Intonation guides sentence processing in the left inferior
Constantijn L van der Burghta†, Tomás Gouchaa†, Angela D Friedericia, Jens Kreitewolfa,b*,
a Department of Neuropsychology, Max Planck Institute for Human Cognitive and Brain
Sciences, 04103 Leipzig, Germany
b Department of Psychology, University of Lübeck, 23562 Lübeck, Germany
†,* Authors contributed equally
Keywords: prosody, syntax, sentence comprehension, lateralisation, fMRI
Declarations of interest:
Constantijn van der Burght
Department of Neuropsychology
Max Planck Institute for Human Cognitive and Brain Sciences
04103 Leipzig, Germany
phone: +49 (0)341 9940 2238
Speech prosody, the variation in sentence melody and rhythm, plays a crucial role in
sentence comprehension. Specifically, changes in intonational pitch along a sentence can
affect our understanding of who did what to whom. To date, it remains unclear how the
brain processes this particular use of intonation and which brain regions are involved. In
particular, one central matter of debate concerns the lateralisation of intonation processing.
To study the role of intonation in sentence comprehension, we designed a functional MRI
experiment in which participants listened to spoken sentences. Critically, the interpretation
of these sentences depended on either intonational or grammatical cues. Our results
showed stronger functional activity in the left inferior frontal gyrus (IFG) when the
intonational cue was crucial for sentence comprehension compared to when it was not.
When instead a grammatical cue was crucial for sentence comprehension, we found
involvement of an overlapping region in the left IFG, as well as in a posterior temporal
region. A further analysis revealed that the lateralisation of intonation processing depends
on its role in syntactic processing: activity in the IFG was lateralised to the left hemisphere
when intonation was the only source of information to comprehend the sentence. In
contrast, activity in the IFG was right-lateralised when intonation did not contribute to
sentence comprehension. Together, these results emphasise the key role of the left IFG in
sentence comprehension, showing the importance of this region when intonation
establishes sentence structure. Furthermore, our results provide evidence for the theory
that the lateralisation of prosodic processing is modulated by its linguistic role.
In everyday conversation, different components of speech need to be integrated to
understand the meaning of what someone is saying. Besides the meaning of each individual
word in the sentence, additional information is crucial to understand who did what to whom
(Everaert, Huybregts, Chomsky, Berwick, & Bolhuis, 2015; Sportiche, Koopman, & Stabler,
2013). Besides syntax, one way in which this information can be conveyed is through
prosody: the changes in rhythm and melody of speech (Cutler, Dahan, & van Donselaar,
1997). An important feature of prosody is intonation, marked by the changes in pitch along a
sentence. Intonation can dramatically change the interpretation of a sentence. For instance,
the sentence “the teacher said the student is mistaken” has two possible interpretations
depending on the particular intonation. One can either say “the teacher said: the student is
mistaken” or “the teacher, said the student, is mistaken”. Here, the particular use of
intonation determines whether the teacher or the student is alleged to be wrong, by
creating boundaries between different parts of the sentence. The prosodic features marking
these boundaries are a pause between the two sentence parts, preceded by a rise in pitch
and a lengthening of the syllable before the pause. Together, these features acoustically
separate two parts of a sentence, and constitute a so-called intonational phrase boundary
(IPB) (Selkirk, 1984).
Despite many years of neurocognitive research on prosody, it remains largely
unknown how exactly intonation contributes to sentence comprehension, and what the
brain implementation of this process is. Although pitch was shown to be preferably
processed in the right hemisphere (Zatorre, 2001), early neurocognitive models on prosody
postulated that the stronger the linguistic function of prosody the larger the leftward
lateralisation (Friederici & Alter, 2004; Van Lancker, 1980). A recent study using intracranial
cortical recordings showed that intonation is processed in specific neural populations in the
temporal lobe that are not involved in processing other speech components such as the
sounds of words (Tang, Hamilton, & Chang, 2017). Yet, intonation is rapidly integrated with
other phonetic components (e.g. consonants and vowels) to interpret a sentence, as has
been shown in early behavioural studies (Marslen-Wilson, Tyler, Warren, Grenier, & Lee,
1992). Furthermore, electrophysiological studies have demonstrated that prosodic
information and information about sentence structure are integrated online during sentence
processing (Friederici, Cramon, & Kotz, 2007; Männel & Friederici, 2009; Sammler, Kotz,
Eckstein, Ott, & Friederici, 2010; Steinhauer, Alter, & Friederici, 1999, see Bögels et al. (2011)
for a review). To date, neuroimaging studies have been inconclusive with regard to the brain
regions involved in this use of intonation for sentence comprehension. Functional
neuroimaging research has focused on particular parts of prosody rather than its actual
contribution to sentence comprehension. For instance, a network of superior temporal and
fronto-opercular regions in the right hemisphere has been found to be involved in speech
processing depending on the presence of pitch information (Meyer, Alter, Friederici,
Lohmann, & Cramon, 2002; Meyer, Steinhauer, Alter, Friederici, & Cramon, 2004; Plante,
Creusere, & Sabin, 2002). A similar fronto-temporal network has been found in the
perception of natural compared to hummed speech (Ischebeck, Friederici, & Alter, 2008).
However, since these previous studies compared various types of filtered speech, they
focused on the acoustic processing of intonation rather than its use for sentence
comprehension. Only few neuroimaging studies have investigated which brain regions are
involved when prosodic information guides sentence comprehension. These studies either
involved a rather quantitative analysis of the intonational cue (e.g., the presence of two
intonational phrase boundaries versus a single one (Ischebeck et al., 2008)) or compared
conditions in which the stimuli were not lexically matched (Strelnikov, Vorobyev,
Chernigovskaya, & Medvedev, 2006). Consequently, several brain regions have been found
to support the processing of intonational contours in speech, but it is unknown whether
these regions also play a role in guiding sentence comprehension.
Aside from intonation, grammatical cues can guide sentence comprehension, by
means of a particular word form (morphosyntax). For example, in the sentence “The
teachers said the student is mistaken” the word form of teachers and said (both signal
plural) and student and is (both singular) establishes that the teachers describe the student’s
behaviour, not the other way around. This example shows how the sentence structure can
be established by grammatical cues. Previous work has shown that these grammatical cues
are processed in a left-hemisphere network of frontal and temporal regions (see Friederici
(2011) and Hagoort (2014) for reviews). Specifically, when grammatical cues are the only
informative elements available to interpret a sentence structure, the posterior part of the
left inferior frontal gyrus (IFG) has been shown to be engaged (Goucha & Friederici, 2015).
Functional imaging and lesion studies have further shown that successful processing of
grammatical cues relies on an intact left superior temporal cortex (Bornkessel, Zysset,
Friederici, Cramon, & Schlesewsky, 2005; Regel, Kotz, Henseler, & Friederici, 2017; Rolheiser,
Stamatakis, & Tyler, 2011).
Taken together, grammatical as well as intonational cues can guide sentence
comprehension by resolving ambiguities and establishing the structure of a sentence.
Whereas the cortical network supporting processing of grammatical cues has been
extensively studied, it remains poorly understood how exactly intonation contributes to
sentence comprehension, and what the neural correlate of this contribution is. We aimed to
fill this gap by studying processing of spoken sentences in which either intonational cues or
grammatical cues are fundamental to understand what is being said. We designed a
functional magnetic resonance imaging (fMRI) paradigm to achieve two goals. The first goal
was to investigate the contribution of intonational and grammatical cues for sentence
comprehension. The second goal was to study the hemispheric lateralisation of intonation
processing. To achieve this, participants had to comprehend specific sentence types (see
Figure 1). Across conditions, the sentence structure was established by different language
cues: sentences could be interpreted in two possible ways until a point at which the cue
ensured only one possible interpretation (Marslen-Wilson et al., 1992). This cue was either
intonational (an IPB) or grammatical. The grammatical cue was established by morphological
case marking of a personal pronoun, such that it matched only one of the two verbs in the
sentence. The paradigm centred around the following sentence, which is open to two
Peter verspricht Nick dafür zu bezahlen
(i) Peter promises Nick to pay for it
(ii) Peter promises to pay Nick for it
In these sentences, intonational and/or grammatical cues are required to convey who did
what to whom. Without them, listeners cannot identify whether Nick was promised
something or paid for something instead. In our key conditions, the position of an
intonational phrase boundary (marked with #) helped the listener to identify one of the two
(A) Peter verspricht Nick # dafür zu bezahlen
Peter promises Nick to pay for it
(B) Peter verspricht # Nick dafür zu bezahlen
Peter promises to pay Nick for it
Alternatively, the presence of a grammatical cue established a clear interpretation. Here, we
made use of German case marking, which distinguishes between the different roles a word
can have in a sentence. Depending on the word form and its corresponding case (dative or
accusative), the personal pronoun can only be the object of the verb to promise or to pay.
(C) Peter verspricht sie dafür zu bezahlen
Peter promises to pay+ACC herACC for it
As a second goal of our study, we investigated the hemispheric lateralisation of
intonation processing. Although a general consensus exists that processing of linguistic
components such as grammatical cues is predominantly left-lateralised (Friederici, 2011;
Vigneau et al., 2006), it has long been debated whether a similar lateralisation exists for the
processing of intonation (Luks, Nusbaum, & Levy, 1998; van Lancker, 1980; Wildgruber et al.,
2004). In general, the right hemisphere is seen as dominant in the processing of pitch
information, including intonation (Poeppel, 2003). However, dichotic listening (Luks et al.,
1998) and lesion studies (reviewed in Witteman et al. (2011)) have shown that lateralisation
depends on the linguistic function of intonation. Moreover, results from functional
neuroimaging studies suggest that intonation processing in the IFG and temporal cortex is
lateralised, but the contribution of either hemisphere depends on the specific control task
used (Kreitewolf, Friederici, & Kriegstein, 2014). It is likely that processing of linguistic
prosody relies on fronto-temporal networks in both hemispheres (Belyk & Brown, 2014;
Witteman et al., 2011), with a dominance of the left hemisphere when pitch information is
used to signal linguistic aspects (Friederici & Alter, 2004; van Lancker, 1980). However, it
remains elusive whether intonation processing is lateralised when it contributes to sentence
comprehension, and to which hemisphere. We addressed this question by investigating the
neural processing of intonation, focusing specifically on its use for sentence comprehension.
Our paradigm allowed for an investigation of the linguistic importance of prosody, since we
varied the linguistic role of the intonational cue across conditions while keeping acoustical
information identical. To this end, we included a condition in which the IPB was present but
not essential to establish the sentence structure (see Figure 1).
In summary, our study was designed to answer two separate questions. First, we
investigated how the presence of intonational and grammatical cues influences sentence
processing. We hypothesised that processing depends on the availability of the specific cue
type in the sentence, and whether this cue appeared in isolation or in combination with a
second cue. Based on previous studies (Goucha & Friederici, 2015; Kreitewolf et al., 2014),
we expected increased activity in the left IFG (and possibly additional bilateral superior
temporal regions) in conditions in which intonation was the only cue establishing the
structure. When sentence comprehension relied on a grammatical cue only, we also
expected increased activity in the left IFG, possibly with additional recruitment of the left
posterior temporal cortex (Bornkessel et al., 2005; Regel et al., 2017). Our second research
question concerned the lateralisation of intonation processing. Specifically, we investigated
whether the role of the intonational cue in establishing the sentence structure determined
the lateralisation of brain areas involved in intonation processing. Based on previous work
(e.g Kreitewolf et al., 2014), we hypothesised that lateralisation depends on the linguistic
function of the intonational cue. For processing of intonational cues that establish sentence
structure, we expected left-lateralised activity of core language regions (IFG and posterior
superior temporal gyrus (pSTG)). In contrast, we expected a shift towards the right IFG when
prosodic content was present, but not used to establish the structure of the sentence.
Twenty-six native German speakers (15 female; mean age: 26.3 years; age range: 20-33
years) were included in the final analyses. All participants were right-handed (Oldfield, 1971)
and had normal or corrected-to-normal vision. All reported normal hearing and none were
professional musicians. None had a history of neurological or psychiatric illness, drug or
alcohol abuse, chronic medical disease, or any other contraindication against participation in
an MRI experiment. Twelve additional participants had to be excluded because they did not
complete the experiment (n = 2) or because they performed below chance level in at least
one of the six stimulus conditions (n = 10). The sample size was determined based on
previous fMRI studies on sentence processing (e.g. Goucha & Friederici, 2015; Kristensen,
Wang, Petersson, & Hagoort, 2013; Perrone-Bertolotti, Dohen, Lœvenbruck, Sato, Pichat &
Baciu 2013). The exclusion criteria were established prior to data analysis. All participants
gave written consent prior to participating in the experiment, which was approved by the
ethics committee of the University of Leipzig.
2.2. Experimental design
To investigate the effect of intonational and grammatical cues on sentence comprehension,
we used an event-related fMRI design that employed six different sentence types with
varying amount of intonational and grammatical information (see Figure 1A).
The stimulus set was built around the following sentence:
A verspricht B dafür zu bezahlen (literally: A promises B for it to pay)
which has two possible interpretations.
i. [A verspricht B [dafür zu bezahlen]]
[A promises B [to pay for it]]
ii. [A verspricht [B dafür zu bezahlen]]
[A promises [to pay B for it]
In German, the two structures (i) and (ii) can be realised by an identical string of words. This
requires specific language cues to distinguish the two possible meanings and to clarify
whether B is the object of the verb to promise or to pay. One such cue is the position of an
IPB (indicated with “#”), which can create the distinction as follows:
Prosody Only 1 (1) [Peter verspricht Nick # [dafür zu bezahlen]]
(ProsOnly1) [Peter promises Nick # [to pay for it]]
Prosody Only 2 (2) [Peter verspricht # [Nick dafür zu bezahlen]]
(ProsOnly2) [Peter promises # [to pay Nick for it]]
The IPB acoustically divides the sentence and groups the proper noun Nick to either of the
two verbs. Without the IPB, ProsOnly1 and ProsOnly2 are ambiguous. The IPB is defined by a
pitch rise and syllable lengthening, followed by a pause (Selkirk, 1984) (see Figure 1B).
An additional cue can resolve the ambiguity, for example, when a personal pronoun
is used (such as she) instead of a proper noun (Nick). In German, personal pronouns are
inflected, meaning that their morphosyntactic form defines their role in the sentence (i.e.,
by case marking). Similarly, verbs require objects in a specific case. For example,
“versprechen” (to promise) requires objects in the dative case, whereas “bezahlen” (to pay)
requires an accusative. Making use of the German case marking system, we constructed
sentences in which the structure is built by a grammatical cue only:
Grammatical Only (3) [Peter verspricht [sie dafür zu bezahlen]]
(GramOnly) [Peter promises+DAT [to pay+ACC herACC for it]]
Although the position of the word “sie” (her) does not yet clarify to what verb the word
belongs, the case marking of the word ensures that “sie” is necessarily an object of to pay
and cannot belong to the verb to promise. The sentence can only be interpreted in one way
because of the morphosyntactic form of herACC (“sie”).
To investigate sentence processing guided by these cues, we designed control
conditions with additional cues, for example:
Baseline Prosody Only (4) [Peter verspricht Nick # [sie zu bezahlen]]
(BL ProsOnly) [Peter promises+DAT Nick # [to pay+ACC herACC for it]]
In this sentence, identification of the syntactic structure is facilitated by the additional
grammatical cue “sie”, as compared to (ProsOnly1). Thus, it is not necessary to disambiguate
the verb-argument structure because two objects (Nick and her) are present in this
Similarly, as a control condition for the experimental condition GramOnly, we
created sentences that contained an intonational cue in addition to the grammatical cue, for
Baseline Grammatical Only (5) [Peter verspricht # [sie dafür zu bezahlen]]
(BL GramOnly) [Peter promises+DAT # [to pay+ACC herACC for it]]
A final baseline condition was created, in which intonation was not required to
understand who did what to whom the in the sentence. This sentence type had an IPB, as in
ProsOnly1 and ProsOnly2, but the verb-argument structure did not have to be resolved.
Baseline Prosody No Choice (6) [Peter verspricht # [heute dafür zu bezahlen]]
(BL ProsNoChoice) [Peter promises+DAT # [to pay+ACC for it today]]
Each condition consisted of 50 unique verb combinations matched to a variety of
German first names (yielding a total of 300 sentences). In each sentence, the verb in the
main clause required an object in the dative form and the verb in the subordinate clause
required an object in the accusative form, or vice versa. This ensured that the grammatical
cue (the personal pronoun), in either the dative or accusative form, could be unambiguously
assigned to either verb. The matching of dative and accusative verbs to main clause or
subordinate clause was pseudo-randomised across trials.
To confirm that participants were equally likely to attribute the object in the
sentence to the first or the second verb, we calculated if participants had a response bias for
either condition ProsOnly1 or ProsOnly2. Using methods of signal detection theory
(Macmillan & Creelman, 1991) this response bias turned out not to be significant from 0,
suggesting that participants had no intrinsic bias for either syntactic structure (see Meyer et
al., 2016, for a similar approach).
Figure 1. Experimental design. (A) Experimental conditions. Across conditions, a different
combination of language cues established the verb-argument structure; that is, whether the
object in the sentence (“Nick” or “she”) belongs to the first or second verb. Prosodic cue
(indicated by “#”): an acoustic break marking a transition in a sentence (i.e., intonational
phrase boundary; IPB). Grammatical cue (in bold typeface): morphosyntactic case marking of
the personal pronoun “sie” (she), matched to either the verb in the main clause (“verspricht”
(promises)) or sub clause (“zu bezahlen” (to pay)). ‘Prosody’: presence of a prosodic cue.
‘Grammatical’: presence of a grammatical cue. ‘Only’ indicates that that cue was the only
cue present in that sentence. ‘BL’: baseline condition matched to the specific cue.
‘NoChoice’: the prosodic cue did not influence the response choice in the task. (B)
Spectrogram of an example stimulus with intonational and grammatical cues. Here, the
syntactic structure is established by both the grammatical cue “sie” (she) and an intonational
cue in shape of an IPB. The IPB is composed of three acoustic events, indicated with arrows:
(i) a pitch rise and (ii) lengthening of the syllable, followed by (iii) a pause. (C) Overview of an
experimental trial. (D) Overview of the fMRI session.
2.3. Experimental procedure
Each experimental trial (see Figure 1C) started with a white fixation cross, which
turned red 200 ms prior to auditory stimulation to alert the participant. Subsequently, the
spoken sentence was presented. After each sentence, a visual comprehension question was
shown for 1500 ms, to which participants were asked to respond with a right-handed button
press. Participants had to respond within 4 s. The subsequent trial started after the response
phase and an additional delay of 0, 750 or 1500 ms (uniformly jittered).
To ensure active listening, the comprehension question was visually presented after
the sentence. The question probed sentence comprehension by asking whether an object
was present in either the main or subordinate clause: participants were asked Do you know
who Peter promises something? (“Weiß man, wem Peter etwas verspricht?”) or Do you
know, who is paid? (“Weiß man, wer bezahlt wird?”). The question types were presented
pseudo-randomly, with each of the two question types occurring evenly across sentence
During the experimental session (Figure 1D), the six experimental conditions were
presented in a pseudo-random order: two stimuli of the same condition were always
separated by at least two trials so that the conditions were distributed evenly throughout
the experiment. 50 null trials with an average-trial duration of 8.5 s were pseudo-randomly
interspersed with the other conditions throughout the experiment. The experiment was
performed within one session with a total duration of 52 minutes. Each participant
performed a short practice session immediately before the fMRI experiment, which mirrored
the main experiment but consisted of different stimuli.
Auditory stimuli were presented through MR-compatible headphones (MR confon
OPTIME 1; MR confon GmbH, Magdeburg, Germany). Participants additionally wore earplugs
to attenuate scanner noise. Stimulus presentation and response collection were controlled
via Presentation (Neurobehavioural Systems, Inc., Albany, CA, USA), with visual stimuli
presented on an LCD projector (PLC-XP50L, SANYO, Tokyo, Japan). Participants could see the
projection via a mirror that was attached to the head coil.
2.4. Stimulus properties
Sentences were spoken by a male, professional native German speaker and recorded
in a sound-attenuating chamber (IAC – I200 series, Winchester, United Kingdom). The
digitised speech signals (sampling rate 44.1kHz; resolution 16 bits) were adjusted to the
same root mean square value using MATLAB (Mathworks, Inc., Sherborn, MA, USA). To
ensure consistent comparisons between stimuli with matching main or subordinate clauses,
all stimuli were cross-spliced: the clauses before and after the pause were cut in Adobe®
AuditionTM CS5.5 and concatenated to form the stimulus sentences. This procedure ensured
that identical sentence parts across conditions originated from the same recording.
Importantly, in this way we guaranteed that in contrasts between two sentences with
IPBs, those IPBs were acoustically identical. That is, for ProsOnly1 & BL_ProsOnly, and for
ProsOnly2, BL_GramOnly & BL_ProsNoChoice, the onsets of the stimuli up to and including
the IPB originated from the same recordings.
To further improve acoustic consistency across the stimulus set, in those sentences
that contained an IPB (all except GramOnly) we introduced a pause of constant duration
(100 ms): all first parts of the stimuli were cut until the pause, to which a pause of constant
duration was added, followed finally by the second part of the stimuli (which had been cut
after the pause). We chose 100 ms based on pilot study results, showing that such a pause
could be clearly perceived and sounded natural.
The GramOnly condition, containing no IPB, was also constructed by cross-splicing
two elements. To prevent the realisation of an IPB in the first element, we had the speaker
produce a sentence without a syntactic boundary after the verb (where an IPB would be
illegal). Subsequently, we spliced the recording after the verb and concatenated it to the
same sentence ending as in BL_GramOnly. This yielded a sentence with natural prosody but
without any of the three acoustic cues characterising the IPB (pitch rise, syllable lengthening
and a pause). Furthermore, this ensured that the sentence endings of GramOnly and its
baseline equivalent were matched. Spectrograms of all 6 sentence conditions are provided
in Supplementary Figure 1.
Participant debriefings and a pilot study on a separate sample of participants (n = 18)
confirmed that all stimuli were perceived as natural, grammatical, and non-ambiguous.
2.5. fMRI acquisition
Functional imaging was performed on a 3 Tesla Siemens Skyra scanner (Siemens Healthcare,
Erlangen, Germany) using a 20-channel head coil. A gradient-echo echo-planar-imaging (EPI)
sequence was run (acquisition time [TA] = 2s; continuous scanning; echo delay time [TE] =
30ms; flip angle 78°; matrix size 64 × 64; field of view [FOV] = 192 × 192mm; 30 slices of
3mm thickness; in-plane resolution = 3mm x 3mm; gap = 1mm). For anatomical registration,
T1-weighted images were either acquired during the scanning session or obtained from the
in-house database when available.
2.6. Data analysis
2.6.1. Behavioural data
Response times and accuracy data were analysed using SPSS (IBM Corp., Armonk, NY, USA).
Since behavioural measures were not normally distributed (Kolmogorov-Smirnov tests; all p <
0.05), Friedman tests were used as a non-parametric alternative to repeated-measures
analyses of variance. Follow-up Wilcoxon signed-rank tests were performed as post-hoc tests.
Initial p-values lower than α = 0.05 (two-tailed) were considered significant for all
comparisons. To correct for multiple comparisons (a total number of 15), Bonferroni
corrections were applied, yielding a corrected α-level of 0.0033 (0.05/15).
2.6.2. fMRI data
fMRI data were pre-processed and statistically analysed using SPM12
(www.fil.ion.ucl.ac.uk/spm, Wellcome Trust Centre for Neuroimaging). All functional images
were realigned to the first image in the time series to correct head motion and unwarped to
correct distortions caused by inhomogeneity in the magnetic field. After the T1-weighted
image was co-registered to the mean EPI image, it was normalised to the Montreal
Neurological Institute (MNI) template image. The deformation parameters resulting from
this step were used to normalise all EPI images to MNI space. Finally, the data were
smoothed using an isotropic Gaussian kernel of 8mm full-width at half-maximum.
Statistical analysis of the fMRI data was performed using a general linear model in
SPM12. The onset and duration of each sentence were modelled per condition and
convolved with the canonical hemodynamic response function. To account for domain-
general effects of task performance on brain activation, we took into consideration
between-condition differences in reaction times in our model. To this end, we built a
regressor with response onsets and response times for each trial. This regressor was
orthogonalized to the condition regressors and included in the general linear model
(following Grinband, Wager, Lindquist, Ferrera, & Hirsch, 2008). Incorrect trials were
modelled as a separate noise condition. A high-pass filter with 128s cut-off was applied.
Contrast images of each condition of interest and participant were combined in a group
random effects analysis with a factorial design: the six experimental conditions entered as
levels of the factor CONDITION. Results were thresholded at an FWE-corrected cluster level
of p < 0.05, using an initial uncorrected voxel-wise threshold of p < 0.001 (Friston, Worsley,
Frackowiak, Mazziotta, & Evans, 1994). All activation peak coordinates are reported in MNI
space and the SPM anatomy toolbox (version 2.2c) (Eickhoff et al., 2005) was used for
anatomical localisation. Results were visualised using the BrainNet Viewer (Xia, Wang, & He,
Additionally, we performed a lateralisation analysis. This analysis was conducted by
normalizing the raw EPI images to a symmetrical MNI template. The first-level analysis was
run as described above, and the resulting contrast images were left-right flipped (Bozic,
Tyler, Ives, Randall, & Marslen-Wilson, 2010; Josse, Kherif, Flandin, Seghier, & Price, 2009;
Liégeois et al., 2002). On the second level, paired t-tests were run to compare the image of a
particular contrast of interest to its left-right flipped equivalent. We applied the same
statistical thresholds that were used in the activation analysis.
2.7. Availability of study materials
Data from this study have not been publicly archived since the conditions of our ethics
approval do not permit to do so. Analysis code and stimulus materials are available at
https://github.com/CLvanderBurght/prossyn/. No part of the study procedures or analyses
was pre-registered prior to the research being conducted.
Both reaction times (RTs; Figure 2A) and accuracy rates (Figure 2B) differed across
conditions, as shown by Friedman tests (RTs: χ2(5) = 75.87, p < 0.001; accuracy: χ2(5) = 50.41,
p < 0.001). Pair-wise comparisons between conditions showed differences in the difficulty of
sentence comprehension reflected in accuracy and reaction times. In general, participants
showed lower accuracies and higher reaction times in conditions where only one cue was
available compared to the matched baseline conditions. Accuracy decreased and reaction
times increased in ProsOnly1 (1) as compared to BL_ProsOnly (4), indicating that sentence
comprehension was more difficult when only one language cue was present (RTs: Z = -4.457,
p < 0.001; accuracy: Z = -3.523, p < 0.001). Similarly, sentence comprehension was more
difficult in GramOnly (3), which contained only a grammatical cue, compared to
BL_GramOnly (5), which contained both a grammatical and an intonational cue (RTs: Z = -
4.026, p < 0.001; accuracy: Z = -3.760, p < 0.001). Other comparisons between matched
sentences were significant (e.g. conditions 2 vs 6: RTs: Z = -4.178, p < 0.001; accuracy: Z = -
3.816, p < 0.001; see supplementary tables S1 and S2 for complete pair-wise comparisons).
Comparisons between matched conditions of interest (1 versus 2) or control conditions (4
versus 6) were not significant.
Figure 2: Behavioural results: reaction times (A) and accuracy rates (B) per condition. Error
bars indicate ± 1 SEM. Lines between conditions and asterisks indicate pair-wise
Mean Reaction times (s)
Mean accuracy (% correct)
comparisons with p-values smaller than 0.0033 (Bonferroni-corrected). BL stands for
3.2. fMRI – activity analysis
We investigated how the presence of specific language cues shapes sentence
comprehension in the brain. To this end, we contrasted sentences in which only one specific
language cue established the sentence structure to matched control conditions (i.e., 1 vs 4
and 3 vs 5, cf. Figure 1A). In the control sentences, processing of the sentence structure was
facilitated in comparison to the conditions of interest, because multiple cues instead of a
single cue marked the sentence structure. For an overview of significant activation clusters,
see Table 1.
Syntactic structure established by a prosodic cue
To investigate processing of sentence structure established by prosodic information, the
following experimental conditions were contrasted: ProsOnly1 (1), in which a prosodic cue is
the only factor establishing the sentence structure, versus BL_ProsOnly (4), in which
additional cues determined the sentence structure (a grammatical cue). This contrast,
reflecting sentence processing guided by a prosodic cue, yielded increased task-related
activity in the left inferior frontal gyrus (IFG, peak activity at pars triangularis, x, y, z = -45, 29,
-4; T = 4.93; Figure 3A and Table 1).
Syntactic structure established by a grammatical cue
To investigate processing of sentence structure marked by a grammatical cue, we contrasted
GramOnly (3), in which only a grammatical cue marks the sentence structure, to a matched
control condition in which an additional intonational cue establishes the sentence structure.
Since in BL_GramOnly (5) the sentence structure was already established by the intonational
cue, the grammatical cue was less important for resolving the structure. This contrast
resulted in functional activation clusters in the pars opercularis of the left IFG (x, y, z = -51,
11, 8; T = 4.43) and the left superior temporal gyrus and sulcus (x, y, z = -57, -16, 2; T = 5.23;
The reverse of the above described contrasts (3 > 1 and 4 > 5) did not yield significant
activation clusters. Additionally, other contrasts between matched sentences (ProsOnly2 >
BL_GramOnly and ProsOnly2 > BL_ProsNoChoice) did not show significant results at p < 0.05,
Functional MRI results showing significant activation clusters for the different contrasts of
interest. (A) Syntactic structure processing guided by a prosodic cue. (B) Syntactic structure
processing guided by a grammatical cue. Bar plots show contrast estimates for each
condition at activation peaks, indicated by the pointer, in arbitrary units (a.u.). All
comparisons are thresholded on the cluster level at p < 0.05, FWE-corrected. Peak activity
coordinates are in MNI space. BL stands for baseline.
3.3. fMRI – lateralisation analysis
A right-hemispheric dominance for intonation processing is often found in prosody research
(Meyer et al., 2002; Sammler, Grosbras, Anwander, Bestelmeyer, & Belin, 2015). However,
meta-analyses on prosody studies point towards a bilateral network for prosody processing
(Belyk & Brown, 2014; Witteman et al., 2011). It has previously been suggested that
intonation processing is left-lateralised specifically when pitch information is linguistically
relevant (Friederici & Alter, 2004; Kreitewolf et al., 2014; van Lancker, 1980). However, this
has not yet been shown with well-matched sentence stimuli. We therefore investigated
sentence conditions in which the intonation was matched acoustically but differed in terms
of linguistic importance. Specifically, we compared conditions in which intonation guided
sentence comprehension with matched conditions in which intonation was superfluous for
sentence comprehension. The resulting contrast images were compared to their equivalent
images in right-left flipped orientation. Results are summarised in Table 2.
First, we investigated lateralisation of intonation processing when the prosodic cue
was crucial for sentence comprehension, assessed by the contrast ProsOnly1 vs BL_ProsOnly
(same contrast as in the activity analysis in Figure 3A). The results showed that the
functional activation in the IFG was left-lateralised (x, y, z = -54, 29, 5, pars triangularis; T =
6.08; Figure 4A). Other areas that showed left-lateralised activity were the supplementary
motor area (x, y, z = -6, 23, 50; T = 4.87) and the superior temporal gyrus (x, y, z = -51, -34,
2; T = 4.20). Additionally, functional activation was right-lateralised in the pre- and post-
central gyrus (x, y, z = 30, -19, 56; T = 5.53) and in the superior temporal gyrus (x, y, z = 54, -
4, 8; T= 5.20).
In a second contrast, we isolated prosodic processing when the prosodic cue was
superfluous for the sentence structure (BL_ProsNoChoice vs ProsOnly2). In the condition
BL_ProsNoChoice, the task did not require processing of the intonational cue to
disambiguate the sentence structure, whereas condition ProsOnly2 was a matching sentence
in which the IPB was necessary for building the sentence structure. The processing of a
superfluous intonational cue showed an overall pattern of right-lateralised activity.
Functional activation of the inferior frontal gyrus was right-lateralised, with peak activations
in the pars opercularis (x, y, z = 51, 20, 8; T = 5.19) and pars triangularis (x, y, z = 48, 44, 8; T =
5.45) (Figure 4B). Additional right-lateralised activations were found in the superior temporal
sulcus and gyrus (x, y, z = 57, -37, 8; T = 4.34), the supplementary motor area (x, y, z = 9, 23,
50; T = 7.24), and the precuneus (x, y, z = 6, -55, 41; T = 6.01). Activity in the pre/post-central
gyrus (x, y, z = -39, -16, 50; T = 4.94) was stronger in the left than right hemisphere.
Lateralisation analysis showing functional contrasts of interest compared to their left-right
flipped equivalent. (A) Lateralised functional activity evoked by processing of sentence
structure guided by a prosodic cue. (B) Lateralised functional activity evoked by processing a
sentence structure in which the prosodic cue is superfluous. All comparisons are thresholded
on the cluster level at p < 0.05, FWE-corrected. BL stands for baseline.
Table 1 Task-related activity for the comparisons of interest thresholded on the cluster level
at p < 0.05, FWE-corrected.
MNI coordinates (x y z; in mm)
Prosodic cue establishes sentence structure (ProsOnly1 (1) > BL_ProsOnly (4))
Grammatical cue establishes sentence structure (GramOnly (3) > BL_GramOnly (5))
Table 2 Task-related activity for the comparisons of interest in the lateralisation analysis. All
results were thresholded on the cluster level at p < 0.05, FWE-corrected.
MNI coordinates (x y z; in mm)
Prosodic cue establishes sentence structure (ProsOnly1 (1) vs BL_ProsOnly (4))
Inferior frontal gyrus
Prosodic cue superfluous for sentence structure (BL_ProsNoChoice (6) vs ProsOnly2 (2))
Inferior frontal gyrus,
Inferior frontal gyrus,
In this study we show that the left inferior frontal gyrus has a key role in processing prosodic
information that is used for sentence comprehension. By comparing the role of intonational
and grammatical cues in sentence processing, we provide novel evidence that intonation is
processed in the left hemisphere when its function is syntactic.
Our first aim was to investigate whether different types of language cue available to
understand the structure of a sentence determined the recruitment of different brain areas.
As a main finding, we show that the left IFG is involved in sentence processing both when
intonational and grammatical cues establish the sentence structure. As a second finding, we
show that lateralisation of activity depends on whether or not intonation is decisive for the
interpretation of the sentence structure. When intonation was the only cue establishing the
sentence structure, activity in the IFG was left-lateralised. Conversely, activity in the IFG was
lateralised to the right hemisphere when the intonational cue was superfluous for sentence
comprehension, even though the cues were acoustically identical.
Our results substantially extend previous neuroimaging work that emphasised the
importance of the left IFG for sentence comprehension (Bornkessel-Schlesewsky &
Schlesewsky, 2013; Friederici, 2011; Hagoort, 2014). We show that the left IFG plays a major
role when an intonational cue is used to build sentence structure. Previous neuroimaging
studies on prosody processing reported the left IFG as part of a wider network of bilateral
fronto-temporal regions (reviewed in Belyk & Brown (2014)). In contrast, in this study we
find the left IFG in isolation. This difference from previous work is likely due to two aspects
of prosody which were tackled in the present study but have been largely ignored in the
literature to date. First, many previous neuroimaging studies have focused on the acoustic
aspect of prosody processing rather than its role in guiding sentence comprehension. A
predominantly right-hemispheric temporal network has often been identified in prosody
experiments drawing comparisons between normal speech and acoustically manipulated
speech, such as speech with flattened pitch (Meyer et al., 2004) or filtered speech in which
only the pitch contour remained (Hesling, Clément, Bordessoules, & Allard, 2005; Meyer et
al., 2002; 2004). The temporal areas found in these studies are likely to reflect processing of
acoustic properties of linguistic prosody, and in particular of pitch. In contrast to these
previous studies, we presented an intonational cue that was acoustically identical in our
condition of interest (in which the cue was used for sentence comprehension; ProsOnly1)
and in its matched control condition. Our finding of activity in the left IFG without additional
activity in auditory regions can be explained by the acoustic similarity of these two
conditions. As another novel aspect, our study investigated how intonation is used to guide
the interpretation of the sentence. Notably, linguistic prosody can come in various forms
(Cutler et al., 1997), of which marking of a syntactic boundary by an intonational phrase
boundary (IPB) is arguably the most important for sentence structure. The importance of
prosodic information in syntactic phrasing has been demonstrated in electrophysiological
studies (Friederici et al., 2007; Steinhauer et al., 1999). Previous fMRI studies, however, have
not studied this use of prosody. Rather, fMRI research has focused on types of linguistic
prosody which are not as crucial for the syntactic structure of a sentence, such as marking a
question or statement (Kreitewolf et al., 2014; Sammler et al., 2015) or placing stress
(Kristensen et al., 2013; Perrone-Bertolotti et al., 2013). Moreover, studies often used a low-
level or non-linguistic baseline condition rather than a comparable linguistic control task
(Meyer et al., 2002; 2004; Plante et al., 2002). Right fronto-temporal areas have been found
in question/statement versus phoneme discrimination tasks (Kreitewolf et al., 2014;
Sammler et al., 2015), with functional activity switching to the left hemisphere when
contrasted against a non-linguistic task of speaker identification (Kreitewolf et al., 2014). In
turn, processing of pitch focus was shown to involve bilateral frontal and superior temporal
regions (Kristensen et al., 2013; Perrone-Bertolotti et al., 2013), rather than the isolated
recruitment of the left IFG found here. Critically, in the aforementioned studies, analysis of
pitch differences was required to deduce linguistic meaning from the speech signal (i.e., by
distinguishing question/statements or establishing constituent focus), but it was not used for
the interpretation of the syntactic structure. In sum, we argue that the type of linguistic
prosody in the current study (i.e., the use of the IPB) forms a more direct link to sentence
structure processing than the previous studies, thus isolating functional activation in the left
Our results show that the left IFG was also involved when a grammatical cue (i.e.,
word form) guided sentence comprehension. In the grammatical cue condition (GramOnly),
the sentence structure could only be resolved by matching the case of the personal pronoun
(morphosyntactic information) to either one of the verbs in the sentence. Although different
from the intonational cue, both cues had the syntactic function of unambiguously attributing
the object in the sentence to one of two verbs. Case marking ensured that the personal
pronoun could only match one of two verbs, similar to how the position of the IPB
established a single possible interpretation of the sentence. We found that the left IFG was
engaged in the processing of both cue types, which indicates that this area responds to
different kinds of cues resolving ambiguity in sentence structure. This finding points to a
more general involvement of the left IFG in the processing of sentence structure. Activity in
the left STG/STS, on the other hand, was only present when sentence structure was built by
a grammatical cue. This is not surprising given that superior temporal regions have been
associated with morphosyntactic processing in lesions studies (Dronkers, Wilkins, Van Valin,
Redfern, & Jaeger, 2004) and fMRI studies on assigning subject and object to a verb
(Bornkessel et al., 2005). Additionally, the posterior STG has been shown to play a critical
role in the production of the correct morphosyntactic form (D. K. Lee et al., 2018).
To isolate the processing of each type of linguistic cue, we compared an experimental
condition in which only one cue (either intonational or grammatical) was available to resolve
sentence structure ambiguity, to a control condition in which both cues were available.
These comparisons did not only reveal differences in the neural activity but also on the
behavioural level: participants responded faster and more accurately in the control
conditions compared to the experimental conditions. Consequently, one could argue that
the increased activity in the left IFG reflects differences in task difficulty rather than
differences in linguistic processing per se. However, our general linear model included a
regressor which modelled all trial-by-trial responses as a boxcar function with trial-by-trial
reaction times as duration (Grinband et al., 2008). Since one regressor was built modelling
reaction times across all conditions, this regressor should account for variance introduced by
between-condition differences in reaction times and should therefore regress out domain
general effects. Moreover, increased cognitive demand usually relies on a domain-general
network that in the frontal lobe excludes most of the IFG (Fedorenko, Duncan, & Kanwisher,
2013) and rather includes premotor regions, the anterior cingulate cortex, and the middle
frontal gyrus (Duncan, 2010). Taken together, we did not find support for the alternative
explanation that the observed activity in left IFG was due to differences in task difficulty.
With respect to the lateralisation of intonation processing, we set out to advance a
debate that has been held for decades. Although early studies indicated a right-hemispheric
advantage for emotional prosody and left-hemispheric dominance for linguistic prosody
(Heilman, Bowers, Speedie, & Coslett, 1984; Luks et al., 1998), recent meta-analyses suggest
involvement of a more bilateral network (Belyk & Brown, 2014; Witteman et al., 2011). Our
paradigm allowed us to assess the lateralisation of prosody processing in function of its
linguistic importance since we varied the linguistic role of the intonational cue across
conditions while keeping acoustical information identical across conditions. We found that
processing intonation was left-lateralised in the IFG when it guided sentence
comprehension. In contrast, when intonation was superfluous for disambiguation of the
sentence structure, activation in the IFG was shifted to the right hemisphere. The latter
finding can be explained by our manipulation in which the IPB was not relevant for
disambiguating the sentence structure. This resulted in a relative dominance of the right IFG
when prosody was processed without being used to establish the sentence structure. This
interpretation is consistent with previous studies (Kreitewolf et al., 2014; Sammler et al.,
2015), demonstrating right IFG involvement when intonational contours were processed
without requiring integration into a sentence structure. The observed right-hemispheric
lateralisation of the pSTS in our study further converges with previous work (Meyer et al.,
2002; 2004; Sammler et al., 2015) and with models describing a right-hemispheric
dominance of auditory regions in processing pitch information in speech, such as
intonational contours (Poeppel, 2003). Together, the previous and present results suggest
that the right pSTS is preferentially involved in processing of intonational contours as such,
but not in the subsequent integration of this information during sentence comprehension.
In summary, our results provide evidence for a key role of the left IFG in sentence processing
when only intonation conveys the structure of the sentence. Activity in this region
overlapped with the region that was active when the sentence structure was established by
a grammatical cue, i.e. word form. This finding extends previous work on the contribution of
the left IFG in sentence comprehension, highlighting the role of this region in the integration
of prosodic as well as grammatical cues into the sentence structure. Moreover, we found
that lateralisation of intonation processing depends on whether or not intonation is critical
for understanding a sentence structure. This supports the notion that processing of prosodic
information is lateralised in function of its linguistic role (Friederici & Alter, 2004; Kreitewolf
et al., 2014; Luks et al., 1998; van Lancker, 1980), showing this distinction for the first time in
an fMRI study using sentence-level intonation in natural speech.
The authors would like to thank Mandy Jochemko, Anke Kummer, and Simone Wipper for
MRI data acquisition and Kerstin Flake, Stephan Liebig, and Servaas van der Burght for help
with the figure design. This work was funded by the Max Planck Society.
Conflict of interest
The authors declare that there are no conflicts of interest.
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Supplementary materials Van der Burght et al.
Intonation guides sentence processing in the left inferior frontal gyrus
Supplementary table 1 Reaction times. Pair-wise comparisons (Wilcoxon signed-rank tests)
between reaction times in all experimental conditions. Table shows Z-scores followed by the
P-value per comparison. P-values smaller than 0.0033 (Bonferroni-corrected) are marked
with an †.
Supplementary table 2 Accuracy rates. Pair-wise comparisons (Wilcoxon signed-rank tests)
between accuracy rates in all experimental conditions. Table shows Z-scores followed by the
P-value per comparison. P-values smaller than 0.0033 (Bonferroni-corrected) are marked
4 BL ProsodyOnly [Thomas verspricht Nick # [sie zu bezahlen]]
[Thomas promises Nick # [to pay her for it]]
5 BL GrammaticalOnly [Thomas verspricht # [sie dafür zu bezahlen]]
[Thomas promises # [to pay her for it]]
[Thomas verspricht # [heute dafür zu bezahlen]]
[Thomas promises # [to pay for it today]]
Supplementary Figure 1
[Thomas verspricht Nick # [dafür zu bezahlen]]
[Thomas verspricht # [Nick dafür zu bezahlen]]
[Thomas verspricht [sie dafür zu bezahlen]]
[Thomas verspricht Nick # [sie zu bezahlen]]
[Thomas verspricht # [sie dafür zu bezahlen]]
[Thomas verspricht # [heute dafür zu bezahlen]]
3GrammaticalOnly [Thomas verspricht [sie dafür zu bezahlen]]
[Thomas promises [to pay her for it]]
2[Thomas verspricht # [Nick dafür zu bezahlen]]
[Thomas promises # [to pay Nick for it]]
ProsodyOnly1 [Thomas verspricht Nick # [dafür zu bezahlen]]
[Thomas promises Nick # [to pay for it]]
Caption Supplementary Figure 1
Spectrograms with pitch contours for each of the experimental conditions. In the five
conditions with intonational phrase boundary (IPB), three acoustic events can be observed:
(i) a pitch rise and (ii) syllable lengthening, followed by (iii) a pause. Note that in the
Grammatical Only condition the IPB is absent, marked by an absence of pitch rise, syllable
lengthening and pause (indicated with an X).