Article

Dorsal and Ventral Pathways for Prosody

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Abstract

Our vocal tone-the prosody-contributes a lot to the meaning of speech beyond the actual words. Indeed, the hesitant tone of a "yes" may be more telling than its affirmative lexical meaning [1]. The human brain contains dorsal and ventral processing streams in the left hemisphere that underlie core linguistic abilities such as phonology, syntax, and semantics [2-4]. Whether or not prosody-a reportedly right-hemispheric faculty [5, 6]-involves analogous processing streams is a matter of debate. Functional connectivity studies on prosody leave no doubt about the existence of such streams [7, 8], but opinions diverge on whether information travels along dorsal [9] or ventral [10, 11] pathways. Here we show, with a novel paradigm using audio morphing combined with multimodal neuroimaging and brain stimulation, that prosody perception takes dual routes along dorsal and ventral pathways in the right hemisphere. In experiment 1, categorization of speech stimuli that gradually varied in their prosodic pitch contour (between statement and question) involved (1) an auditory ventral pathway along the superior temporal lobe and (2) auditory-motor dorsal pathways connecting posterior temporal and inferior frontal/premotor areas. In experiment 2, inhibitory stimulation of right premotor cortex as a key node of the dorsal stream decreased participants' performance in prosody categorization, arguing for a motor involvement in prosody perception. These data draw a dual-stream picture of prosodic processing that parallels the established left-hemispheric multi-stream architecture of language, but with relative rightward asymmetry.

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... As determined in our study, listening to interjections relative to concrete nouns, activated the left dorso-central insula and the circuit in the right hemisphere comprising the posterior part of superior temporal sulcus (pSTS), the anterior part of the superior temporal sulcus (aSTS), the inferior frontal gyrus (IFG) and the premotor cortex (PMC). These data are fully in agreement with Sammler et al. (2015), who identified this same brain network during a prosody perception task. In their fMRI study, Sammler et al. (2015) asked participants to listen to linguistic stimuli (the words "pear" and "bear") pronounced with different a rhythm and tone such that they conveyed two different communicative intentions (naming and asking). ...
... These data are fully in agreement with Sammler et al. (2015), who identified this same brain network during a prosody perception task. In their fMRI study, Sammler et al. (2015) asked participants to listen to linguistic stimuli (the words "pear" and "bear") pronounced with different a rhythm and tone such that they conveyed two different communicative intentions (naming and asking). In that study, the participants were requested to determine whether the speaker was naming or asking for the object; in a control condition, the same words were pronounced with no prosody and the participants were asked to determine whether the speaker said "bear" or "pear." ...
... In that study, the participants were requested to determine whether the speaker was naming or asking for the object; in a control condition, the same words were pronounced with no prosody and the participants were asked to determine whether the speaker said "bear" or "pear." Sammler et al. (2015) found that the right hemisphere network comprising pSTS, aSTS, IFG and PMC was only activated under the experimental condition. ...
Article
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In previous studies on auditory vitality forms, we found that listening to action verbs pronounced gently or rudely, produced, relative to a neutral robotic voice, activation of the dorso-central insula. One might wonder whether this insular activation depends on the conjunction of action verbs and auditory vitality forms, or whether auditory vitality forms are sufficient per se to activate the insula. To solve this issue, we presented words not related to actions such as concrete nouns (e.g.“ball”), pronounced gently or rudely. No activation of the dorso-central insula was found. As a further step, we examined whether interjections, i.e. speech stimuli conveying communicative intention (e.g. “hello”), pronounced with different vitality forms, would be able to activate, relative to control, the insula. The results showed that stimuli conveying a communicative intention, pronounced with different auditory vitality forms activate the dorsal-central insula. These data deepen our understanding of the vitality forms processing, showing that insular activation is not specific to action verbs, but can be also activated by speech acts conveying communicative intention such as interjections. These findings also show the intrinsic social nature of vitality forms because activation of the insula was not observed in the absence of a communicative intention.
... Specifically, our assumption is based on ear advantage for linguistic and melodic signal processing. In this context, the underlying neural mechanisms suggest that the left hemisphere that connects primarily to the right ear is dominant in the processing of language-bearing signals (e.g., Tervaniemi & Hugdahl, 2003;Vigneau et al., 2006), and the right hemisphere that links to the left ear is predominant in the melodic and speech intonation processing (e.g., Meyer et al., 2002;Sammler et al., 2015). As a consequence, and in order to offer the brain a presumably optimal signal for processing, a dichotic listening approach is adopted. ...
... The L-FR signals elicited larger amplitudes of N400 and P600 components, which indicated that a heavier mental load was required for semantic and syntactic manipulations relative to the NL-NR signals. This may result from the L-FR signals violating the left ear advantage for prosodic information and the right ear advantage for linguistic signals (Meyer et al., 2002;Sammler et al., 2015;Tervaniemi & Hugdahl, 2003;Vigneau et al., 2006). L-FR sent prosodic signals to the right ear and linguistic information to the left ear, which is presumably a non-optimal signal for the brain to process. ...
... Unlike the phonological processing of the unfiltered words, the low-pass filtered sentence signals contain only prosodic information without noticeable segmental features of the speech sounds. Thus, the FL-FR signals induced activation in the bilateral fronto-temporal areas for intonation and rhythm processing, in addition, a pathway linking posterior temporal to IFG in the right hemisphere was prominently involved in the prosodic processing (Meyer et al., 2002;Sammler et al., 2015). ...
Article
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The quality of the physical language signals to which learners are exposed and which result in neurobiological activity leading to perception constitutes a variable that is rarely, if ever, considered in the context of language learning. It deserves some attention. The current study identifies an optimal audio language input signal for Chinese EFL/ESL learners generated by modifying the physical features of language-bearing audio signals. This is achieved by applying the principles of verbotonalism in a dichotic listening context. Low-pass filtered (320 Hz cut-off) and unfiltered speech signals were dichotically and diotically directed to each hemisphere of the brain through the contralateral ear. Temporal and spatial neural signatures for the processing of the signals were detected in a combined Event-Related Potential (ERP) and functional Magnetic Resonance Imaging (fMRI) experiment. Results showed that the filtered stimuli in the left ear and unfiltered in the right ear (FL-R) configuration provided optimal auditory language input by actively exploiting left-hemispheric dominance for language processing and right-hemispheric dominance for melodic processing, i.e., each hemisphere was fed the signals that it should be best equipped to process - and it actually did so effectively. In addition, the filtered stimuli in the right ear and unfiltered in the left ear (L-FR) configuration was identified as entirely non-optimal for language learners. Other outcomes included significant load reduction through exposure to both-ear-filtered FL-FR signals as well as the confirmation that non-language signals were recognized by the brain as irrelevant to language and did not trigger any language processing. These various outcomes will necessarily entail further research.
... Linguistic and affective prosody perception has been proposed to rely on dorsal and ventral streams in the right hemisphere [8,9], that is, frontotemporal regions interconnected dorsally via the arcuate fasciculus (AF) and ventrally via the inferior fronto-occipital fasciculus (IFOF). These streams stand in dynamic exchange with the left hemisphere language networks via the corpus callosum (CC) [10][11][12]. ...
... The DTI analyses focused on the AF, IFOF, CC, and tapetum, as these tracts have been implicated in both prosody perception and amusia [8,10,11,15,19,23]. Furthermore, based on our study hypotheses and the expected right-lateralization of results, right inferior longitudinal fasciculus and uncinate fasciculus were also dissected. ...
... These combined data highlight the damage and disconnection of right IFOF, and associated areas, as the most likely causes of poststroke deficits in prosody perception. The right ventral stream has been suggested to play a critical role in prosody perception [8,15,16], but larger DTI studies evaluating the necessity of WM tracts have been lacking [17]. The right IFOF interconnects right frontal, temporal, and inferior parietal/occipital areas and is a major anatomical ventral stream pathway [34,35]. ...
Article
Background: To determine and compare lesion patterns and structural dysconnectivity underlying post-stroke aprosodia and amusia, using a data-driven multimodal neuroimaging approach. Methods: Thirty-nine patients with right or left hemisphere stroke were enrolled in a cohort study and tested for linguistic and affective prosody perception and musical pitch and rhythm perception at subacute and 3-month post-stroke stages. Participants listened to words spoken with different prosodic stress that changed their meaning, and to words spoken with six different emotions, and chose which meaning or emotion was expressed. In the music tasks, participants judged pairs of short melodies as same or different in terms of pitch or rhythm. Structural MRI data was acquired at both stages, and machine learning-based lesion-symptom mapping and deterministic tractography were used to identify lesion patterns and damaged white matter pathways giving rise to aprosodia and amusia. Results: Both aprosodia and amusia were behaviorally strongly correlated and associated with similar lesion patterns in right fronto-insular and striatal areas. In multiple regression models, reduced fractional anisotropy (FA) and lower tract volume of the right inferior fronto-occipital fasciculus were the strongest predictors for both disorders, over time. Conclusions: These results highlight a common origin of aprosodia and amusia, both arising from damage and disconnection of the right ventral auditory stream integrating rhythmic-melodic acoustic information in prosody and music. Comorbidity of these disabilities may worsen the prognosis and affect rehabilitation success.
... Furthermore, the right hemisphere contributes to some important aspect of language such as pitch prosody comprehension and production [38][39][40], supported by the right hemispheric white matter bundles connecting the right lateralized ventral attention network (VAN) [41]. ...
... Indeed, this is the case as evidenced by Fig. 1, which shows the increased recruitment of the right superior temporal sulcus/superior temporal gyrus in the Overt condition. This appears to be related to the fact that prosodic pitch control involves the right hemisphere along ventral and dorsal pathways connected by white matter bundles similar to those found in the language network in the left hemisphere [38,41]. ...
... We referred to this network as the VAN [58,59], involved in the detection of behaviorally relevant stimuli [60]. Interestingly, within this network, the pSTS appears to be activated by the pitch prosody [38]. FMRI studies have indicated that this area serves as an input region transferring prosodic acoustic information to frontal areas [61], but is also involved in facial action units recognition [62] and more generally activated by observed intentional actions [63]. ...
Article
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Background Pre-surgical mapping of language using functional MRI aimed principally to determine the dominant hemisphere. This mapping is currently performed using covert linguistic task in way to avoid motion artefacts potentially biasing the results. However, overt task is closer to natural speaking, allows a control on the performance of the task, and may be easier to perform for stressed patients and children. However, overt task, by activating phonological areas on both hemispheres and areas involved in pitch prosody control in the non-dominant hemisphere, is expected to modify the determination of the dominant hemisphere by the calculation of the lateralization index (LI). Objective Here, we analyzed the modifications in the LI and the interactions between cognitive networks during covert and overt speech task. Methods Thirty-three volunteers participated in this study, all but four were right-handed. They performed three functional sessions consisting of (1) covert and (2) overt generation of a short sentence semantically linked with an audibly presented word, from which we estimated the “Covert” and “Overt” contrasts, and a (3) resting-state session. The resting-state session was submitted to spatial independent component analysis to identify language network at rest (LANG), cingulo-opercular network (CO), and ventral attention network (VAN). The LI was calculated using the bootstrapping method. Results The LI of the LANG was the most left-lateralized (0.66 ± 0.38). The LI shifted from a moderate leftward lateralization for the Covert contrast (0.32 ± 0.38) to a right lateralization for the Overt contrast (− 0.13 ± 0.30). The LI significantly differed from each other. This rightward shift was due to the recruitment of right hemispheric temporal areas together with the nodes of the CO. Conclusion Analyzing the overt speech by fMRI allowed improvement in the physiological knowledge regarding the coordinated activity of the intrinsic connectivity networks. However, the rightward shift of the LI in this condition did not provide the basic information on the hemispheric language dominance. Overt linguistic task cannot be recommended for clinical purpose when determining hemispheric dominance for language.
... The temporal-spectral dissociation is in line with the finding that the perception of lower-level prosodic information relies on a righthemispheric network that is also recruited during music perception, including song ( Gandour et al., 2004 ;Kreitewolf et al., 2014 ;Merrill et al., 2012 ;Sammler et al., 2015 ;Tong et al., 2005 ). This network encompasses primary and secondary auditory areas along the temporal gyrus, including Heschl's gyrus and planum temporale, and frontal areas, most prominently the inferior frontal gyrus. ...
... We performed our study under the following hypotheses: (a) pitchautocorrelations should scale with activity in the right-hemispheric prosodic fronto-temporal processing areas (see Sammler et al. 2015 ); (b) poems and songs should recruit areas along the superior and middle temporal gyrus in both the left and the right hemisphere; (c) liking ratings should correspond to activity within the reward network, including orbitofrontal cortex, cingulate cortex, the striatum and further subcortical areas. Overall, we set out to show that a direct comparison of poems and their musical settings allows to further our understanding of the affinities and differences in the prosodic processing of language and music and of the neurobiological bases of (aesthetic) perception/evaluation. ...
... The supramarginal gyrus is considered to support pitch memory ( Schaal et al., 2017 ;Schaal et al., 2015 ), thus playing an integral role in perceiving familiar melodies. The right IFG, on the other hand, is part of the prosodic network ( Sammler et al., 2015 ), yet also plays a crucial role for "musical syntax ", i.e., the processing of non-local, structural and hierarchical dependencies between tones of a melody ( Bianco et al., 2016 ;Cheung et al., 2018 ;Koelsch, 2006Koelsch, , 2011Kunert et al., 2015 ;Maess et al., 2001 ;Patel, 2005 ). Again, we interpret this finding as evidence that melodic properties of songs can well be captured by pitch autocorrelations in a neurobiologically plausible way. ...
Article
The neural processing of speech and music is still a matter of debate. A long tradition that assumes shared processing capacities for the two domains contrasts with views that assume domain-specific processing. We here contribute to this topic by investigating, in a functional magnetic imaging (fMRI) study, ecologically valid stimuli that are identical in wording and differ only in that one group is typically spoken (or silently read), whereas the other is sung: poems and their respective musical settings. We focus on the melodic properties of spoken poems and their sung musical counterparts by looking at proportions of significant autocorrelations (PSA) based on pitch values extracted from their recordings. Following earlier studies, we assumed a bias of poem-processing towards the left and a bias for song-processing on the right hemisphere. Furthermore, PSA values of poems and songs were expected to explain variance in left- vs. right-temporal brain areas, while continuous liking ratings obtained in the scanner should modulate activity in the reward network. Overall, poem processing compared to song processing relied on left temporal regions, including the superior temporal gyrus, while song processing compared to poem processing recruited more right temporal areas, including Heschl's gyrus and the superior temporal gyrus. PSA values co-varied with activation in bilateral temporal regions for poems, and in right-dominant fronto-temporal regions for songs, while continuous liking ratings were correlated with activity in the default mode network for both poems and songs. The pattern of results suggests that the neural processing of poems and their musical settings is based on processing their melodic properties in bilateral temporal auditory areas and an additional right fronto-temporal network supporting the processing of melodies in songs. These findings take a middle ground in providing evidence for specific processing circuits for speech and music processing in the left and right hemisphere, but simultaneously for shared processing of melodic aspects of both poems and their musical settings in the right temporal cortex. Thus, we demonstrate the neurobiological plausibility of assuming the importance of melodic properties in spoken and sung aesthetic language alike, along with the involvement of the default mode network in the aesthetic appreciation of these properties in both domains.
... Recent models of propositional language organization have moved from discussing functional localization in terms of neural substrate "islands" (with damage to cortical regions resulting in mutually exclusive aphasia subtypes) to models comprising these cortical regions as well as their connections via white matter pathways, representing ventral and dorsal processing streams [10][11][12]. A similar topography is observed in the right hemisphere for affective prosody, wherein more dorsally situated structures are implicated in production and more ventrally located regions in recognition [13][14][15]. Focusing specifically on affective prosody recognition, right hemisphere temporoparietal regions [12,[16][17][18][19][20][21][22] as well as frontal [2,14], subcortical [15,[21][22][23], and white matter [24] structures have been implicated. ...
... It would follow that even finer specialization may occur for distinct affective prosody recognition subprocesses such as that observed in propositional language (e.g., lexical selection, phonological decoding, etc.). Leading models of affective prosody recognition broadly divide subprocesses into sensory/perceptual extraction (Stage 1), sensory/perceptual-conceptual integration (Stage 2), and cognitive evaluation (Stage 3) [2,13,14,23,25]. The model proposed by Wright and colleagues [2], and further refined by Sheppard and colleagues [23] describes a threestage model of receptive affective prosody that elaborates upon how acoustic-prosodic features are mapped onto semantic representations of emotions. ...
... Thirty-eight adults with acute RHD (14 female, 24 male; 16 African American, 22 Caucasian; 31 right-handed, 5 left-handed, 2 unknown) were identified for the study. Participants were 63.7 ± 13.3 years of age (28-87) and had 14.0 ± 3.6 years of education (7)(8)(9)(10)(11)(12)(13)(14)(15)(16)(17)(18)(19)(20)(21)(22). Behavioral testing was initiated within 3.3 ± 1.9 days (1-7) and a routine clinical MRI of the brain was obtained for 35/38 participants within 1.2 ± 1.5 days (0-7) of hospital admission for acute stroke. ...
Article
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Difficulty recognizing affective prosody (receptive aprosodia) can occur following right hemisphere damage (RHD). Not all individuals spontaneously recover their ability to recognize affective prosody, warranting behavioral intervention. However, there is a dearth of evidence-based receptive aprosodia treatment research in this clinical population. The purpose of the current study was to investigate an explicit training protocol targeting affective prosody recognition in adults with RHD and receptive aprosodia. Eighteen adults with receptive aprosodia due to acute RHD completed affective prosody recognition before and after a short training session that targeted proposed underlying perceptual and conceptual processes. Behavioral impairment and lesion characteristics were investigated as possible influences on training effectiveness. Affective prosody recognition improved following training, and recognition accuracy was higher for pseudo- vs. real-word sentences. Perceptual deficits were associated with the most posterior infarcts, conceptual deficits were associated with frontal infarcts, and a combination of perceptual-conceptual deficits were related to temporoparietal and subcortical infarcts. Several right hemisphere ventral stream regions and pathways along with frontal and parietal hypoperfusion predicted training effectiveness. Explicit acoustic-prosodic-emotion training improves affective prosody recognition, but it may not be appropriate for everyone. Factors such as linguistic context and lesion location should be considered when planning prosody training.
... This observation converges with similar findings in LH homologues: frontal regions, particularly the inferior frontal lobe, are critical for speech-language production (e.g., Fridriksson et al., 2018;Hickock & Poeppel, 2007;Indefrey & Levelt, 2004). According to some models of receptive aprosodia (Grandjean, 2020;Sammler et al., 2015;Schirmer & Kotz, 2006), this region is also critical for (affective) prosody recognition processes, which aligns with cases of receptive aprosodia following damage to the inferior right frontal lobe. Thus, findings suggest that the connections to frontal lobe structures are also important to consider. ...
... White matter pathway involvement was reported by only three articles, with Patel et al. (2018) reporting SLF, IFOF, and UF involvement in impaired affective prosody use in discourse. Sammler et al. (2015) used multi-fiber probabilistic tractography to estimate white matter pathways connecting cortical RH regions activated during linguistic prosody recognition. They observed a ventral connection between posterior superior temporal sulcus and inferior frontal gyrus via the middle longitudinal fasciculus as well as a dorsal connection between the two regions via the arcuate fasciculus and SLF. ...
Article
Affective prosody, or the changes in rate, rhythm, pitch, and loudness that convey emotion, has long been implicated as a function of the right hemisphere (RH), yet there is a dearth of literature identifying the specific neural regions associated with its processing. The current systematic review aimed to evaluate the evidence on affective prosody localization in the RH. One hundred and ninety articles from 1970 to February 2020 investigating affective prosody comprehension and production in patients with focal brain damage were identified via database searches. Eleven articles met inclusion criteria, passed quality reviews, and were analyzed for affective prosody localization. Acute, subacute, and chronic lesions demonstrated similar profile characteristics. Localized right antero-superior (i.e., dorsal stream) regions contributed to affective prosody production impairments, whereas damage to more postero-lateral (i.e., ventral stream) regions resulted in affective prosody comprehension deficits. This review provides support that distinct RH regions are vital for affective prosody comprehension and production, aligning with literature reporting RH activation for affective prosody processing in healthy adults as well. The impact of study design on resulting interpretations is discussed.
... In contrast, other neurocognitive studies have focused on the role of prosody in expressing speaker's emotional states (Ethofer et al. 2006;Schirmer and Kotz 2006;Alter 2011;Brück et al. 2011;Alba-Ferrara et al. 2012;Witteman et al. 2012;Grandjean 2021). Only a few functional magnetic resonance imaging (fMRI) studies (Sammler et al. 2015;Hellbernd and Sammler 2018) have investigated the effect of intonation on the listener in communicative function understanding. Sammler et al. (2015) reported the existence in the right hemisphere of a ventral auditory pathway along the superior temporal lobe and a dorsal auditory-motor one during the comprehension of single words that function as statement or question types. ...
... Only a few functional magnetic resonance imaging (fMRI) studies (Sammler et al. 2015;Hellbernd and Sammler 2018) have investigated the effect of intonation on the listener in communicative function understanding. Sammler et al. (2015) reported the existence in the right hemisphere of a ventral auditory pathway along the superior temporal lobe and a dorsal auditory-motor one during the comprehension of single words that function as statement or question types. Functional interactions between auditory and "social" brain areas have also been documented in the processing of single words differing in prosody that express criticisms, wishes, and suggestions, whereby the latter areas are thought to signify the processing of information about theory of mind (ToM), including the attribution of mental states to oneself and others (Hellbernd and Sammler 2018). ...
Article
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During conversations, speech prosody provides important clues about the speaker’s communicative intentions. In many languages, a rising vocal pitch at the end of a sentence typically expresses a question function, whereas a falling pitch suggests a statement. Here, the neurophysiological basis of intonation and speech act understanding were investigated with high-density electroencephalography (EEG) to determine whether prosodic features are reflected at the neurophysiological level. Already approximately 100 ms after the sentence-final word differing in prosody, questions, and statements expressed with the same sentences led to different neurophysiological activity recorded in the event-related potential. Interestingly, low-pass filtered sentences and acoustically matched nonvocal musical signals failed to show any neurophysiological dissociations, thus suggesting that the physical intonation alone cannot explain this modulation. Our results show rapid neurophysiological indexes of prosodic communicative information processing that emerge only when pragmatic and lexico-semantic information are fully expressed. The early enhancement of question-related activity compared with statements was due to sources in the articulatory-motor region, which may reflect the richer action knowledge immanent to questions, namely the expectation of the partner action of answering the question. The present findings demonstrate a neurophysiological correlate of prosodic communicative information processing, which enables humans to rapidly detect and understand speaker intentions in linguistic interactions.
... Basal ganglia and cerebellum also play a major role in the adaptation in response to the emotional perception of a sound (Sammler et al., 2015). Specifically, the ventral basal ganglia is involved in the processing of musical emotions and affective voices (Paulmann et al., 2005;Paulmann et al., 2008;Pell & Leonard, 2003). ...
... Specifically, in the medial frontal cortex, emotional prosody seems to activate more strongly the dorso-caudal part (Frühholz et al., 2016). In addition, emotional prosody processing involves the right anterior superior temporal gyrus Liebenthal et al., 2016;Sammler et al., 2015). ...
Thesis
The aim of this PhD thesis is to further characterize non-verbal auditory cognition, in particular when it is impaired. The first axis of this research focused on the investigation of emotion perception in two populations with potential deficits: in individuals with congenital amusia and patients with brain-damage. In congenital amusia, Study 1 demonstrated a deficit of emotional prosody perception. This deficit was specifically present for short vowels (versus long sentences). However, the deficit was only present in the explicit recognition task and not in the implicit intensity ratings of the same emotions. Moreover, Study 2 allowed us to relate this explicit recognition deficit with early automatic brain processing decrease by using electroencephalography. In brain-damaged patients, Study 3 demonstrated a deficit for musical emotion perception in relation with the side of the lesion. Based on these results and previous studies, we decided to design a new rehabilitation strategy for the training of non-verbal auditory cognition. In the second axis of this thesis, we focused on developing a new training strategy and chose to test this new training with cochlear implant (CI) users as this population is in high demand for better auditory cognition. We first designed a new short assessment battery for non-verbal auditory cognition. Study 4 demonstrated its efficiency to reveal specific deficits in CI users and in normal-hearing participants by using vocoded sounds. Moreover, findings revealed some evidence that audiovisual cues might help CI users to enhance their non-verbal auditory perception, as previously suggested with verbal material. We then designed a new training strategy by using multisensory integration and more specifically audiovisual stimulation. We suggest that this training could enhance non-verbal auditory abilities of CI users, but also of control participants. We aim to demonstrate the efficiency of this training in a long-term implementation by acquiring both behavioral measures (with the assessment battery developed in Study 4) in all populations with a deficit, but also magnetoencephalographic measures in control participants (Study 5). Overall, this PhD research brings further insight in the field of non-verbal auditory cognition and its associated deficits, and provides a new tool aiming to measure and remediate these deficits, which will then be useable for evaluation in clinical settings
... Additionally, it should be noted that recently, converging evidence has also supported a right hemisphere dual stream organization for emotional prosody processing analogous to the left hemisphere dual streams for language processing (Sammler et al., 2015;Sheppard et al., 2020;Wright et al., 2018). A great deal of evidence exists supporting the left hemisphere dual stream model of language processing, with a dorsal stream for sound-to-articulation mapping and a ventral stream for sound-to-meaning processing (Fridriksson et al., 2016;Hickok & Poeppel, 2004Saur et al., 2008;Scott et al., 2000). ...
... For example, in a study of 23 patients following acute right hemisphere stroke, Sheppard et al. (2020) found evidence using multivariable regression that a model including percent damage to ventral stream regions predicted impaired receptive emotional prosody, but a model including dorsal stream regions did not. In a study using both functional imaging and tractography in unimpaired participants, Sammler et al. (2015) also found evidence for strong right-lateralization (with a substantially smaller level of left-hemisphere activation) of a dorsal stream for expressive emotional prosody generation and ventral stream for comprehension. In contrast, Seydell-Greenwald et al. (2020) evaluated receptive emotional prosody using functional imaging in neurologically unimpaired individuals and found strong right lateralized activation in frontotemporal areas, with bilateral activation in IFG pars orbitalis, anterior insula, and amygdala. ...
Article
Introduction Speakers naturally produce prosodic variations depending on their emotional state. Receptive prosody has several processing stages. We aimed to conduct lesion-symptom mapping to determine whether damage (core infarct or hypoperfusion) to specific brain areas was associated with receptive aprosodia or with impairment at different processing stages in individuals with acute right hemisphere stroke. We also aimed to determine whether different subtypes of receptive aprosodia exist that are characterized by distinctive behavioral performance patterns. Methods Twenty patients with receptive aprosodia following right hemisphere ischemic stroke were enrolled within five days of stroke; clinical imaging was acquired. Participants completed tests of receptive emotional prosody, and tests of each stage of prosodic processing (Stage 1: acoustic analysis; Stage 2: analyzing abstract representations of acoustic characteristics that convey emotion; Stage 3: semantic processing). Emotional facial recognition was also assessed. LASSO regression was used to identify predictors of performance on each behavioral task. Predictors entered into each model included 14 right hemisphere regions, hypoperfusion in four vascular territories as measured using FLAIR hyperintense vessel ratings, lesion volume, age, and education. A k-medoid cluster analysis was used to identify different subtypes of receptive aprosodia based on performance on the behavioral tasks. Results Impaired receptive emotional prosody and impaired emotional facial expression recognition were both predicted by greater percent damage to the caudate. The k-medoid cluster analysis identified three different subtypes of aprosodia. One group was primarily impaired on Stage 1 processing and primarily had frontotemporal lesions. The second group had a domain-general emotion recognition impairment and maximal lesion overlap in subcortical areas. Finally, the third group was characterized by a Stage 2 processing deficit and had lesion overlap in posterior regions. Conclusions Subcortical structures, particularly the caudate, play an important role in emotional prosody comprehension. Receptive aprosodia can result from impairments at different processing stages.
... Eckstein and Friederici (2005), who first cited the RAN effect in German sentences, referred to this anterior negativity as reflecting a mismatch for prosodically incongruent words or an absence of prosodic information in a sentence. Prosodic information is discussed for the right-hemispheric pathway (e.g., Meyer, Alter, Friederici, Lohmann, & von Cramon, 2002;Sammler, Grosbras, Anwander, Bestelmeyer, & Belin, 2015). Since RAN reflects a pure aspect of auditory sentence comprehension processing, a RAN in the ERPs followed by a late positivity effect in Turkish for the main effect of prosodic incongruity on post-verbal position was presented. ...
... Since the focus marking processing in a language has both a relation with prosody and syntax (See Eckstein, & Friederici, 2005, 2006Sammler et al., 2015), one of the main purposes of the current study was to investigate the interaction of these components. In this study, the neurophysiological markers of prosody and syntax could be significant for the processing of prosodic boundary in Turkish sentences. ...
Article
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Focused elements are generally marked with syntactic canonicity and prosody. Being a scrambled language, Turkish uses both syntactic and prosodic information to mark the focus. However, it does not allow for focus marking in post-verbal position. In this study, the neurophysiological processes of the focus in Turkish are examined by using prosodic and syntactic information. Recent psycholinguistics studies assume that there is an interaction between prosody and syntax through the focus in the online sentence comprehension process. Thirty participants (16 female and 14 male between the ages of 19 and 33), whose native language was Turkish and who spoke monolingual Turkish, and who did not have any neurological, hearing, or linguistic impairments, took part in the experiments measured with Electroencephalogram (EEG). Using an event-related potentials (ERPs) design, this study provides evidence for an interaction between prosody and syntax in Turkish. The experimental design of the study consisted of prosodic, syntactic, and prosodic-syntactic violations. Participants were asked to listen 300 auditory stimuli (100 filler sentences) including sentences with both congruent and incongruent focus. The stimuli consisted of 50 sentences for each experimental condition. All critical words occurred in the sentence-final positions. For the prosodic violation critical words were focused via incongruent focusing on post-verbal position, and for the syntactic violation critical words were manipulated with case marking manipulation (i.e., accusative case versus dative case violations). In addition, for the interaction of prosodic and syntactic violations, critical words were incongruent focused and incongruent case was marked. The results revealed that prosodic incongruity elicited a broadly distributed positivity in posterior regions (400-1200 ms) lateralized to the left hemisphere and a right anterior negativity (RAN) (300-500 ms) effect. Syntactic violations also indicated a distributed anterior negativity (300-500 ms) effect. Supportive evidence for the late interaction of prosodic and syntactic processing in the neural integration of positive 600 (P600) and Closure Positive Shift (CPS) was observed. The findings provide support for recent neurocognitive approaches for late interaction between prosody and syntax in the sentence-final position in Turkish sentences.
... More rapid temporal transitions, related to phonemic information, appear to be preferentially processed in the left hemisphere (Boemio et al., 2005). Slower auditory modulations, those associated with prosodic and syllabic processing, may be preferentially processed in the right hemisphere, which is also associated with rhythm perception (Sammler et al., 2015). In the current study, we take a scalp-level approach due to the difficulty in attaining accurate source-localised results in infants, especially across different ages (Lew et al., 2013;Noreika et al., 2020). ...
... Although the broader neural AM literature suggests that rise time processing may differ developmentally between left and right brain regions, the differential responses of the left and right frontotemporal regions reported here could be due to physiological factors affecting how electrophysiological fluctuations are transmitted from the brain to the scalp (Lew et al., 2013;Noreika et al., 2020). The prior auditory neural literature suggests that the right and left hemispheres play different roles in auditory and linguistic processing, with rapid, phonemic-rate transitions processed in the left (Boemio et al., 2005) and slower modulations processed in the right (Sammler et al., 2015). Pre-reading children show this hemispheric specialisation for "syllabic" but not "phonemic" rates of AM SSN (4 vs 80 Hz; Vanvooren et al., 2014), and both children and adults with dyslexia show atypical right hemisphere synchronisation in response to 4 Hz AM noise (Lizarau et al., 2015). ...
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Amplitude rise times play a crucial role in the perception of rhythm in speech, and reduced perceptual sensitivity to differences in rise time is related to developmental language difficulties. Amplitude rise times also play a mechanistic role in neural entrainment to the speech amplitude envelope. Using an ERP paradigm, here we examined for the first time whether infants at the ages of seven and eleven months exhibit an auditory mismatch response to changes in the rise times of simple repeating auditory stimuli. We found that infants exhibited a mismatch response (MMR) to all of the oddball rise times used for the study. The MMR was more positive at seven than eleven months of age. At eleven months, there was a shift to a mismatch negativity (MMN) that was more pronounced over left fronto-central electrodes. The MMR over right fronto-central electrodes was sensitive to the size of the difference in rise time. The results indicate that neural processing of changes in rise time is present at seven months, supporting the possibility that early speech processing is facilitated by neural sensitivity to these important acoustic cues.
... Moreover, previous studies also showed that the right IFG, especially the right BA 44, was critical for L2 acoustic feature differentiation , prosodic processing (Sammler et al., 2015), and more broadly, speech comprehension and production (e.g., Gajardo-Vidal et al., 2018;Hartwigsen et al., 2010;Harvey et al., 2019;Naeser et al., 2011). However, restricted to the visual domain, this study provides extra evidence that the right IFG, including right BA 44, might function beyond mere phonological processing aspects. ...
... As to the right STG/STS, multiple functions such as the theory of mind, audiovisual integration, motion processing, speech processing, and face processing, have been assigned to this area (Hein & Knight, 2008). The right STG was also suggested to be involved in encoding spatial-temporal properties of sign language (Blunmenthal-Dramé & Malaia, 2018), and word retrieval (Riès et al., 2016), and its connection to right IFG was identified as crucial prerequisite for the categorization of speech stimuli that gradually varied in their prosodic cues (Sammler et al., 2015). However, the structural connection between the right STG and the right frontal lobe via the right arcuate fasciculus was also evidenced to support above-word-level reading comprehension, and its connection to the right occipital lobe through the right inferior longitudinal fasciculus was proposed to be critical for word-level reading (e.g. ...
Article
Word category information (WCI) is proposed to be fundamental for syntactic learning and processing. However, it remains largely unclear how left-hemispheric key regions for language, including BA 44 in the inferior frontal gyrus (IFG) and superior temporal gyrus (STG), interact with their right-hemispheric homologues to support the WCI-based syntactic learning. To address this question, this study employed a unified structural equation modeling (uSEM) approach to explore both the intra- and inter-hemispheric effective connectivity among these areas, to specify the neural underpinnings of handling WCI for syntactic learning. Modeling results identified a distinctive intra-left hemispheric connection from left BA 44 to left STG, a more integrated intra-right hemispheric network, and a particular frontal right-to-left hemispheric connectivity pattern for WCI-based syntactic learning. Further analyses revealed a selective positive correlation between task performance and the lagged effect in left BA 44. These results converge on a critical left fronto-temporal language network with left BA 44 and its connectivity to left STG for WCI-based syntactic learning, which is also facilitated in a domain-general fashion by the right homologues. Together, these results provide novel insights into crucial neural network(s) for syntactic learning on the basis of WCI.
... Previous research has reported motor involvement in prosody and vocal emotion perception. For example, studies have provided causal evidence in adults (Banissy et al. 2010;Sammler et al. 2015) and correlative evidence in children (Correia et al. 2019), supporting a role for the motor system in the perception of prosodic and vocal-emotional cues. In particular, left ventral motor cortex activation has been reported during the perception of positive vocal emotions and has been interpreted as preparation for responsive gestures (e.g. ...
Article
During social interactions, speakers signal information about their emotional state through their voice, which is known as emotional prosody. Little is known regarding the precise brain systems underlying emotional prosody decoding in children and whether accurate neural decoding of these vocal cues is linked to social skills. Here, we address critical gaps in the developmental literature by investigating neural representations of prosody and their links to behavior in children. Multivariate pattern analysis revealed that representations in the bilateral middle and posterior superior temporal sulcus (STS) divisions of voice-sensitive auditory cortex decode emotional prosody information in children. Crucially, emotional prosody decoding in middle STS was correlated with standardized measures of social communication abilities; more accurate decoding of prosody stimuli in the STS was predictive of greater social communication abilities in children. Moreover, social communication abilities were specifically related to decoding sadness, highlighting the importance of tuning in to negative emotional vocal cues for strengthening social responsiveness and functioning. Findings bridge an important theoretical gap by showing that the ability of the voice-sensitive cortex to detect emotional cues in speech is predictive of a child's social skills, including the ability to relate and interact with others.
... Similar approaches have been adopted in other aspects of music and language processing, such as prosody (Sammler, Grosbras, Anwander, Bestelmeyer, & Belin, 2015), linguistic and musical syntax (Musso et al., 2015) and singing (Loui, 2015). Extending the dual-route account (Griffiths, 2008) into a 'Linked Dual Representation' model, Hutchins and Moreno (2013) proposed that vocal information can be encoded through distinct symbolic and motoric mechanisms, but, in addition, motoric representations can be mediated by symbolic representations. ...
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Congenital amusia is a neurogenetic disorder of pitch perception that may also compromise pitch production. We explored whether vocal imitation of song in amusia is influenced by melody familiarity and phonetic content. Thirteen Mandarin-speaking amusics and 13 matched controls imitated novel song segments with lyrics and on the syllable /la/ while their output was recorded. Eleven out of these participants in each group also imitated segments of a familiar song. Subsequent acoustic analysis was conducted to measure pitch and timing matching accuracy. While amusics’ performance was facilitated by melody familiarity in terms of pitch interval deviation, signed interval deviation and number of contour errors, they showed compromised performance compared to controls in absolute pitch deviation, number of pitch interval errors, duration difference, interonset interval difference, and number of time errors in both familiar and novel song conditions. The presence of lyrics did not affect either group’s performance substantially. While a stronger relationship between music perception and novel melody imitation was observed in amusics as opposed to familiar melody imitation, controls showed the opposite pattern. We discuss the implications of the study in terms of music familiarity, memory demands, the relevance of lexical information and the link between perception and production.
... It is plausible that rise time processing may differ developmentally between left and right brain regions if the neural literature is considered. The right and left hemispheres are thought to play different roles in auditory and linguistic processing, with the right hemisphere associated with rhythm perception (Sammler et al., 2015). Slower auditory modulations, those associated with the syllabic rate of speech, may be preferentially processed in the right hemisphere, while more rapid temporal transitions, such as phonemic rate modulations, are processed in the left (Boemio et al., 2005). ...
Preprint
Amplitude rise times play a crucial role in the perception of rhythm in speech, and reduced perceptual sensitivity to differences in rise time is related to developmental language difficulties. Amplitude rise times also play a mechanistic role in neural entrainment to the speech amplitude envelope. Using an ERP paradigm, here we examined for the first time whether infants at the ages of seven and eleven months exhibit an auditory mismatch response to changes in the rise times of simple repeating auditory stimuli. We found that infants exhibited a mismatch response to the oddball rise time that was more positive at seven than eleven months of age. At eleven months, there was a left-lateralised shift to a mismatch negativity. Infants’ ability to detect changes in rise time was generally robust, with a range of oddball stimuli with different rise times each eliciting a mismatch response from 85% of infants. A lateralised effect indicated that the size of the mismatch response varied as the change in rise time became easier to detect. The mismatch response to the different rise time oddballs also stabilised as infants got older. The results indicate that neural processing of changes in rise time develops early in life, supporting the possibility that early speech processing is facilitated by neural sensitivity to these acoustic cues to rhythm.
... However, differential activation patterns emerge in CI patients reflecting compensatory mechanisms due to the degraded auditory information conveyed by the cochlear implant ( Giraud and Truy, 2002 ;Giraud et al., 20 0 0 , 20 01 ). To our knowledge, no study has specifically analyzed the brain network involved in auditory prosodic information in the CIP group, while in NH individuals it is supposed to involve an extensive bilateral cortical and subcortical network ( Hellbernd and Sammler, 2016 ;Sammler et al., 2015 ;Wildgruber et al., 2004 ;Witteman et al., 2011 ). However, there is now a strong assertion that in CI patients, the processing of visuo-auditory speech information is supported by a different cortical network compared to hearing counterparts Stropahl and Debener, 2017 ). ...
Article
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Cochlear implanted (CI) adults with acquired deafness are known to depend on multisensory integration skills (MSI) for speech comprehension through the fusion of speech reading skills and their deficient auditory perception. But, little is known on how CI patients perceive prosodic information relating to speech content. Our study aimed to identify how CI patients use MSI between visual and auditory information to process paralinguistic prosodic information of multimodal speech and the visual strategies employed. A psychophysics assessment was developed, in which CI patients and hearing controls (NH) had to distinguish between a question and a statement. The controls were separated into two age groups (young and aged-matched) to dissociate any effect of aging. In addition, the oculomotor strategies used when facing a speaker in this prosodic decision task were recorded using an eye-tracking device and compared to controls. This study confirmed that prosodic processing is multisensory but it revealed that CI patients showed significant supra-normal audiovisual integration for prosodic information compared to hearing controls irrespective of age. This study clearly showed that CI patients had a visuo-auditory gain more than 3 times larger than that observed in hearing controls. Furthermore, CI participants performed better in the visuo-auditory situation through a specific oculomotor exploration of the face as they significantly fixate the mouth region more than young NH participants who fixate the eyes, whereas the aged-matched controls presented an intermediate exploration pattern equally reported between the eyes and mouth. To conclude, our study demonstrated that CI patients have supra-normal skills MSI when integrating visual and auditory linguistic prosodic information, and a specific adaptive strategy developed as it participates directly in speech content comprehension.
... This is a core effector region for predicting actions and preparing behavioural responses to salient and arousing events, in music, vocalizations and other cognitive domains. 65,92,137,[140][141][142] This region is intimately linked to the salience network 90 and plays an essential role in processing auditory expectations, especially when these have been established through sensorimotor integration as is generally the case for music. 92,143,144 This study suggests that major dementias have distinct profiles of sensory 'surprise' processing, as instantiated in music. ...
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Making predictions about the world and responding appropriately to unexpected events are essential functions of the healthy brain. In neurodegenerative disorders such as frontotemporal dementia and Alzheimer’s disease, impaired processing of ‘surprise’ may underpin a diverse array of symptoms, particularly abnormalities of social and emotional behaviour, but is challenging to characterise. Here we addressed this issue using a novel paradigm: music. We studied 62 patients (24 female; aged 53–88) representing major syndromes of frontotemporal dementia (behavioural variant, semantic variant primary progressive aphasia, nonfluent-agrammatic variant primary progressive aphasia) and typical amnestic Alzheimer’s disease, in relation to 33 healthy controls (18 female; aged 54–78). Participants heard famous melodies containing no deviants or one of three types of deviant note—acoustic (white-noise burst), syntactic (key-violating pitch change) or semantic (key-preserving pitch change). Using a regression model that took elementary perceptual, executive and musical competence into account, we assessed accuracy detecting melodic deviants and simultaneously-recorded pupillary responses and related these to deviant surprise value (information-content) and carrier melody predictability (entropy), calculated using an unsupervised machine-learning model of music. Neuroanatomical associations of deviant detection accuracy and coupling of detection to deviant surprise value were assessed using voxel-based morphometry of patients’ brain MR images. Whereas Alzheimer’s disease was associated with normal deviant detection accuracy, behavioural and semantic variant frontotemporal dementia syndromes were associated with strikingly similar profiles of impaired syntactic and semantic deviant detection accuracy and impaired behavioural and autonomic sensitivity to deviant information-content (all p < 0.05). On the other hand, nonfluent-agrammatic primary progressive aphasia was associated with generalised impairment of deviant discriminability (p < 0.05) due to excessive false-alarms, despite retained behavioural and autonomic sensitivity to deviant information-content and melody predictability. Across the patient cohort, grey matter correlates of acoustic deviant detection accuracy were identified in precuneus, mid and mesial temporal regions; correlates of syntactic deviant detection accuracy and information-content processing, in inferior frontal and anterior temporal cortices, putamen and nucleus accumbens; and a common correlate of musical salience coding in supplementary motor area (all p < 0.05, corrected for multiple comparisons in pre-specified regions of interest). Our findings suggest that major dementias have distinct profiles of sensory ‘surprise’ processing, as instantiated in music. Music may be a useful and informative paradigm for probing the predictive decoding of complex sensory environments in neurodegenerative proteinopathies, with implications for understanding and measuring the core pathophysiology of these diseases.
... It has long been known that language related tasks such as sentence generation activate and require a functioning Broca's area, Wernick's area, and the basal ganglia (Brown et al., 2006). However, less appreciated is the fact that music tasks such as melody generation activate the exact same network (Koelsch et al., 2002;Brown et al., 2006), although with a bias towards the right hemisphere, while language is biased towards the left hemisphere (Sammler et al., 2015). Consistent with a common neural implementation, children with language impairments also show musical impairments (Jentschke et al., 2008). ...
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Retracing the evolutionary steps by which human brains evolved can offer insights into the underlying mechanisms of human brain function as well as the phylogenetic origin of various features of human behavior. To this end, this article presents a model for interpreting the physical and behavioral modifications throughout major milestones in human brain evolution. This model introduces the concept of a “breakthrough” as a useful tool for interpreting suites of brain modifications and the various adaptive behaviors these modifications enabled. This offers a unique view into the ordered steps by which human brains evolved and suggests several unique hypotheses on the mechanisms of human brain function.
... For example, the mapping rules for European Portuguese establish that parenthetical phrases are signaled by intonational phrase breaks [12]. Interestingly, besides the complexity of the neural networks for prosody described in recent research, it has been suggested that the processing of pitch contour differences between statements and questions involves both the ventral and dorsal pathways in the right hemisphere [73], while the processing of prosodic phrasing involves the right fronto-temporal and dorsal networks, together with inter-hemispheric pathways to map morphosyntactic and phonological structures [74]. The different effects of medication and disease duration on nuclear contours and prosodic phrasing found in the current study raise new questions that future research on PD prosody should address to better understand the mechanisms underlying prosodic impairments and inform therapeutic planning. ...
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The phonology of prosody has received little attention in studies of motor speech disorders. The present study investigates the phonology of intonation (nuclear contours) and speech chunking (prosodic phrasing) in Parkinson's disease (PD) as a function of medication intake and duration of the disease. Following methods of the prosodic and intonational phonology frameworks , we examined the ability of 30 PD patients to use intonation categories and prosodic phrasing structures in ways similar to 20 healthy controls to convey similar meanings. Speech data from PD patients were collected before and after a dopaminomimetic drug intake and were phonologi-cally analyzed in relation to nuclear contours and intonational phrasing. Besides medication, disease duration and the presence of motor fluctuations were also factors included in the analyses. Overall, PD patients showed a decreased ability to use nuclear contours and prosodic phrasing. Medication improved intonation regardless of disease duration but did not help with dysprosodic phrasing. In turn, disease duration and motor fluctuations affected phrasing patterns but had no impact on intonation. Our study demonstrated that the phonology of prosody is impaired in PD, and prosodic categories and structures may be differently affected, with implications for the understanding of PD neurophysiology and therapy.
... Nevertheless, emerging evidences have suggested the recruitment of bilateral sensorimotor system during speech production (Stephens et al. 2010;Silbert et al. 2014b) and speech-in-noise comprehension (Du et al. 2014(Du et al. , 2016. Recent studies have speculated that the left and right sensorimotorrelated regions could have different functional roles for speech processing: while the phonemic processing is strongly left-lateralized, the sensorimotor processing of prosody has been suggested to be lateralized to the right hemisphere (Hickok and Poeppel 2007;Sammler et al. 2015;Tang et al. 2021). Hereby, it is plausible to assume that the right-lateralized neural couplings over the sensorimotor regions on the listener's side could reflect a shared representation of the prosodic information. ...
Article
While the increasingly globalized world has brought more and more demands for non-native language communication, the prevalence of background noise in everyday life poses a great challenge to non-native speech comprehension. The present study employed an interbrain approach based on functional near-infrared spectroscopy (fNIRS) to explore how people adapt to comprehend non-native speech information in noise. A group of Korean participants who acquired Chinese as their non-native language was invited to listen to Chinese narratives at 4 noise levels (no noise, 2 dB, −6 dB, and − 9 dB). These narratives were real-life stories spoken by native Chinese speakers. Processing of the non-native speech was associated with significant fNIRS-based listener-speaker neural couplings mainly over the right hemisphere at both the listener's and the speaker's sides. More importantly, the neural couplings from the listener's right superior temporal gyrus, the right middle temporal gyrus, as well as the right postcentral gyrus were found to be positively correlated with their individual comprehension performance at the strongest noise level (−9 dB). These results provide interbrain evidence in support of the right-lateralized mechanism for non-native speech processing and suggest that both an auditory-based and a sensorimotor-based mechanism contributed to the non-native speech-in-noise comprehension.
... fMRI studies have shown that the right temporal and frontal areas are activated when individuals perform tasks involving prosody. In particular, for the time-sensitive evaluation of prosody, the premotor cortex and inferior frontal gyrus in the right hemisphere are activated [63]. Impairments of prosody might also have affected the conversational task performance of patients with RHD in the current study. ...
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Patients with right hemisphere damage (RHD) occasionally complain of difficulties in conversation. A conversation is a type of communication between the speaker and listener, and several elements are required for a conversation to take place. However, it is unclear which of those elements affect communication in patients with RHD. Therefore, we prospectively enrolled 11 patients with right hemispheric damage due to acute cerebral infarction, within 1 week of onset. To evaluate patients’ conversational abilities, we used a structured conversation task, namely, the “Hallym Conversation and Pragmatics Protocol”. The topics of conversation were “family”, “leisure”, and “other/friends”. The conversation characteristics were classified according to three indices: the “conversational participation index”, “topic manipulation index”, and “conversational breakdown index”. Patients with RHD were compared with 11 age-, sex-, and years of education-matched healthy adults. The most common site of damage in the patients with RHD was the periventricular white matter. There was no significant difference in performance between the two groups according to the conversation participation index and in the discontinuance rate assessed with the conversational breakdown index. However, patients with RHD showed a lower topic maintenance rate and higher topic initiation and topic switching rates, according to the topic manipulation index. Therefore, we explored the characteristics of impaired conversation abilities in patients with RHD by assessing their ability to converse and manage topics during structured conversations, and found difficulties with pragmatics and communication discourse in these patients.
... A decreased FC within the VAN was detected in our study, which indicates that the function of attention in patients with SLE may not be easily influenced by the outside world. Moreover, the specifically decreased regions inside the VAN, including the supramarginal gyrus and the pars triangularis, are relevant to attention, sentence processing, and phonetic processing (Friederici & Gierhan, 2013;Sammler et al., 2015). Hence, the perception ability of SLE patients may be impaired. ...
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The aim of this study was to investigate the abnormities in functional connectivity (FC) within each modular network and between modular networks in patients with systemic lupus erythematosus (SLE). Twelve meaningful modular networks were identified via independent component analysis from 41 patients and 40 volunteers. Parametric tests were used to compare the intra- and intermodular FC between the groups. Partial correlation analysis was used to seek the relationships between abnormal FCs and the clinical data. Compared to the controls, SLE patients showed decreased intramodular FC in the anterior default mode network (aDMN), posterior default mode network (pDMN), ventral attention network (VAN), and sensorimotor network (SMN) and increased intramodular FC in the medial visual network (mVN) and left frontoparietal network. In addition, SLE patients showed decreased intermodular FC between the SMN and the lateral visual network (lVN), between the SMN and the VAN, and between the pDMN and the lVN and exhibited increased intermodular FC between the SMN and the salience network (SAN), between the pDMN and the SAN, and between the aDMN and the VAN. Moreover, we found several correlations among the abnormal FCs and the Mini-Mental State Examination in SLE patients. Mild cognitive impairment is compensated by the hyperconnectivity between the aDMN and the VAN, while severe cognitive impairment tends to be compensated by the hyperconnectivity between the SMN and the SAN. The FC value between the SMN and the SAN and between the aDMN and the VAN may serve as neuroimaging markers for monitoring cognitive progression in SLE patients.
... 14 Similar to the dual-stream model of language processing in the left hemisphere, 17,18 analogous dual ventral and dorsal streams in the right hemisphere have been proposed for prosody comprehension. 19 Two different models for lateralization of emotional processing have been proposed. One of these, the "right hemisphere hypothesis", claims that the right hemisphere is dominant for every kind of emotional perception and expression. ...
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Why do we schedule romantic meetings and trysts for dinner and not for breakfast? How many people do you know who prefer a romantic walk in the morning instead of in the evening? Why are the moon, sunset, night, and candle-light universally a part of the romantic lexicon? Population based data revealed that people usually make love when they go to bed at night and most sexual activities occur between 10 pm and 2 am. Given these universal patterns of behavior, what answers does neuroscience give to these questions? It appears that brain lateralization and its specific network processes offer a useful framework. In this hypothesis article we offer a view of night-oriented romantic processing based on findings concerning circadian rhythm, brain lateralization, and sleep evolution. Previous studies revealed a smooth change in brain lateralization from the left in the morning to the right hemisphere in the evening among right-handed people. Based on the hemispheric lateralization for emotional processing and evidence from evolutionary studies, we will try to draw a framework for analyzing night-oriented romantic behavior in humans. To this end, we review findings on brain lateralization and it s specific network processes with respect to emotion processing, sleep, waking and vigilance.
... I will develop this question in more details in the next chapter.A common correlate of syntactic and acoustic surprise processing at a cognitive level was identified in supplementary motor cortex. This is a core effector region for predicting actions and preparing behavioural responses to salient and arousing events, in music, vocalisations and other cognitive domains(Warren et al., 2006;Jacobsen et al., 2015;Sammler et al., 2015;Gordon et al., 2018;Slattery et al., 2019). Its involvement here might plausibly signal preparedness to react to incongruence in familiar melodies. ...
Thesis
The clinical complexity and pathological diversity of neurodegenerative diseases impose immense challenges for diagnosis and the design of rational interventions. To address these challenges, there is a need to identify new paradigms and biomarkers that capture shared pathophysiological processes and can be applied across a range of diseases. One core paradigm of brain function is predictive coding: the processes by which the brain establishes predictions and uses them to minimise prediction errors represented as the difference between predictions and actual sensory inputs. The processes involved in processing unexpected events and responding appropriately are vulnerable in common dementias but difficult to characterise. In my PhD work, I have exploited key properties of music – its universality, ecological relevance and structural regularity – to model and assess predictive cognition in patients representing major syndromes of frontotemporal dementia – non-fluent variant PPA (nfvPPA), semantic-variant PPA (svPPA) and behavioural-variant FTD (bvFTD) - and Alzheimer’s disease relative to healthy older individuals. In my first experiment, I presented patients with well-known melodies containing no deviants or one of three types of deviant - acoustic (white-noise burst), syntactic (key-violating pitch change) or semantic (key-preserving pitch change). I assessed accuracy detecting melodic deviants and simultaneously-recorded pupillary responses to these deviants. I used voxel-based morphometry to define neuroanatomical substrates for the behavioural and autonomic processing of these different types of deviants, and identified a posterior temporo-parietal network for detection of basic acoustic deviants and a more anterior fronto-temporo-striatal network for detection of syntactic pitch deviants. In my second chapter, I investigated the ability of patients to track the statistical structure of the same musical stimuli, using a computational model of the information dynamics of music to calculate the information-content of deviants (unexpectedness) and entropy of melodies (uncertainty). I related these information-theoretic metrics to performance for detection of deviants and to ‘evoked’ and ‘integrative’ pupil reactivity to deviants and melodies respectively and found neuroanatomical correlates in bilateral dorsal and ventral striatum, hippocampus, superior temporal gyri, right temporal pole and left inferior frontal gyrus. Together, chapters 3 and 4 revealed new hypotheses about the way FTD and AD pathologies disrupt the integration of predictive errors with predictions: a retained ability of AD patients to detect deviants at all levels of the hierarchy with a preserved autonomic sensitivity to information-theoretic properties of musical stimuli; a generalized impairment of surprise detection and statistical tracking of musical information at both a cognitive and autonomic levels for svPPA patients underlying a diminished precision of predictions; the exact mirror profile of svPPA patients in nfvPPA patients with an abnormally high rate of false-alarms with up-regulated pupillary reactivity to deviants, interpreted as over-precise or inflexible predictions accompanied with normal cognitive and autonomic probabilistic tracking of information; an impaired behavioural and autonomic reactivity to unexpected events with a retained reactivity to environmental uncertainty in bvFTD patients. Chapters 5 and 6 assessed the status of reward prediction error processing and updating via actions in bvFTD. I created pleasant and aversive musical stimuli by manipulating chord progressions and used a classic reinforcement-learning paradigm which asked participants to choose the visual cue with the highest probability of obtaining a musical ‘reward’. bvFTD patients showed reduced sensitivity to the consequence of an action and lower learning rate in response to aversive stimuli compared to reward. These results correlated with neuroanatomical substrates in ventral and dorsal attention networks, dorsal striatum, parahippocampal gyrus and temporo-parietal junction. Deficits were governed by the level of environmental uncertainty with normal learning dynamics in a structured and binarized environment but exacerbated deficits in noisier environments. Impaired choice accuracy in noisy environments correlated with measures of ritualistic and compulsive behavioural changes and abnormally reduced learning dynamics correlated with behavioural changes related to empathy and theory-of-mind. Together, these experiments represent the most comprehensive attempt to date to define the way neurodegenerative pathologies disrupts the perceptual, behavioural and physiological encoding of unexpected events in predictive coding terms.
... The left STS plays a particular function in this network, as it is not only a multisensory area (Specht and Wigglesworth, 2018), but also an area that is very sensitive to phonetic information in an acoustic signal, as demonstrated by the "sound morphing" paradigms (Specht et al., , 2009Osnes et al., 2011a). Later, Sammler et al. (2015) could demonstrate with a similar paradigm a corresponding effect for prosody processing for the right STS. This study also reflects this differential responsiveness and division of labor between the left and right STS since both paradigms activate the left STS, but only the auditory variant activates the right STS. ...
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The present study replicates a known visual language paradigm, and extends it to a paradigm that is independent from the sensory modality of the stimuli and, hence, could be administered either visually or aurally, such that both patients with limited sight or hearing could be examined. The stimuli were simple sentences, but required the subject not only to understand the content of the sentence but also to formulate a response that had a semantic relation to the content of the presented sentence. Thereby, this paradigm does not only test perception of the stimuli, but also to some extend sentence and semantic processing, and covert speech production within one task. When the sensory base-line condition was subtracted, both the auditory and visual version of the paradigm demonstrated a broadly overlapping and asymmetric network, comprising distinct areas of the left posterior temporal lobe, left inferior frontal areas, left precentral gyrus, and supplementary motor area. The consistency of activations and their asymmetry was evaluated with a conjunction analysis, probability maps, and intraclass correlation coefficients (ICC). This underlying network was further analyzed with dynamic causal modeling (DCM) to explore whether not only the same brain areas were involved, but also the network structure and information flow were the same between the sensory modalities. In conclusion, the paradigm reliably activated the most central parts of the speech and language network with a great consistency across subjects, and independently of whether the stimuli were administered aurally or visually. However, there was individual variability in the degree of functional asymmetry between the two sensory conditions.
... The right hemisphere is also activated in subprocesses, such as metaphor comprehension (Ferstl et al., 2008). Although not exclusive, increased connectivity with the perisylvian regions in the right hemisphere suggests affective prosody (Sammler et al., 2015;Hertrich et al., 2020), which is important for singing. Note that there was also an increase in the connectivity with the superior temporal gyrus, including the planum temporale, in the right hemisphere. ...
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Performing an opera requires singers on stage to process mental imagery and theory of mind tasks in conjunction with singing and action control. Although it is conceivable that the precuneus, as a posterior hub of the default mode network, plays an important role in opera performance, how the precuneus contributes to opera performance has not been elucidated yet. In this study, we aimed to investigate the contribution of the precuneus to singing in an opera. Since the precuneus processes mental scenes, which are multimodal and integrative, we hypothesized that it is involved in opera performance by integrating multimodal information required for performing a character in an opera. We tested this hypothesis by analyzing the functional connectivity of the precuneus during imagined singing and rest. This study included 42 opera singers who underwent functional magnetic resonance imaging when performing “imagined operatic singing” with their eyes closed. During imagined singing, the precuneus showed increased functional connectivity with brain regions related to language, mirror neuron, socio-cognitive/emotional, and reward processing. Our findings suggest that, with the aid of its widespread connectivity, the precuneus and its network allow embodiment and multimodal integration of mental scenes. This information processing is necessary for imagined singing as well as performing an opera. We propose a novel role of the precuneus in opera performance.
... However, SMA ( Benson et al., 2001 ;Scott et al., 2004 ;Warren et al., 2006 ;Jardri et al., 2007 ) and insula ( Ackermann et al., 2001 ;Benson et al., 2001 ;Steinbrink et al., 2009 ;Hervais-Adelman et al., 2012 ) activation has also been reported during passive listening under various conditions. Further, there are studies showing involvement of SMA and insula during auditory tasks, which goes beyond merely motor-execution related activity, suggesting a role of these regions in accent imitation ( Adank et al., 2013 ), vocal affect differentiation ( Bestelmeyer et al., 2014 ), and categorization of different prosodic contours (statement vs question) ( Sammler et al., 2015 ). Further, our results show that the regions which discriminate different syllable reports co-localize with areas activated during passive listening (in the left and right STG, in the left vMC, in the left inferior frontal operculum and the left AI). ...
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Which processes in the human brain lead to the categorical perception of speech sounds? Investigation of this question is hampered by the fact that categorical speech perception is normally confounded by acoustic differences in the stimulus. By using ambiguous sounds, however, it is possible to dissociate acoustic from perceptual stimulus representations. Twenty-seven normally hearing individuals took part in an fMRI study in which they were presented with an ambiguous syllable (intermediate between /da/ and /ga/) in one ear and with disambiguating acoustic feature (third formant, F3) in the other ear. Multi-voxel pattern searchlight analysis was used to identify brain areas that consistently differentiated between response patterns associated with different syllable reports. By comparing responses to different stimuli with identical syllable reports and identical stimuli with different syllable reports, we disambiguated whether these regions primarily differentiated the acoustics of the stimuli or the syllable report. We found that BOLD activity patterns in left perisylvian regions (STG, SMG), left inferior frontal regions (vMC, IFG, AI), left supplementary motor cortex (SMA/pre-SMA), and right motor and somatosensory regions (M1/S1) represent listeners’ syllable report irrespective of stimulus acoustics. Most of these regions are outside of what is traditionally regarded as auditory or phonological processing areas. Our results indicate that the process of speech sound categorization implicates decision-making mechanisms and auditory-motor transformations.
... In particular, the modulation of pitch in speech and singing has been suggested to be primarily controlled via the dorsal laryngeal motor cortex (Dichter et al., 2018;Eichert et al., 2020). And causal inference with transcranial magnetic stimulation demonstrated, for example, that in particular the laryngeal representation in the right hemisphere is involved in vocal pitch regulation (Finkel et al., 2019) and auditory pitch discrimination (Sammler et al., 2015). However, currently the ground truth of the anatomical connectivity of LMC to laryngeal motor neurons and cortical and subcortical brain areas results from tracing studies of a single cortical motor representation in mammals and humans (Kuypers, 1958;Kirzinger and Jürgens, 1982;Simonyan and Jürgens, 2002;Simonyan, 2014). ...
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Recent studies have identified two distinct cortical representations of voice control in humans, the ventral and the dorsal laryngeal motor cortex. Strikingly, while persistent developmental stuttering has been linked to a white matter deficit in the ventral laryngeal motor cortex, intensive fluency shaping intervention modulated the functional connectivity of the dorsal laryngeal motor cortical network. Currently, it is unknown whether the underlying structural network organization of these two laryngeal representations is distinct or differently shaped by stuttering intervention. Using probabilistic diffusion tractography in 22 individuals who stutter and participated in a fluency shaping intervention, in 18 individuals who stutter and did not participate in the intervention, and in 28 control participants, we here compare structural networks of the dorsal laryngeal motor cortex and the ventral laryngeal motor cortex and test intervention-related white matter changes. We show (i) that all participants have weaker ventral laryngeal motor cortex connections compared to the dorsal laryngeal motor cortex network, regardless of speech fluency, (ii) connections of the ventral laryngeal motor cortex were stronger in fluent speakers, (iii) the connectivity profile of the ventral laryngeal motor cortex predicted stuttering severity, (iv) but the ventral laryngeal motor cortex network is resistant to a fluency shaping intervention. Our findings substantiate a weaker structural organization of the ventral laryngeal motor cortical network in developmental stuttering and imply that assisted recovery supports neural compensation rather than normalization. Moreover, the resulting dissociation provides evidence for functionally segregated roles of the ventral laryngeal motor cortical and dorsal laryngeal motor cortical networks.
... The dual-stream model of language processing, one of the most used neuropsychological models of speech and language organization (Kreisler et al., 2000;Poeppel, 2004, 2007;Fridriksson et al., 2016), has been used frequently as the classic template (Kümmerer et al., 2013;Fridriksson et al., 2016;Zündorf et al., 2016;Goucha et al., 2017;McKinnon et al., 2018;Northam et al., 2018) in order to evaluate language dysfunction concerning brain damage in PSA. This model describes two large-scale processing streams, namely, the dorsal stream is crucial for producing fluent speech and auditory-motor integration processes, and the ventral stream supports the mapping between sound and meaning and thus it is related to auditory comprehension (Saur et al., 2008;Chang et al., 2015;Sammler et al., 2015;Fridriksson et al., 2016;Garrod and Pickering, 2016). Most previous studies mainly focused on the location and extension of the brain damage after stroke, and less attention was paid to the differences in the native languages (Anglade et al., 2014). ...
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The role of the right hemisphere (RH) in post-stroke aphasia (PSA) has not been completely understood. In general, the language alterations in PSA are normally evaluated from the perspective of the language processing models developed from Western languages such as English. However, the successful application of the models for assessing Chinese-language functions in patients with PSA has not been reported. In this study, the features of specific language-related lesion distribution and early variations of structure in RH in Chinese patients with PSA were investigated. Forty-two aphasic patients (female: 13, male: 29, mean age: 58 ± 12 years) with left hemisphere (LH) injury between 1 and 6 months after stroke were included. The morphological characteristics, both at the levels of gray matter (GM) and white matter (WM), were quantified by 3T multiparametric brain MRI. The Fridriksson et al.’s dual-stream model was used to compare language-related lesion regions. Voxel-based lesion-symptom mapping (VLSM) analysis has been performed. Our results showed that lesions in the precentral, superior frontal, middle frontal, and postcentral gyri were responsible for both the production and comprehension dysfunction of Chinese patients with PSA and were quite different from the lesions described by using the dual-stream model of Fridriksson et al. Furthermore, gray matter volume (GMV) was found significantly decreased in RH, and WM integrity was disturbed in RH after LH injury in Chinese patients with PSA. The different lesion patterns between Chinese patients with PSA and English-speaking patients with PSA may indicate that the dual-stream model of Fridriksson et al. is not suitable for the assessment of Chinese-language functions in Chinese patients with PSA in subacute phase of recovery. Moreover, decreased structural integrity in RH was found in Chinese patients with PSA.
... Also note that at the group level, the distinction between these clusters is much more clearly delineated in analyses at a strict level of significance (FWE-corrected, p<0.05) than at less strict significance levels (Fig. 3). Furthermore, the homotopic right hemispheric IFC regions, which are also assumed to be organized in a dual-stream architecture (45), show a different dynamic across filter sizes and significance levels. If we had analyzed the data at the usual singlefilter level, say, at the SPM standard 8 mm filter, and reported (and/or based our conclusions upon) the analysis on a strict significance level, the dual-stream architecture of right hemispheric speech processing would have remained unnoticed. ...
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In the past three decades, neuroimaging has provided important insights into structure-function relationships in the human brain. Recently, however, the methods for analyzing functional magnetic resonance imaging (fMRI) data have come under scrutiny, with studies questioning cross-software comparability, the validity of statistical inference and interpretation, and the influence of the spatial filter size on neuroimaging analyses. As most fMRI studies only use a single filter for analysis, much information on the size and shape of the BOLD signal in Gaussian scale space remains hidden and constrains the interpretation of fMRI studies. To investigate the influence of the spatial observation scale on fMRI analysis, we use a spatial multiscale analysis with a range of Gaussian filters from 1-20 mm (full width at half maximum) to analyze fMRI data from a speech repetition paradigm in 25 healthy subjects. We show that analyzing the fMRI data over a range of Gaussian filter kernels reveals substantial variability in the neuroanatomical localization and the average signal strength and size of suprathreshold clusters depending on the filter size. We also demonstrate how small spatial filters bias the results towards subcortical and cerebellar clusters. Furthermore, we describe substantially different scale-dependent cluster size dynamics between cortical and cerebellar clusters. We discuss how spatial multiscale analysis may substantially improve the interpretation of fMRI data. We propose to further develop a spatial multiscale analysis to fully explore the deep structure of the BOLD signal in Gaussian scale space.
... The lefthemispheric ventral pathway is involved in processing semantic content ( DeWitt and Rauschecker, 2012 ; Pylkkänen, 2019 ; Rauschecker and Scott, 2009 ) and local syntactic structure building ( Friederici et al., 2006 ), while the dorsal route is linked to speech production ( Hickok and Poeppel, 2007 ;Rauschecker and Scott, 2009 ) and the building of complex syntactic structures ( Wilson et al., 2011 ). Right-hemispheric ventral and dorsal pathways are both involved in processing speech prosody ( Fruhholz et al., 2015 ;Sammler et al., 2015 ). Furthermore, the integrity of the right-hemispheric dorsal pathway has been found to be crucial for building linguistic prosodic structure ( Sammler et al., 2018 ). ...
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Chunking language has been proposed to be vital for comprehension enabling the extraction of meaning from a continuous stream of speech. However, neurocognitive mechanisms of chunking are poorly understood. The present study investigated neural correlates of chunk boundaries intuitively identified by listeners in natural speech drawn from linguistic corpora using magneto- and electroencephalography (MEEG). In a behavioral experiment, subjects marked chunk boundaries in the excerpts intuitively, which revealed highly consistent chunk boundary markings across the subjects. We next recorded brain activity to investigate whether chunk boundaries with high and medium agreement rates elicit distinct evoked responses compared to non-boundaries. Pauses placed at chunk boundaries elicited a closure positive shift with the sources over bilateral auditory cortices. In contrast, pauses placed within a chunk were perceived as interruptions and elicited a biphasic emitted potential with sources located in the bilateral primary and non-primary auditory areas with right-hemispheric dominance, and in the right inferior frontal cortex. Furthermore, pauses placed at stronger boundaries elicited earlier and more prominent activation over the left hemisphere suggesting that brain responses to chunk boundaries of natural speech can be modulated by the relative strength of different linguistic cues, such as syntactic structure and prosody.
... However, despite a left-hemispheric dominance, the right hemisphere also contributes to language (Pujol et al., 1999). For instance, several right-hemispheric regions, including the inferior frontal gyrus and premotor cortex as well as posterior temporal gyrus and sulcus play a role in prosody processing (Sammler et al., 2015). In summary, while previous work strongly argues for a lefthemispheric dominance of language, the right hemisphere also makes a substantial contribution to language function. ...
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Language is a cognitive function that is asymmetrically distributed across both hemispheres, with left dominance for most linguistic operations. One key question of interest in cognitive neuroscience studies is related to the contribution of both hemispheres in bilingualism. Previous work shows a difference of both hemispheres for auditory processing of emotional and non-emotional words in bilinguals and monolinguals. In this study, we examined the differences between both hemispheres in the processing of emotional and non-emotional words of mother tongue language and foreign language. Sixty university students with Persian mother tongue and English as their second language were included. Differences between hemispheres were compared using the dichotic listening test. We tested the effect of hemisphere, language and emotion and their interaction. The right ear (associated with the left hemisphere) showed an advantage for the processing of all words in the first language, and positive words in the second language. Overall, our findings support previous studies reporting left-hemispheric dominance in late bilinguals for processing auditory stimuli.
... One of these, the right-hemisphere hypothesis, postulates that emotion is processed predominantly in the right hemisphere (Rohr et al., 2013;Gainotti, 2018). Emotional prosodic processing is also preferentially right-lateralized (Kotz et al., 2006;Sammler et al., 2015). Another hypothesis, the valence hypothesis, suggests that both hemispheres are involved in emotion processing (Rohr et al., 2013). ...
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Although the primary role of the auditory cortical areas is to process actual sounds, these areas are also activated by tasks that process imagined music, suggesting that the auditory cortical areas are involved in the processes underlying musical imagery. However, the mechanism by which these areas are involved in such processes is unknown. To elucidate this feature of the auditory cortical areas, we analyzed their functional networks during imagined music performance in comparison with those in the resting condition. While imagined music performance does not produce any musical sounds, the participants heard the same actual sounds from the MRI equipment in both experimental conditions. Therefore, if the functional connectivity between these conditions differs significantly, one can infer that the auditory cortical areas are actively involved in imagined music performance. Our functional connectivity analysis revealed a significant enhancement in the auditory network during imagined music performance relative to the resting condition. The reconfiguration profile of the auditory network showed a clear right-lateralized increase in the connectivity of the auditory cortical areas with brain regions associated with cognitive, memory, and emotional information processing. On the basis of these results, we hypothesize that auditory cortical areas and their networks are actively involved in imagined music performance through the integration of auditory imagery into mental imagery associated with music performance.
... The dorsal pathway is confined to the left hemisphere and is responsible for transformation of time-dependent speech signals into a motor format. The ventral pathway is organized more bilaterally and converts complex auditory features (e.g., prosody) into time-invariant speech representations [144,145]. The stronger lateralization and frequency dependency of the dorsal route further supports the prediction of the magnocellular pathway involvement in the impaired perception and motor computation of speech with intact prosody in DD. ...
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To construct a coherent multi-modal percept, vertebrate brains extract low-level features (such as spatial and temporal frequencies) from incoming sensory signals. However, because frequency processing is lateralized with the right hemisphere favouring low frequencies while the left favours higher frequencies, this introduces asymmetries between the hemispheres. Here, we describe how this lateralization shapes the development of several cognitive domains, ranging from visuo-spatial and numerical cognition to language, social cognition, and even aesthetic appreciation, and leads to the emergence of asymmetries in behaviour. We discuss the neuropsychological and educational implications of these emergent asymmetries and suggest future research approaches.
Article
People with right hemisphere brain lesions (PRHL) are often faced with impairments in their communicative abilities, which can impact their participation. These impairments may be related to diminished executive functions and/or reduced attentional or socio-cognitive capacities. PRHL can also be a!ected by deficits in processing of linguistic prosody. Linguistic prosody refers to prosodic information that primarily fulfils linguistic functions, such as marking syntactic structure by prosodic boundaries. These boundaries are realized by changes of the pitch contour, lengthening of segments, or the insertion or lengthening of pauses in the speech signal. In the present study, the receptive and productive prosodic abilities of a group of PRHL were investigated. They had to identify, read aloud, or repeat prosodic boundaries in coordinate three-name sequences with or without an internal prosodic boundary. We found heterogeneous patterns in all tasks. A large number of participants showed impairments in prosodic processing – either in one modality or in both modalities (perception, production). Some participants showed no or only minor impairments. The data are discussed with respect to previous literature on prosodic processing in PRHL. We consider it important for speech and language therapists to individually assess prosodic abilities in PRHL.
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The mismatch negativity (MMN) is an event related brain potential (ERP) elicited by unpredicted sounds presented in a sequence of repeated auditory stimuli. The neural sources of the MMN have been previously attributed to a fronto-temporo-parietal network which crucially overlaps with the so-called auditory dorsal stream, involving inferior and middle frontal, inferior parietal, and superior and middle temporal regions. These cortical areas are structurally connected by the arcuate fasciculus (AF), a three-branch pathway supporting the feedback-feedforward loop involved in auditory-motor integration, auditory working memory, storage of acoustic templates, as well as comparison and update of those templates. Here, we characterized the individual differences in the white-matter macrostructural properties of the AF and explored their link to the electrophysiological marker of passive change detection gathered in a melodic multifeature MMN-EEG paradigm in 26 healthy young adults without musical training. Our results show that left fronto-temporal white-matter connectivity plays an important role in the pre-attentive detection of rhythm modulations within a melody. Previous studies have shown that this AF segment is also critical for language processing and learning. This strong coupling between structure and function in auditory change detection might be related to life-time linguistic (and possibly musical) exposure and experiences, as well as to timing processing specialization of the left auditory cortex. To the best of our knowledge, this is the first time in which the relationship between neurophysiological (EEG) and brain white-matter connectivity indexes using DTI-tractography are studied together. Thus, the present results, although still exploratory, add to the existing evidence on the importance of studying the constraints imposed on cognitive functions by the underlying structural connectivity.
Thesis
The present thesis reports on interdisciplinary attempts to elucidate the hitherto unexplored neural correlates of linguistic processing in conditions of non-experimental, natural overt speech production. To this end, the author of this thesis, supported by colleagues, built up a multimodal neurolinguistic corpus (“The Freiburg/First Neurolinguistic Corpus”), composed of synchronized audio, video, electrocorticographic (ECoG) materials and linguistic annotations on different levels of linguistic abstraction. This corpus allowed us to study the neural effects related to several aspects of natural language, each of whom was treated in a separate study. Study 1 was dedicated to prosody and addressed the neural activity related to the production of the focus accent, Study 2 was dedicated to questions related to the neural representation of word complexity, and Study 3 investigated the syntactic processing accompanying natural clause production. The psycholinguistic methods we used consisted of (i) application of a matching procedure to select controlled word categories out of the natural language data (Study 1), (ii) orthogonalization of the linguistic parameters with the help of a linear regression model to overcome the problem of collinearity between correlated linguistic parameters, and (iii) the usage of a principal component analysis to extract most informative components of the linguistic material. The neuroscientific approach consisted either of group comparisons, in which neural effects underlying linguistically distinctive groups of words were compared (Study 1) or of correlation of neural activity with individual linguistic parameters (Study 2) and with principal components explaining most of the variances in the linguistic data (Study 3). We were interested in neural effects reflecting differences in the effort related to the production of speech units of different linguistic complexity. We were looking for neural effects which would be spatially focalized and which would be manifested in gamma activity (>35 Hz), since it known as a reliable and functionally specific marker of event-related effort. Based on knowledge from previous research, we were expecting neural effects to be spatially focalized, extended in time and extended over a broad range of gamma frequencies. We were, indeed, able to observe anatomically local effects. In Study 1, activity in postcentral areas proved to convey information about prosodic properties of content words. In Study 2, the proportional relation between the number of consonants and vowels, which was the most informative parameter with regard to the neural representation of word complexity, showed effects in two anatomically focal areas: the frontal one was located at the junction of the premotor cortex, the prefrontal cortex, and the posterior portion of Broca’s area (Brodmann area 44). The postcentral one lay directly above the lateral sulcus and comprised effects on the ventral central sulcus, in the parietal operculum and in the adjacent inferior parietal cortex. The preliminary findings of Study 3 indicate that pericentral cortical areas implicated in mouth motor functions show correlations with syntax-relevant information, reflected in the first principal component explaining over 30% of the variances in the linguistic data. While Study 3 yielded temporo-frequentially extended effects in high gamma frequencies, as we had expected, Studies 1 and 2 showed temporally and frequentially narrow effects with little reproducibility in terms of these neural characteristics. This may indicate moderate representation of the phenomena investigated in Studies 1 and 2 in the investigated neural signals. Alternatively, since the spectrum of gamma frequencies is a composite phenomenon relying on multiple cell types, it is also conceivable that these temporo-frequentially narrow effects may point to the functionally specific activation of small local populations of neurons, whose signal properties vary between linguistic parameters and subjects. Since we are not aware of published ECoG works investigating neural effects of linguistic processing during natural speech production, further validation of these observations and speculations is required. Beyond the here summarized findings, the overall contribution of this work to the field of neurolinguistics is that we have developed ways to study the neural effects related to natural language production, to obtain control over potentially confounding parameters in such data, and to surmount the problem of collinearity between multiple linguistic features of the neurolinguistic material.
Article
Musical training is associated with increased structural and functional connectivity between auditory sensory areas and higherorder brain networks involved in speech and motor processing. Whether such changed connectivity patterns facilitate the cortical propagation of speech information in musicians remains poorly understood. We here used magnetoencephalography (MEG) source imaging and a novel seed-based intersubject phase-locking approach to investigate the effects of musical training on the interregional synchronization of stimulus-driven neural responses during listening to naturalistic continuous speech presented in silence. MEG data were obtained from 20 young human subjects (both sexes) with different degrees of musical training. Our data show robust bilateral patterns of stimulus-driven interregional phase synchronization between auditory cortex and frontotemporal brain regions previously associated with speech processing. Stimulus-driven phase locking was maximal in the delta band, but was also observed in the theta and alpha bands. The individual duration of musical training was positively associated with the magnitude of stimulus-driven alpha-band phase locking between auditory cortex and parts of the dorsal and ventral auditory processing streams. These findings provide evidence for a positive relationship between musical training and the propagation of speech-related information between auditory sensory areas and higher-order processing networks, even when speech is presented in silence. We suggest that the increased synchronization of higher-order cortical regions to auditory cortex may contribute to the previously described musician advantage in processing speech in background noise.
Article
Auditory sentence comprehension involves processing content (semantics), grammar (syntax), and intonation (prosody). The left inferior frontal gyrus (IFG) is involved in sentence comprehension guided by these different cues, with neuroimaging studies preferentially locating syntactic and semantic processing in separate IFG subregions. However, this regional specialisation has not been confirmed with a neurostimulation method. Consequently, the causal role of such a specialisation remains unclear. This study probed the role of the posterior IFG (pIFG) for syntactic processing and the anterior IFG (aIFG) for semantic processing with repetitive transcranial magnetic stimulation (rTMS) in a task that required the interpretation of the sentence's prosodic realisation. Healthy participants performed a sentence completion task with syntactic and semantic decisions, while receiving 10 Hz rTMS over either left aIFG, pIFG, or vertex (control). Initial behavioural analyses showed an inhibitory effect on accuracy without task‐specificity. However, electric field simulations revealed differential effects for both subregions. In the aIFG, stronger stimulation led to slower semantic processing, with no effect of pIFG stimulation. In contrast, we found a facilitatory effect on syntactic processing in both aIFG and pIFG, where higher stimulation strength was related to faster responses. Our results provide first evidence for the functional relevance of left aIFG in semantic processing guided by intonation. The stimulation effect on syntactic responses emphasises the importance of the IFG for syntax processing, without supporting the hypothesis of a pIFG‐specific involvement. Together, the results support the notion of functionally specialised IFG subregions for diverse but fundamental cues for language processing. We investigated using transcranial magnetic stimulation whether subregions of the inferior frontal gyrus (IFG) underlie specific subdomains of language processing. The results suggest a domain‐specific role for the posterior IFG for grammatical processing, and a cross‐domain role for anterior IFG across language functions.
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Whether reading in different writing systems recruits language-unique or language-universal neural processes is a long-standing debate. Many studies have shown the left Arcuate Fasciculus (AF) to be involved in phonological and reading processes. In contrast, little is known about the role of the right AF in reading, but some have suggested that it may play a role in visual spatial aspects of reading or the prosodic components of language. The right AF may be more important for reading in Chinese due to its logographic and tonal properties, but this hypothesis has yet to be tested. We recruited a group of Chinese-English bilingual children (8.2 to 12.0 years old) to explore the common and unique relation of reading skill in English and Chinese to Fractional Anisotropy (FA) in the bilateral AF. We found that both English and Chinese reading skills were positively correlated with FA in the rostral part of the left AF-direct segment. Additionally, English reading skill was positively correlated with FA in the caudal part of the left AF-direct segment, which was also positively correlated with phonological awareness. In contrast, Chinese reading skill was positively correlated with FA in certain segments of the right AF, which was positively correlated with visual spatial ability, but not tone discrimination ability. Our results suggest that there are language universal substrates of reading across languages, but that certain left AF nodes support phonological mechanisms important for reading in English, whereas certain right AF nodes support visual spatial mechanisms important for reading in Chinese.
Article
Primary Objective We hypothesized that, in patients with acute severe traumatic brain injury (TBI) who recover basic language function, speech-evoked blood-oxygen-level-dependent (BOLD) functional MRI (fMRI) responses within the canonical language network increase over the first 6 months post-injury. Research Design We conducted a prospective, longitudinal fMRI pilot study of adults with acute severe TBI admitted to the intensive care unit. We also enrolled age- and sex-matched healthy subjects. Methods and Procedures We evaluated BOLD signal in bilateral superior temporal gyrus (STG) and inferior frontal gyrus (IFG) regions of interest acutely and approximately 6 months post-injury. Given evidence that regions outside the canonical language network contribute to language processing, we also performed exploratory whole-brain analyses. Main Outcomes and Results Of the 16 patients enrolled, eight returned for follow-up fMRI, all of whom recovered basic language function. We observed speech-evoked longitudinal BOLD increases in the left STG, but not in the right STG, right IFG, or left IFG. Whole-brain analysis revealed increases in the right supramarginal and middle temporal gyri but no differences between patients and healthy subjects (n = 16). Conclusion This pilot study suggests that, in patients with severe TBI who recover language function,speech-evoked BOLD responses in bihemispheric language-processing cortex reemerge by 6 months post-injury.
Chapter
Music and language processing have been repeatedly compared but similarities and differences between domains are challenging to quantify. This chapter takes a step back and focuses specifically on the role of fine-grained changes in pitch, which play a role in both domains but are not widely studied. In addition to describing the units, we provide empirical evidence for the specific role of small units in music: scoops, which are small dynamic pitch change at the start or end of sung notes within a melody. We report results from a new experiment that builds on a recent study that addressed two distinct processes for the evaluation of pitch accuracy (Larrouy-Maestri & Pfordresher, 2018). The present study compared accuracy ratings to a more ecologically valid listening task: preference judgments. By replicating and extending previous findings, we describe the processing of small units in music perception and propose research directions to further investigate such units in speech perception, and ultimately gain the necessary insight to make meaningful cross-domain comparisons.
Article
The present study aimed to examine the perception of music and prosody in patients who had undergone a severe traumatic brain injury (TBI). Our second objective was to describe the association between music and prosody impairments in clinical individual presentations. Thirty-six patients who were out of the acute phase underwent a set of music and prosody tests: two subtests of the Montreal Battery for Evaluation of Amusia evaluating respectively melody (scale) and rhythm perception, two subtests of the Montreal Evaluation of Communication on prosody understanding in sentences, and two other tests evaluating prosody understanding in vowels. Forty-two percent of the patients were impaired in the melodic test, 51% were impaired in the rhythmic test, and 71% were impaired in at least one of the four prosody tests. The amusic patients performed significantly worse than non-amusics on the four prosody tests. This descriptive study shows for the first time the high prevalence of music deficits after severe TBI. It also suggests associations between prosody and music impairments, as well as between linguistic and emotional prosody impairments. Causes of these impairments remain to be explored.
Article
The polyvagal theory introduces the premise that the autonomic nervous system (ANS) has an essential role in detecting threats and calming social engagement. In busy courtrooms, the subtle nuances of communication can be missed or overlooked. Messages of threat activate an autonomic state from calm to agitated beneath conscious awareness. When shifts occur, information can be misunderstood or misinterpreted, resulting in overly defensive responses from all parties. The polyvagal theory suggests a perspective that encourages self‐reflection and awareness to counterbalance the inevitable physical and mental stress of managing todays’ courtrooms.
Article
Congenital amusia is a neurogenetic disorder of pitch perception that may also compromise pitch production. Despite amusics’ long documented difficulties with pitch, previous evidence suggests that familiar music may have an implicit facilitative effect on their performance. It remains, however, unknown whether vocal imitation of song in amusia is influenced by melody familiarity and the presence of lyrics. To address this issue, thirteen Mandarin speaking amusics and 13 matched controls imitated novel song segments with lyrics and on the syllable /la/. Eleven out of these participants in each group also imitated segments of a familiar song. Subsequent acoustic analysis was conducted to measure pitch and timing matching accuracy based on eight acoustic measures. While amusics showed worse imitation performance than controls across seven out of the eight pitch and timing measures, melody familiarity was found to have a favorable effect on their performance on three pitch-related acoustic measures. The presence of lyrics did not affect either group’s performance substantially. Correlations were observed between amusics’ performance on the Montreal Battery of Evaluation of Amusia and imitation of the novel song. We discuss implications in terms of music familiarity, memory demands, the relevance of lexical information, and the link between perception and production.
Article
Functional magnetic resonance imaging (fMRI) studies have revealed a functional reorganization in patients with sensorineural hearing loss (SNHL). The structural basement of functional changes has also been investigated recently. Graph theory analysis brings a new understanding of the structural connectome and topological features in central neural system diseases. However, little is known about the structural network connectome changes in SNHL patients, especially in children. We explored the differences in topologic organization, rich-club organization, and structural connection between children with congenital bilateral profound SNHL and normal hearing under the age of three using graph theory analysis and probabilistic tractography. Compared with the normal-hearing (NH) group, the SNHL group showed no difference in global and nodal topological parameters. Increased structural connection strength were found in the right cortico-striatal-thalamus-cortical circuity. Decreased cross-hemisphere connections were found between the right precuneus and the left auditory cortex as well as the left subcortical regions. Rich-club organization analysis found increased local connection in the SNHL group. These results revealed structural organizations after hearing deprivation in congenital bilateral profound SNHL children.
Thesis
L’hémisphère non dominant supporte la cognition visuo-spatiale et sociale. Sont atteinte est à l’origine du syndrome d’héminégligence. Les objectifs préliminaires de cette thèse étaient (1) de proposer une taxonomie des fonctions cérébrales supportées par l’hémisphère droit ; (2) d’harmoniser les données d’anatomie fonctionnelle avec les données neuropsychologiques et de sémiologie clinique de ces fonctions. (3) d’harmoniser la nomenclature anatomique entre les études de dissections et d’imagerie de l’hémisphère droit. (4) d’harmoniser la nomenclature anatomique du principal faisceau de substance blanche : le faisceau arqué. Le principal but de cette thèse était (5) d’approfondir nos connaissances sur l’anatomie du réseau ventral de l’attention, qui est un réseau fonctionnel cortical identifié par l’IRM de repos, impliqué dans la cognition visuo-spatiale. En traitant les données d’IRM fonctionnelle de repos de 50 sujets, nous avons pu décrire les localisations corticales du réseau ventral de l’attention dans notre cohorte de sujets sains. Nous avons ensuite utilisé les zones corticales du réseau ventral de l’attention comme régions germes, pour l’étude des fibres blanches in-vivo (tractographie), ainsi que sur des cerveaux par méthode de Klingler (ex-vivo). Nous avons pu ainsi définir les faisceaux de fibres blanches du réseau ventral de l’attention. Ces faisceaux comprennent le faisceau longitudinal dans sa troisième portion (SLF III) et le faisceau arqué droits. Le développement d’un test neuropsychologique spécifique stimulant le réseau ventral de l’attention reste à concrétiser pour pouvoir réaliser cette chirurgie éveillée. Dans ce sens, nous avons publié des travaux (6) visant à l’utilisation des lunettes de réalité virtuelle en condition de chirurgie éveillée, pour réaliser de nouveaux tests neuropsychologiques dédiés à l’exploration des fonctions cognitives de l’hémisphère droit. Ce travail de Thèse a été valorisé par 7 publications, 1 article soumis et le développement d’un test neuropsychologique utilisant la réalité virtuelle en chirurgie éveillée cérébrale.
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Speech and language are considered uniquely human abilities: animals have communication systems, but they do not match human linguistic skills in terms of recursive structure and combinatorial power. Yet, in evolution, spoken language must have emerged from neural mechanisms at least partially available in animals. In this paper, we will demonstrate how our understanding of speech perception, one important facet of language, has profited from findings and theory in nonhuman primate studies. Chief among these are physiological and anatomical studies showing that primate auditory cortex, across species, shows patterns of hierarchical structure, topographic mapping and streams of functional processing. We will identify roles for different cortical areas in the perceptual processing of speech and review functional imaging work in humans that bears on our understanding of how the brain decodes and monitors speech. A new model connects structures in the temporal, frontal and parietal lobes linking speech perception and production.
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Built on an analogy between the visual and auditory systems, the following dual stream model for language processing was suggested recently: a dorsal stream is involved in mapping sound to articulation, and a ventral stream in mapping sound to meaning. The goal of the study presented here was to test the neuroanatomical basis of this model. Combining functional magnetic resonance imaging (fMRI) with a novel diffusion tensor imaging (DTI)-based tractography method we were able to identify the most probable anatomical pathways connecting brain regions activated during two prototypical language tasks. Sublexical repetition of speech is subserved by a dorsal pathway, connecting the superior temporal lobe and premotor cortices in the frontal lobe via the arcuate and superior longitudinal fascicle. In contrast, higher-level language comprehension is mediated by a ventral pathway connecting the middle temporal lobe and the ventrolateral prefrontal cortex via the extreme capsule. Thus, according to our findings, the function of the dorsal route, traditionally considered to be the major language pathway, is mainly restricted to sensory-motor mapping of sound to articulation, whereas linguistic processing of sound to meaning requires temporofrontal interaction transmitted via the ventral route. • DTI • extreme capsule • fMRI • language networks • arcuate fascicle • extreme capsule
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Experimental and imaging studies in monkeys have outlined various long association fiber bundles within the temporoparietal region. In the present study the trajectory of the middle longitudinal fascicle (MdLF) has been delineated in 4 human subjects using diffusion tensor magnetic resonance imaging segmentation and tractography. The MdLF seems to extend from the inferior parietal lobule (IPL), specifically the angular gyrus, to the temporal pole remaining within the white matter of the superior temporal gyrus (STG). Comparison of the superior longitudinal fascicle II-arcuate fascicle (SLF II-AF) with the MdLF in the same subjects revealed that MdLF is located in a medial and caudal position relative to SLF II-AF and that it extends more rostrally. Given the location of MdLF between the IPL (angular gyrus) and the STG, it is suggested that MdLF could have a role in language and attention functions.
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Social interaction relies on the ability to react to communication signals. Although cortical sensory-motor "mirror" networks are thought to play a key role in visual aspects of primate communication, evidence for a similar generic role for auditory-motor interaction in primate nonverbal communication is lacking. We demonstrate that a network of human premotor cortical regions activated during facial movement is also involved in auditory processing of affective nonverbal vocalizations. Within this auditory-motor mirror network, distinct functional subsystems respond preferentially to emotional valence and arousal properties of heard vocalizations. Positive emotional valence enhanced activation in a left posterior inferior frontal region involved in representation of prototypic actions, whereas increasing arousal enhanced activation in presupplementary motor area cortex involved in higher-order motor control. Our findings demonstrate that listening to nonverbal vocalizations can automatically engage preparation of responsive orofacial gestures, an effect that is greatest for positive-valence and high-arousal emotions. The automatic engagement of responsive orofacial gestures by emotional vocalizations suggests that auditory-motor interactions provide a fundamental mechanism for mirroring the emotional states of others during primate social behavior. Motor facilitation by positive vocal emotions suggests a basic neural mechanism for establishing cohesive bonds within primate social groups.
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Despite decades of research, the functional neuroanatomy of speech processing has been difficult to characterize. A major impediment to progress may have been the failure to consider task effects when mapping speech-related processing systems. We outline a dual-stream model of speech processing that remedies this situation. In this model, a ventral stream processes speech signals for comprehension, and a dorsal stream maps acoustic speech signals to frontal lobe articulatory networks. The model assumes that the ventral stream is largely bilaterally organized--although there are important computational differences between the left- and right-hemisphere systems--and that the dorsal stream is strongly left-hemisphere dominant.
Chapter
Evidence indicates that speech movements are planned to follow auditory trajectories. This is followed by a description of the Directions Into Velocities of Articulators (DIVA) model, which provides a detailed account of the role of auditory feedback in speech motor development and control. A brief description of the higher-order brain areas involved in speech sequencing (including the pre-SMA and inferior frontal sulcus) is then provided, followed by a description of the Hierarchical State Feedback Control (HSFC) model, which posits internal error detection and correction processes that can detect and correct speech production errors prior to articulation. The chapter closes with a treatment of promising future directions of research into auditory-motor interactions in speech, including the use of intracranial recording techniques such as electrocorticography in humans, the investigation of the potential roles of various large-scale brain rhythms in speech perception and production, and the development of brain–computer interfaces that use auditory feedback to allow profoundly paralyzed users to learn to produce speech using a speech synthesizer.
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Recent neurophysiological studies of speaking are beginning to elucidate the neural mechanisms underlying auditory feedback processing during vocalizations. Here we review how research findings impact our state feedback control (SFC) model of speech motor control. We will discuss the evidence for cortical computations that compare incoming feedback with predictions derived from motor efference copy. We will also review observations from auditory feedback perturbation studies that demonstrate clear evidence for a state estimate correction process, which drives compensatory motor behavioral responses. While there is compelling support for cortical computations in the SFC model, there are still several outstanding questions that await resolution by future neural investigations. Copyright © 2015 Elsevier Ltd. All rights reserved.
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This paper reviews the major findings and hypotheses to emerge in the literature concerned with speech prosody. Both production and perception of prosody are considered. Evidence from studies of patients with lateralized left or right hemisphere damage are presented, as well as relevant data from anatomical and functional imaging studies.
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Just as eyes are often considered a gateway to the soul, the human voice offers a window through which we gain access to our fellow human beings' minds - their attitudes, intentions and feelings. Whether in talking or singing, crying or laughing, sighing or screaming, the sheer sound of a voice communicates a wealth of information that, in turn, may serve the observant listener as valuable guidepost in social interaction. But how do human beings extract information from the tone of a voice? In an attempt to answer this question, the present article reviews empirical evidence detailing the cerebral processes that underlie our ability to decode emotional information from vocal signals. The review will focus primarily on two prominent classes of vocal emotion cues: laughter and speech prosody (i.e. the tone of voice while speaking). Following a brief introduction, behavioral as well as neuroimaging data will be summarized that allows to outline cerebral mechanisms associated with the decoding of emotional voice cues, as well as the influence of various context variables (e.g. co-occurring facial and verbal emotional signals, attention focus, person-specific parameters such as gender and personality) on the respective processes. Building on the presented evidence, a cerebral network model will be introduced that proposes a differential contribution of various cortical and subcortical brain structures to the processing of emotional voice signals both in isolation and in context of accompanying (facial and verbal) emotional cues.