When Elephants Fly: Differential Sensitivity of Right - 麻省理工学院人工智能研究专业

When Elephants Fly: Differential Sensitivity of Right
and Left Inferior Frontal Gyri to Discourse
and World Knowledge

Laura Menenti1, Karl Magnus Petersson1,2, Rene´ Scheeringa1,
and Peter Hagoort1,2

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抽象的

& Both local discourse and world knowledge are known to in-
fluence sentence processing. We investigated how these two
sources of information conspire in language comprehension. 二
types of critical sentences, correct and world knowledge anom-
alies, were preceded by either a neutral or a local context. 这
latter made the world knowledge anomalies more acceptable or
plausible. We predicted that the effect of world knowledge anom-
alies would be weaker for the local context. World knowledge
effects have previously been observed in the left inferior frontal

地区 (布罗德曼区 45/47). In the current study, an effect of
world knowledge was present in this region in the neutral context.
We also observed an effect in the right inferior frontal gyrus, 哪个
was more sensitive to the discourse manipulation than the left
inferior frontal gyrus. 此外, the left angular gyrus reacted
strongly to the degree of discourse coherence between the con-
text and critical sentence. 全面的, both world knowledge and the
discourse context affect the process of meaning unification, 但
do so by recruiting partly different sets of brain areas. &

介绍

‘‘The elephant flies’’ is a sentence you can immediately
understand, although you know that elephants typically do
not fly. Maybe you thought of a circus traveling by air-
plane or maybe you thought of Dumbo. To comprehend a
句子, we rapidly make use of the knowledge we have
about the world, and even if what we read or hear does
not fit with what we typically take for granted we usually
comprehend utterances as long as they are well-formed.
Sentence comprehension can be further facilitated if the
reader or hearer is provided an adequate context. If we
first read the sentence, ‘‘Dumbo is a fantasy animal.’’ then
‘‘The elephant flies.’’ becomes a coherent sentence in its
语境. The question then is, how understanding of the
sentence ‘‘The elephant flies’’ is modulated depending on
what we just read or heard. When reading texts or listen-
ing to speech, we are constantly integrating the current
word meaning with the preceding information. 抓握
the neural implementation of contextual integration is an
essential part of understanding the neural organization of
language comprehension.

Modulations of sentence comprehension by discourse
have previously been investigated with various techniques.
In an eye-movement study, Duffy and Keir (2004) 在-
vestigated the influence of disambiguating discourse on

1Donders Institute for Brain, 认识, and Behavior, 奈梅亨,
荷兰人, 2Max Planck Institute for Psycholinguistics,
奈梅亨, 荷兰人

processing of gender stereotypes. Sentences such as ‘‘The
electrician taught herself. . .’’ evoke longer reading times
of the word ‘‘herself’’ because electricians are stereo-
typically men and readers therefore take longer time in-
tegrating the incoming word ‘‘herself.’’ However, after a
disambiguating context explaining that the electrician was
a woman, the effect of stereotype anomaly on reading
times at the critical word disappeared. Hald, Steenbeek-
Planting, and Hagoort (2007) investigated modulation of
world knowledge comprehension by discourse context
with ERPs. Hagoort, Hald, Bastiaansen, and Petersson
(2004) had already established that world knowledge
anomalies, just like semantic anomalies, elicit an N400
影响, which indexes ease of semantic integration of con-
tent words. Hald et al. (2007) presented world knowl-
edge sentences (correct or anomalies: ‘‘Dutch trains are
yellow/white and blue.’’) that were preceded by two types
of contexts: (我) neutral contexts introduced the topic
of the world knowledge sentence and were in line with
the world knowledge of the listener/reader (例如, ‘‘The
Netherlands are famous for their designers. 此外,
the Dutch railways have chosen a very conspicuous color
scheme, which makes them recognizable everywhere.’’);
(二) local contexts introduced information that made de-
viations from the default world knowledge more accept-
有能力的 (例如, ‘‘The coming world championships are one
big national spectacle. The Dutch railways have painted
the Dutch flag on their trains.’’) They found that the
N400 effect to world knowledge anomalies was weaker
when the sentences were preceded by the local context,

D 2008 麻省理工学院

认知神经科学杂志 21:12, PP. 2358–2368

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thereby supporting the idea that ongoing discourse can
modulate the integration of world knowledge information
retrieved from long-term memory. 相似地, Nieuwland
and Van Berkum (2006) investigated discourse modula-
tion of animacy anomalies such as, ‘‘The peanut was in
love.’’ Such animacy anomalies presented in isolation elic-
ited a clear N400 effect compared to neutral sentences,
as ‘‘The peanut was salted,’’ but when these sentences
were preceded by a supporting discourse introducing an
amorous peanut, the N400 effect not only disappeared,
but was reversed: ‘‘The peanut was salted’’ now elicited
a larger N400 than ‘‘The peanut was in love.’’ Taken
一起, this suggests that local discourse can override
semantic and world knowledge anomalies and that the
anomaly effect is thus inherently context-dependent.

To form a coherent representation of a multiword ut-
terance, the reader must not only retrieve and integrate
lexical information and world knowledge that is stored
in long-term memory. She also has to form a represen-
tation of the ongoing discourse itself. Here it is impor-
tant to make a distinction between three different types
of discourse representation: the surface code, 文本
根据, and the situation model (van Dijk & Kintsch, 1983;
see also Graesser, Millis, and Zwaan, 1997, for an ex-
tensive and more recent review). The surface code is a
literal representation of the exact wording and syntax of
文本 (‘‘The animal tamer gently patted the elephant.
The animal was afraid of entering the plane.’’), 文本
base contains explicit text propositions in a stripped-
down format that preserves meaning (例如, PAT [动物
tamer, elephant], AFRAID [elephant], ENTER [elephant,
飞机]), and the situation model is the content or the
conceptual structure of the microworld that the text de-
scribes (Circus traveling by airplane). In the present
文章, by discourse representation, we mean the situa-
tion model level of representation, which determines
how additional incoming information is interpreted.

A recurring issue in fMRI research on discourse com-
prehension is the involvement of the right hemisphere
in language processing and more specifically in discourse
加工. Jung-Beeman (2005; and Beeman 1998) 亲-
posed a division of labor between the right and left hemi-
spheres in language processing. In this proposal, 正确的
and left hemispheres perform similar functions in lan-
guage processing but they encode information in a different
方式, with the right hemisphere performing coarser seman-
tic coding than the left. 换句话说, it is suggested that
semantic fields are more focused in the left hemisphere,
whereas they are more diffuse in the right hemisphere.
例如, the semantic field for the word foot might
include hand, leg, toe in the left hemisphere, 但在
添加, bare, step, ball in the right hemisphere. Reading
foot will then activate a few related words quite strongly in
the left hemisphere, and many words more weakly in the
正确的. 此外, the right hemisphere has been shown to
be involved in processing jokes (Bartolo, Benuzzi, Nocetti,
Baraldi, & Nichelli, 2006) and metaphors (Stringaris et al.,

2006; but see Stringaris, 梅德福, Giampietro, Brammer,
& 大卫, 2007; see Mitchell & Crow, 2005, for a review).
This has been taken to suggest that the right hemisphere
is involved in processing nonliteral language. Recent evi-
登塞, 然而, seems to suggest that the division of
labor is different, in that the left hemisphere is more sen-
sitive to salient (dominant) meanings and direct semantic
关系, whereas the right hemisphere is more sen-
sitive to nonsalient (novel and inferred) meanings and
more remote relationships, irrespective of whether they
are literal or not (Giora, 2007; Mashal, Faust, 活动, &
Jung-Beeman, 2007; 施密特, DeBuse, & Seger, 2007).

A few studies indicate that regions in the right hemi-
sphere are sensitive to discourse coherence. Kuperberg,
Lakshmanan, Caplan, and Holcomb (2006) found that
the right inferior frontal gyrus (RIFG) showed stron-
ger responses to sentences unrelated to preceding two-
sentence contexts than to related sentences. In line with
this finding, St George, Kutas, Martinez, and Sereno
(1999) found that untitled (and therefore less coherent)
paragraphs elicited more activation than titled paragraphs
in the right but not in the left hemisphere. 此外,
Caplan and Dapretto (2001) found that such coherence
violation effects depend on the precise type of anomaly.
Their subjects read question–answer dialogues where the
answer was on- or off-topic, versus logical or illogical. 为了
例子, for the question, ‘‘Do you believe in angels?'' 这
on-topic answer was, ‘‘Yes, I have my own special angel,’’
whereas the off-topic answer was, ‘‘Yes, I like to go to
camp.’’ For the question, ‘‘Do you like having fun?'' 这
logical answer was, ‘‘Yes, because it makes me happy,’’
whereas the illogical answer was, ‘‘No, because it makes
me happy.’’ They found that the effect of discourse anom-
alies was more left-lateralized for logicality and more right-
lateralized for topicality.

Hagoort (2005) proposes that the left inferior frontal
gyrus (LIFG) is an active workspace for language pro-
cessing. This region is involved in unifying structured
pieces of knowledge that are stored in long-term mem-
奥里 ( 杰肯道夫, 2007; 狐狸 & 活动, 2000). 它也是
integrates linguistic with nonlinguistic information, 在-
包括, 例如, world knowledge (Hagoort et al.,
2004) or gestures ( Willems, Ozyurek, & Hagoort, 2007).
Hagoort et al. (2004) compared integration of semantic
and world knowledge. Semantic knowledge is knowl-
edge about the words in our language. World knowl-
边缘, 另一方面, refers to our knowledge about
the state of affairs in the world. An example will clarify
the distinction: the sentence, ‘‘Dutch trains are sour,’’
violates our knowledge about the meaning of the word
‘‘sour,’’ which is not applicable to trains. On the other
手, the sentence, ‘‘Dutch trains are white,’’ is an in-
terpretable sentence, but it just so happens that Dutch
trains are yellow and, 所以, most Dutch people will
know this sentence to be false. Hagoort et al. found that
compared to correct sentences, the LIFG, 具体来说
BA 45 和 47, showed stronger BOLD responses to both

Menenti et al.

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semantic and world knowledge anomalies. They con-
cluded that this result provided evidence for a role of
the LIFG in the integration of world knowledge as well
as semantic knowledge, in line with linguistic accounts
that have argued for the absence of a clear distinction
between semantic and world knowledge regarding lexi-
cal items (杰肯道夫, 2002).

The aim of the present fMRI study is to investigate
whether the LIFG unifies information from long-term
memory and discourse. We investigated whether ongo-
ing discourse could modulate the integration of world
knowledge sentences in the LIFG. If a sentence that
departs from our world knowledge (‘‘The elephant
flies’’) is preceded by a discourse context that supports
它 (Circus traveling by airplane), unification should be
更轻松. 如上所述, there are a few fMRI studies
that shed light on how the brain processes multisen-
tence utterances. Several of these have found that the
right hemisphere is involved in discourse processing (看
米切尔 & Crow, 2005; Bookheimer, 2002 for reviews).
Because we expected right hemisphere involvement to
be stronger in our study with multisentence texts than it
was in Hagoort et al. (2004), where single sentences were
提出, we investigated modulations of world knowl-
edge unification not only in the left hemisphere regions
where they found increased responses to anomalies but
also in their right hemisphere counterparts. To avoid con-
融合, we do not think a true dichotomy exists between
‘‘correct’’ sentences and ‘‘anomalies.’’ Rather, 有一个
continuum from sentences that fit very well with our
world knowledge to sentences that are odd in the light
of what we know. 然而, for ease of reading, 我们用
the terms ‘‘correct’’ and ‘‘anomaly’’ as shorthand.

Our paradigm was similar to the one used by Hald et al.
(2007), with either neutral or local contexts preceding
correct world knowledge sentences or anomalies. 这
local contexts were designed to make the world knowl-
edge anomalies more plausible. The predictions were as
如下: 第一的, left BA 45 和 47 were expected to show a
stronger response to world knowledge anomalies than
to correct sentences in the neutral context. 第二, 这
local context should facilitate integration of the anomaly.
The effect of world knowledge should therefore be re-
duced in the local context compared to the neutral con-
文本. 第三, because the right hemisphere is known to
be involved in discourse processing, we predicted that
the right BA 45 和 47 would also show modulation of
world knowledge processing by discourse context. Fi-
nally, we investigated the effect of both world knowl-
edge and discourse context at the whole-brain level to
see which additional regions might be involved.

方法

Subjects

Thirty-two healthy right-handed Dutch native speakers
with normal or corrected-to-normal vision (13 男人,

19 女性; mean age = 22 ± 3) participated in the ex-
periment. All subjects had attended or were attending
university education in the Netherlands. All subjects gave
written informed consent prior to the experiment and
received a fee for their participation.

Stimuli and Design

One hundred thirty-two sets of Dutch-language stimuli
were created for this experiment. Like Hald et al. (2007),
we used a 2 (西德:1) 2 factorial design with the factors discourse
语境 (neutral/local) and world knowledge (correct/
anomaly). The contexts were either in accordance with
world knowledge (neutral context), or contained a moti-
vation for the world knowledge anomaly in the critical
句子, making the anomaly more acceptable (当地的
语境). The critical sentences were either in accordance
with world knowledge or departed from world knowledge
(anomaly). 有, 然而, one important difference
compared to the ERP study by Hald et al. In the present
学习, the contexts were much more tightly matched:
The two versions of a particular context had an identical
syntactic structure, were matched on sentence and word
length and on word frequency, and differed in as few
words as possible. This was necessary due to the lower
temporal resolution of fMRI compared to EEG. In an fMRI
学习, contexts differing as widely as they did in the ERP
study could lead to strong but hard-to-interpret effects on
BOLD responses to the critical sentences (见表 1 为了
translated examples of our stimuli).

All contexts consisted of three sentences. The first sen-
tence was equal between the two versions of the context
and introduced the subject of the text. The other two
sentences were matched on syntactic structure, word fre-
quency, and number of words, and differed on as few
words as possible. The three context sentences were fol-
lowed by the critical sentence. This critical sentence had
two versions, which only differed in one word, the critical
word. The critical word never appeared in sentence-final
位置, and the contexts did not contain either of the
critical words.

Two hundred twenty-five sets of stimuli were pre-
tested with 75 subjects in total. Nineteen subjects were
given the single critical sentences only, with no prior
语境. The sentences were truncated before the crit-
ical word and subjects were asked to complete them
to make a complete, grammatical sentence. The partici-
pants were instructed to fill in the first thing that came
to mind, and to keep completions as short as possible.
Thirty-one subjects were given a list of items consisting
of half neutral and half local contexts in a random or-
这, again followed by the truncated critical sentences.
Stimuli were selected for the experiment if none of the
subjects completed the critical sentence with the world
knowledge anomaly in the single sentence condition.
After the neutral context, not more than two subjects
should fill in the critical word constituting the anomaly,

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认知神经科学杂志

体积 21, 数字 12

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桌子 1. Example Stimuli

Examples

Neutral contexts

(1) Carl Barks wrote many Donald Duck stories and invented Duckburg. In his early sketches we see
Huey, Dewey and Louie as young well-behaved boys with hats and scarves. They often go out to
help old ladies.

(2) More and more lamp posts are placed in the Netherlands. This way it is easier to see the road.

This is nice for drivers.

(3) There now is a vaccination against measles. The disease is more frequent the lower the age of

patients is. They are more vulnerable to contagion.

Local contexts

(1) Carl Barks wrote many Donald Duck stories and invented Duckburg. In his early sketches we see
Huey, Dewey and Louie as young bad boys with striped sweaters and masks. They often go out to
rob old ladies.

(2) More and more lamp posts are placed in the Netherlands. This way it is harder to see the night sky.

This is sad for astronomers.

(3) There now is a vaccination against measles. The disease is more dangerous the higher the age of

patients is. They are more vulnerable to complications.

Critical sentences

(1) Donald Duck’s nephews are boy scouts/thieves and very smart.

(2) With the lights on you can see more/less at night.

(3) Measles is a disease that especially affects children/elderly.

Neutral or local contexts were followed by one out of two critical sentences, which only differed on one word (下划线的).

whereas after the local context, at least three subjects
should respond with the anomaly. The difference be-
tween the cloze probabilities of the critical word consti-
tuting the world knowledge anomaly after the local and
neutral contexts was required to be at least 20%. Paired-
sample t tests showed that the two contexts differed
significantly in terms of cloze probability of the correct
critical word [t(133) = 18.3, p < .001] and of the anom- alies [t(133) = (cid:2)27.2, p < .001]. The characteristics of the selected items are reported in Table 2. The cloze probability of the correct critical word is higher after the neutral context than in the single sentence, and lower after the local context. The cloze probability of the anomaly is higher after the local context than after the neutral context, or in the single sentence. Local contexts do not succeed in completely overriding world knowl- edge, shown by the residual cloze probability of the Table 2. Results of the Pretest for the Selected Items Cloze Probability Condition Correct Anomaly Neutral context Local context Single sentence 77% 34% 59% 1% 43% 0% Reported are the cloze probabilities of correct completions and anom- alies after the truncated critical sentence (Measles is a disease that especially affects_________ ). The truncated critical sentence was pre- sented after the neutral context, the local context, or in isolation. correct word in the local context condition (34%). In the experiment, four different lists each contained one ver- sion of every item. The items were presented in four dif- ferent pseudorandom orders. No more than three items of the same condition were presented in a row. Procedure Each participant saw 132 trials (33/condition). The ex- perimental session was split in four subsequent runs of (cid:3)15 min. To ensure that the subjects remained atten- tive, they were given a content question about the pre- ceding text on 10% of the trials (e.g., ‘‘Who wrote the Donald Duck stories?’’), and two possible answers from which to choose with a button press (‘‘Carl Barks/ Walt Disney’’). After two runs participants underwent the ana- tomical scan. A trial consisted of the following elements: The dis- course contexts were presented sentence-by-sentence. Each sentence was presented for a total duration of 900 msec + (250 msec times the number of words). The interstimulus interval (ISI) between sentences was 750 msec and, after the context had been presented, an ISI between 2500 and 3500 msec occurred in which a blank screen was presented. Subsequently, the critical sentence was presented word-by-word, with 500 msec per word and a 100-msec ISI between words. The in- tertrial interval (ITI) between the critical sentence and the next discourse context was jittered between 4000 and 6000 msec. When there was a question, it was pre- sented 2000 msec after the last word of the critical Menenti et al. 2361 D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D h o t w t n p o : a / d / e m d i t f r p o m r c h . s p i l d v i e r e r c c t h . m a i r e . d c u o o m c / n j a o r c t i n c / e a - p r d t i 2 c 1 l e 1 2 - p 2 d 3 f 5 / 8 2 1 1 9 / 3 1 7 2 6 / 0 2 5 3 o 5 c 8 n / 2 1 0 7 0 5 8 9 8 2 4 1 1 4 6 / 3 j o p c d n . b y 2 0 g 0 u 8 e . s t 2 o 1 n 1 6 0 3 7 . S p e d p f e m b y b e r g u 2 0 e 2 s 3 t / j . f . / . t o n 1 8 M a y 2 0 2 1 sentence. The two possible answers were presented below the sentence 500 msec after its onset and stayed on screen for 3500 msec. At the end of this question– answer period, there was a normal ITI of 4000–6000 msec before the next context was presented. Scanning Parameters Subjects were scanned with a Siemens 3-T Tim-Trio MRI scanner, using a 12-channel surface coil. The TR was 1.86 sec, each volume contained 28 slices of 3 mm thickness with a slice gap of 10%. The voxel size was 3 (cid:1) 3 (cid:1) 3 mm3, the field of view was 224 mm. We kept the TR shorter than usual to allow for subsequent dynamic causal modeling analyses, which require a shorter TR. The number of slices, therefore, did not allow acquisi- tion of a full brain volume in most subjects. The exper- imenter always made sure that the entire temporal and frontal lobes were scanned, where the activations of in- terest were expected. However, this implied that in many subjects data from a part of the parietal lobe were not acquired. A whole-brain, high-resolution, structural T1- weighted MP-RAGE sequence was also performed to characterize the subjects’ anatomy (TR = 2.30 sec, 192 slices with voxel size of 1 mm3, FOV = 256 mm). Data Analysis The data were preprocessed using SPM5 (Friston et al., 1995). Images were realigned to correct for movement artifacts, were slice-timing corrected, and the mean func- tional image for each subject was coregistered to the EPI template provided by SPM5 and to the subjects’ ana- tomical image. Finally, images were anatomically nor- malized to the MNI space and spatially filtered with an isotropic 3-D Gaussian kernel (FWHM = 10 mm). Data were analyzed using the general linear model and sta- tistical parametric mapping (Friston et al., 1995). In a 2 (cid:1) 2 design, with the factors context (neutral/local) and world knowledge (correct/anomaly), all four regressors were modeled from the onset of the critical word to the end of the sentence and were subsequently convolved with the canonical hemodynamic response function pro- vided by SPM5. Additional regressors were included for the discourse context, the first part of the critical sen- tence, the intertrial and interstimulus intervals, and fi- nally, six movement parameters. At the single-subject level, the four conditions of interest were contrasted separately against the first part of the critical sentences of all conditions, which formed a high-level baseline. The four resulting contrast images were included in a second-level random effects analysis as well as the fac- tor subject to account for between-subject variability. For the ROI analysis small-volume correction was used for two 10-mm spheres centered around the local max- ima for the world knowledge effect reported by Hagoort et al. (2004) (MNI coordinates: left BA 47 [(cid:2)48 30 (cid:2)13] and left BA 45 [(cid:2)44 30 10]) and their right homotopic regions (see Figure 1 for the location of the ROIs). For both the ROI and the whole-brain analysis, images were thresholded at t = 3.16 ( p < .001, uncorrected). The cluster size was used as the test statistic and only clusters significant at p < .05, corrected for multiple nonindepen- dent comparisons, are reported. Local maxima are also reported for all clusters with their respective family-wise error (FWE) corrected p values. RESULTS ROI Analysis In a small-volume corrected analysis for both ROIs in the LIFG, clusters were found that showed a significant effect of world knowledge in the neutral context. Signif- icant clusters were also present in both RIFG ROIs. For the local context, no clusters were found that showed a significant world knowledge effect in three of the four regions (left BA 45, right BA 45, and right BA 47). In the left BA 47, a cluster was present showing a residual effect of world knowledge in the local context, although it was smaller than in the neutral context. We predicted that the effect of world knowledge would be weakened by the local context because the world knowledge anomaly would be easier to integrate after such a context. Clus- ters showing the predicted interaction between world knowledge and context were found in right BA 47 and BA 45, but in neither of the left ROIs (see Table 3 for all cluster coordinates and p values). For the average beta values in each ROI, the interac- tion between context and world knowledge was signif- icant in both right-hemisphere ROIs, but in neither of the left-hemisphere ROIs, although it showed a trend D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D h o t w t n p o : a / d / e m d i t f r p o m r c h . s p i l d v i e r e r c c t h . m a i r e . d c u o o m c / n j a o r c t i n c / e a - p r d t i 2 c 1 l e 1 2 - p 2 d 3 f 5 / 8 2 1 1 9 / 3 1 7 2 6 / 0 2 5 3 o 5 c 8 n / 2 1 0 7 0 5 8 9 8 2 4 1 1 4 6 / 3 j o p c d n . b y 2 0 g 0 u 8 e . s t 2 o 1 n 1 6 0 3 7 . S p e d p f e m b y b e r g u 2 0 e 2 s 3 t / j f . / . t . o n 1 8 M a y 2 0 2 1 Figure 1. ROIs in the current study. Two 10-mm spheres centered at MNI coordinates [(cid:2)44, 30, 10] (BA 45, top) and [(cid:2)48, 30, (cid:2)13] (BA 47, bottom), and their right hemisphere equivalents. 2362 Journal of Cognitive Neuroscience Volume 21, Number 12 Table 3. Results for the ROI Analyses ROI Cluster Size (kE) p (Cluster) t ( Voxel) pFWE ( Voxel) x World Knowledge Effect in Neutral Context L BA 45 L BA 47 R BA 45 R BA 47 103 300 271 307 .003 .000 .000 .000 4.300 5.330 5.630 5.800 .001 .000 .000 .000 (cid:2)48 (cid:2)46 50 44 World Knowledge Effect in Local Context L BA 47 25 .015 4.270 .000 (cid:2)48 Interaction World Knowledge (cid:1) Context R BA 47 R BA 45 1 5 8 1 .032 .026 .024 .032 3.360 3.410 3.300 3.260 .022 .019 .025 .028 48 50 50 40 y 24 34 28 38 38 36 32 30 34 z 16 (cid:2)16 16 (cid:2)12 (cid:2)16 (cid:2)4 4 12 2 Small-volume correction was performed for the four ROIs located in bilateral BA 45 and 47. Reported are all clusters significant at cluster level with p < .05, corrected for multiple comparisons. The MNI coordinates of the maxima of all clusters and their respective voxelwise t and p values (FWE corrected for multiple nonindependent comparisons) are also reported. toward an interaction in left BA 47 [left BA 45: F(1, 31) < 1; left BA 47: F(1, 31) = 3.4, p < .074; right BA 45: F(1, 31) = 6.3, p < .017; right BA 47: F(1, 31) = 9.5, p < .008]. Figure 2 shows the difference between correct sen- tences and anomalies in all ROIs, clearly showing that the world knowledge effect is more reduced in the RIFG than in the LIFG. To further specify the interaction between context and world knowledge in the RIFG, we extracted the average beta values for the clusters showing a significant inter- action in the small-volume corrected analysis in the two right-hemisphere ROIs. We inserted these in a 2 (cid:1) 2 (cid:1) 2 (Brain region (cid:1) Context (cid:1) World knowledge) repeated measures general linear model. A significant main effect Figure 2. Difference bar graphs between correct and anomalous sentences in neutral and local contexts, in four ROIs (right/left BA 45 and 47). Displayed are the differences in average beta values between correct sentences and anomalies for both types of contexts for the entire ROIs. D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D h o t w t n p o : a / d / e m d i t f r p o m r c h . s p i l d v i e r e r c c t h . m a i r e . d c u o o m c / n j a o r c t i n c / e a - p r d t i 2 c 1 l e 1 2 - p 2 d 3 f 5 / 8 2 1 1 9 / 3 1 7 2 6 / 0 2 5 3 o 5 c 8 n / 2 1 0 7 0 5 8 9 8 2 4 1 1 4 6 / 3 j o p c d n . b y 2 0 g 0 u 8 e . s t 2 o 1 n 1 6 0 3 7 . S p e d p f e m b y b e r g u 2 0 e 2 s 3 t / j / . f . . t o n 1 8 M a y 2 0 2 1 Menenti et al. 2363 was found for world knowledge [F(1, 31) = 17.86, p < .0001], which as expected was qualified by a Context (cid:1) World knowledge interaction [F(1, 31) = 14.92, p < .001], but also by an Area (cid:1) World knowledge interaction [F(1, 31) = 6.27, p < .018], with a stronger effect of world knowledge in BA 45 than in BA 47. For the Context (cid:1) World knowledge interaction, in which we were mainly interested, we tested for the simple effects of both world knowledge and context. The effect of world knowledge was significant within the neutral (F = 31.9, p < .0001) but not the local context (F < 1). The effect of context was significant for both correct sentences (F = 11.9, p < .002) and world knowledge anomalies (F = 4.3, p < .047). The effect was reversed in the latter condition: The beta values were higher for world knowledge anoma- lies in the neutral than in the local context. This suggests that the local context facilitated integration of the world knowledge anomalies. We did not find the interaction between context and world knowledge in the small-volume corrected data in LIFG. Because, however, we found a trend toward an inter- action in the mean beta values for left BA 47, we performed the same simple effects tests outlined above for the av- erage beta values of this ROI. The effect of world knowl- edge was significant for the neutral context (F = 20.770, p < .0001) and showed a trend toward significance in the local context (F = 3.572, p < .068). The effect of context was only significant for the correct sentences (F = 7.925, p < .008; anomalies: F < 1). The trend toward an in- teraction effect in left BA 47 is therefore characterized by a reduced effect of world knowledge in the local context. Figure 3 shows the differences between local and neutral contexts for the two world knowledge conditions in all ROIs, clearly showing that the context effect is reversed in the right- but not in the left-hemisphere ROIs. Whole-brain Analysis To further investigate the modulation of world knowl- edge effects by preceding discourse, we also performed a whole-brain analysis. Results for all following compari- sons are reported in Table 4. Clusters are reported if sig- nificant at p < .05 at cluster-level, corrected for multiple nonindependent comparisons. World Knowledge Effect in the Neutral Context Seven clusters showed a stronger response to world knowledge anomalies than to correct sentences. These were bilateral inferior frontal (BA 45 and 47), right mid- dle temporal (BA 21), the bilateral angular gyri (BA 39/ 40), the right caudate nucleus, and a cluster spreading across the bilateral superior frontal gyri (BA 9). All clus- ters are shown in Figure 4. In the whole-brain analysis, there was no significant effect of world knowledge in the local context. Main Effect of Context One cluster showed a marginally significant effect at the cluster level (but highly significant at the voxel level). This cluster was located in the left middle temporal gyrus and showed a stronger response to local than to Figure 3. Difference bar graphs between neutral and local contexts for correct and anomalous sentences, in four ROIs (right/left BA 45 and 47). Displayed are the differences in average beta values between neutral and local context contexts for each type of sentence. 2364 Journal of Cognitive Neuroscience Volume 21, Number 12 D o w n l o a d e d l l / / / / j t t f / i t . : / / f r o m D h o t w t n p o : a / d / e m d i t f r p o m r c h . s p i l d v i e r e r c c t h . m a i r e . d c u o o m c / n j a o r c t i n c / e a - p r d t i 2 c 1 l e 1 2 - p 2 d 3 f 5 / 8 2 1 1 9 / 3 1 7 2 6 / 0 2 5 3 o 5 c 8 n / 2 1 0 7 0 5 8 9 8 2 4 1 1 4 6 / 3 j o p c d n . b y 2 0 g 0 u 8 e . s t 2 o 1 n 1 6 0 3 7 . S p e d p f e m b y b e r g u 2 0 e 2 s 3 t / j f . t . . / o n 1 8 M a y 2 0 2 1 Table 4. Results for All Tested Contrasts in Whole-brain Analysis Effect x y z Label Cluster Size (kE) p (Clusters) t pFWE ( Voxel) World knowledge effect in neutral context 46 42 (cid:2)14 R BA 47 (cid:2)46 34 (cid:2)16 L BA 47 14 54 24 R BA 9 64 (cid:2)42 (cid:2)8 R BA 21 56 (cid:2)56 26 R BA 39/40 (cid:2)58 (cid:2)64 24 L BA 39/40 12 4 10 R caudate 2412 1502 1612 944 347 815 394 World knowledge effect in local context No significant clusters Main effect context (local > neutral)

(西德:2)60 (西德:2)22

20 L BA 21

Interaction World knowledge (西德:1) Context

(西德:2)64 (西德:2)48

28 BA 39/40

266

375

.000

.023

.000

.015

.054

.018

6.52

5.33

4.94

4.87

5.74

4.19

5.46

4.83

.000

.004

.020

.034

.026

.001

.237

.003

.030

Listed are clusters with a cluster size (kE) significant at p < .05, corrected for multiple comparisons. The local maxima of all clusters and their respective voxelwise t and p values (FWE corrected for multiple nonindependent comparisons) are also reported. neutral contexts. Simple main effects analyses revealed that for this cluster the effect was independent of whether the final sentence was correct or constituted a world knowledge anomaly (correct: F = 19.26, p < .001, anom- aly: F = 11.69, p < .002). Interaction World Knowledge and Context This comparison resulted in one significant cluster lo- cated in the left angular gyrus (see Figure 5A). Two clus- ters showed a marginally significant response. These were the right hemisphere equivalent of the cluster in the left angular gyrus as well as a cluster in the RIFG. This last result is similar to the effect observed in the ROI analysis. Comparison of the beta values in the left angular gyrus revealed a crossover interaction between context and world knowledge, suggesting that this re- gion is predominantly sensitive to the coherence be- tween critical sentence and context (see Figure 5B). DISCUSSION The first main result of this study was to replicate the world knowledge effect of Hagoort et al. (2004): Sentences with world knowledge anomalies triggered significantly greater activation of the left inferior frontal gyrus (LIFG; BA 45 and 47) compared to correct sentences, particularly for the neu- tral context. A similar effect was observed in the RIFG (BA 45 and 47; cf. Figure 2). The second main finding of this study relates to the modulation of the world knowledge effect by context. More specifically, we predicted that the effect of world knowledge would be reduced in the local context com- pared to the neutral context. Consistent with this, there was a trend toward an interaction between context and world knowledge in the LIFG (in BA 47, but not in BA 45), suggesting that this region supports on-line unification of linguistic and extralinguistic information (cf. Figure 2). The residual effect of world knowledge in the local con- text observed in the LIFG suggests that the ongoing D o w n l o a d e d l l / / / / j t t f / i t . : / / f r o m D h o t w t n p o : a / d / e m d i t f r p o m r c h . s p i l d v i e r e r c c t h . m a i r e . d c u o o m c / n j a o r c t i n c / e a - p r d t i 2 c 1 l e 1 2 - p 2 d 3 f 5 / 8 2 1 1 9 / 3 1 7 2 6 / 0 2 5 3 o 5 c 8 n / 2 1 0 7 0 5 8 9 8 2 4 1 1 4 6 / 3 j o p c d n . b y 2 0 g 0 u 8 e . s t 2 o 1 n 1 6 0 3 7 . S p e d p f e m b y b e r g u 2 0 e 2 s 3 t / j f . / . t . o n 1 8 M a y 2 0 2 1 Figure 4. Effect of world knowledge in the neutral context. The map is thresholded at p < .001, uncorrected; only clusters with p < .05, corrected at cluster level, are shown. (A) Significant clusters rendered onto the cortical surface. (B) Transversal slice at z = 10 showing the significant cluster in the caudate nucleus. Menenti et al. 2365 Figure 5. Interaction of world knowledge and context in the angular gyrus. (A) Location of the cluster showing a significant interaction. (B) Difference bar graph for the beta values in this cluster. D o w n l o a d e d l l / / / / j t t f / i t . : / / f r o m D h o t w t n p o : a / d / e m d i t f r p o m r c h . s p i l d v i e r e r c c t h . m a i r e . d c u o o m c / n j a o r c t i n c / e a - p r d t i 2 c 1 l e 1 2 - p 2 d 3 f 5 / 8 2 1 1 9 / 3 1 7 2 6 / 0 2 5 3 o 5 c 8 n / 2 1 0 7 0 5 8 9 8 2 4 1 1 4 6 / 3 j o p c d n . b y 2 0 g 0 u 8 e . s t 2 o 1 n 1 6 0 3 7 . S p e d p f e m b y b e r g u 2 0 e 2 s 3 t / j . / . t . f o n 1 8 M a y 2 0 2 1 discourse (i.e., the local context) does not fully neutralize the world knowledge effect during comprehension of the critical sentence but the trend toward an interaction be- tween context and world knowledge in the left BA 47 indicates that the context does affect processing of the world knowledge sentences in this area at least to some extent. In contrast to the LIFG, in the RIFG, the interaction between world knowledge and context was significant, consistent with our third prediction that the right hemi- sphere is involved in discourse processing. Indeed, spec- ification of the interaction in the RIFG revealed that after the local context, the BOLD response to anomalies was weaker in the local than in the neutral context. This sug- gests that the RIFG is more sensitive to the discourse modulation than the LIFG. These results are in line with several studies showing a strong involvement of the right hemisphere in dis- course processing (for a review, see Bookheimer, 2002). The RIFG, therefore, also seems to play an important role in semantic unification, in addition to the LIFG (an effect already noticed in the data of Hagoort et al., 2004, albeit nonsignificant). Moreover, the RIFG might be rela- tively more involved than the LIFG in forming an inte- grated representation of ongoing discourse. Our results are also in line with the idea that the left hemisphere is more sensitive to dominant semantic relationships, whereas the right hemisphere is more sensitive to novel, inferred semantic relationships (Giora, 2007; see Intro- duction). The LIFG continues to detect the discrepancy in the sentence, ‘‘Measles is a disease that especially affects elderly,’’ due to the strong association between measles and children, whereas the RIFG has inferred the novel association between measles and elderly from the context (There now is a vaccination against measles. The disease is more dangerous the higher the age of pa- tients is. They are more vulnerable to complications.) What the precise division of labor is between the LIFG and the RIFG remains to be explored. The fact that we did not observe a significant interaction between con- text and world knowledge in the LIFG suggests that this region remains sensitive to semantic unification of incom- ing information with prior world knowledge, even if pre- ceding discourse context overrides this knowledge. The interpretation of the world knowledge effect in the RIFG is less straightforward: The interaction between context and world knowledge and the absence of a world knowl- edge effect in the local context suggest that this area is predominantly sensitive to the congruency between dis- course context and target sentence. However, because the effect of world knowledge is not completely reversed in the local context, the region is also sensitive to infor- mation from long-term memory. The main new finding in this study is that semantic unification at the level investigated here is more bilateral than was earlier observed. Clearly, the RIFG is involved in forming a representation of the ongoing discourse. This result fits with previous claims that the right hemi- sphere is involved in processing multisentence texts. As outlined in the Introduction, the right hemisphere is sensitive to text coherence (Kuperberg et al., 2006; Caplan & Dapretto, 2001; St George et al., 1999). Long, Baynes, and Prat (2005) used divided visual field experi- ments to determine what kind of discourse representa- tion each hemisphere contributes to. They concluded that only the left hemisphere forms a propositional rep- resentation (the textbase in the description of van Dijk & Kintsch, 1983; see Introduction), evidenced by stronger priming effects for words previously presented in the same propositions than in different propositions when presented in the right visual field. The right hemisphere shows no differential priming for words in same or differ- ent propositions and, therefore, does not appear to form a propositional representation. However, because it does show stronger priming for words in same passages than in different passages, it seems to form a more general discourse representation. We suggest that this general discourse representation is a situation model. Because the target sentences in our paradigm contain distinct propositions from the preceding passages, they cause no local anomalies in the propositional structure. However, at level, they violate the content of the a more global 2366 Journal of Cognitive Neuroscience Volume 21, Number 12 preceding discourse. Our results therefore suggest that the RIFG is more sensitive to discourse anomalies than the LIFG. In addition to the ROI analysis, we also performed a whole-brain analysis to further investigate effects of world knowledge and the modulations of these effects by context. We observed a main effect of context in the left middle temporal gyrus. Note, however, that this ef- fect occurs at the critical part of the final sentence and is independent of the final sentence type which the sub- jects read. The contexts were tightly matched on word frequency, number of words, and syntactic structure, although the local contexts were, by definition, more novel than the neutral ones, which may have increased the overall integration load, irrespective of the type of sentence that had to be integrated. For the effect of world knowledge in the neutral context, we observed several significant clusters, located in the IFG bilaterally, the bilateral medial superior frontal gyri, the angular gyrus bilaterally, the middle temporal gyrus, and the right caudate nucleus. Compared to Hagoort et al. (2004), there were three modifications in the current study. First, we scanned 32 subjects instead of 14, leading to greater statistical power in our analyses. Second, we used a more high-level baseline, which may have reduced the noise in the data, thereby increasing sensitivity. Finally and crucially, our world knowledge sentences were pre- ceded by contexts. This meant that a stronger response to world knowledge anomalies than to correct sentences is very likely induced by a mismatch between the critical sentence and the preceding context. The world knowl- edge anomalies in this condition do not only violate world knowledge, but also the content of the neutral discourse context. In the local context, it is the correct sentence that violates the previous discourse. In regions where congruency between the previous context and the critical sentence is what matters, an interaction between world knowledge and context would be expected. This interaction was present in the left angular gyrus (and was present just below threshold in the right angular gyrus). These regions therefore appear to be more sensitive to the coherence between the discourse context and the critical sentence than to world knowledge per se. Different functions have been hypothesized for the angular gyrus. For example, it has been implicated in visual word form recognition (Deleon et al., 2007). Our critical words were designed to have a higher cloze probability in precisely the two conditions that show the lowest activation in the left angular gyrus. These highly constraining contexts may have led to top–down influences on word form retrieval, which would explain the interaction effect. Price (2000) argues that the angular gyrus is not specific to visual word forms but rather is engaged when semantic associations are made. Mechelli, Josephs, Lambon Ralph, McClelland, and Price (2007) lend empirical support to the suggestion that the left angular gyrus is sensitive to semantic associations between words by showing that the left angular gyrus shows a stronger BOLD response to semantically related word pairs than to unrelated and phonologically related word pairs. Humphries, Binder, Medler, and Liebenthal (2007) propose a similar but slightly more complex role of the left angular gyrus in se- mantic integration. In a study which compared congruent sentences, sentences with unrelated words and pseudo- word sentences, and reshuffled word list versions of all these, they found stronger activation in the left angular gyrus for congruent sentences than for all other condi- tions. They therefore suggested that this area is involved in integrating incoming semantic information with an on- going context. At first sight, our results do not match those of Humphries et al. (2007), as they find weaker activations in the random sentences (which would seem the hardest to integrate) than in the congruent sentence. Like us, Ni et al. (2000) also found stronger activations in the angular gyrus in sentences containing semantic anom- alies than congruent sentences. One possibility is that the random sentences in the Humphries et al. study fail to generate semantic integration in the first place, whereas sentences with an anomaly (present study, Ni et al., 2000) do generate a context in which the word meaning em- bodying the anomaly is then harder to integrate. The inferior parietal region has also been implicated in the phonological loop component of working memory (Baddeley, 2003). The phonological loop is a component of working memory that temporarily stores sound and lan- guage. It consists of two components, a phonological store and an articulatory rehearsal process. Baddeley (2003) and others have suggested that the inferior parietal region is involved in the storage component. This might explain the interaction between context and world knowledge that we observed for this region. It is possible that when subjects encounter a sentence that does not easily combine with the information that they just read, they retrieve the sur- face form and check whether their interpretation was cor- rect. However, because our experiment was not designed to distinguish between the different possibilities outlined above, we will not draw strong conclusions about the role of the left angular gyrus at this point. In conclusion, we suggest that the RIFG is specifically involved in forming a general representation (a situation model) of ongoing discourse and in unifying discourse information with previously stored knowledge in long- term memory. Based on our findings, we suggest that both the LIFG and the RIFG are recruited in unification, but with a division of labor when it comes to discourse comprehension. What precisely distinguishes the dif- ferent types of context that recruit the LIFG and the RIFG, respectively, for unification is a question for fur- ther research. 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When language meets action: The neural integration of gesture and speech. Cerebral Cortex, 17, 2322–2333. 2368 Journal of Cognitive Neuroscience Volume 21, Number 12 D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D h o t w t n p o : a / d / e m d i t f r p o m r c h . s p i l d v i e r e r c c t h . m a i r e . d c u o o m c / n j a o r c t i n c / e a - p r d t i 2 c 1 l e 1 2 - p 2 d 3 f 5 / 8 2 1 1 9 / 3 1 7 2 6 / 0 2 5 3 o 5 c 8 n / 2 1 0 7 0 5 8 9 8 2 4 1 1 4 6 / 3 j o p c d n . b y 2 0 g 0 u 8 e . s t 2 o 1 n 1 6 0 3 7 . S p e d p f e m b y b e r g u 2 0 e 2 s 3 t / j . t f . . / o n 1 8 M a y 2 0 2 1 When Elephants Fly: Differential Sensitivity of Right image

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