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The key assumption of the predictive coding framework is that internal representations are used to generate predictions on how the sensory input will look like in the immediate future. These predictions are tested against the actual input by the so-called prediction error units, which encode the residuals of the predictions. What happens to prediction errors, however, if predictions drawn by different stages of the sensory hierarchy contradict each other? To answer this question, we conducted two fMRI experiments while female and male human participants listened to sequences of sounds: pure tones in the first experiment and frequency-modulated sweeps in the second experiment. In both experiments, we used repetition to induce predictions based on stimulus statistics (stats-informed predictions) and abstract rules disclosed in the task instructions to induce an orthogonal set of (task-informed) predictions. We tested three alternative scenarios: neural responses in the auditory sensory pathway encode prediction error with respect to (1) the stats-informed predictions, (2) the task-informed predictions, or (3) a combination of both. Results showed that neural populations in all recorded regions (bilateral inferior colliculus, medial geniculate body, and primary and secondary auditory cortices) encode prediction error with respect to a combination of the two orthogonal sets of predictions. The findings suggest that predictive coding exploits the non-linear architecture of the auditory pathway for the transmission of predictions. Such non-linear transmission of predictions might be crucial for the predictive coding of complex auditory signals like speech. Significance Statement Sensory systems exploit our subjective expectations to make sense of an overwhelming influx of sensory signals. It is still unclear how expectations at each stage of the processing pipeline are used to predict the representations at the other stages. The current view is that this transmission is hierarchical and linear. Here we measured fMRI responses in auditory cortex, sensory thalamus, and midbrain while we induced two sets of mutually inconsistent expectations on the sensory input, each putatively encoded at a different stage. We show that responses at all stages are concurrently shaped by both sets of expectations. The results challenge the hypothesis that expectations are transmitted linearly and provide for a normative explanation of the non-linear physiology of the corticofugal sensory system., (Copyright © 2023 the authors.)

During face-to-face communication, the perception and recognition of facial movements can facilitate individuals' understanding of what is said. Facial movements are a form of complex biological motion. Separate neural pathways are thought to processing (1) simple, nonbiological motion with an obligatory waypoint in the motion-sensitive visual middle temporal area (V5/MT); and (2) complex biological motion. Here, we present findings that challenge this dichotomy. Neuronavigated offline transcranial magnetic stimulation (TMS) over V5/MT on 24 participants (17 females and 7 males) led to increased response times in the recognition of simple, nonbiological motion as well as visual speech recognition compared with TMS over the vertex, an active control region. TMS of area V5/MT also reduced practice effects on response times, that are typically observed in both visual speech and motion recognition tasks over time. Our findings provide the first indication that area V5/MT causally influences the recognition of visual speech. SIGNIFICANCE STATEMENT In everyday face-to-face communication, speech comprehension is often facilitated by viewing a speaker's facial movements. Several brain areas contribute to the recognition of visual speech. One area of interest is the motion-sensitive visual medial temporal area (V5/MT), which has been associated with the perception of simple, nonbiological motion such as moving dots, as well as more complex, biological motion such as visual speech. Here, we demonstrate using noninvasive brain stimulation that area V5/MT is causally relevant in recognizing visual speech. This finding provides new insights into the neural mechanisms that support the perception of human communication signals, which will help guide future research in typically developed individuals and populations with communication difficulties., (Copyright © 2023 the authors.)

Introduction: Autistic individuals often have difficulties with recognizing what another person is saying in noisy conditions such as in a crowded classroom or a restaurant. The underlying neural mechanisms of this speech perception difficulty are unclear. In typically developed individuals, three cerebral cortex regions are particularly related to speech-in-noise perception: the left inferior frontal gyrus (IFG), the right insula, and the left inferior parietal lobule (IPL). Here, we tested whether responses in these cerebral cortex regions are altered in speech-in-noise perception in autism., Methods: Seventeen autistic adults and 17 typically developed controls (matched pairwise on age, sex, and IQ) performed an auditory-only speech recognition task during functional magnetic resonance imaging (fMRI). Speech was presented either with noise (noise condition) or without noise (no noise condition, i.e., clear speech)., Results: In the left IFG, blood-oxygenation-level-dependent (BOLD) responses were higher in the control compared to the autism group for recognizing speech-in-noise compared to clear speech. For this contrast, both groups had similar response magnitudes in the right insula and left IPL. Additionally, we replicated previous findings that BOLD responses in speech-related and auditory brain regions (including bilateral superior temporal sulcus and Heschl's gyrus) for clear speech were similar in both groups and that voice identity recognition was impaired for clear and noisy speech in autism., Discussion: Our findings show that in autism, the processing of speech is particularly reduced under noisy conditions in the left IFG-a dysfunction that might be important in explaining restricted speech comprehension in noisy environments., (© 2022 The Authors. Brain and Behavior published by Wiley Periodicals LLC.)

Expectations substantially influence perception, but the neural mechanisms underlying this influence are not fully understood. A prominent view is that sensory neurons encode prediction error with respect to expectations on upcoming sensory input. Although the encoding of prediction error has been previously demonstrated in the human auditory cortex (AC), previous studies often induced expectations using stimulus repetition, potentially confounding prediction error with neural habituation. These studies also measured AC as a single population, failing to consider possible predictive specializations of different AC fields. Moreover, the few studies that considered prediction error to stimuli other than pure tones yielded conflicting results. Here, we used functional magnetic resonance imaging (fMRI) to systematically investigate prediction error to subjective expectations in auditory cortical fields Te1.0, Te1.1, Te1.2, and Te3, and two types of stimuli: pure tones and frequency modulated (FM) sweeps. Our results show that prediction error is elicited with respect to the participants' expectations independently of stimulus repetition and similarly expressed across auditory fields. Moreover, despite the radically different strategies underlying the decoding of pure tones and FM-sweeps, both stimulus modalities were encoded as prediction error in most fields of AC. Altogether, our results provide unequivocal evidence that predictive coding is the general encoding mechanism in AC., (© The Author(s) 2022. Published by Oxford University Press.)

Autism spectrum disorder (ASD) is characterised by social communication difficulties. These difficulties have been mainly explained by cognitive, motivational, and emotional alterations in ASD. The communication difficulties could, however, also be associated with altered sensory processing of communication signals. Here, we assessed the functional integrity of auditory sensory pathway nuclei in ASD in three independent functional magnetic resonance imaging experiments. We focused on two aspects of auditory communication that are impaired in ASD: voice identity perception, and recognising speech-in-noise. We found reduced processing in adults with ASD as compared to typically developed control groups (pairwise matched on sex, age, and full-scale IQ) in the central midbrain structure of the auditory pathway (inferior colliculus [IC]). The right IC responded less in the ASD as compared to the control group for voice identity, in contrast to speech recognition. The right IC also responded less in the ASD as compared to the control group when passively listening to vocal in contrast to non-vocal sounds. Within the control group, the left and right IC responded more when recognising speech-in-noise as compared to when recognising speech without additional noise. In the ASD group, this was only the case in the left, but not the right IC. The results show that communication signal processing in ASD is associated with reduced subcortical sensory functioning in the midbrain. The results highlight the importance of considering sensory processing alterations in explaining communication difficulties, which are at the core of ASD., (© 2022 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

The human lateral geniculate nucleus (LGN) of the visual thalamus is a key subcortical processing site for visual information analysis. Due to its small size and deep location within the brain, a non-invasive characterization of the LGN and its microstructurally distinct magnocellular (M) and parvocellular (P) subdivisions in humans is challenging. Here, we investigated whether structural quantitative MRI (qMRI) methods that are sensitive to underlying microstructural tissue features enable MR-based mapping of human LGN M and P subdivisions. We employed high-resolution 7 Tesla in-vivo qMRI in N = 27 participants and ultra-high resolution 7 Tesla qMRI of a post-mortem human LGN specimen. We found that a quantitative assessment of the LGN and its subdivisions is possible based on microstructure-informed qMRI contrast alone. In both the in-vivo and post-mortem qMRI data, we identified two components of shorter and longer longitudinal relaxation time (T 1 ) within the LGN that coincided with the known anatomical locations of a dorsal P and a ventral M subdivision, respectively. Through ground-truth histological validation, we further showed that the microstructural MRI contrast within the LGN pertains to cyto- and myeloarchitectonic tissue differences between its subdivisions. These differences were based on cell and myelin density, but not on iron content. Our qMRI-based mapping strategy paves the way for an in-depth understanding of LGN function and microstructure in humans. It further enables investigations into the selective contributions of LGN subdivisions to human behavior in health and disease., (Copyright © 2021. Published by Elsevier Inc.)

The role of the motor cortex in perceptual and cognitive functions is highly controversial. Here, we investigated the hypothesis that the motor cortex can be instrumental for translating foreign language vocabulary. Human participants of both sexes were trained on foreign language (L2) words and their native language translations over 4 consecutive days. L2 words were accompanied by complementary gestures (sensorimotor enrichment) or pictures (sensory enrichment). Following training, participants translated the auditorily presented L2 words that they had learned. During translation, repetitive transcranial magnetic stimulation was applied bilaterally to a site within the primary motor cortex (Brodmann area 4) located in the vicinity of the arm functional compartment. Responses within the stimulated motor region have previously been found to correlate with behavioral benefits of sensorimotor-enriched L2 vocabulary learning. Compared to sham stimulation, effective perturbation by repetitive transcranial magnetic stimulation slowed down the translation of sensorimotor-enriched L2 words, but not sensory-enriched L2 words. This finding suggests that sensorimotor-enriched training induced changes in L2 representations within the motor cortex, which in turn facilitated the translation of L2 words. The motor cortex may play a causal role in precipitating sensorimotor-based learning benefits, and may directly aid in remembering the native language translations of foreign language words following sensorimotor-enriched training. These findings support multisensory theories of learning while challenging reactivation-based theories. SIGNIFICANCE STATEMENT Despite the potential for sensorimotor enrichment to serve as a powerful tool for learning in many domains, its underlying brain mechanisms remain largely unexplored. Using transcranial magnetic stimulation and a foreign language (L2) learning paradigm, we found that sensorimotor-enriched training can induce changes in L2 representations within the motor cortex, which in turn causally facilitate the translation of L2 words. The translation of recently acquired L2 words may therefore rely not only on auditory information stored in memory or on modality-independent L2 representations, but also on the sensorimotor context in which the words have been experienced., (Copyright © 2021 the authors.)

Recognizing speech in background noise is a strenuous daily activity, yet most humans can master it. An explanation of how the human brain deals with such sensory uncertainty during speech recognition is to-date missing. Previous work has shown that recognition of speech without background noise involves modulation of the auditory thalamus (medial geniculate body; MGB): there are higher responses in left MGB for speech recognition tasks that require tracking of fast-varying stimulus properties in contrast to relatively constant stimulus properties (e.g., speaker identity tasks) despite the same stimulus input. Here, we tested the hypotheses that (1) this task-dependent modulation for speech recognition increases in parallel with the sensory uncertainty in the speech signal, i.e., the amount of background noise; and that (2) this increase is present in the ventral MGB, which corresponds to the primary sensory part of the auditory thalamus. In accordance with our hypothesis, we show, by using ultra-high-resolution functional magnetic resonance imaging (fMRI) in male and female human participants, that the task-dependent modulation of the left ventral MGB (vMGB) for speech is particularly strong when recognizing speech in noisy listening conditions in contrast to situations where the speech signal is clear. The results imply that speech in noise recognition is supported by modifications at the level of the subcortical sensory pathway providing driving input to the auditory cortex. SIGNIFICANCE STATEMENT Speech recognition in noisy environments is a challenging everyday task. One reason why humans can master this task is the recruitment of additional cognitive resources as reflected in recruitment of non-language cerebral cortex areas. Here, we show that also modulation in the primary sensory pathway is specifically involved in speech in noise recognition. We found that the left primary sensory thalamus (ventral medial geniculate body; vMGB) is more involved when recognizing speech signals as opposed to a control task (speaker identity recognition) when heard in background noise versus when the noise was absent. This finding implies that the brain optimizes sensory processing in subcortical sensory pathway structures in a task-specific manner to deal with speech recognition in noisy environments., (Copyright © 2021 the authors.)

Recognising the identity of voices is a key ingredient of communication. Visual mechanisms support this ability: recognition is better for voices previously learned with their corresponding face (compared to a control condition). This so-called 'face-benefit' is supported by the fusiform face area (FFA), a region sensitive to facial form and identity. Behavioural findings indicate that the face-benefit increases in noisy listening conditions. The neural mechanisms for this increase are unknown. Here, using functional magnetic resonance imaging, we examined responses in face-sensitive regions while participants recognised the identity of auditory-only speakers (previously learned by face) in high (SNR -4 dB) and low (SNR +4 dB) levels of auditory noise. We observed a face-benefit in both noise levels, for most participants (16 of 21). In high-noise, the recognition of face-learned speakers engaged the right posterior superior temporal sulcus motion-sensitive face area (pSTS-mFA), a region implicated in the processing of dynamic facial cues. The face-benefit in high-noise also correlated positively with increased functional connectivity between this region and voice-sensitive regions in the temporal lobe in the group of 16 participants with a behavioural face-benefit. In low-noise, the face-benefit was robustly associated with increased responses in the FFA and to a lesser extent the right pSTS-mFA. The findings highlight the remarkably adaptive nature of the visual network supporting voice-identity recognition in auditory-only listening conditions., (© 2021 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

Predictive processing, a leading theoretical framework for sensory processing, suggests that the brain constantly generates predictions on the sensory world and that perception emerges from the comparison between these predictions and the actual sensory input. This requires two distinct neural elements: generative units, which encode the model of the sensory world; and prediction error units, which compare these predictions against the sensory input. Although predictive processing is generally portrayed as a theory of cerebral cortex function, animal and human studies over the last decade have robustly shown the ubiquitous presence of prediction error responses in several nuclei of the auditory, somatosensory, and visual subcortical pathways. In the auditory modality, prediction error is typically elicited using so-called oddball paradigms, where sequences of repeated pure tones with the same pitch are at unpredictable intervals substituted by a tone of deviant frequency. Repeated sounds become predictable promptly and elicit decreasing prediction error; deviant tones break these predictions and elicit large prediction errors. The simplicity of the rules inducing predictability make oddball paradigms agnostic about the origin of the predictions. Here, we introduce two possible models of the organizational topology of the predictive processing auditory network: (1) the global view, that assumes that predictions on the sensory input are generated at high-order levels of the cerebral cortex and transmitted in a cascade of generative models to the subcortical sensory pathways; and (2) the local view, that assumes that independent local models, computed using local information, are used to perform predictions at each processing stage. In the global view information encoding is optimized globally but biases sensory representations along the entire brain according to the subjective views of the observer. The local view results in a diminished coding efficiency, but guarantees in return a robust encoding of the features of sensory input at each processing stage. Although most experimental results to-date are ambiguous in this respect, recent evidence favors the global model., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Tabas and von Kriegstein.)

Frequency modulation (FM) is a basic constituent of vocalisation in many animals as well as in humans. In human speech, short rising and falling FM-sweeps of around 50 ms duration, called formant transitions, characterise individual speech sounds. There are two representations of FM in the ascending auditory pathway: a spectral representation, holding the instantaneous frequency of the stimuli; and a sweep representation, consisting of neurons that respond selectively to FM direction. To-date computational models use feedforward mechanisms to explain FM encoding. However, from neuroanatomy we know that there are massive feedback projections in the auditory pathway. Here, we found that a classical FM-sweep perceptual effect, the sweep pitch shift, cannot be explained by standard feedforward processing models. We hypothesised that the sweep pitch shift is caused by a predictive feedback mechanism. To test this hypothesis, we developed a novel model of FM encoding incorporating a predictive interaction between the sweep and the spectral representation. The model was designed to encode sweeps of the duration, modulation rate, and modulation shape of formant transitions. It fully accounted for experimental data that we acquired in a perceptual experiment with human participants as well as previously published experimental results. We also designed a new class of stimuli for a second perceptual experiment to further validate the model. Combined, our results indicate that predictive interaction between the frequency encoding and direction encoding neural representations plays an important role in the neural processing of FM. In the brain, this mechanism is likely to occur at early stages of the processing hierarchy., Competing Interests: The authors have declared that no competing interests exist.

Despite a rise in the use of "learning by doing" pedagogical methods in praxis, little is known as to how the brain benefits from these methods. Learning by doing strategies that utilize complementary information ("enrichment") such as gestures have been shown to optimize learning outcomes in several domains including foreign language (L2) training. Here we tested the hypothesis that behavioral benefits of gesture-based enrichment are critically supported by integrity of the biological motion visual cortices (bmSTS). Prior functional neuroimaging work has implicated the visual motion cortices in L2 translation following sensorimotor-enriched training; the current study is the first to investigate the causal relevance of these structures in learning by doing contexts. Using neuronavigated transcranial magnetic stimulation and a gesture-enriched L2 vocabulary learning paradigm, we found that the bmSTS causally contributed to behavioral benefits of gesture-enriched learning. Visual motion cortex integrity benefitted both short- and long-term learning outcomes, as well as the learning of concrete and abstract words. These results adjudicate between opposing predictions of two neuroscientific learning theories: While reactivation-based theories predict no functional role of specialized sensory cortices in vocabulary learning outcomes, the current study supports the predictive coding theory view that these cortices precipitate sensorimotor-based learning benefits., (© The Author(s) 2020. Published by Oxford University Press.)

The subcortical sensory pathways are the fundamental channels for mapping the outside world to our minds. Sensory pathways efficiently transmit information by adapting neural responses to the local statistics of the sensory input. The long-standing mechanistic explanation for this adaptive behaviour is that neural activity decreases with increasing regularities in the local statistics of the stimuli. An alternative account is that neural coding is directly driven by expectations of the sensory input. Here, we used abstract rules to manipulate expectations independently of local stimulus statistics. The ultra-high-field functional-MRI data show that abstract expectations can drive the response amplitude to tones in the human auditory pathway. These results provide first unambiguous evidence of abstract processing in a subcortical sensory pathway. They indicate that the neural representation of the outside world is altered by our prior beliefs even at initial points of the processing hierarchy., Competing Interests: AT, GM, SK, RT, Kv No competing interests declared, (© 2020, Tabas et al.)

Atypical eye contact in communication is a common characteristic in autism spectrum disorders. Autistic traits vary along a continuum extending into the neurotypical population. The relation between autistic traits and brain mechanisms underlying spontaneous eye contact during verbal communication remains unexplored. Here, we used simultaneous functional magnetic resonance imaging and eye tracking to investigate this relation in neurotypical people within a naturalistic verbal context. Using multiple regression analyses, we found that brain response in the posterior superior temporal sulcus (pSTS) and its connectivity with the fusiform face area (FFA) during eye contact with a speaker predicted the level of autistic traits measured by Autism-spectrum Quotient (AQ). Further analyses for different AQ subclusters revealed that these two predictors were negatively associated with attention to detail. The relation between FFA-pSTS connectivity and the attention to detail ability was mediated by individuals' looking preferences for speaker's eyes. This study identified the role of an individual eye contact pattern in the relation between brain mechanisms underlying natural eye contact during verbal communication and autistic traits in neurotypical people. The findings may help to increase our understanding of the mechanisms of atypical eye contact behavior during natural communication.

Faces and voices are prominent cues for person-identity recognition. Face recognition behavior and associated brain responses can be enhanced by intranasal administration of oxytocin. It is unknown whether oxytocin can also augment voice-identity recognition mechanisms. To find it out is particularly relevant for individuals who have difficulties recognizing voice identity such as individuals diagnosed with autism spectrum disorder (ASD). We conducted a combined behavioral and functional magnetic resonance imaging (fMRI) study to investigate voice-identity recognition following intranasal administration of oxytocin or placebo in a group of adults diagnosed with ASD (full-scale intelligence quotient > 85) and pairwise-matched typically developing (TD) controls. A single dose of 24 IU oxytocin was administered in a randomized, double-blind, placebo-controlled and cross-over design. In the control group, but not in the ASD group, administration of oxytocin compared to placebo increased responses to recognition of voice identity in contrast to speech in the right posterior superior temporal sulcus/gyrus (pSTS/G) - a region implicated in the perceptual analysis of voice-identity information. In the ASD group, the right pSTS/G responses were positively correlated with voice-identity recognition accuracy in the oxytocin condition, but not in the placebo condition. Oxytocin did not improve voice-identity recognition performance at the group level. The ASD compared to the control group had lower right pSTS/G responses to voice-identity recognition. Since ASD is known to have atypical pSTS/G, the results indicate that the potential of intranasal oxytocin to enhance mechanisms for voice-identity recognition might be variable and dependent on the functional integrity of this brain region.

Faces convey social information such as emotion and speech. Facial emotion processing is supported via interactions between dorsal-movement and ventral-form visual cortex regions. Here, we explored, for the first time, whether similar dorsal-ventral interactions (assessed via functional connectivity), might also exist for visual-speech processing. We then examined whether altered dorsal-ventral connectivity is observed in adults with high-functioning autism spectrum disorder (ASD), a disorder associated with impaired visual-speech recognition. We acquired functional magnetic resonance imaging (fMRI) data with concurrent eye tracking in pairwise matched control and ASD participants. In both groups, dorsal-movement regions in the visual motion area 5 (V5/MT) and the temporal visual speech area (TVSA) were functionally connected to ventral-form regions (i.e., the occipital face area [OFA] and the fusiform face area [FFA]) during the recognition of visual speech, in contrast to the recognition of face identity. Notably, parts of this functional connectivity were decreased in the ASD group compared to the controls (i.e., right V5/MT-right OFA, left TVSA-left FFA). The results confirmed our hypothesis that functional connectivity between dorsal-movement and ventral-form regions exists during visual-speech processing. Its partial dysfunction in ASD might contribute to difficulties in the recognition of dynamic face information relevant for successful face-to-face communication., (© 2019 The Authors. Human Brain Mapping published by Wiley Periodicals, Inc.)

In perceptual decision making the brain extracts and accumulates decision evidence from a stimulus over time and eventually makes a decision based on the accumulated evidence. Several characteristics of this process have been observed in human electrophysiological experiments, especially an average build-up of motor-related signals supposedly reflecting accumulated evidence, when averaged across trials. Another recently established approach to investigate the representation of decision evidence in brain signals is to correlate the within-trial fluctuations of decision evidence with the measured signals. We here report results of this approach for a two-alternative forced choice reaction time experiment measured using magnetoencephalography (MEG) recordings. Our results show: (1) that decision evidence is most strongly represented in the MEG signals in three consecutive phases and (2) that posterior cingulate cortex is involved most consistently, among all brain areas, in all three of the identified phases. As most previous work on perceptual decision making in the brain has focused on parietal and motor areas, our findings therefore suggest that the role of the posterior cingulate cortex in perceptual decision making may be currently underestimated., (Copyright © 2020 Bitzer, Park, Maess, von Kriegstein and Kiebel.)

Sensory thalami are central sensory pathway stations for information processing. Their role for human cognition and perception, however, remains unclear. Recent evidence suggests an involvement of the sensory thalami in speech recognition. In particular, the auditory thalamus (medial geniculate body, MGB) response is modulated by speech recognition tasks and the amount of this task-dependent modulation is associated with speech recognition abilities. Here, we tested the specific hypothesis that this behaviorally relevant modulation is present in the MGB subsection that corresponds to the primary auditory pathway (i.e., the ventral MGB [vMGB]). We used ultra-high field 7T fMRI to identify the vMGB, and found a significant positive correlation between the amount of task-dependent modulation and the speech recognition performance across participants within left vMGB, but not within the other MGB subsections. These results imply that modulation of thalamic driving input to the auditory cortex facilitates speech recognition., Competing Interests: PM, MM, Fd, RT, SK, Kv No competing interests declared, (© 2019, Mihai et al.)

Developmental dyslexia is characterized by the inability to acquire typical reading and writing skills. Dyslexia has been frequently linked to cerebral cortex alterations; however, recent evidence also points toward sensory thalamus dysfunctions: dyslexics showed reduced responses in the left auditory thalamus (medial geniculate body, MGB) during speech processing in contrast to neurotypical readers. In addition, in the visual modality, dyslexics have reduced structural connectivity between the left visual thalamus (lateral geniculate nucleus, LGN) and V5/MT, a cerebral cortex region involved in visual movement processing. Higher LGN-V5/MT connectivity in dyslexics was associated with the faster rapid naming of letters and numbers (RANln), a measure that is highly correlated with reading proficiency. Here, we tested two hypotheses that were directly derived from these previous findings. First, we tested the hypothesis that dyslexics have reduced structural connectivity between the left MGB and the auditory-motion-sensitive part of the left planum temporale (mPT). Second, we hypothesized that the amount of left mPT-MGB connectivity correlates with dyslexics RANln scores. Using diffusion tensor imaging-based probabilistic tracking, we show that male adults with developmental dyslexia have reduced structural connectivity between the left MGB and the left mPT, confirming the first hypothesis. Stronger left mPT-MGB connectivity was not associated with faster RANln scores in dyslexics, but was in neurotypical readers. Our findings provide the first evidence that reduced cortico-thalamic connectivity in the auditory modality is a feature of developmental dyslexia and it may also affect reading-related cognitive abilities in neurotypical readers. SIGNIFICANCE STATEMENT Developmental dyslexia is one of the most widespread learning disabilities. Although previous neuroimaging research mainly focused on pathomechanisms of dyslexia at the cerebral cortex level, several lines of evidence suggest an atypical functioning of subcortical sensory structures. By means of diffusion tensor imaging, we here show that dyslexic male adults have reduced white matter connectivity in a cortico-thalamic auditory pathway between the left auditory motion-sensitive planum temporale and the left medial geniculate body. Connectivity strength of this pathway was associated with measures of reading fluency in neurotypical readers. This is novel evidence on the neurocognitive correlates of reading proficiency, highlighting the importance of cortico-subcortical interactions between regions involved in the processing of spectrotemporally complex sound., (Copyright © 2019 the authors 0270-6474/19/391720-13$15.00/0.)

Recognizing the identity of others by their voice is an important skill for social interactions. To date, it remains controversial which parts of the brain are critical structures for this skill. Based on neuroimaging findings, standard models of person-identity recognition suggest that the right temporal lobe is the hub for voice-identity recognition. Neuropsychological case studies, however, reported selective deficits of voice-identity recognition in patients predominantly with right inferior parietal lobe lesions. Here, our aim was to work towards resolving the discrepancy between neuroimaging studies and neuropsychological case studies to find out which brain structures are critical for voice-identity recognition in humans. We performed a voxel-based lesion-behaviour mapping study in a cohort of patients (n = 58) with unilateral focal brain lesions. The study included a comprehensive behavioural test battery on voice-identity recognition of newly learned (voice-name, voice-face association learning) and familiar voices (famous voice recognition) as well as visual (face-identity recognition) and acoustic control tests (vocal-pitch and vocal-timbre discrimination). The study also comprised clinically established tests (neuropsychological assessment, audiometry) and high-resolution structural brain images. The three key findings were: (i) a strong association between voice-identity recognition performance and right posterior/mid temporal and right inferior parietal lobe lesions; (ii) a selective association between right posterior/mid temporal lobe lesions and voice-identity recognition performance when face-identity recognition performance was factored out; and (iii) an association of right inferior parietal lobe lesions with tasks requiring the association between voices and faces but not voices and names. The results imply that the right posterior/mid temporal lobe is an obligatory structure for voice-identity recognition, while the inferior parietal lobe is only a facultative component of voice-identity recognition in situations where additional face-identity processing is required., (© The Author (2017). Published by Oxford University Press on behalf of the Guarantors of Brain.)

Speech information inherent in face movements is important for understanding what is said in face-to-face communication. Individuals with autism spectrum disorders (ASD) have difficulties in extracting speech information from face movements, a process called visual-speech recognition. Currently, it is unknown what dysfunctional brain regions or networks underlie the visual-speech recognition deficit in ASD. We conducted a functional magnetic resonance imaging (fMRI) study with concurrent eye tracking to investigate visual-speech recognition in adults diagnosed with high-functioning autism and pairwise matched typically developed controls. Compared to the control group (n = 17), the ASD group (n = 17) showed decreased Blood Oxygenation Level Dependent (BOLD) response during visual-speech recognition in the right visual area 5 (V5/MT) and left temporal visual speech area (TVSA) - brain regions implicated in visual-movement perception. The right V5/MT showed positive correlation with visual-speech task performance in the ASD group, but not in the control group. Psychophysiological interaction analysis (PPI) revealed that functional connectivity between the left TVSA and the bilateral V5/MT and between the right V5/MT and the left IFG was lower in the ASD than in the control group. In contrast, responses in other speech-motor regions and their connectivity were on the neurotypical level. Reduced responses and network connectivity of the visual-movement regions in conjunction with intact speech-related mechanisms indicate that perceptual mechanisms might be at the core of the visual-speech recognition deficit in ASD. Communication deficits in ASD might at least partly stem from atypical sensory processing and not higher-order cognitive processing of socially relevant information., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

Developmental dyslexia is a highly prevalent reading disorder affecting about 5%-10% of children [1]. It is characterized by slow and/or inaccurate word recognition skills as well as by poor spelling and decoding abilities [2]. Partly due to technical challenges with investigating subcortical sensory structures, current research on dyslexia in humans by and large focuses on the cerebral cortex [3-7]. These studies found that dyslexia is typically associated with functional and structural alterations of a distributed left-hemispheric cerebral cortex network (e.g., [8, 9]). However, findings from animal models and post mortem studies in humans suggest that dyslexia might also be associated with structural alterations in subcortical sensory pathways [10-14] (reviewed in [7]). Whether these alterations also exist in dyslexia in vivo and how they relate to dyslexia symptoms is currently unknown. Here, we used ultra-high-resolution structural magnetic resonance imaging (MRI), diffusion MRI, and probabilistic tractography to investigate the structural connections of the visual sensory pathway in dyslexia in vivo. We discovered that individuals with dyslexia have reduced structural connections in the direct pathway between the left visual thalamus (lateral geniculate nucleus [LGN]) and left middle temporal area V5/MT, but not between the left LGN and left primary visual cortex. In addition, left V5/MT-LGN connectivity strength correlated with rapid naming abilities-a key deficit in dyslexia [15]. These findings provide the first evidence of specific structural alterations in the connections between the sensory thalamus and cortex in developmental dyslexia. The results challenge current standard models and provide novel evidence for the importance of cortico-thalamic interactions in explaining dyslexia., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Previous studies have shown that listeners are better able to understand speech when they are familiar with the talker's voice. In most of these studies, talker familiarity was ensured by explicit voice training; that is, listeners learned to identify the familiar talkers. In the real world, however, the characteristics of familiar talkers are learned incidentally, through communication. The present study investigated whether speech comprehension benefits from implicit voice training; that is, through exposure to talkers' voices without listeners explicitly trying to identify them. During four training sessions, listeners heard short sentences containing a single verb (e.g., "he writes"), spoken by one talker. The sentences were mixed with noise, and listeners identified the verb within each sentence while their speech-reception thresholds (SRT) were measured. In a final test session, listeners performed the same task, but this time they heard different sentences spoken by the familiar talker and three unfamiliar talkers. Familiar and unfamiliar talkers were counterbalanced across listeners. Half of the listeners performed a test session in which the four talkers were presented in separate blocks (blocked paradigm). For the other half, talkers varied randomly from trial to trial (interleaved paradigm). The results showed that listeners had lower SRT when the speech was produced by the familiar talker than the unfamiliar talkers. The type of talker presentation (blocked vs. interleaved) had no effect on this familiarity benefit. These findings suggest that listeners implicitly learn talker-specific information during a speech-comprehension task, and exploit this information to improve the comprehension of novel speech material from familiar talkers.

In the native language, abstract and concrete nouns are represented in distinct areas of the cerebral cortex. Currently, it is unknown whether this is also the case for abstract and concrete nouns of a foreign language. Here, we taught adult native speakers of German 45 abstract and 45 concrete nouns of a foreign language. After learning the nouns for 5 days, participants performed a vocabulary translation task during functional magnetic resonance imaging. Translating abstract nouns in contrast to concrete nouns elicited responses in regions that are also responsive to abstract nouns in the native language: the left inferior frontal gyrus and the left middle and superior temporal gyri. Concrete nouns elicited larger responses in the angular gyri bilaterally and the left parahippocampal gyrus than abstract nouns. The cluster in the left angular gyrus showed psychophysiological interaction (PPI) with the left lingual gyrus. The left parahippocampal gyrus showed PPI with the posterior cingulate cortex. Similar regions have been previously found for concrete nouns in the native language. The results reveal similarities in the cortical representation of foreign language nouns with the representation of native language nouns that already occur after 5 days of vocabulary learning. Furthermore, we showed that verbal and enriched learning methods were equally suitable to teach foreign abstract and concrete nouns. Hum Brain Mapp 38:4398-4412, 2017. © 2017 Wiley Periodicals, Inc., (© 2017 Wiley Periodicals, Inc.)

Eye contact occurs frequently and voluntarily during face-to-face verbal communication. However, the neural mechanisms underlying eye contact when it is accompanied by spoken language remain unexplored to date. Here we used a novel approach, fixation-based event-related functional magnetic resonance imaging (fMRI), to simulate the listener making eye contact with a speaker during verbal communication. Participants' eye movements and fMRI data were recorded simultaneously while they were freely viewing a pre-recorded speaker talking. The eye tracking data were then used to define events for the fMRI analyses. The results showed that eye contact in contrast to mouth fixation involved visual cortical areas (cuneus, calcarine sulcus), brain regions related to theory of mind/intentionality processing (temporoparietal junction, posterior superior temporal sulcus, medial prefrontal cortex) and the dorsolateral prefrontal cortex. In addition, increased effective connectivity was found between these regions for eye contact in contrast to mouth fixations. The results provide first evidence for neural mechanisms underlying eye contact when watching and listening to another person talking. The network we found might be well suited for processing the intentions of communication partners during eye contact in verbal communication., (© The Author (2016). Published by Oxford University Press. For Permissions, please email: journals.permissions@oup.com.)

The ability to recognise the identity of others is a key requirement for successful communication. Brain regions that respond selectively to voices exist in humans from early infancy on. Currently, it is unclear whether dysfunction of these voice-sensitive regions can explain voice identity recognition impairments. Here, we used two independent functional magnetic resonance imaging studies to investigate voice processing in a population that has been reported to have no voice-sensitive regions: autism spectrum disorder (ASD). Our results refute the earlier report that individuals with ASD have no responses in voice-sensitive regions: Passive listening to vocal, compared to non-vocal, sounds elicited typical responses in voice-sensitive regions in the high-functioning ASD group and controls. In contrast, the ASD group had a dysfunction in voice-sensitive regions during voice identity but not speech recognition in the right posterior superior temporal sulcus/gyrus (STS/STG)-a region implicated in processing complex spectrotemporal voice features and unfamiliar voices. The right anterior STS/STG correlated with voice identity recognition performance in controls but not in the ASD group. The findings suggest that right STS/STG dysfunction is critical for explaining voice recognition impairments in high-functioning ASD and show that ASD is not characterised by a general lack of voice-sensitive responses., (© The Author (2016). Published by Oxford University Press.)

Decisions in everyday life are prone to error. Standard models typically assume that errors during perceptual decisions are due to noise. However, it is unclear how noise in the sensory input affects the decision. Here we show that there are experimental tasks for which one can analyse the exact spatio-temporal details of a dynamic sensory noise and better understand variability in human perceptual decisions. Using a new experimental visual tracking task and a novel Bayesian decision making model, we found that the spatio-temporal noise fluctuations in the input of single trials explain a significant part of the observed responses. Our results show that modelling the precise internal representations of human participants helps predict when perceptual decisions go wrong. Furthermore, by modelling precisely the stimuli at the single-trial level, we were able to identify the underlying mechanism of perceptual decision making in more detail than standard models.

At present, it is largely unclear how the human brain optimally learns foreign languages. We investigated teaching strategies that utilize complementary information ("enrichment"), such as pictures or gestures, to optimize vocabulary learning outcome. We found that learning while performing gestures was more efficient than the common practice of learning with pictures and that both enrichment strategies were better than learning without enrichment ("verbal learning"). We tested the prediction of an influential cognitive neuroscience theory that provides explanations for the beneficial behavioral effects of enrichment: the "multisensory learning theory" attributes the benefits of enrichment to recruitment of brain areas specialized in processing the enrichment. To test this prediction, we asked participants to translate auditorily presented foreign words during fMRI. Multivariate pattern classification allowed us to decode from the brain activity under which enrichment condition the vocabulary had been learned. The visual-object-sensitive lateral occipital complex (LOC) represented auditory words that had been learned with pictures. The biological motion superior temporal sulcus (bmSTS) and motor areas represented auditory words that had been learned with gestures. Importantly, brain activity in these specialized visual and motor brain areas correlated with behavioral performance. The cortical activation pattern found in the present study strongly supports the multisensory learning theory in contrast to alternative explanations. In addition, the results highlight the importance of learning foreign language vocabulary with enrichment, particularly with self-performed gestures., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The human voice is the primary carrier of speech but also a fingerprint for person identity. Previous neuroimaging studies have revealed that speech and identity recognition is accomplished by partially different neural pathways, despite the perceptual unity of the vocal sound. Importantly, the right STS has been implicated in voice processing, with different contributions of its posterior and anterior parts. However, the time point at which vocal and speech processing diverge is currently unknown. Also, the exact role of the right STS during voice processing is so far unclear because its behavioral relevance has not yet been established. Here, we used the high temporal resolution of magnetoencephalography and a speech task control to pinpoint transient behavioral correlates: we found, at 200 msec after stimulus onset, that activity in right anterior STS predicted behavioral voice recognition performance. At the same time point, the posterior right STS showed increased activity during voice identity recognition in contrast to speech recognition whereas the left mid STS showed the reverse pattern. In contrast to the highly speech-sensitive left STS, the current results highlight the right STS as a key area for voice identity recognition and show that its anatomical-functional division emerges around 200 msec after stimulus onset. We suggest that this time point marks the speech-independent processing of vocal sounds in the posterior STS and their successful mapping to vocal identities in the anterior STS.

Recognizing the identity of other individuals across different sensory modalities is critical for successful social interaction. In the human brain, face- and voice-sensitive areas are separate, but structurally connected. What kind of information is exchanged between these specialized areas during cross-modal recognition of other individuals is currently unclear. For faces, specific areas are sensitive to identity and to physical properties. It is an open question whether voices activate representations of face identity or physical facial properties in these areas. To address this question, we used functional magnetic resonance imaging in humans and a voice-face priming design. In this design, familiar voices were followed by morphed faces that matched or mismatched with respect to identity or physical properties. The results showed that responses in face-sensitive regions were modulated when face identity or physical properties did not match to the preceding voice. The strength of this mismatch signal depended on the level of certainty the participant had about the voice identity. This suggests that both identity and physical property information was provided by the voice to face areas. The activity and connectivity profiles differed between face-sensitive areas: (i) the occipital face area seemed to receive information about both physical properties and identity, (ii) the fusiform face area seemed to receive identity, and (iii) the anterior temporal lobe seemed to receive predominantly identity information from the voice. We interpret these results within a prediction coding scheme in which both identity and physical property information is used across sensory modalities to recognize individuals., (© 2014 Wiley Periodicals, Inc.)

Recognizing other individuals is an essential skill in humans and in other species. Over the last decade, it has become increasingly clear that person-identity recognition abilities are highly variable. Roughly 2% of the population has developmental prosopagnosia, a congenital deficit in recognizing others by their faces. It is currently unclear whether developmental phonagnosia, a deficit in recognizing others by their voices, is equally prevalent, or even whether it actually exists. Here, we aimed to identify cases of developmental phonagnosia. We collected more than 1,000 data sets from self-selected German individuals by using a web-based screening test that was designed to assess their voice-recognition abilities. We then examined potentially phonagnosic individuals by using a comprehensive laboratory test battery. We found two novel cases of phonagnosia: AS, a 32-year-old female, and SP, a 32-year-old male; both are otherwise healthy academics, have normal hearing, and show no pathological abnormalities in brain structure. The two cases have comparable patterns of impairments: both performed at least 2 SDs below the level of matched controls on tests that required learning new voices, judging the familiarity of famous voices, and discriminating pitch differences between voices. In both cases, only voice-identity processing per se was affected: face recognition, speech intelligibility, emotion recognition, and musical ability were all comparable to controls. The findings confirm the existence of developmental phonagnosia as a modality-specific impairment and allow a first rough prevalence estimate., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Over a typical career piano tuners spend tens of thousands of hours exploring a specialized acoustic environment. Tuning requires accurate perception and adjustment of beats in two-note chords that serve as a navigational device to move between points in previously learned acoustic scenes. It is a two-stage process that depends on the following: first, selective listening to beats within frequency windows, and, second, the subsequent use of those beats to navigate through a complex soundscape. The neuroanatomical substrates underlying brain specialization for such fundamental organization of sound scenes are unknown. Here, we demonstrate that professional piano tuners are significantly better than controls matched for age and musical ability on a psychophysical task simulating active listening to beats within frequency windows that is based on amplitude modulation rate discrimination. Tuners show a categorical increase in gray matter volume in the right frontal operculum and right superior temporal lobe. Tuners also show a striking enhancement of gray matter volume in the anterior hippocampus, parahippocampal gyrus, and superior temporal gyrus, and an increase in white matter volume in the posterior hippocampus as a function of years of tuning experience. The relationship with gray matter volume is sensitive to years of tuning experience and starting age but not actual age or level of musicality. Our findings support a role for a core set of regions in the hippocampus and superior temporal cortex in skilled exploration of complex sound scenes in which precise sound "templates" are encoded and consolidated into memory over time in an experience-dependent manner.

Developmental dyslexia, a severe and persistent reading and spelling impairment, is characterized by difficulties in processing speech sounds (i.e., phonemes). Here, we test the hypothesis that these phonological difficulties are associated with a dysfunction of the auditory sensory thalamus, the medial geniculate body (MGB). By using functional MRI, we found that, in dyslexic adults, the MGB responded abnormally when the task required attending to phonemes compared with other speech features. No other structure in the auditory pathway showed distinct functional neural patterns between the two tasks for dyslexic and control participants. Furthermore, MGB activity correlated with dyslexia diagnostic scores, indicating that the task modulation of the MGB is critical for performance in dyslexics. These results suggest that deficits in dyslexia are associated with a failure of the neural mechanism that dynamically tunes MGB according to predictions from cortical areas to optimize speech processing. This view on task-related MGB dysfunction in dyslexics has the potential to reconcile influential theories of dyslexia within a predictive coding framework of brain function.

Auditory object analysis requires two fundamental perceptual processes: the definition of the boundaries between objects, and the abstraction and maintenance of an object's characteristic features. Although it is intuitive to assume that the detection of the discontinuities at an object's boundaries precedes the subsequent precise representation of the object, the specific underlying cortical mechanisms for segregating and representing auditory objects within the auditory scene are unknown. We investigated the cortical bases of these two processes for one type of auditory object, an "acoustic texture," composed of multiple frequency-modulated ramps. In these stimuli, we independently manipulated the statistical rules governing (1) the frequency-time space within individual textures (comprising ramps with a given spectrotemporal coherence) and (2) the boundaries between textures (adjacent textures with different spectrotemporal coherences). Using functional magnetic resonance imaging, we show mechanisms defining boundaries between textures with different coherences in primary and association auditory cortices, whereas texture coherence is represented only in association cortex. Furthermore, participants' superior detection of boundaries across which texture coherence increased (as opposed to decreased) was reflected in a greater neural response in auditory association cortex at these boundaries. The results suggest a hierarchical mechanism for processing acoustic textures that is relevant to auditory object analysis: boundaries between objects are first detected as a change in statistical rules over frequency-time space, before a representation that corresponds to the characteristics of the perceived object is formed.

We understand speech from different speakers with ease, whereas artificial speech recognition systems struggle with this task. It is unclear how the human brain solves this problem. The conventional view is that speech message recognition and speaker identification are two separate functions and that message processing takes place predominantly in the left hemisphere, whereas processing of speaker-specific information is located in the right hemisphere. Here, we distinguish the contribution of specific cortical regions, to speech recognition and speaker information processing, by controlled manipulation of task and resynthesized speaker parameters. Two functional magnetic resonance imaging studies provide evidence for a dynamic speech-processing network that questions the conventional view. We found that speech recognition regions in left posterior superior temporal gyrus/superior temporal sulcus (STG/STS) also encode speaker-related vocal tract parameters, which are reflected in the amplitude peaks of the speech spectrum, along with the speech message. Right posterior STG/STS activated specifically more to a speaker-related vocal tract parameter change during a speech recognition task compared with a voice recognition task. Left and right posterior STG/STS were functionally connected. Additionally, we found that speaker-related glottal fold parameters (e.g., pitch), which are not reflected in the amplitude peaks of the speech spectrum, are processed in areas immediately adjacent to primary auditory cortex, i.e., in areas in the auditory hierarchy earlier than STG/STS. Our results point to a network account of speech recognition, in which information about the speech message and the speaker's vocal tract are combined to solve the difficult task of understanding speech from different speakers.