Variability in white matter structure relates to hallucination proneness.

Hallucinations are a prominent transdiagnostic psychiatric symptom but are also prevalent in individuals who do not require clinical care. Moreover, persistent psychosis-like experience in otherwise healthy individuals may be related to an increased risk to transition to a psychotic disorder. This suggests a common etiology across clinical and non-clinical individuals along a multidimensional psychosis continuum that may be detectable in structural variations of the brain. The current diffusion tensor imaging study assessed 50 healthy individuals (35 females) to identify possible differences in white matter associated with hallucination proneness (HP). This approach circumvents potential confounds related to medication, hospitalization, and disease progression common in clinical individuals. We determined how HP relates to white matter structure in selected association, commissural, and projection fiber pathways putatively linked to psychosis. Increased HP was associated with enhanced fractional anisotropy (FA) in the right uncinate fasciculus, the right anterior and posterior arcuate fasciculus, and the corpus callosum. These findings support the notion of a psychosis continuum, providing first evidence of structural white matter variability associated with HP in healthy individuals. Furthermore, alterations in the targeted pathways likely indicate an association between HP-related structural variations and the putative salience and attention mechanisms that these pathways subserve.

An open-source toolbox for measuring vocal tract shape from real-time magnetic resonance images.

Real-time magnetic resonance imaging (rtMRI) is a technique that provides high-contrast videographic data of human anatomy in motion. Applied to the vocal tract, it is a powerful method for capturing the dynamics of speech and other vocal behaviours by imaging structures internal to the mouth and throat. These images provide a means of studying the physiological basis for speech, singing, expressions of emotion, and swallowing that are otherwise not accessible for external observation. However, taking quantitative measurements from these images is notoriously difficult. We introduce a signal processing pipeline that produces outlines of the vocal tract from the lips to the larynx as a quantification of the dynamic morphology of the vocal tract. Our approach performs simple tissue classification, but constrained to a researcher-specified region of interest. This combination facilitates feature extraction while retaining the domain-specific expertise of a human analyst. We demonstrate that this pipeline generalises well across datasets covering behaviours such as speech, vocal size exaggeration, laughter, and whistling, as well as producing reliable outcomes across analysts, particularly among users with domain-specific expertise. With this article, we make this pipeline available for immediate use by the research community, and further suggest that it may contribute to the continued development of fully automated methods based on deep learning algorithms.

Speech With Pauses Sounds Deceptive to Listeners With and Without Hearing Impairment.

PURPOSE: Communication is as much persuasion as it is the transfer of information. This creates a tension between the interests of the speaker and those of the listener, as dishonest speakers naturally attempt to hide deceptive speech and listeners are faced with the challenge of sorting truths from lies. Listeners with hearing impairment in particular may have differing levels of access to the acoustical cues that give away deceptive speech. A greater tendency toward speech pauses has been hypothesized to result from the cognitive demands of lying convincingly. Higher vocal pitch has also been hypothesized to mark the increased anxiety of a dishonest speaker. METHOD: Listeners with or without hearing impairments heard short utterances from natural conversations, some of which had been digitally manipulated to contain either increased pausing or raised vocal pitch. Listeners were asked to guess whether each statement was a lie in a two-alternative forced-choice task. Participants were also asked explicitly which cues they believed had influenced their decisions. RESULTS: Statements were more likely to be perceived as a lie when they contained pauses, but not when vocal pitch was raised. This pattern held regardless of hearing ability. In contrast, both groups of listeners self-reported using vocal pitch cues to identify deceptive statements, though at lower rates than pauses. CONCLUSIONS: Listeners may have only partial awareness of the cues that influence their impression of dishonesty. Listeners with hearing impairment may place greater weight on acoustical cues according to the differing degrees of access provided by hearing aids. SUPPLEMENTAL MATERIAL: https://doi.org/10.23641/asha.24052446.

Real-time magnetic resonance imaging reveals distinct vocal tract configurations during spontaneous and volitional laughter.

A substantial body of acoustic and behavioural evidence points to the existence of two broad categories of laughter in humans: spontaneous laughter that is emotionally genuine and somewhat involuntary, and volitional laughter that is produced on demand. In this study, we tested the hypothesis that these are also physiologically distinct vocalizations, by measuring and comparing them using real-time magnetic resonance imaging (rtMRI) of the vocal tract. Following Ruch and Ekman (Ruch and Ekman 2001 In Emotions, qualia, and consciousness (ed. A Kaszniak), pp. 426-443), we further predicted that spontaneous laughter should be relatively less speech-like (i.e. less articulate) than volitional laughter. We collected rtMRI data from five adult human participants during spontaneous laughter, volitional laughter and spoken vowels. We report distinguishable vocal tract shapes during the vocalic portions of these three vocalization types, where volitional laughs were intermediate between spontaneous laughs and vowels. Inspection of local features within the vocal tract across the different vocalization types offers some additional support for Ruch and Ekman's predictions. We discuss our findings in light of a dual pathway hypothesis for the neural control of human volitional and spontaneous vocal behaviours, identifying tongue shape and velum lowering as potential biomarkers of spontaneous laughter to be investigated in future research. This article is part of the theme issue 'Cracking the laugh code: laughter through the lens of biology, psychology and neuroscience'.

Hypersensitivity to passive voice hearing in hallucination proneness.

Voices are a complex and rich acoustic signal processed in an extensive cortical brain network. Specialized regions within this network support voice perception and production and may be differentially affected in pathological voice processing. For example, the experience of hallucinating voices has been linked to hyperactivity in temporal and extra-temporal voice areas, possibly extending into regions associated with vocalization. Predominant self-monitoring hypotheses ascribe a primary role of voice production regions to auditory verbal hallucinations (AVH). Alternative postulations view a generalized perceptual salience bias as causal to AVH. These theories are not mutually exclusive as both ascribe the emergence and phenomenology of AVH to unbalanced top-down and bottom-up signal processing. The focus of the current study was to investigate the neurocognitive mechanisms underlying predisposition brain states for emergent hallucinations, detached from the effects of inner speech. Using the temporal voice area (TVA) localizer task, we explored putative hypersalient responses to passively presented sounds in relation to hallucination proneness (HP). Furthermore, to avoid confounds commonly found in in clinical samples, we employed the Launay-Slade Hallucination Scale (LSHS) for the quantification of HP levels in healthy people across an experiential continuum spanning the general population. We report increased activation in the right posterior superior temporal gyrus (pSTG) during the perception of voice features that positively correlates with increased HP scores. In line with prior results, we propose that this right-lateralized pSTG activation might indicate early hypersensitivity to acoustic features coding speaker identity that extends beyond own voice production to perception in healthy participants prone to experience AVH.

Emotional salience but not valence impacts anterior cingulate cortex conflict processing.

Stimuli that evoke emotions are salient, draw attentional resources, and facilitate situationally appropriate behavior in complex or conflicting environments. However, negative and positive emotions may motivate different response strategies. For example, a threatening stimulus might evoke avoidant behavior, whereas a positive stimulus may prompt approaching behavior. Therefore, emotional stimuli might either elicit differential behavioral responses when a conflict arises or simply mark salience. The present study used functional magnetic resonance imaging to investigate valence-specific emotion effects on attentional control in conflict processing by employing an adapted flanker task with neutral, negative, and positive stimuli. Slower responses were observed for incongruent than congruent trials. Neural activity in the dorsal anterior cingulate cortex was associated with conflict processing regardless of emotional stimulus quality. These findings confirm that both negative and positive emotional stimuli mark salience in both low (congruent) and high (incongruent) conflict scenarios. Regardless of the conflict level, emotional stimuli deployed greater attentional resources in goal directed behavior.

Beyond language: The unspoken sensory-motor representation of the tongue in non-primates, non-human and human primates.

The English idiom "on the tip of my tongue" commonly acknowledges that something is known, but it cannot be immediately brought to mind. This phrase accurately describes sensorimotor functions of the tongue, which are fundamental for many tongue-related behaviors (e.g., speech), but often neglected by scientific research. Here, we review a wide range of studies conducted on non-primates, non-human and human primates with the aim of providing a comprehensive description of the cortical representation of the tongue's somatosensory inputs and motor outputs across different phylogenetic domains. First, we summarize how the properties of passive non-noxious mechanical stimuli are encoded in the putative somatosensory tongue area, which has a conserved location in the ventral portion of the somatosensory cortex across mammals. Second, we review how complex self-generated actions involving the tongue are represented in more anterior regions of the putative somato-motor tongue area. Finally, we describe multisensory response properties of the primate and non-primate tongue area by also defining how the cytoarchitecture of this area is affected by experience and deafferentation.

Speech timing cues reveal deceptive speech in social deduction board games.

The faculty of language allows humans to state falsehoods in their choice of words. However, while what is said might easily uphold a lie, how it is said may reveal deception. Hence, some features of the voice that are difficult for liars to control may keep speech mostly, if not always, honest. Previous research has identified that speech timing and voice pitch cues can predict the truthfulness of speech, but this evidence has come primarily from laboratory experiments, which sacrifice ecological validity for experimental control. We obtained ecologically valid recordings of deceptive speech while observing natural utterances from players of a popular social deduction board game, in which players are assigned roles that either induce honest or dishonest interactions. When speakers chose to lie, they were prone to longer and more frequent pauses in their speech. This finding is in line with theoretical predictions that lying is more cognitively demanding. However, lying was not reliably associated with vocal pitch. This contradicts predictions that increased physiological arousal from lying might increase muscular tension in the larynx, but is consistent with human specialisations that grant Homo sapiens sapiens an unusual degree of control over the voice relative to other primates. The present study demonstrates the utility of social deduction board games as a means of making naturalistic observations of human behaviour from semi-structured social interactions.

Individual differences in vocal size exaggeration.

The human voice carries socially relevant information such as how authoritative, dominant, and attractive the speaker sounds. However, some speakers may be able to manipulate listeners by modulating the shape and size of their vocal tract to exaggerate certain characteristics of their voice. We analysed the veridical size of speakers' vocal tracts using real-time magnetic resonance imaging as they volitionally modulated their voice to sound larger or smaller, corresponding changes to the size implied by the acoustics of their voice, and their influence over the perceptions of listeners. Individual differences in this ability were marked, spanning from nearly incapable to nearly perfect vocal modulation, and was consistent across modalities of measurement. Further research is needed to determine whether speakers who are effective at vocal size exaggeration are better able to manipulate their social environment, and whether this variation is an inherited quality of the individual, or the result of life experiences such as vocal training.

Singers show enhanced performance and neural representation of vocal imitation.

Humans have a remarkable capacity to finely control the muscles of the larynx, via distinct patterns of cortical topography and innervation that may underpin our sophisticated vocal capabilities compared with non-human primates. Here, we investigated the behavioural and neural correlates of laryngeal control, and their relationship to vocal expertise, using an imitation task that required adjustments of larynx musculature during speech. Highly trained human singers and non-singer control participants modulated voice pitch and vocal tract length (VTL) to mimic auditory speech targets, while undergoing real-time anatomical scans of the vocal tract and functional scans of brain activity. Multivariate analyses of speech acoustics, larynx movements and brain activation data were used to quantify vocal modulation behaviour and to search for neural representations of the two modulated vocal parameters during the preparation and execution of speech. We found that singers showed more accurate task-relevant modulations of speech pitch and VTL (i.e. larynx height, as measured with vocal tract MRI) during speech imitation; this was accompanied by stronger representation of VTL within a region of the right somatosensory cortex. Our findings suggest a common neural basis for enhanced vocal control in speech and song. This article is part of the theme issue 'Voice modulation: from origin and mechanism to social impact (Part I)'.

A dual larynx motor networks hypothesis.

Humans are vocal modulators par excellence. This ability is supported in part by the dual representation of the laryngeal muscles in the motor cortex. Movement, however, is not the product of motor cortex alone but of a broader motor network. This network consists of brain regions that contain somatotopic maps that parallel the organization in motor cortex. We therefore present a novel hypothesis that the dual laryngeal representation is repeated throughout the broader motor network. In support of the hypothesis, we review existing literature that demonstrates the existence of network-wide somatotopy and present initial evidence for the hypothesis' plausibility. Understanding how this uniquely human phenotype in motor cortex interacts with broader brain networks is an important step toward understanding how humans evolved the ability to speak. We further suggest that this system may provide a means to study how individual components of the nervous system evolved within the context of neuronal networks. This article is part of the theme issue 'Voice modulation: from origin and mechanism to social impact (Part I)'.

Paradoxical facilitation alongside interhemispheric inhibition.

Neurophysiological experiments using transcranial magnetic stimulation (TMS) have sought to probe the function of the motor division of the corpus callosum. Primary motor cortex sends projections via the corpus callosum with a net inhibitory influence on the homologous region of the opposite hemisphere. Interhemispheric inhibition (IHI) experiments probe this inhibitory pathway. A test stimulus (TS) delivered to the motor cortex in one hemisphere elicits motor evoked potentials (MEPs) in a target muscle, while a conditioning stimulus (CS) applied to the homologous region of the opposite hemisphere modulates the effect of the TS. We predicted that large CS MEPs would be associated with increased IHI since they should be a reliable index of how effectively contralateral motor cortex was stimulated and therefore of the magnitude of interhemispheric inhibition. However, we observed a strong tendency for larger CS MEPs to be associated with reduced interhemispheric inhibition which in the extreme lead to a net effect of facilitation. This surprising effect was large, systematic, and observed in nearly all participants. We outline several hypotheses for mechanisms which may underlie this phenomenon to guide future research.

Human larynx motor cortices coordinate respiration for vocal-motor control.

Vocal flexibility is a hallmark of the human species, most particularly the capacity to speak and sing. This ability is supported in part by the evolution of a direct neural pathway linking the motor cortex to the brainstem nucleus that controls the larynx the primary sound source for communication. Early brain imaging studies demonstrated that larynx motor cortex at the dorsal end of the orofacial division of motor cortex (dLMC) integrated laryngeal and respiratory control, thereby coordinating two major muscular systems that are necessary for vocalization. Neurosurgical studies have since demonstrated the existence of a second larynx motor area at the ventral extent of the orofacial motor division (vLMC) of motor cortex. The vLMC has been presumed to be less relevant to speech motor control, but its functional role remains unknown. We employed a novel ultra-high field (7T) magnetic resonance imaging paradigm that combined singing and whistling simple melodies to localise the larynx motor cortices and test their involvement in respiratory motor control. Surprisingly, whistling activated both 'larynx areas' more strongly than singing despite the reduced involvement of the larynx during whistling. We provide further evidence for the existence of two larynx motor areas in the human brain, and the first evidence that laryngeal-respiratory integration is a shared property of both larynx motor areas. We outline explicit predictions about the descending motor pathways that give these cortical areas access to both the laryngeal and respiratory systems and discuss the implications for the evolution of speech.

Expectancy changes the self-monitoring of voice identity.

Self-voice attribution can become difficult when voice characteristics are ambiguous, but functional magnetic resonance imaging (fMRI) investigations of such ambiguity are sparse. We utilized voice-morphing (self-other) to manipulate (un-)certainty in self-voice attribution in a button-press paradigm. This allowed investigating how levels of self-voice certainty alter brain activation in brain regions monitoring voice identity and unexpected changes in voice playback quality. FMRI results confirmed a self-voice suppression effect in the right anterior superior temporal gyrus (aSTG) when self-voice attribution was unambiguous. Although the right inferior frontal gyrus (IFG) was more active during a self-generated compared to a passively heard voice, the putative role of this region in detecting unexpected self-voice changes during the action was demonstrated only when hearing the voice of another speaker and not when attribution was uncertain. Further research on the link between right aSTG and IFG is required and may establish a threshold monitoring voice identity in action. The current results have implications for a better understanding of the altered experience of self-voice feedback in auditory verbal hallucinations.

Evolution of the speech-ready brain: The voice/jaw connection in the human motor cortex.

A prominent model of the origins of speech, known as the "frame/content" theory, posits that oscillatory lowering and raising of the jaw provided an evolutionary scaffold for the development of syllable structure in speech. Because such oscillations are nonvocal in most nonhuman primates, the evolution of speech required the addition of vocalization onto this scaffold in order to turn such jaw oscillations into vocalized syllables. In the present functional MRI study, we demonstrate overlapping somatotopic representations between the larynx and the jaw muscles in the human primary motor cortex. This proximity between the larynx and jaw in the brain might support the coupling between vocalization and jaw oscillations to generate syllable structure. This model suggests that humans inherited voluntary control of jaw oscillations from ancestral species, but added voluntary control of vocalization onto this via the evolution of a new brain area that came to be situated near the jaw region in the human motor cortex.

Whistling shares a common tongue with speech: bioacoustics from real-time MRI of the human vocal tract.

Most human communication is carried by modulations of the voice. However, a wide range of cultures has developed alternative forms of communication that make use of a whistled sound source. For example, whistling is used as a highly salient signal for capturing attention, and can have iconic cultural meanings such as the catcall, enact a formal code as in boatswain's calls or stand as a proxy for speech in whistled languages. We used real-time magnetic resonance imaging to examine the muscular control of whistling to describe a strong association between the shape of the tongue and the whistled frequency. This bioacoustic profile parallels the use of the tongue in vowel production. This is consistent with the role of whistled languages as proxies for spoken languages, in which one of the acoustical features of speech sounds is substituted with a frequency-modulated whistle. Furthermore, previous evidence that non-human apes may be capable of learning to whistle from humans suggests that these animals may have similar sensorimotor abilities to those that are used to support speech in humans.

The role of the cerebellum in adaptation: ALE meta-analyses on sensory feedback error.

It is widely accepted that unexpected sensory consequences of self-action engage the cerebellum. However, we currently lack consensus on where in the cerebellum, we find fine-grained differentiation to unexpected sensory feedback. This may result from methodological diversity in task-based human neuroimaging studies that experimentally alter the quality of self-generated sensory feedback. We gathered existing studies that manipulated sensory feedback using a variety of methodological approaches and performed activation likelihood estimation (ALE) meta-analyses. Only half of these studies reported cerebellar activation with considerable variation in spatial location. Consequently, ALE analyses did not reveal significantly increased likelihood of activation in the cerebellum despite the broad scientific consensus of the cerebellum's involvement. In light of the high degree of methodological variability in published studies, we tested for statistical dependence between methodological factors that varied across the published studies. Experiments that elicited an adaptive response to continuously altered sensory feedback more frequently reported activation in the cerebellum than those experiments that did not induce adaptation. These findings may explain the surprisingly low rate of significant cerebellar activation across brain imaging studies investigating unexpected sensory feedback. Furthermore, limitations of functional magnetic resonance imaging to probe the cerebellum could play a role as climbing fiber activity associated with feedback error processing may not be captured by it. We provide methodological recommendations that may guide future studies.

Accessory to dissipate heat from transcranial magnetic stimulation coils.

BACKGROUND: Transcranial magnetic stimulation (TMS) produces magnetic pulses by passing a strong electrical current through coils of wire. Repeated stimulation accumulates heat, which places practical constraints on experimental design. New method: We designed a condensation-free pre-chilled heat sink to extend the operational duration of transcranial magnetic stimulation coils. RESULTS: The application of a pre-chilled heat sink reduced the rate of heating across all tests and extended the duration of stimulation before coil overheating, particularly in conditions where heat management was problematic. Comparison with existing method: Applying an external heat sink had the practical effect of extending the operational time of TMS coils by 5.8-19.3 minutes compared to standard operating procedures. CONCLUSION: Applying an external heat sink increases the quantity of data that can be collected within a single experimental session.

How does human motor cortex regulate vocal pitch in singers?

Vocal pitch is used as an important communicative device by humans, as found in the melodic dimension of both speech and song. Vocal pitch is determined by the degree of tension in the vocal folds of the larynx, which itself is influenced by complex and nonlinear interactions among the laryngeal muscles. The relationship between these muscles and vocal pitch has been described by a mathematical model in the form of a set of 'control rules'. We searched for the biological implementation of these control rules in the larynx motor cortex of the human brain. We scanned choral singers with functional magnetic resonance imaging as they produced discrete pitches at four different levels across their vocal range. While the locations of the larynx motor activations varied across singers, the activation peaks for the four pitch levels were highly consistent within each individual singer. This result was corroborated using multi-voxel pattern analysis, which demonstrated an absence of patterned activations differentiating any pairing of pitch levels. The complex and nonlinear relationships between the multiple laryngeal muscles that control vocal pitch may obscure the neural encoding of vocal pitch in the brain.

Poor neuro-motor tuning of the human larynx: a comparison of sung and whistled pitch imitation.

Vocal imitation is a hallmark of human communication that underlies the capacity to learn to speak and sing. Even so, poor vocal imitation abilities are surprisingly common in the general population and even expert vocalists cannot match the precision of a musical instrument. Although humans have evolved a greater degree of control over the laryngeal muscles that govern voice production, this ability may be underdeveloped compared with control over the articulatory muscles, such as the tongue and lips, volitional control of which emerged earlier in primate evolution. Human participants imitated simple melodies by either singing (i.e. producing pitch with the larynx) or whistling (i.e. producing pitch with the lips and tongue). Sung notes were systematically biased towards each individual's habitual pitch, which we hypothesize may act to conserve muscular effort. Furthermore, while participants who sung more precisely also whistled more precisely, sung imitations were less precise than whistled imitations. The laryngeal muscles that control voice production are under less precise control than the oral muscles that are involved in whistling. This imprecision may be due to the relatively recent evolution of volitional laryngeal-motor control in humans, which may be tuned just well enough for the coarse modulation of vocal-pitch in speech.