Joint population coding and temporal coherence link an attended talker's voice and location features in naturalistic multi-talker scenes.

Listeners readily extract multi-dimensional auditory objects such as a 'localized talker' from complex acoustic scenes with multiple talkers. Yet, the neural mechanisms underlying simultaneous encoding and linking of different sound features - for example, a talker's voice and location - are poorly understood. We analyzed invasive intracranial recordings in neurosurgical patients attending to a localized talker in real-life cocktail party scenarios. We found that sensitivity to an individual talker's voice and location features was distributed throughout auditory cortex and that neural sites exhibited a gradient from sensitivity to a single feature to joint sensitivity to both features. On a population level, cortical response patterns of both dual-feature sensitive sites but also single-feature sensitive sites revealed simultaneous encoding of an attended talker's voice and location features. However, for single-feature sensitive sites, the representation of the primary feature was more precise. Further, sites which selective tracked an attended speech stream concurrently encoded an attended talker's voice and location features, indicating that such sites combine selective tracking of an attended auditory object with encoding of the object's features. Finally, we found that attending a localized talker selectively enhanced temporal coherence between single-feature voice sensitive sites and single-feature location sensitive sites, providing an additional mechanism for linking voice and location in multi-talker scenes. These results demonstrate that a talker's voice and location features are linked during multi-dimensional object formation in naturalistic multi-talker scenes by joint population coding as well as by temporal coherence between neural sites. SIGNIFICANCE STATEMENT: Listeners effortlessly extract auditory objects from complex acoustic scenes consisting of multiple sound sources in naturalistic, spatial sound scenes. Yet, how the brain links different sound features to form a multi-dimensional auditory object is poorly understood. We investigated how neural responses encode and integrate an attended talker's voice and location features in spatial multi-talker sound scenes to elucidate which neural mechanisms underlie simultaneous encoding and linking of different auditory features. Our results show that joint population coding as well as temporal coherence mechanisms contribute to distributed multi-dimensional auditory object encoding. These findings shed new light on cortical functional specialization and multidimensional auditory object formation in complex, naturalistic listening scenes. HIGHLIGHTS: Cortical responses to an single talker exhibit a distributed gradient, ranging from sites that are sensitive to both a talker's voice and location (dual-feature sensitive sites) to sites that are sensitive to either voice or location (single-feature sensitive sites).Population response patterns of dual-feature sensitive sites encode voice and location features of the attended talker in multi-talker scenes jointly and with equal precision.Despite their sensitivity to a single feature at the level of individual cortical sites, population response patterns of single-feature sensitive sites also encode location and voice features of a talker jointly, but with higher precision for the feature they are primarily sensitive to.Neural sites which selectively track an attended speech stream concurrently encode the attended talker's voice and location features.Attention selectively enhances temporal coherence between voice and location selective sites over time.Joint population coding as well as temporal coherence mechanisms underlie distributed multi-dimensional auditory object encoding in auditory cortex.

Modality-specific brain representations during automatic processing of face, voice and body expressions.

A central question in affective science and one that is relevant for its clinical applications is how emotions provided by different stimuli are experienced and represented in the brain. Following the traditional view emotional signals are recognized with the help of emotion concepts that are typically used in descriptions of mental states and emotional experiences, irrespective of the sensory modality. This perspective motivated the search for abstract representations of emotions in the brain, shared across variations in stimulus type (face, body, voice) and sensory origin (visual, auditory). On the other hand, emotion signals like for example an aggressive gesture, trigger rapid automatic behavioral responses and this may take place before or independently of full abstract representation of the emotion. This pleads in favor specific emotion signals that may trigger rapid adaptative behavior only by mobilizing modality and stimulus specific brain representations without relying on higher order abstract emotion categories. To test this hypothesis, we presented participants with naturalistic dynamic emotion expressions of the face, the whole body, or the voice in a functional magnetic resonance (fMRI) study. To focus on automatic emotion processing and sidestep explicit concept-based emotion recognition, participants performed an unrelated target detection task presented in a different sensory modality than the stimulus. By using multivariate analyses to assess neural activity patterns in response to the different stimulus types, we reveal a stimulus category and modality specific brain organization of affective signals. Our findings are consistent with the notion that under ecological conditions emotion expressions of the face, body and voice may have different functional roles in triggering rapid adaptive behavior, even if when viewed from an abstract conceptual vantage point, they may all exemplify the same emotion. This has implications for a neuroethologically grounded emotion research program that should start from detailed behavioral observations of how face, body, and voice expressions function in naturalistic contexts.

Imaging-based frequency mapping for cochlear implants - Evaluated using a daily randomized controlled trial.

Background: Due to variation in electrode design, insertion depth and cochlear morphology, patients with a cochlear implant (CI) often have to adapt to a substantial mismatch between the characteristic response frequencies of cochlear neurons and the stimulus frequencies assigned to electrode contacts. We introduce an imaging-based fitting intervention, which aimed to reduce frequency-to-place mismatch by aligning frequency mapping with the tonotopic position of electrodes. Results were evaluated in a novel trial set-up where subjects crossed over between intervention and control using a daily within-patient randomized approach, immediately from the start of CI rehabilitation. Methods: Fourteen adult participants were included in this single-blinded, daily randomized clinical trial. Based on a fusion of pre-operative imaging and a post-operative cone beam CT scan (CBCT), mapping of electrical input was aligned to natural place-pitch arrangement in the individual cochlea. That is, adjustments to the CI's frequency allocation table were made so electrical stimulation of frequencies matched as closely as possible with corresponding acoustic locations in the cochlea. For a period of three months, starting at first fit, a scheme was implemented whereby the blinded subject crossed over between the experimental and standard fitting program using a daily randomized wearing schedule, and thus effectively acted as their own control. Speech outcomes (such as speech intelligibility in quiet and noise, sound quality and listening effort) were measured with both settings throughout the study period. Results: On a group level, standard fitting obtained subject preference and showed superior results in all outcome measures. In contrast, two out of fourteen subjects preferred the imaging-based fitting and correspondingly had better speech understanding with this setting compared to standard fitting. Conclusion: On average, cochlear implant fitting based on individual tonotopy did not elicit higher speech intelligibility but variability in individual results strengthen the potential for individualized frequency fitting. The novel trial design proved to be a suitable method for evaluation of experimental interventions in a prospective trial setup with cochlear implants.

Interaction of bottom-up and top-down neural mechanisms in spatial multi-talker speech perception.

How the human auditory cortex represents spatially separated simultaneous talkers and how talkers' locations and voices modulate the neural representations of attended and unattended speech are unclear. Here, we measured the neural responses from electrodes implanted in neurosurgical patients as they performed single-talker and multi-talker speech perception tasks. We found that spatial separation between talkers caused a preferential encoding of the contralateral speech in Heschl's gyrus (HG), planum temporale (PT), and superior temporal gyrus (STG). Location and spectrotemporal features were encoded in different aspects of the neural response. Specifically, the talker's location changed the mean response level, whereas the talker's spectrotemporal features altered the variation of response around response's baseline. These components were differentially modulated by the attended talker's voice or location, which improved the population decoding of attended speech features. Attentional modulation due to the talker's voice only appeared in the auditory areas with longer latencies, but attentional modulation due to location was present throughout. Our results show that spatial multi-talker speech perception relies upon a separable pre-attentive neural representation, which could be further tuned by top-down attention to the location and voice of the talker.

Reorganization of Sound Location Processing in the Auditory Cortex of Blind Humans.

Auditory spatial tasks induce functional activation in the occipital-visual-cortex of early blind humans. Less is known about the effects of blindness on auditory spatial processing in the temporal-auditory-cortex. Here, we investigated spatial (azimuth) processing in congenitally and early blind humans with a phase-encoding functional magnetic resonance imaging (fMRI) paradigm. Our results show that functional activation in response to sounds in general-independent of sound location-was stronger in the occipital cortex but reduced in the medial temporal cortex of blind participants in comparison with sighted participants. Additionally, activation patterns for binaural spatial processing were different for sighted and blind participants in planum temporale. Finally, fMRI responses in the auditory cortex of blind individuals carried less information on sound azimuth position than those in sighted individuals, as assessed with a 2-channel, opponent coding model for the cortical representation of sound azimuth. These results indicate that early visual deprivation results in reorganization of binaural spatial processing in the auditory cortex and that blind individuals may rely on alternative mechanisms for processing azimuth position.

Cortical mechanisms of spatial hearing.

Humans and other animals use spatial hearing to rapidly localize events in the environment. However, neural encoding of sound location is a complex process involving the computation and integration of multiple spatial cues that are not represented directly in the sensory organ (the cochlea). Our understanding of these mechanisms has increased enormously in the past few years. Current research is focused on the contribution of animal models for understanding human spatial audition, the effects of behavioural demands on neural sound location encoding, the emergence of a cue-independent location representation in the auditory cortex, and the relationship between single-source and concurrent location encoding in complex auditory scenes. Furthermore, computational modelling seeks to unravel how neural representations of sound source locations are derived from the complex binaural waveforms of real-life sounds. In this article, we review and integrate the latest insights from neurophysiological, neuroimaging and computational modelling studies of mammalian spatial hearing. We propose that the cortical representation of sound location emerges from recurrent processing taking place in a dynamic, adaptive network of early (primary) and higher-order (posterior-dorsal and dorsolateral prefrontal) auditory regions. This cortical network accommodates changing behavioural requirements and is especially relevant for processing the location of real-life, complex sounds and complex auditory scenes.

Active Sound Localization Sharpens Spatial Tuning in Human Primary Auditory Cortex.

Spatial hearing sensitivity in humans is dynamic and task-dependent, but the mechanisms in human auditory cortex that enable dynamic sound location encoding remain unclear. Using functional magnetic resonance imaging (fMRI), we assessed how active behavior affects encoding of sound location (azimuth) in primary auditory cortical areas and planum temporale (PT). According to the hierarchical model of auditory processing and cortical functional specialization, PT is implicated in sound location ("where") processing. Yet, our results show that spatial tuning profiles in primary auditory cortical areas (left primary core and right caudo-medial belt) sharpened during a sound localization ("where") task compared with a sound identification ("what") task. In contrast, spatial tuning in PT was sharp but did not vary with task performance. We further applied a population pattern decoder to the measured fMRI activity patterns, which confirmed the task-dependent effects in the left core: sound location estimates from fMRI patterns measured during active sound localization were most accurate. In PT, decoding accuracy was not modulated by task performance. These results indicate that changes of population activity in human primary auditory areas reflect dynamic and task-dependent processing of sound location. As such, our findings suggest that the hierarchical model of auditory processing may need to be revised to include an interaction between primary and functionally specialized areas depending on behavioral requirements.SIGNIFICANCE STATEMENT According to a purely hierarchical view, cortical auditory processing consists of a series of analysis stages from sensory (acoustic) processing in primary auditory cortex to specialized processing in higher-order areas. Posterior-dorsal cortical auditory areas, planum temporale (PT) in humans, are considered to be functionally specialized for spatial processing. However, this model is based mostly on passive listening studies. Our results provide compelling evidence that active behavior (sound localization) sharpens spatial selectivity in primary auditory cortex, whereas spatial tuning in functionally specialized areas (PT) is narrow but task-invariant. These findings suggest that the hierarchical view of cortical functional specialization needs to be extended: our data indicate that active behavior involves feedback projections from higher-order regions to primary auditory cortex.