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Contains fulltext : 291625.pdf (Publisher’s version ) (Open Access) Many cochlear implant users with binaural residual (acoustic) hearing benefit from combining electric and acoustic stimulation (EAS) in the implanted ear with acoustic amplification in the other. These bimodal EAS listeners can potentially use low-frequency binaural cues to localize sounds. However, their hearing is generally asymmetric for mid- and high-frequency sounds, perturbing or even abolishing binaural cues. Here, we investigated the effect of a frequency-dependent binaural asymmetry in hearing thresholds on sound localization by seven bimodal EAS listeners. Frequency dependence was probed by presenting sounds with power in low-, mid-, high-, or mid-to-high-frequency bands. Frequency-dependent hearing asymmetry was present in the bimodal EAS listening condition (when using both devices) but was also induced by independently switching devices on or off. Using both devices, hearing was near symmetric for low frequencies, asymmetric for mid frequencies with better hearing thresholds in the implanted ear, and monaural for high frequencies with no hearing in the non-implanted ear. Results show that sound-localization performance was poor in general. Typically, localization was strongly biased toward the better hearing ear. We observed that hearing asymmetry was a good predictor for these biases. Notably, even when hearing was symmetric a preferential bias toward the ear using the hearing aid was revealed. We discuss how frequency dependence of any hearing asymmetry may lead to binaural cues that are spatially inconsistent as the spectrum of a sound changes. We speculate that this inconsistency may prevent accurate sound-localization even after long-term exposure to the hearing asymmetry.

Poster presentation for the Conference on Implantable Auditory Prosthesis (CIAP),Lake Tahoe, USA (July, 2023).

We tested whether sensitivity to acoustic spectrotemporal modulations can be observed from reaction times for normal-hearing and impaired-hearing conditions. In a manual reaction-time task, normal-hearing listeners had to detect the onset of a ripple (with density between 0-8 cycles/octave and a fixed modulation depth of 50%), that moved up or down the log-frequency axis at constant velocity (between 0-64 Hz), in an otherwise-unmodulated broadband white-noise. Spectral and temporal modulations elicited band-pass filtered sensitivity characteristics, with fastest detection rates around 1 cycle/oct and 32 Hz for normal-hearing conditions. These results closely resemble data from other studies that typically used the modulation-depth threshold as a sensitivity criterion. To simulate hearing-impairment, stimuli were processed with a 6-channel cochlear-implant vocoder, and a hearing-aid simulation that introduced separate spectral smearing and low-pass filtering. Reaction times were always much slower compared to normal hearing, especially for the highest spectral densities. Binaural performance was predicted well by the benchmark race model of binaural independence, which models statistical facilitation of independent monaural channels. For the impaired-hearing simulations this implied a “best-of-both-worlds” principle in which the listeners relied on the hearing-aid ear to detect spectral modulations, and on the cochlear-implant ear for temporal-modulation detection. Although singular-value decomposition indicated that the joint spectrotemporal sensitivity matrix could be largely reconstructed from independent temporal and spectral sensitivity functions, in line with time-spectrum separability, a substantial inseparable spectral-temporal interaction was present in all hearing conditions. These results suggest that the reaction-time task yields a valid and effective objective measure of acoustic spectrotemporal-modulation sensitivity.

Cochlear implants are neuroprostheses that stimulate the auditory nerve using electric pulses, allowing people experiencing a sensorineural hearing loss to recover hearing sensations. The auditory nerve can integrate current over time, potentially affecting feature representations. This integration may also depend on auditory nerve conditions. Additionally, setups testing temporal interactions use discrimination or subjective rating paradigms to assess differences at supra-threshold levels, not allowing a continuum of responses from threshold, and requiring additional time to get responses. An evaluation of temporal integration can help us identify channels that are more susceptible to temporal interactions, providing further information for device fitting. We tested temporal integration in cochlear implant recipients by measuring reaction times to simple stimulus manipulations, namely: amplitude changes, polarity and separation between pulses.

Sound localization and identification are challenging in acoustically rich environments. The relation between these two processes is still poorly understood. As natural sound-sources rarely occur exactly simultaneously, we wondered whether the auditory system could identify (“what”) and localize (“where”) two spatially separated sounds with synchronous onsets. While listeners typically report hearing a single source at an average location, one study found that both sounds may be accurately localized if listeners are explicitly being told two sources exist. We here tested whether simultaneous source identification (one vs. two) and localization is possible, by letting listeners choose to make either one or two head-orienting saccades to the perceived location(s). Results show that listeners could identify two sounds only when presented on different sides of the head, and that identification accuracy increased with their spatial separation. Notably, listeners were unable to accurately localize either sound, irrespective of whether one or two sounds were identified. Instead, the first (or only) response always landed near the average location, while second responses were unrelated to the targets. We conclude that localization of synchronous sounds in the absence of prior information is impossible. We discuss that the putative cortical ‘what’ pathway may not transmit relevant information to the ‘where’ pathway. We examine how a broadband interaural correlation cue could help to correctly identify the presence of two sounds without being able to localize them. We propose that the persistent averaging behavior reveals that the ‘where’ system intrinsically assumes that synchronous sounds originate from a single source.Significance StatementIt is poorly understood whether identification (‘what’) of sounds and their localization (‘where’) are inter-related, or independent neural processes. We measured sound-localization responses towards synchronous sounds to examine potential coupling of these processes. We varied the spatial configurations of two sounds and found that although identification improved considerably with larger spatial separation, their localization was unaffected: responses were always directed towards the average location. This shows absence of mutual coupling of information between the ‘what’ and ‘where’ streams in the auditory system. We also show how broadband interaural correlation could explain the improved identification results, without affecting localization performance, and explain how the persistent spatial averaging could be understood from strong internal priors regarding sound synchronicity.

The ventriloquism aftereffect (VAE) describes the persistent shift of perceived sound location after having been adapted to a ventriloquism condition, in which the sound was repeatedly paired with a displaced visual stimulus. In the latter case, participants consistently mislocalize the sound in the direction of the visual stimulus (ventriloquism effect, VE). Previous studies provide conflicting reports regarding the strength of the VAE, ranging from 0 to nearly 100%. Moreover, there is controversy about its generalization to different sounds than the one inducing the VE, ranging from no transfer at all, to full transfer across different sound spectra. Here, we imposed the VE for three different sounds: a low-frequency and a high-frequency narrow-band noise, and a broadband Gaussian white noise (GWN). In the adaptation phase, listeners generated fast goal-directed head movements to localize the sound, presented across a 70 deg range in the horizontal plane, while ignoring a visual distracter that was consistently displaced 10 deg to the right of the sound. In the post-adaptation phase, participants localized narrow-band sounds with center frequencies from 0.5 to 8 kHz, as well as GWN, without the visual distracter. Our results show that the VAE amounted to approximately 40% of the VE and generalized well across the entire frequency domain. We also found that the strength of the VAE correlated with the pre-adaptation sound-localization performance. We compare our results with previous reports and discuss different hypotheses regarding optimal audio-visual cue integration.Significance statementWhere we believe sounds to emanate from is heavily influenced by what we see. We experience this illusion - the ventriloquism effect - regularly. For example, when watching movies, we perceive the voices of actors to originate from the actors’ lips rather than from the speakers. Long exposure to a spatial mismatch between what we hear and what we see, leads to a recalibration of our localization of sounds - the ventriloquism aftereffect. Unclear was whether exposure to an audiovisual mismatch for one sound type would also lead to adaptive changes for other sounds. The present study shows that the ventriloquism aftereffect generalizes, implying that it does not act at an early auditory stage, but at a late multisensory stage.

The cochlear implant (CI) allows profoundly deaf individuals to partially recover hearing. Still, due to the coarse acoustic information provided by the implant, CI users have considerable difficulties in recognizing speech, especially in noisy environments. CI users therefore rely heavily on visual cues to augment speech comprehension, more so than normal-hearing individuals. However, it is unknown how attention to one (focused) or both (divided) modalities plays a role in multisensory speech recognition. Here we show that unisensory speech listening and reading were negatively impacted in divided-attention tasks for CI users - but not for normal-hearing individuals. Our psychophysical experiments revealed that, as expected, listening thresholds were consistently better for the normal-hearing, while lipreading thresholds were largely similar for the two groups. Moreover, audiovisual speech recognition for normal-hearing individuals could be described well by probabilistic summation of auditory and visual speech recognition, while CI users were better integrators than expected from statistical facilitation alone. Our results suggest that this benefit in integration comes at a cost. Unisensory speech recognition is degraded for CI users when attention needs to be divided across modalities. We conjecture that CI users exhibit an integration-attention trade-off. They focus solely on a single modality during focused-attention tasks, but need to divide their limited attentional resources in situations with uncertainty about the upcoming stimulus modality. We argue that in order to determine the benefit of a CI for speech comprehension, situational factors need to be discounted by presenting speech in realistic or complex audiovisual environments.Significance statementDeaf individuals using a cochlear implant require significant amounts of effort to listen in noisy environments due to their impoverished hearing. Lipreading can benefit them and reduce the burden of listening by providing an additional source of information. Here we show that the improved speech recognition for audiovisual stimulation comes at a cost, however, as the cochlear-implant users now need to listen and speech-read simultaneously, paying attention to both modalities. The data suggests that cochlear-implant users run into the limits of their attentional resources, and we argue that they, unlike normal-hearing individuals, always need to consider whether a multisensory benefit outweighs the unisensory cost in everyday environments.

An interesting problem for the human saccadic eye-movement system is how to deal with the degrees-of-freedom problem: the six extra-ocular muscles provide three rotational degrees of freedom, while only two are needed to point gaze at any direction. Measurements show that 3D eye orientations during head-fixed saccades in far-viewing conditions lie in Listing’s plane (LP), in which the eye’s cyclotorsion is zero (Listing’s law, LL). Moreover, while saccades are executed as single-axis rotations around a stable eye-angular velocity axis, they follow straight trajectories in LP. Another distinctive saccade property is their nonlinear main-sequence dynamics: the affine relationship between saccade size and movement duration, and the saturation of peak velocity with amplitude. To explain all these properties, we developed a computational model, based on a simplified and upscaled robotic prototype of an eye with 3 degrees of freedom, driven by three independent motor commands, coupled to three antagonistic elastic muscle pairs. As the robotic prototype was not intended to faithfully mimic the detailed biomechanics of the human eye, we did not impose specific prior mechanical constraints on the ocular plant that could, by themselves, generate Listing’s law and the main-sequence. Instead, our goal was to study how these properties can emerge from the application of optimal control principles to simplified eye models. We performed a numerical linearization of the nonlinear system dynamics around the origin using system identification techniques, and developed open-loop controllers for 3D saccade generation. Applying optimal control to the simulated model, could reproduce both Listing’s law and and the main-sequence. We verified the contribution of different terms in the cost optimization functional to realistic 3D saccade behavior, and identified four essential terms: total energy expenditure by the motors, movement duration, gaze accuracy, and the total static force exerted by the muscles during fixation. Our findings suggest that Listing’s law, as well as the saccade dynamics and their trajectories, may all emerge from the same common mechanism that aims to optimize speed-accuracy trade-off for saccades, while minimizing the total muscle force during eccentric fixation., Author summary Saccades are rapid eye movements that humans and other animals perform three to four times per second to scan and perceive the environment around them. These movements orient the eye in space with high precision and in a highly stereotyped fashion. Existing studies on animal models advocate that both mechanical and neuronal functions play an important role in the control of the saccades, but some facts are still not fully understood due to difficulties in experimenting and measuring the variables in living animals. Instead, robots are computational and physical models of reality that expose all its variables and can be programmed in interpretable ways. We have built a robotic model of an artificial eye containing the basic ingredients of human eyes: full 3D rotations, viscous friction and 6 muscle-like actuators connected to the eyeball in a geometry similar to the biological system. By synthesizing robotic eye control systems we found that important characteristics of the movements become similar to human saccades when the control relies on few simple fundamental principles: the maximization of saccade accuracy and the minimization of saccade duration, energy in control, and force in the muscles during fixation.

To program a goal-directed response in the presence of acoustic reflections, the audio-motor system should suppress the detection of time-delayed sources. We examined the effects of spatial separation and interstimulus delay on the ability of human listeners to localize a pair of broadband sounds in the horizontal plane. Participants indicated how many sounds were heard and where these were perceived by making one or two head-orienting localization responses. Results suggest that perceptual fusion of the two sounds depends on delay and spatial separation. Leading and lagging stimuli in close spatial proximity required longer stimulus delays to be perceptually separated than those further apart. Whenever participants heard one sound, their localization responses for synchronous sounds were oriented to a weighted average of both source locations. For short delays, responses were directed toward the leading stimulus location. Increasing spatial separation enhanced this effect. For longer delays, responses were again directed toward a weighted average. When participants perceived two sounds, the first and the second response were directed to either of the leading and lagging source locations. Perceived locations were interchanged often in their temporal order (in ∼40% of trials). We show that the percept of two sounds occurring requires sufficient spatiotemporal separation, after which localization can be performed with high accuracy. We propose that the percept of temporal order of two concurrent sounds results from a different process than localization and discuss how dynamic lateral excitatory-inhibitory interactions within a spatial sensorimotor map could explain the findings.

Although moving sound-sources abound in natural auditory scenes, it is not clear how the human brain processes auditory motion. Previous studies have indicated that, although ocular localization responses to stationary sounds are quite accurate, ocular smooth pursuit of moving sounds is very poor. We here demonstrate that human subjects faithfully track a sound’s unpredictable movements in the horizontal plane with smooth-pursuit responses of the head. Our analysis revealed that the stimulus–response relation was well described by an under-damped passive, second-order low-pass filter in series with an idiosyncratic, fixed, pure delay. The model contained only two free parameters: the system’s damping coefficient, and its central (resonance) frequency. We found that the latter remained constant at about 0.6 Hz throughout the experiment for all subjects. Interestingly, the damping coefficient systematically increased with trial number, suggesting the presence of an adaptive mechanism in the auditory pursuit system. This mechanism functions even for unpredictable sound-motion trajectories endowed with fixed, but covert, frequency characteristics in open-loop tracking conditions. We conjecture that the auditory pursuit system optimizes a trade-off between response speed and effort. Taken together, our data support the existence of a pursuit system for auditory head-tracking, which would suggest the presence of a neural representation of a spatial auditory fovea. Significance statement Inspired by the visual ocular smooth-pursuit system, several studies have used eye movements to track moving sounds, but obtained poor pursuit performance, which led to the idea that the auditory system lacks sensitivity to sound velocity. We here demonstrate accurate head-pursuit of sounds, moving along unpredictable trajectories in the horizontal plane. Interestingly, the auditory pursuit responses adapted to the covert movement spectrum of the stimulus ensemble, from which we infer that the system may optimize a trade-off between movement speed and effort. Our results support the existence of an auditory pursuit system, and we discuss its implications for the neural mechanisms that represent and track moving sounds.

The latency of the auditory steady-state response (ASSR) may provide valuable information regarding the integrity of the auditory system, as it could potentially reveal the presence of multiple intracerebral sources. To estimate multiple latencies from high-order ASSRs, we propose a novel two-stage procedure that consists of a nonparametric estimation method, called apparent latency from phase coherence (ALPC), followed by a heuristic sequential forward selection algorithm (SFS). Compared with existing methods, ALPC-SFS requires few prior assumptions, and is straightforward to implement for higher-order nonlinear responses to multi-cosine sound complexes with their initial phases set to zero. It systematically evaluates the nonlinear components of the ASSRs by estimating multiple latencies, automatically identifies involved ASSR components, and reports a latency consistency index (LCI). To verify the proposed method, we performed simulations for several scenarios: two nonlinear subsystems with different or overlapping outputs. We compared the results from our method with predictions from existing, parametric methods. We also recorded the EEG from ten normal-hearing adults by bilaterally presenting superimposed tones with four frequencies that evoke a unique set of ASSRs. From these ASSRs, two major latencies were found to be stable across subjects on repeated measurement days. The two latencies are dominated by low-frequency (LF) (near 40 Hz, at around 41-52 ms) and high-frequency (HF) (>80 Hz, at around 21-27 ms) ASSR components. The frontal-central (FC) brain region showed longer latencies on LF components, but shorter latencies on HF components, when compared with temporal-lobe regions. In conclusion, the proposed nonparametric ALPC-SFS method, applied to zero-phase, multi-cosine sound complexes is more suitable for evaluating embedded nonlinear systems underlying ASSRs than existing methods. It may therefore be a promising objective measure for hearing performance and auditory cortex (dys)function. The Matlab scripts for the ALPC-SFS method is available here: https://github.com/ieeeWang/ALPC-SFS-method-Matlab-scripts.

Chronic single-sided deaf (CSSD) listeners lack the availability of binaural difference cues to localize sounds in the horizontal plane. Hence, for directional hearing they have to rely on different types of monaural cues: loudness perceived in their hearing ear, which is affected in a systematic way by the acoustic head shadow, on spectral cues provided by the low-pass filtering characteristic of the head, and on high-frequency spectral-shape cues from the pinna of their hearing ear. Presumably, these cues are differentially weighted against prior assumptions on the properties of sound sources in the environment. The rules guiding this weighting process are not well understood. In this preliminary study, we trained three CSSD listeners to localize a fixed intensity, high-pass filtered sound source at ten locations in the horizontal plane with visual feedback. After training, we compared their localization performance to sounds with different intensities, presented in the two-dimensional frontal hemifield to their pre-training results. We show that the training had rapidly readjusted the contributions of monaural cues and internal priors, which resulted to be imposed by the multisensory information provided during the training. We compare the results with the strategies found for the acute monaural hearing condition of normal-hearing listeners, described in an earlier study [1].

Functional near-infrared spectroscopy (fNIRS) is an optical, non-invasive neuroimaging technique that investigates human brain activity by calculating concentrations of oxy- and deoxyhemoglobin. The aim of this publication is to review the current state of the art as to how fNIRS has been used to study auditory function. We address temporal and spatial characteristics of the hemodynamic response to auditory stimulation as well as experimental factors that affect fNIRS data such as acoustic and stimulus-driven effects. The rising importance that fNIRS is generating in auditory neuroscience underlines the strong potential of the technology, and it seems likely that fNIRS will become a useful clinical tool.

The midbrain superior colliculus (SC) is a crucial sensorimotor interface in the generation of rapid saccadic gaze shifts. For every saccade it recruits a large population of cells in its vectorial motor map. Supra-threshold electrical microstimulation in the SC reveals that the stimulated site produces the saccade vector specified by the motor map. Electrically evoked saccades (E-saccades) have kinematic properties that strongly resemble natural, visual-evoked saccades (V-saccades), with little influence of the stimulation parameters. Moreover, synchronous stimulation at two sites yields eye movements that resemble a weighted vector average of the individual stimulation effects. Single-unit recordings have indicated that the SC population acts as a vectorial pulse generator by specifying the instantaneous gaze-kinematics through dynamic summation of the movement effects of all SC spike trains. But how to reconcile the a-specific stimulation pulses with these intricate saccade properties? We recently developed a spiking neural network model of the SC, in which microstimulation initially activates a relatively small set of (~50) neurons around the electrode tip, which subsequently sets up a large population response (~5,000 neurons) through lateral synaptic interactions. Single-site microstimulation in this network thus produces the saccade properties and firing rate profiles as seen in single-unit recording experiments. We here show that this mechanism also accounts for many results of simultaneous double stimulation at different SC sites. The resulting E-saccade trajectories resemble a weighted average of the single-site effects, in which stimulus current strength of the electrode pulses serve as weighting factors. We discuss under which conditions the network produces effects that deviate from experimental results.

Contains fulltext : 177716.pdf (Publisher’s version ) (Open Access) OBJECTIVES: This study aimed to improve access to high-frequency interaural level differences (ILD), by applying extreme frequency compression (FC) in the hearing aid (HA) of 13 bimodal listeners, using a cochlear implant (CI) and conventional HA in opposite ears. DESIGN: An experimental signal-adaptive frequency-lowering algorithm was tested, compressing frequencies above 160 Hz into the individual audible range of residual hearing, but only for consonants (adaptive FC), thus protecting vowel formants, with the aim to preserve speech perception. In a cross-over design with at least 5 weeks of acclimatization between sessions, bimodal performance with and without adaptive FC was compared for horizontal sound localization, speech understanding in quiet and in noise, and vowel, consonant and voice-pitch perception. RESULTS: On average, adaptive FC did not significantly affect any of the test results. Yet, two subjects who were fitted with a relatively weak frequency compression ratio, showed improved horizontal sound localization. After the study, four subjects preferred adaptive FC, four preferred standard frequency mapping, and four had no preference. Noteworthy, the subjects preferring adaptive FC were those with best performance on all tasks, both with and without adaptive FC. CONCLUSION: On a group level, extreme adaptive FC did not change sound localization and speech understanding in bimodal listeners. Possible reasons are too strong compression ratios, insufficient residual hearing or that the adaptive switching, although preserving vowel perception, may have been ineffective to produce consistent ILD cues. Individual results suggested that two subjects were able to integrate the frequency-compressed HA input with that of the CI, and benefitted from enhanced binaural cues for horizontal sound localization. 12 p.

The auditory system relies on binaural differences and spectral pinna cues to localize sounds in azimuth and elevation. However, the acoustic input can be unreliable, due to uncertainty about the environment, and neural noise. A possible strategy to reduce sound-location uncertainty is to integrate the sensory observations with sensorimotor information from previous experience, to infer where sounds are more likely to occur. We investigated whether and how human sound localization performance is affected by the spatial distribution of target sounds, and changes thereof. We tested three different open-loop paradigms, in which we varied the spatial range of sounds in different ways. For the narrowest ranges, target-response gains were highly idiosyncratic and deviated from an optimal gain predicted by error-minimization; in the horizontal plane the deviation typically consisted of a response overshoot. Moreover, participants adjusted their behavior by rapidly adapting their gain to the target range, both in elevation and in azimuth, yielding behavior closer to optimal for larger target ranges. Notably, gain changes occurred without any exogenous feedback about performance. We discuss how the findings can be explained by a sub-optimal model in which the motor-control system reduces its response error across trials to within an acceptable range, rather than strictly minimizing the error.

The brain estimates the two-dimensional direction of sounds from the pressure-induced displacements of the eardrums. Accurate localization along the horizontal plane (azimuth angle) is enabled by binaural difference cues in timing and intensity. Localization along the vertical plane (elevation angle), including frontal and rear directions, relies on spectral cues made possible by the elevation dependent filtering in the idiosyncratic pinna cavities. However, the problem of extracting elevation from the sensory input is ill-posed, since the spectrum results from a convolution between source spectrum and the particular head-related transfer function (HRTF) associated with the source elevation, which are both unknown to the system. It is not clear how the auditory system deals with this problem, or which implicit assumptions it makes about source spectra. By varying the spectral contrast of broadband sounds around the 6–9 kHz band, which falls within the human pinna’s most prominent elevation-related spectral notch, we here suggest that the auditory system performs a weighted spectral analysis across different frequency bands to estimate source elevation. We explain our results by a model, in which the auditory system weighs the different spectral bands, and compares the convolved weighted sensory spectrum with stored information about its own HRTFs, and spatial prior assumptions.

Single-unit recordings in head-restrained monkeys indicated that the population of saccade-related cells in the midbrain Superior Colliculus (SC) encodes the kinematics of desired straight saccade trajectories by the cumulative number of spikes. In addition, the nonlinear main sequence of saccades (their amplitude–peak velocity saturation) emerges from a spatial gradient of peak-firing rates of collicular neurons, rather than from neural saturation at brain-stem burst generators. We here extend this idea to eye-head gaze shifts and illustrate how the cumulative spike-count in head-unrestrained monkeys relates to the desired gaze trajectory and its kinematics. We argue that the output of the motor SC is an abstract desired gaze-motor signal, which drives in a feedforward way the instantaneous kinematics of ongoing gaze shifts, including the strong influence of initial eye position on gaze kinematics. We propose that the neural population acts as a vectorial gaze pulse-generator for eye-head saccades, which is sub-sequently decomposed into signals that drive both motor systems in appropriate craniocentric reference frames within a dynamic gaze-velocity feedback loop.

The midbrain superior colliculus (SC) generates a rapid saccadic eye movement to a sensory stimulus by recruiting a population of cells in its topographically organized motor map. Supra-threshold electrical microstimulation in the SC reveals that the site of stimulation produces a normometric saccade vector with little effect of the stimulation parameters. Moreover, electrically evoked saccades (E-saccades) have kinematic properties that strongly resemble natural, visual-evoked saccades (V-saccades). These findings support models in which the saccade vector is determined by a center-of-gravity computation of activated neurons, while its trajectory and kinematics arise from downstream feedback circuits in the brainstem. Recent single-unit recordings, however, have indicated that the SC population also specifies instantaneous kinematics. These results support an alternative model, in which the desired saccade trajectory, including its kinematics, follows from instantaneous summation of movement effects of all SC spike trains. But how to reconcile this model with microstimulation results? Although it is thought that microstimulation activates a large population of SC neurons, the mechanism through which it arises is unknown. We developed a spiking neural network model of the SC, in which microstimulation directly activates a relatively small set of neurons around the electrode tip, which subsequently sets up a large population response through lateral synaptic interactions. We show that through this mechanism the population drives an E-saccade with near-normal kinematics that are largely independent of the stimulation parameters. Only at very low stimulus intensities the network recruits a population with low firing rates, resulting in abnormally slow saccades., Author summary The midbrain Superior Colliculus (SC) is crucial for generating rapid saccadic eye movements. It contains a topographically organized map of visuomotor space, in which a large population of recruited cells determines the metrics and kinematics of saccades. The dynamic spike-counting model explains how this population encodes the ensuing eye movement through linear dynamic summation of the spike-effects of each recruited neuron. Electrical microstimulation in the motor map produces saccades with a vector that corresponds with the location of the electrode in the map, and with very similar kinematics as normal visually-evoked saccades. Although the summation model accounts for the kinematics of visually-evoked saccades, it could, so far, not be reconciled with the effects of microstimulation. Here we modeled the SC motor map with a spiking neural network, in which cells are connected through tuned local excitatory and global inhibitory synapses. The network was tuned such that stimulation directly recruits only a small subset of neurons, from which activity rapidly spreads across the motor map to set up a (near-)normal population. Simulations with this computational model show that this scheme explains the metrics and kinematics of electrically evoked saccades.

In dynamic visual or auditory gaze double-steps, a brief target flash or sound burst is presented in midflight of an ongoing eye-head gaze shift. Behavioral experiments in humans and monkeys have indicated that the subsequent eye and head movements to the target are goal-directed, regardless of stimulus timing, first gaze shift characteristics, and initial conditions. This remarkable behavior requires that the gaze-control system 1) has continuous access to accurate signals about eye-in-head position and ongoing eye-head movements, 2) that it accounts for different internal signal delays, and 3) that it is able to update the retinal ( TE) and head-centric ( TH) target coordinates into appropriate eye-centered and head-centered motor commands on millisecond time scales. As predictive, feedforward remapping of targets cannot account for this behavior, we propose that targets are transformed and stored into a stable reference frame as soon as their sensory information becomes available. We present a computational model, in which recruited cells in the midbrain superior colliculus drive eyes and head to the stored target location through a common dynamic oculocentric gaze-velocity command, which is continuously updated from the stable goal and transformed into appropriate oculocentric and craniocentric motor commands. We describe two equivalent, yet conceptually different, implementations that both account for the complex, but accurate, kinematic behaviors and trajectories of eye-head gaze shifts under a variety of challenging multisensory conditions, such as in dynamic visual-auditory multisteps.

F.C. Donders was born on May 27, 1818 in Tilburg, the Netherlands. This Summer, the Donders Institute for Brain, Cognition and Behaviour at Radboud University Nijmegen commemorated the 200th annive...

How the human auditory system learns to map complex pinna-induced spectral-shape cues onto veridical estimates of sound-source elevation in the median plane is still unclear. Earlier studies demonstrated considerable sound-localisation plasticity after applying pinna moulds, and to altered vision. Several factors may contribute to auditory spatial learning, like visual or motor feedback, or updated priors. We here induced perceptual learning for sounds with degraded spectral content, having weak, but consistent, elevation-dependent cues, as demonstrated by low-gain stimulus-response relations. During training, we provided visual feedback for only six targets in the midsagittal plane, to which listeners gradually improved their response accuracy. Interestingly, listeners’ performance also improved without visual feedback, albeit less strongly. Post-training results showed generalised improved response behaviour, also to non-trained locations and acoustic spectra, presented throughout the two-dimensional frontal hemifield. We argue that the auditory system learns to reweigh contributions from low-informative spectral bands to update its prior elevation estimates, and explain our results with a neuro-computational model.

Contains fulltext : 200557.pdf (Publisher’s version ) (Open Access) Sensory representations are typically endowed with intrinsic noise, leading to variability and inaccuracies in perceptual responses. The Bayesian framework accounts for an optimal strategy to deal with sensory-motor uncertainty, by combining the noisy sensory input with prior information regarding the distribution of stimulus properties. The maximum-a-posteriori (MAP) estimate selects the perceptual response from the peak (mode) of the resulting posterior distribution that ensure optimal accuracy-precision trade-off when the underlying distributions are Gaussians (minimal mean-squared error, with minimum response variability). We tested this model on human eye- movement responses toward broadband sounds, masked by various levels of background noise, and for head movements to sounds with poor spectral content. We report that the response gain (accuracy) and variability (precision) of the elevation response components changed systematically with the signal-to-noise ratio of the target sound: gains were high for high SNRs and decreased for low SNRs. In contrast, the azimuth response components maintained high gains for all conditions, as predicted by maximum-likelihood estimation. However, we found that the elevation data did not follow the MAP prediction. Instead, results were better described by an alternative decision strategy, in which the response results from taking a random sample from the posterior in each trial. We discuss two potential implementations of a simple posterior sampling scheme in the auditory system that account for the results and argue that although the observed response strategies for azimuth and elevation are sub-optimal with respect to their variability, it allows the auditory system to actively explore the environment in the absence of adequate sensory evidence.

Two synchronous sounds at different locations in the midsagittal plane induce a fused percept at a weighted-average position, with weights depending on relative sound intensities. In the horizontal plane, sound fusion (stereophony) disappears with a small onset asynchrony of 1–4 ms. The leading sound then fully determines the spatial percept (the precedence effect). Given that accurate localisation in the median plane requires an analysis of pinna-related spectral-shape cues, which takes ~25–30 ms of sound input to complete, we wondered at what time scale a precedence effect for elevation would manifest. Listeners localised the first of two sounds, with spatial disparities between 10–80 deg, and inter-stimulus delays between 0–320 ms. We demonstrate full fusion (averaging), and largest response variability, for onset asynchronies up to at least 40 ms for all spatial disparities. Weighted averaging persisted, and gradually decayed, for delays >160 ms, suggesting considerable backward masking. Moreover, response variability decreased with increasing delays. These results demonstrate that localisation undergoes substantial spatial blurring in the median plane by lagging sounds. Thus, the human auditory system, despite its high temporal resolution, is unable to spatially dissociate sounds in the midsagittal plane that co-occur within a time window of at least 160 ms.

Contains fulltext : 136102.pdf (Publisher’s version ) (Closed access) Primate sensory systems subserve complex neurocomputational functions. Consequently, these systems are organised anatomically in a distributed fashion, commonly linking areas to form specialised processing streams. Each stream is related to a specific function, as evidenced from studies of the visual cortex, which features rather prominent segregation into spatial and non-spatial domains. It has been hypothesised that other sensory systems, including auditory, are organised in a similar way on the cortical level. Recent studies offer rich qualitative evidence for the dual stream hypothesis. Here we provide a new paradigm to quantitatively uncover these patterns in the auditory system, based on an analysis of multiple anatomical studies using multivariate techniques. As a test case, we also apply our assessment techniques to more ubiquitously-explored visual system. Importantly, the introduced framework opens the possibility for these techniques to be applied to other neural systems featuring a dichotomised organisation, such as language or music perception.

After hearing a tone, the human auditory system becomes more sensitive to similar tones than to other tones. Current auditory models explain this phenomenon by a simple bandpass attention filter. Here, we demonstrate that auditory attention involves multiple pass-bands around octave-related frequencies above and below the cued tone. Intriguingly, this "octave effect" not only occurs for physically presented tones, but even persists for the missing fundamental in complex tones, and for imagined tones. Our results suggest neural interactions combining octave-related frequencies, likely located in nonprimary cortical regions. We speculate that this connectivity scheme evolved from exposure to natural vibrations containing octave-related spectral peaks, e.g., as produced by vocal cords. ispartof: Proceedings of the National Academy of Sciences of the United States of America vol:110 issue:38 pages:15225-15230 ispartof: location:United States status: published

Human sound localization in the mid-saggital plane (elevation) relies on an analysis of the idiosyncratic spectral shape cues provided by the head and pinnae. However, because the actual free-field stimulus spectrum is a-priori unknown to the auditory system, the problem of extracting the elevation angle from the sensory spectrum is ill-posed. Here we test different spectral localization models by eliciting head movements toward broad-band noise stimuli with randomly shaped, rippled amplitude spectra emanating from a speaker at a fixed location, while varying the ripple bandwidth between 1.5 and 5.0 cycles/octave. Six listeners participated in the experiments. From the distributions of localization responses toward the individual stimuli, we estimated the listeners' spectral-shape cues underlying their elevation percepts, by applying maximum-likelihood estimation. The reconstructed spectral cues resulted to be invariant to the considerable variation in ripple bandwidth, and for each listener they had a remarkable resemblance to the idiosyncratic head-related transfer functions (HRTFs). These results are not in line with models that rely on the detection of a single peak or notch in the amplitude spectrum, nor with a local analysis of first- and second-order spectral derivatives. Instead, our data support a model in which the auditory system performs a cross-correlation between the sensory input at the eardrum-auditory nerve, and stored representations of HRTF spectral shapes, to extract the perceived elevation angle.

To program a goal-directed response in the presence of multiple sounds, the audiomotor system should separate the sound sources. The authors examined whether the brain can segregate synchronous broadband sounds in the midsagittal plane, using amplitude modulations as an acoustic discrimination cue. To succeed in this task, the brain has to use pinna-induced spectral-shape cues and temporal envelope information. The authors tested spatial segregation performance in the midsagittal plane in two paradigms in which human listeners were required to localize, or distinguish, a target amplitude-modulated broadband sound when a non-modulated broadband distractor was played simultaneously at another location. The level difference between the amplitude-modulated and distractor stimuli was systematically varied, as well as the modulation frequency of the target sound. The authors found that participants were unable to segregate, or localize, the synchronous sounds. Instead, they invariably responded toward a level-weighted average of both sound locations, irrespective of the modulation frequency. An increased variance in the response distributions for double sounds of equal level was also observed, which cannot be accounted for by a segregation model, or by a probabilistic averaging model.

Single-unit recordings suggest that the midbrain superior colliculus (SC) acts as an optimal controller for saccadic gaze shifts. The SC is proposed to be the site within the visuomotor system where the nonlinear spatial-to-temporal transformation is carried out: the population encodes the intended saccade vector by its location in the motor map (spatial), and its trajectory and velocity by the distribution of firing rates (temporal). The neurons' burst profiles vary systematically with their anatomical positions and intended saccade vectors, to account for the nonlinear main-sequence kinematics of saccades. Yet, the underlying collicular mechanisms that could result in these firing patterns are inaccessible to current neurobiological techniques. Here, we propose a simple spiking neural network model that reproduces the spike trains of saccade-related cells in the intermediate and deep SC layers during saccades. The model assumes that SC neurons have distinct biophysical properties for spike generation that depend on their anatomical position in combination with a center-surround lateral connectivity. Both factors are needed to account for the observed firing patterns. Our model offers a basis for neuronal algorithms for spatiotemporal transformations and bio-inspired optimal controllers.

Contains fulltext : 117242.pdf (Publisher’s version ) (Closed access) It is unclear whether top-down processing in the auditory cortex (AC) interferes with its bottom-up analysis of sound. Recent studies indicated non-acoustic modulations of AC responses, and that attention changes a neuron's spectrotemporal tuning. As a result, the AC would seem ill-suited to represent a stable acoustic environment, which is deemed crucial for auditory perception. To assess whether top-down signals influence acoustic tuning in tasks without directed attention, we compared monkey single-unit AC responses to dynamic spectrotemporal sounds under different behavioral conditions. Recordings were mostly made from neurons located in primary fields (primary AC and area R of the AC) that were well tuned to pure tones, with short onset latencies. We demonstrated that responses in the AC were substantially modulated during an auditory detection task and that these modulations were systematically related to top-down processes. Importantly, despite these significant modulations, the spectrotemporal receptive fields of all neurons remained remarkably stable. Our results suggest multiplexed encoding of bottom-up acoustic and top-down task-related signals at single AC neurons. This mechanism preserves a stable representation of the acoustic environment despite strong non-acoustic modulations. 13 p.

Contains fulltext : 167811.pdf (Publisher’s version ) (Open Access) Conclusion In users of a cochlear implant (CI) and a hearing aid (HA) in contralateral ears, frequency-dependent loudness balancing between devices did, on average, not lead to improved speech understanding as compared to broadband balancing. However, nine out of 15 bimodal subjects showed significantly better speech understanding with either one of the fittings. Objectives Sub-optimal fittings and mismatches in loudness are possible explanations for the large individual differences seen in listeners using bimodal stimulation. Methods HA gain was adjusted for soft and loud input sounds in three frequency bands (0-548, 548-1000, and >1000 Hz) to match loudness with the CI. This procedure was compared to a simple broadband balancing procedure that reflected current clinical practice. In a three-visit cross-over design with 4 weeks between sessions, speech understanding was tested in quiet and in noise and questionnaires were administered to assess benefit in real world. Results Both procedures resulted in comparable HA gains. For speech in noise, a marginal bimodal benefit of 0.3 +/- 4 dB was found, with large differences between subjects and spatial configurations. Speech understanding in quiet and in noise did not differ between the two loudness balancing procedures. 7 p.

Contains fulltext : 208416.pdf (Publisher’s version ) (Open Access) Bilateral cochlear-implant (CI) users and single-sided deaf listeners with a CI are less effective at localizing sounds than normal-hearing (NH) listeners. This performance gap is due to the degradation of binaural and monaural sound localization cues, caused by a combination of device-related and patient-related issues. In this study, we targeted the device-related issues by measuring sound localization performance of 11 NH listeners, listening to free-field stimuli processed by a real-time CI vocoder. The use of a real-time vocoder is a new approach, which enables testing in a free-field environment. For the NH listening condition, all listeners accurately and precisely localized sounds according to a linear stimulus-response relationship with an optimal gain and a minimal bias both in the azimuth and in the elevation directions. In contrast, when listening with bilateral real-time vocoders, listeners tended to orient either to the left or to the right in azimuth and were unable to determine sound source elevation. When listening with an NH ear and a unilateral vocoder, localization was impoverished on the vocoder side but improved toward the NH side. Localization performance was also reflected by systematic variations in reaction times across listening conditions. We conclude that perturbation of interaural temporal cues, reduction of interaural level cues, and removal of spectral pinna cues by the vocoder impairs sound localization. Listeners seem to ignore cues that were made unreliable by the vocoder, leading to acute reweighting of available localization cues. We discuss how current CI processors prevent CI users from localizing sounds in everyday environments.

We examined horizontal directional hearing in patients with acquired severe unilateral conductive hearing loss (UCHL). All patients (n = 12) had been fitted with a bone conduction device (BCD) to restore bilateral hearing. The patients were tested in the unaided (monaural) and aided (binaural) hearing condition. Five listeners without hearing loss were tested as a control group while listening with a monaural plug and earmuff, or with both ears (binaural). We randomly varied stimulus presentation levels to assess whether listeners relied on the acoustic head-shadow effect (HSE) for horizontal (azimuth) localization. Moreover, to prevent sound localization on the basis of monaural spectral shape cues from head and pinna, subjects were exposed to narrow band (1/3 octave) noises. We demonstrate that the BCD significantly improved sound localization in 8/12 of the UCHL patients. Interestingly, under monaural hearing (BCD off), we observed fairly good unaided azimuth localization performance in 4/12 of the patients. Our multiple regression analysis shows that all patients relied on the ambiguous HSE for localization. In contrast, acutely plugged control listeners did not employ the HSE. Our data confirm and further extend results of recent studies on the use of sound localization cues in chronic and acute monaural listening.