Intermediate acoustic-to-semantic representations link behavioral and neural responses to natural sounds.

Recognizing sounds implicates the cerebral transformation of input waveforms into semantic representations. Although past research identified the superior temporal gyrus (STG) as a crucial cortical region, the computational fingerprint of these cerebral transformations remains poorly characterized. Here, we exploit a model comparison framework and contrasted the ability of acoustic, semantic (continuous and categorical) and sound-to-event deep neural network representation models to predict perceived sound dissimilarity and 7 T human auditory cortex functional magnetic resonance imaging responses. We confirm that spectrotemporal modulations predict early auditory cortex (Heschl's gyrus) responses, and that auditory dimensions (for example, loudness, periodicity) predict STG responses and perceived dissimilarity. Sound-to-event deep neural networks predict Heschl's gyrus responses similar to acoustic models but, notably, they outperform all competing models at predicting both STG responses and perceived dissimilarity. Our findings indicate that STG entails intermediate acoustic-to-semantic sound representations that neither acoustic nor semantic models can account for. These representations are compositional in nature and relevant to behavior.

Decoding sounds depicting hand-object interactions in primary somatosensory cortex.

Neurons, even in the earliest sensory regions of cortex, are subject to a great deal of contextual influences from both within and across modality connections. Recent work has shown that primary sensory areas can respond to and, in some cases, discriminate stimuli that are not of their target modality: for example, primary somatosensory cortex (SI) discriminates visual images of graspable objects. In the present work, we investigated whether SI would discriminate sounds depicting hand-object interactions (e.g. bouncing a ball). In a rapid event-related functional magnetic resonance imaging experiment, participants listened attentively to sounds from 3 categories: hand-object interactions, and control categories of pure tones and animal vocalizations, while performing a one-back repetition detection task. Multivoxel pattern analysis revealed significant decoding of hand-object interaction sounds within SI, but not for either control category. Crucially, in the hand-sensitive voxels defined from an independent tactile localizer, decoding accuracies were significantly higher for hand-object interactions compared to pure tones in left SI. Our findings indicate that simply hearing sounds depicting familiar hand-object interactions elicit different patterns of activity in SI, despite the complete absence of tactile stimulation. These results highlight the rich contextual information that can be transmitted across sensory modalities even to primary sensory areas.

What do we mean with sound semantics, exactly? A survey of taxonomies and ontologies of everyday sounds.

Taxonomies and ontologies for the characterization of everyday sounds have been developed in several research fields, including auditory cognition, soundscape research, artificial hearing, sound design, and medicine. Here, we surveyed 36 of such knowledge organization systems, which we identified through a systematic literature search. To evaluate the semantic domains covered by these systems within a homogeneous framework, we introduced a comprehensive set of verbal sound descriptors (sound source properties; attributes of sensation; sound signal descriptors; onomatopoeias; music genres), which we used to manually label the surveyed descriptor classes. We reveal that most taxonomies and ontologies were developed to characterize higher-level semantic relations between sound sources in terms of the sound-generating objects and actions involved (what/how), or in terms of the environmental context (where). This indicates the current lack of a comprehensive ontology of everyday sounds that covers simultaneously all semantic aspects of the relation between sounds. Such an ontology may have a wide range of applications and purposes, ranging from extending our scientific knowledge of auditory processes in the real world, to developing artificial hearing systems.

A functional magnetic resonance imaging examination of audiovisual observation of a point-light string quartet using intersubject correlation and physical feature analysis.

We use functional Magnetic Resonance Imaging (fMRI) to explore synchronized neural responses between observers of audiovisual presentation of a string quartet performance during free viewing. Audio presentation was accompanied by visual presentation of the string quartet as stick figures observed from a static viewpoint. Brain data from 18 musical novices were obtained during audiovisual presentation of a 116 s performance of the allegro of String Quartet, No. 14 in D minor by Schubert played by the 'Quartetto di Cremona.' These data were analyzed using intersubject correlation (ISC). Results showed extensive ISC in auditory and visual areas as well as parietal cortex, frontal cortex and subcortical areas including the medial geniculate and basal ganglia (putamen). These results from a single fixed viewpoint of multiple musicians are greater than previous reports of ISC from unstructured group activity but are broadly consistent with related research that used ISC to explore listening to music or watching solo dance. A feature analysis examining the relationship between brain activity and physical features of the auditory and visual signals yielded findings of a large proportion of activity related to auditory and visual processing, particularly in the superior temporal gyrus (STG) as well as midbrain areas. Motor areas were also involved, potentially as a result of watching motion from the stick figure display of musicians in the string quartet. These results reveal involvement of areas such as the putamen in processing complex musical performance and highlight the potential of using brief naturalistic stimuli to localize distinct brain areas and elucidate potential mechanisms underlying multisensory integration.

Group-level inference of information-based measures for the analyses of cognitive brain networks from neurophysiological data.

The reproducibility crisis in neuroimaging and in particular in the case of underpowered studies has introduced doubts on our ability to reproduce, replicate and generalize findings. As a response, we have seen the emergence of suggested guidelines and principles for neuroscientists known as Good Scientific Practice for conducting more reliable research. Still, every study remains almost unique in its combination of analytical and statistical approaches. While it is understandable considering the diversity of designs and brain data recording, it also represents a striking point against reproducibility. Here, we propose a non-parametric permutation-based statistical framework, primarily designed for neurophysiological data, in order to perform group-level inferences on non-negative measures of information encompassing metrics from information-theory, machine-learning or measures of distances. The framework supports both fixed- and random-effect models to adapt to inter-individuals and inter-sessions variability. Using numerical simulations, we compared the accuracy in ground-truth retrieving of both group models, such as test- and cluster-wise corrections for multiple comparisons. We then reproduced and extended existing results using both spatially uniform MEG and non-uniform intracranial neurophysiological data. We showed how the framework can be used to extract stereotypical task- and behavior-related effects across the population covering scales from the local level of brain regions, inter-areal functional connectivity to measures summarizing network properties. We also present an open-source Python toolbox called Frites1 that includes the proposed statistical pipeline using information-theoretic metrics such as single-trial functional connectivity estimations for the extraction of cognitive brain networks. Taken together, we believe that this framework deserves careful attention as its robustness and flexibility could be the starting point toward the uniformization of statistical approaches.

Functionally homologous representation of vocalizations in the auditory cortex of humans and macaques.

How the evolution of speech has transformed the human auditory cortex compared to other primates remains largely unknown. While primary auditory cortex is organized largely similarly in humans and macaques,1 the picture is much less clear at higher levels of the anterior auditory pathway,2 particularly regarding the processing of conspecific vocalizations (CVs). A "voice region" similar to the human voice-selective areas3,4 has been identified in the macaque right anterior temporal lobe with functional MRI;5 however, its anatomical localization, seemingly inconsistent with that of the human temporal voice areas (TVAs), has suggested a "repositioning of the voice area" in recent human evolution.6 Here we report a functional homology in the cerebral processing of vocalizations by macaques and humans, using comparative fMRI and a condition-rich auditory stimulation paradigm. We find that the anterior temporal lobe of both species possesses cortical voice areas that are bilateral and not only prefer conspecific vocalizations but also implement a representational geometry categorizing them apart from all other sounds in a species-specific but homologous manner. These results reveal a more similar functional organization of higher-level auditory cortex in macaques and humans than currently known.

The representational dynamics of perceived voice emotions evolve from categories to dimensions.

Long-standing affective science theories conceive the perception of emotional stimuli either as discrete categories (for example, an angry voice) or continuous dimensional attributes (for example, an intense and negative vocal emotion). Which position provides a better account is still widely debated. Here we contrast the positions to account for acoustics-independent perceptual and cerebral representational geometry of perceived voice emotions. We combined multimodal imaging of the cerebral response to heard vocal stimuli (using functional magnetic resonance imaging and magneto-encephalography) with post-scanning behavioural assessment of voice emotion perception. By using representational similarity analysis, we find that categories prevail in perceptual and early (less than 200 ms) frontotemporal cerebral representational geometries and that dimensions impinge predominantly on a later limbic-temporal network (at 240 ms and after 500 ms). These results reconcile the two opposing views by reframing the perception of emotions as the interplay of cerebral networks with different representational dynamics that emphasize either categories or dimensions.

The perception of caricatured emotion in voice.

Affective vocalisations such as screams and laughs can convey strong emotional content without verbal information. Previous research using morphed vocalisations (e.g. 25% fear/75% anger) has revealed categorical perception of emotion in voices, showing sudden shifts at emotion category boundaries. However, it is currently unknown how further modulation of vocalisations beyond the veridical emotion (e.g. 125% fear) affects perception. Caricatured facial expressions produce emotions that are perceived as more intense and distinctive, with faster recognition relative to the original and anti-caricatured (e.g. 75% fear) emotions, but a similar effect using vocal caricatures has not been previously examined. Furthermore, caricatures can play a key role in assessing how distinctiveness is identified, in particular by evaluating accounts of emotion perception with reference to prototypes (distance from the central stimulus) and exemplars (density of the stimulus space). Stimuli consisted of four emotions (anger, disgust, fear, and pleasure) morphed at 25% intervals between a neutral expression and each emotion from 25% to 125%, and between each pair of emotions. Emotion perception was assessed using emotion intensity ratings, valence and arousal ratings, speeded categorisation and paired similarity ratings. We report two key findings: 1) across tasks, there was a strongly linear effect of caricaturing, with caricatured emotions (125%) perceived as higher in emotion intensity and arousal, and recognised faster compared to the original emotion (100%) and anti-caricatures (25%-75%); 2) our results reveal evidence for a unique contribution of a prototype-based account in emotion recognition. We show for the first time that vocal caricature effects are comparable to those found previously with facial caricatures. The set of caricatured vocalisations provided open a promising line of research for investigating vocal affect perception and emotion processing deficits in clinical populations.

Contributions of local speech encoding and functional connectivity to audio-visual speech perception.

Seeing a speaker's face enhances speech intelligibility in adverse environments. We investigated the underlying network mechanisms by quantifying local speech representations and directed connectivity in MEG data obtained while human participants listened to speech of varying acoustic SNR and visual context. During high acoustic SNR speech encoding by temporally entrained brain activity was strong in temporal and inferior frontal cortex, while during low SNR strong entrainment emerged in premotor and superior frontal cortex. These changes in local encoding were accompanied by changes in directed connectivity along the ventral stream and the auditory-premotor axis. Importantly, the behavioral benefit arising from seeing the speaker's face was not predicted by changes in local encoding but rather by enhanced functional connectivity between temporal and inferior frontal cortex. Our results demonstrate a role of auditory-frontal interactions in visual speech representations and suggest that functional connectivity along the ventral pathway facilitates speech comprehension in multisensory environments.

Vibrotactile Sensitivity in Active Touch: Effect of Pressing Force.

An experiment was conducted to study the effects of force produced by active touch on vibrotactile perceptual thresholds. The task consisted in pressing the fingertip against a flat rigid surface that provided either sinusoidal or broadband vibration. Three force levels were considered, ranging from light touch to hard press. Finger contact areas were measured during the experiment, showing positive correlation with the respective applied forces. Significant effects on thresholds were found for vibration type and force level. Moreover, possibly due to the concurrent effect of large (unconstrained) finger contact areas, active pressing forces, and long duration stimuli, the measured perceptual thresholds are considerably lower than what previously reported in the literature.

A statistical framework for neuroimaging data analysis based on mutual information estimated via a gaussian copula.

We begin by reviewing the statistical framework of information theory as applicable to neuroimaging data analysis. A major factor hindering wider adoption of this framework in neuroimaging is the difficulty of estimating information theoretic quantities in practice. We present a novel estimation technique that combines the statistical theory of copulas with the closed form solution for the entropy of Gaussian variables. This results in a general, computationally efficient, flexible, and robust multivariate statistical framework that provides effect sizes on a common meaningful scale, allows for unified treatment of discrete, continuous, unidimensional and multidimensional variables, and enables direct comparisons of representations from behavioral and brain responses across any recording modality. We validate the use of this estimate as a statistical test within a neuroimaging context, considering both discrete stimulus classes and continuous stimulus features. We also present examples of analyses facilitated by these developments, including application of multivariate analyses to MEG planar magnetic field gradients, and pairwise temporal interactions in evoked EEG responses. We show the benefit of considering the instantaneous temporal derivative together with the raw values of M/EEG signals as a multivariate response, how we can separately quantify modulations of amplitude and direction for vector quantities, and how we can measure the emergence of novel information over time in evoked responses. Open-source Matlab and Python code implementing the new methods accompanies this article. Hum Brain Mapp 38:1541-1573, 2017. © 2016 Wiley Periodicals, Inc.

The dominance of haptics over audition in controlling wrist velocity during striking movements.

Skilled interactions with sounding objects, such as drumming, rely on resolving the uncertainty in the acoustical and tactual feedback signals generated by vibrating objects. Uncertainty may arise from mis-estimation of the objects' geometry-independent mechanical properties, such as surface stiffness. How multisensory information feeds back into the fine-tuning of sound-generating actions remains unexplored. Participants (percussionists, non-percussion musicians, or non-musicians) held a stylus and learned to control their wrist velocity while repeatedly striking a virtual sounding object whose surface stiffness was under computer control. Sensory feedback was manipulated by perturbing the surface stiffness specified by audition and haptics in a congruent or incongruent manner. The compensatory changes in striking velocity were measured as the motor effects of the sensory perturbations, and sensory dominance was quantified by the asymmetry of congruency effects across audition and haptics. A pronounced dominance of haptics over audition suggested a superior utility of somatosensation developed through long-term experience with object exploration. Large interindividual differences in the motor effects of haptic perturbation potentially arose from a differential reliance on the type of tactual prediction error for which participants tend to compensate: vibrotactile force versus object deformation. Musical experience did not have much of an effect beyond a slightly greater reliance on object deformation in mallet percussionists. The bias toward haptics in the presence of crossmodal perturbations was greater when participants appeared to rely on object deformation feedback, suggesting a weaker association between haptically sensed object deformation and the acoustical structure of concomitant sound during everyday experience of actions upon objects.

Predicting the timing of dynamic events through sound: Bouncing balls.

Dynamic information in acoustical signals produced by bouncing objects is often used by listeners to predict the objects' future behavior (e.g., hitting a ball). This study examined factors that affect the accuracy of motor responses to sounds of real-world dynamic events. In experiment 1, listeners heard 2-5 bounces from a tennis ball, ping-pong, basketball, or wiffle ball, and would tap to indicate the time of the next bounce in a series. Across ball types and number of bounces, listeners were extremely accurate in predicting the correct bounce time (CT) with a mean prediction error of only 2.58% of the CT. Prediction based on a physical model of bouncing events indicated that listeners relied primarily on temporal cues when estimating the timing of the next bounce, and to a lesser extent on the loudness and spectral cues. In experiment 2, the timing of each bounce pattern was altered to correspond to the bounce timing pattern of another ball, producing stimuli with contradictory acoustic cues. Nevertheless, listeners remained highly accurate in their estimates of bounce timing. This suggests that listeners can adopt their estimates of bouncing-object timing based on acoustic cues that provide most veridical information about dynamic aspects of object behavior.

A language-familiarity effect for speaker discrimination without comprehension.

The influence of language familiarity upon speaker identification is well established, to such an extent that it has been argued that "Human voice recognition depends on language ability" [Perrachione TK, Del Tufo SN, Gabrieli JDE (2011) Science 333(6042):595]. However, 7-mo-old infants discriminate speakers of their mother tongue better than they do foreign speakers [Johnson EK, Westrek E, Nazzi T, Cutler A (2011) Dev Sci 14(5):1002-1011] despite their limited speech comprehension abilities, suggesting that speaker discrimination may rely on familiarity with the sound structure of one's native language rather than the ability to comprehend speech. To test this hypothesis, we asked Chinese and English adult participants to rate speaker dissimilarity in pairs of sentences in English or Mandarin that were first time-reversed to render them unintelligible. Even in these conditions a language-familiarity effect was observed: Both Chinese and English listeners rated pairs of native-language speakers as more dissimilar than foreign-language speakers, despite their inability to understand the material. Our data indicate that the language familiarity effect is not based on comprehension but rather on familiarity with the phonology of one's native language. This effect may stem from a mechanism analogous to the "other-race" effect in face recognition.

Gender differences in the temporal voice areas.

There is not only evidence for behavioral differences in voice perception between female and male listeners, but also recent suggestions for differences in neural correlates between genders. The fMRI functional voice localizer (comprising a univariate analysis contrasting stimulation with vocal vs. non-vocal sounds) is known to give robust estimates of the temporal voice areas (TVAs). However, there is growing interest in employing multivariate analysis approaches to fMRI data (e.g., multivariate pattern analysis; MVPA). The aim of the current study was to localize voice-related areas in both female and male listeners and to investigate whether brain maps may differ depending on the gender of the listener. After a univariate analysis, a random effects analysis was performed on female (n = 149) and male (n = 123) listeners and contrasts between them were computed. In addition, MVPA with a whole-brain searchlight approach was implemented and classification maps were entered into a second-level permutation based random effects models using statistical non-parametric mapping (SnPM; Nichols and Holmes, 2002). Gender differences were found only in the MVPA. Identified regions were located in the middle part of the middle temporal gyrus (bilateral) and the middle superior temporal gyrus (right hemisphere). Our results suggest differences in classifier performance between genders in response to the voice localizer with higher classification accuracy from local BOLD signal patterns in several temporal-lobe regions in female listeners.

Automatic domain-general processing of sound source identity in the left posterior middle frontal gyrus.

Identifying sound sources is fundamental to developing a stable representation of the environment in the face of variable auditory information. The cortical processes underlying this ability have received little attention. In two fMRI experiments, we investigated passive adaptation to (Exp. 1) and explicit discrimination of (Exp. 2) source identities for different categories of auditory objects (voices, musical instruments, environmental sounds). All cortical effects of source identity were independent of high-level category information, and were accounted for by sound-to-sound differences in low-level structure (e.g., loudness). A conjunction analysis revealed that the left posterior middle frontal gyrus (pMFG) adapted to identity repetitions during both passive listening and active discrimination tasks. These results indicate that the comparison of sound source identities in a stream of auditory stimulation recruits the pMFG in a domain-general way, i.e., independent of the sound category, based on information contained in the low-level acoustical structure. pMFG recruitment during both passive listening and explicit identity comparison tasks also suggests its automatic engagement in sound source identity processing.

The production and perception of emotionally expressive walking sounds: similarities between musical performance and everyday motor activity.

Several studies have investigated the encoding and perception of emotional expressivity in music performance. A relevant question concerns how the ability to communicate emotions in music performance is acquired. In accordance with recent theories on the embodiment of emotion, we suggest here that both the expression and recognition of emotion in music might at least in part rely on knowledge about the sounds of expressive body movements. We test this hypothesis by drawing parallels between musical expression of emotions and expression of emotions in sounds associated with a non-musical motor activity: walking. In a combined production-perception design, two experiments were conducted, and expressive acoustical features were compared across modalities. An initial performance experiment tested for similar feature use in walking sounds and music performance, and revealed that strong similarities exist. Features related to sound intensity, tempo and tempo regularity were identified as been used similarly in both domains. Participants in a subsequent perception experiment were able to recognize both non-emotional and emotional properties of the sound-generating walkers. An analysis of the acoustical correlates of behavioral data revealed that variations in sound intensity, tempo, and tempo regularity were likely used to recognize expressed emotions. Taken together, these results lend support the motor origin hypothesis for the musical expression of emotions.

Singing numbers in cognitive space--a dual-task study of the link between pitch, space, and numbers.

We assessed the automaticity of spatial-numerical and spatial-musical associations by testing their intentionality and load sensitivity in a dual-task paradigm. In separate sessions, 16 healthy adults performed magnitude and pitch comparisons on sung numbers with variable pitch. Stimuli and response alternatives were identical, but the relevant stimulus attribute (pitch or number) differed between tasks. Concomitant tasks required retention of either color or location information. Results show that spatial associations of both magnitude and pitch are load sensitive and that the spatial association for pitch is more powerful than that for magnitude. These findings argue against the automaticity of spatial mappings in either stimulus dimension.

Abstract encoding of auditory objects in cortical activity patterns.

The human brain is thought to process auditory objects along a hierarchical temporal "what" stream that progressively abstracts object information from the low-level structure (e.g., loudness) as processing proceeds along the middle-to-anterior direction. Empirical demonstrations of abstract object encoding, independent of low-level structure, have relied on speech stimuli, and non-speech studies of object-category encoding (e.g., human vocalizations) often lack a systematic assessment of low-level information (e.g., vocalizations are highly harmonic). It is currently unknown whether abstract encoding constitutes a general functional principle that operates for auditory objects other than speech. We combined multivariate analyses of functional imaging data with an accurate analysis of the low-level acoustical information to examine the abstract encoding of non-speech categories. We observed abstract encoding of the living and human-action sound categories in the fine-grained spatial distribution of activity in the middle-to-posterior temporal cortex (e.g., planum temporale). Abstract encoding of auditory objects appears to extend to non-speech biological sounds and to operate in regions other than the anterior temporal lobe. Neural processes for the abstract encoding of auditory objects might have facilitated the emergence of speech categories in our ancestors.

Perceptual evaluation of violins: a quantitative analysis of preference judgments by experienced players.

The overall goal of the research presented here is to better understand how players evaluate violins within the wider context of finding relationships between measurable vibrational properties of instruments and their perceived qualities. In this study, the reliability of skilled musicians to evaluate the qualities of a violin was examined. In a first experiment, violinists were allowed to freely play a set of different violins and were then asked to rank the instruments by preference. Results showed that players were self-consistent, but a large amount of inter-individual variability was present. A second experiment was then conducted to investigate the origin of inter-individual differences in the preference for violins and to measure the extent to which different attributes of the instrument influence preference. Again, results showed large inter-individual variations in the preference for violins, as well as in assessing various characteristics of the instruments. Despite the significant lack of agreement in preference and the variability in how different criteria are evaluated between individuals, violin players tend to agree on the relevance of sound "richness" and, to a lesser extent, "dynamic range" for determining preference.