-long-correct
49 The Xoor is being barely but steadily polished by us this very evening.
50 The boy is being sternly and eternally scolded by the impatient angry
parents.
51 The crew is being really only mildly entertained because the movie
has subtitles.
52 The mail is being shockingly but eYciently delivered even during the
winter holidays.
53 The concert is being thankfully only brieXy postponed until the conductor’s return tomorrow.
54 The soldier is being regularly and thoroughly examined by the serious
drill sergeant.
55 The maid is being duly and cunningly watched because of her bad
attitude.
56 The plumbing is being hopefully by tomorrow Wxed by my landlord
and him.
57 The basement is being hastily and ruthlessly destroyed due to the rain
storm.
58 The oYce is being quietly and cautiously observed during the faithful
employee’s exhibit.
59 The book is being carefully and closely picked by the four curious
students.
60 The lesson is being usually not completely prepared in time for the
class.
61 The manager is being daily quite visibly confronted by the crazy company president.
62 The glass is being skillfully and gingerly selected because of its
romantic value.
63 The computer is being eagerly but earnestly taken for a full cash
refund.
64 The biker is being by next week imprisoned for drunken and disorderly behavior.
65 The gymnast is being not too fairly evaluated in the Xoor exercise
Wnals.
66 The lamb is being gently and gracefully bathed by the sick little girl.
67 The zoo is being all too frequently walked through by groups of children.
68
69
70
71
72
17
The jack is being preferably quite quickly installed by the capable
telephone workman.
The ballerina is being seemingly quite often stalked by the insanely
jealous fan.
The wig is being expertly and neatly created for the famous blonde
actress.
The test is being vaguely yet slowly explained to excited new college
students.
The family is being against all reason evicted by the sleazy realty
agents.
InXectional morphology-long-violation
73 The maid is being duly and cunningly watch because of her bad attitude.
74 The plumbing is being hopefully by tomorrow Wx by my landlord and
him.
75 The basement is being hastily and ruthlessly destroy due to the rain
storm.
76 The oYce is being quietly and cautiously observe during the faithful
employee’s exhibit.
77 The book is being carefully and closely pick by the four curious students.
78 The lesson is being usually not completely prepare in time for the
class.
79 The Xoor is being barely but steadily polish by us this very evening.
80 The boy is being sternly and eternally scold by the impatient angry
parents.
81 The crew is being really only mildly entertain because the movie has
subtitles.
82 The mail is being shockingly but eYciently deliver even during the
winter holidays.
83 The concert is being thankfully only brieXy postpone until the conductor’s return tomorrow.
84 The soldier is being regularly and thoroughly examine by the serious
drill sergeant.
85 The zoo is being all too frequently walk through by groups of children.
86 The jack is being preferably quite quickly install by the capable telephone workman.
87 The ballerina is being seemingly quite often stalk by the insanely jealous fan.
88 The wig is being expertly and neatly create for the famous blonde
actress.
89 The test is being vaguely yet slowly explain to excited new college students.
90 The family is being against all reason evict apartments by sleazy
realty agents.
91 The manager is being daily quite visibly confront by the crazy company president.
92 The glass is being skillfully and gingerly select because of its romantic
value.
93 The computer is being eagerly but earnestly replace by a fancy new
machine.
94 The biker is being by next week imprison for drunken and disorderly
behavior.
95 The gymnast is being not too fairly evaluate in the Xoor exercise
Wnals.
96 The lamb is being gently and gracefully bathe by the sick
little girl.
Grammatical category-short-correct
97
The weather is frigidly and harshly being worsened by the cold
mountain air.
98
The coVee is plainly but freshly being brewed through the gourmet shop Wlter.
99
The café is discreetly but crudely being renovated just in time for
summer.
ARTICLE IN PRESS
18
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
The essay is playfully very tastefully being printed by the rich
magazine editor.
The Wre is safely and intensely being tended by Boy Scout troop
leaders.
The fruit is freshly and pleasingly being displayed during the Wrst
big sale.
The dance is not too seriously being rehearsed prior to the debut
performance.
The movie is critically yet kindly being reviewed by the skinny
bald man.
The shelf is sturdily and Wrmly being repaired outside in the back
yard.
The porch is barely and shabbily being completed before she
goes to work.
The farm is proWtably and greedily being sold at the school’s
charity auction.
The quilt is warmly and lovingly being designed by my friends
from church.
The friend is casually but happily being met near the shopping
mall entrance.
Sarah is willingly and cheaply being taught piano lessons by her
sister Nancy.
The metal is somehow quite specially being reWned by the old village goldsmith.
The politician is silently and solemnly being released today
despite the war crimes.
The sheep is secretly and savagely being killed by the processed
meat company.
The water is abundantly and clearly being Wltered because of the
customer’s threats.
The proposal is illegally and massively being copied by the
deceitful paper company.
The cat is easily and quickly being declawed as requested by the
owner.
The model is morally and physically being starved because of stiV
job rivalry.
The employee is swiftly and rapidly being Wred after the city
paper scandal.
The car is safely but frugally being towed away from the Wre
lane.
The book is insanely and unwisely being burned at the ultra
political rally.
Grammatical category-short-violation
121 The dance is not too seriously being rehearsal prior to the debut performance.
122 The movie is critically yet kindly being newspaper by the skinny bald
man.
123 The shelf is sturdily and Wrmly being hammer outside in the back
yard.
124 The porch is barely and shabbily being completion before she goes to
work.
125 The farm is proWtably and greedily being sale at the school’s charity
auction.
126 The quilt is warmly and lovingly being project by my friends from
church.
127 The weather is frigidly and harshly being blizzard by the cold mountain air.
128 The coVee is plainly but freshly being coVeepot through the gourmet
shop Wlter.
129 The café is discreetly but crudely being renovation just in time for
summer.
130 The essay is playfully and tastefully being printer by the rich magazine editor.
131 The Wre is safely but intensely being forest by Boy Scout troop leaders.
132 The fruit is freshly and pleasingly being exposition during the Wrst
big sale.
133 The proposal is illegally and massively being copier by the deceitful
paper company.
134 The cat is easily and quickly being surgery as requested by the owner.
135 The model is morally and physically being starvation because of stiV
job rivalry.
136 The employee is swiftly and rapidly being harassment after the city
paper scandal.
137 The car is safely but frugally being garage away from the Wre lane.
138 The book is insanely and unwisely being bonWre at the ultra political
rally.
139 The friend is casually but happily being reunion near the shopping
mall entrance.
140 Sarah is willingly and cheaply being class piano lessons by her sister
Nancy.
141 The metal is somehow quite specially being reWnement by the old village goldsmith.
142 The politician is silently and solemnly being freedom today despite
the war crimes.
143 The sheep is secretly and savagely being killer by the processed meat
company.
144 The water is abundantly and clearly being Wltration because of the
customer’s threats.
Grammatical category-long-correct
145 The friend is being casually and happily met near the shopping mall
entrance.
146 Sarah is being willingly and cheaply taught piano lessons by her sister Nancy.
147 The metal is being somehow quite specially reWned by the old village
goldsmith.
148 The politician is being silently and solemnly released today despite
the war crimes.
149 The sheep is being secretly and savagely killed by the processed meat
company.
150 The water is being abundantly and clearly Wltered because of the customer’s threats.
151 The proposal is being illegally and massively copied by the deceitful
paper company.
152 The cat is being easily and quickly declawed as requested by the
owner.
153 The model is being morally and physically starved because of stiV
job rivalry.
154 The employee is being swiftly and rapidly Wred after the city paper
scandal.
155 The car is being safely but frugally towed away from the Wre lane.
156 The book is being insanely and unwisely burned at the ultra political
rally.
157 The weather is being frigidly and harshly worsened by the cold
mountain air.
158 The coVee is being plainly but freshly brewed through the gourmet
shop Wlter.
159 The café is being discreetly but crudely renovated just in time for
summer.
160 The essay is being playfully and tastefully printed by the rich magazine editor.
161 The Wre is being safely but intensely tended by Boy Scout troop leaders.
162 The fruit is being freshly and pleasingly discounted during the Wrst
big sale.
163 The dance is being not too seriously rehearsed prior to the debut performance.
164 The movie is being critically yet kindly reviewed by the skinny bald
man.
165 The shelf is being sturdily and Wrmly repaired outside in the back
yard.
166 The porch is being barely and shabbily completed before she goes to
work.
167 The farm is being proWtably and greedily sold at the school’s charity
auction.
ARTICLE IN PRESS
A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
168 The quilt is being warmly and lovingly designed by my friends from
church.
Grammatical category-long-violation
169 The proposal is being illegal and massive copier by the deceitful
paper company.
170 The cat is being easy and quick surgery as requested by the owner.
171 The model is being moral and physical starvation because of stiV job
rivalry.
172 The employee is being swift and rapid harassment after the city
paper scandal.
173 The car is being safe but frugal garage away from the Wre lane.
174 The book is being insane and unwise bonWre at the ultra political
rally.
175 The friend is being casual but happy reunion near the shopping mall
entrance.
176 Sarah is being willing and cheap class piano lessons by her sister
Nancy.
177 The metal is being somehow quite special reWnement by the old village goldsmith.
178 The politician is being silent and solemn freedom today despite the
war crimes.
179 The sheep is being secret and savage killer by the processed meat
company.
180 The water is being abundant and clear Wltration because of the customer’s threats.
181 The dance is being not too serious rehearsal prior to the debut performance.
182 The movie is being critical yet kind newspaper by the skinny bald
man.
183 The shelf is being sturdy and Wrm hammer outside in the back yard.
184 The porch is being bare and shabby completion before she goes to
work.
185 The farm is being proWtable and greedy sale at the school’s charity
auction.
186 The quilt is being warm and loving project by my friends from
church.
187 The weather is being frigid and harsh blizzard by the cold mountain
air.
188 The coVee is being plain but fresh coVeepot through the gourmet
shop Wlter.
189 The café is being discreet but crude renovation just in time for summer.
190 The essay is being playful and tasteful printer by the rich magazine
editor.
191 The Wre is being safe but intense forest by Boy Scout troop leaders.
192 The fruit is being fresh and pleasing exposition during the Wrst big
sale.
Transitivity-short-correct
193 The door is repeatedly and anxiously being opened quickly by the
nervous salesman.
194 The farmer is admiringly and excitedly being given gifts from his
grateful staV.
195 The Earth is rapidly and harshly being hit by asteroids at high velocity.
196 The prisoner is quietly and covertly being followed through the forgotten gutter system.
197 The singer is overtly and illegally being exploited by her greedy stage
manager.
198 The eagle is deftly but swiftly being lifted through the beautiful
mountain air.
199 The toddler is cautiously but properly being supervised in the day
care center.
200 The truth is quickly and joyously being uncovered because of the
policeman’s toil.
201 The patient is slowly yet forcefully being led outside for a long walk.
19
202 The jelly is completely and utterly being pushed out of the powdered
doughnut.
203 The Xight is Wnally and deWnitely being delayed today after many
unanswered questions.
204 The crisis is nastily but needfully being resolved in the stately President’s oYce.
205 The President is indirectly yet forcefully being dunked into the fountain in May.
206 The girl is softly and stealthily being whisked through the back door
soundlessly.
207 The couple is secretly and eagerly being married because of their
friend’s remarks.
208 The baby is sweetly and constantly being rocked until morning by its
parents.
209 The garbage is aversely and suddenly being smelled because of the
rotten eggs.
210 The play is weekly and monthly being performed by the troop of
actors.
211 The guest is eagerly and excitedly being greeted in the fancy hotel
lobby.
212 The student is often to always being asked questions by the thesis
committee.
213 The woman is brazenly and lewdly being harassed by the building
workers outside.
214 The bride is elegantly and slowly being led down the steep spiral
stairs.
215 The dog is endlessly yet happily being chased up the neighbor’s tall
tree.
216 The festival is almost most deWnitely being brought to the city’s huge
auditorium.
Transitivity-short-violation
217 The toddler is cautiously but properly being slept in the day care center.
218 The truth is quickly and joyously being triumphed because of the
policeman’s toil.
219 The patient is slowly yet forcefully being gone outside for a long
walk.
220 The jelly is completely and utterly being oozed out of the powdered
doughnut.
221 The Xight is Wnally and deWnitely being arrived today after many
unanswered questions.
222 The crisis is nastily but needfully being occurred in the stately President’s oYce.
223 The door is repeatedly and anxiously being looked by the extremely
nervous salesman.
224 The farmer is admiringly and excitedly being emerged gifts from his
grateful staV.
225 The Earth is rapidly and harshly being collided by asteroids at high
velocity.
226 The prisoner is quietly and covertly being escaped through the forgotten gutter system.
227 The singer is overtly and illegally being evolved by her greedy stage
manager.
228 The eagle is deftly but swiftly being soared through the beautiful
mountain air.
229 The guest is eagerly and excitedly being fainted in the fancy hotel
lobby.
230 The student is often to always being struggled questions by the thesis
committee.
231 The woman is brazenly and lewdly being winked by the building
workers outside.
232 The bride is elegantly and slowly being descended down the steep
stairs.
233 The dog is endlessly yet happily being barked up the neighbor’s tall
tree.
234 The festival is almost most deWnitely being came to the city’s huge
auditorium.
ARTICLE IN PRESS
20
A. Cooke et al. / Brain and Language xxx (2005) xxx–xxx
235 The President is indirectly yet forcefully being fallen into the fountain in May.
236 The girl is softly and stealthily being exited through the back door
soundlessly.
237 The couple is secretly and eagerly being eloped because of their
friend’s remarks.
238 The baby is sweetly and constantly being snoozed until morning by
its parents.
239 The garbage is aversely and suddenly being reeked because of the
rotten eggs.
240 The play is weekly and monthly being appeared by the troop of
actors.
Transitivity-long-correct
241 The President is being indirectly yet forcefully dunked into the fountain in May.
242 The girl is being softly and stealthily whisked through the back door
soundlessly.
243 The couple is being secretly and eagerly married because of their
friend’s remarks.
244 The baby is being sweetly and constantly rocked until morning by its
parents.
245 The garbage is being aversely and suddenly smelled because of the
rotten eggs.
246 The play is being weekly and monthly performed by the troop of
actors.
247 The guest is being eagerly and excitedly greeted in the fancy hotel
lobby.
248 The student is being often to always asked questions by the thesis
committee.
249 The woman is being brazenly and lewdly harassed by the building
workers outside.
250 The bride is being elegantly and slowly led down the steep spiral
stairs.
251 The dog is being endlessly yet happily chased up the neighbor’s tall
tree.
252 The festival is being almost most deWnitely brought to the city’s huge
auditorium.
253 The door is being repeatedly and anxiously opened quickly by the
nervous salesman.
254 The farmer is being admiringly and excitedly given gifts from his
grateful staV.
255 The Earth is being rapidly and harshly hit by asteroids at high velocity.
256 The prisoner is being quietly and covertly followed through the forgotten gutter system.
257 The singer is being overtly and illegally exploited by her greedy stage
manager.
258 The eagle is being deftly but swiftly lifted through the beautiful
mountain air.
259 The toddler is being cautiously but properly supervised in the day
care center.
260 The truth is being quickly and joyously uncovered because of the
policeman’s toil.
261 The patient is being slowly yet forcefully led outside for a long walk.
262 The jelly is being completely and utterly pushed out of the powdered
doughnut.
263 The Xight is being Wnally and deWnitely delayed today after many
unanswered questions.
264 The crisis is being nastily but needfully resolved in the stately President’s oYce.
Transitivity-long-violation
265 The guest is being eagerly and excitedly fainted in the lobby’s fancy
hotel.
266 The student is being often to always struggled questions by the thesis
committee.
267 The woman is being brazenly and lewdly winked by the building
workers outside.
268 The bride is being elegantly and slowly descended down the steep
spiral stairs.
269 The dog is being endlessly yet happily barked up the neighbor’s tall
tree.
270 The festival is being almost most deWnitely came to the city’s huge
auditorium.
271 The President is being indirectly yet forcefully fallen into the fountain in May.
272 The girl is being softly and stealthily exited through the back door
soundlessly.
273 The couple is being secretly and eagerly eloped until morning by its
parents.
274 The baby is being sweetly and constantly snoozed until morning by
its parents.
275 The garbage is being aversely and suddenly reeked because of the
rotten eggs.
276 The play is being weekly and monthly appeared by the troop of
actors.
277 The toddler is being cautiously but properly slept in the day care center.
278 The truth is being quickly and joyously triumphed because of the
policeman’s toil.
279 The patient is being slowly yet forcefully gone outside for a long
walk.
280 The jelly is being completely and utterly oozed out of the powdered
doughnut.
281 The Xight is being Wnally and deWnitely arrived today after many
unanswered questions.
282 The crisis is being nastily but needfully occurred in the stately President’s oYce.
283 The door is being repeatedly and anxiously looked by the extremely
nervous salesman.
284 The farmer is being admiringly and excitedly emerged gifts from his
grateful staV.
285 The Earth is being rapidly and harshly collided by asteroids at high
velocity.
286 The prisoner is being quietly and covertly escaped through the forgotten gutter system.
287 The singer is being overtly and illegally evolved by her greedy stage
manager.
288 The eagle is being deftly but swiftly soared through the beautiful
mountain air.
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