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J E Hind

Publications and source records attributed to J E Hind.

At least 19 recordsLinked to original sources

Auditory cortex on the human posterior superior temporal gyrus.

The human superior temporal cortex plays a critical role in hearing, speech, and language, yet its functional organization is poorly understood. Evoked potentials (EPs) to auditory click-train stimulation presented binaurally were recorded chronically from penetrating electrodes implanted in Heschl's gyrus (HG), from pial-surface electrodes placed on the lateral superior temporal gyrus (STG), or from both simultaneously, in awake humans undergoing surgery for medically intractable epilepsy. The distribution of averaged EPs was restricted to a relatively small area on the lateral surface of the posterior STG. In several cases, there were multiple foci of high amplitude EPs lying along this acoustically active portion of STG. EPs recorded simultaneously from HG and STG differed in their sensitivities to general anesthesia and to changes in rate of stimulus presentation. Results indicate that the acoustically active region on the STG is a separate auditory area, functionally distinct from the HG auditory field(s). We refer to this acoustically sensitive area of the STG as the posterior lateral superior temporal area (PLST). Electrical stimulation of HG resulted in short-latency EPs in an area that overlaps PLST, indicating that PLST receives a corticocortical input, either directly or indirectly, from HG. These physiological findings are in accord with anatomic evidence in humans and in nonhuman primates that the superior temporal cortex contains multiple interconnected auditory areas.

Acoustic Stimulation↗

Spatial receptive fields of primary auditory cortical neurons in quiet and in the presence of continuous background noise.

Spatial receptive fields of primary auditory (AI) neurons were studied by delivering, binaurally, synthesized virtual-space signals via earphones to cats under barbiturate anesthesia. Signals were broadband or narrowband transients presented in quiet anechoic space or in acoustic space filled with uncorrelated continuous broadband noise. In the absence of background noise, AI virtual space receptive fields (VSRFs) are typically large, representing a quadrant or more of acoustic space. Within the receptive field, onset latency and firing strength form functional gradients. We hypothesized earlier that functional gradients in the receptive field provide information about sound-source direction. Previous studies indicated that spatial gradients could remain relatively constant across changes in signal intensity. In the current experiments we tested the hypothesis that directional sensitivity to a transient signal, as reflected in the gradient structure of VSRFs of AI neurons, is also retained in the presence of a continuous background noise. When background noise was introduced three major affects on VSRFs were observed. 1) The size of the VSRF was reduced, accompanied by a reduction of firing strength and lengthening of response latency for signals at an acoustic axis and on-lines of constant azimuth and elevation passing through the acoustic axis. These effects were monotonically related to the intensity of the background noise over a noise intensity range of approximately 30 dB. 2) The noise intensity-dependent changes in VSRFs were mirrored by the changes that occurred when the signal intensity was changed in signal-alone conditions. Thus adding background noise was equivalent to a shift in the threshold of a directional signal, and this shift was seen across the spatial receptive field. 3) The spatial gradients of response strength and latency remained evident over the range of background noise intensity that reduced spike count and lengthened onset latency. Those gradients along the azimuth that spanned the frontal midline tended to remain constant in slope and position in the face of increasing intensity of background noise. These findings are consistent with our hypothesis that, under background noise conditions, information that underlies directional acuity and accuracy is retained within the spatial receptive fields of an ensemble of AI neurons.

Acoustic Stimulation↗

Modeling of auditory spatial receptive fields with spherical approximation functions.

A spherical approximation technique is presented that affords a mathematical characterization of a virtual space receptive field (VSRF) based on first-spike latency in the auditory cortex of cat. Parameterizing directional sensitivity in this fashion is much akin to the use of difference-of-Gaussian (DOG) functions for modeling neural responses in visual cortex. Artificial neural networks and approximation techniques typically have been applied to problems conforming to a multidimensional Cartesian input space. The problem with using classical planar Gaussians is that radial symmetry and consistency on the plane actually translate into directionally dependent distortion on spherical surfaces. An alternative set of spherical basis functions, the von Mises basis function (VMBF), is used to eliminate spherical approximation distortion. Unlike the Fourier transform or spherical harmonic expansions, the VMBFs are nonorthogonal, and hence require some form of gradient-descent search for optimal estimation of parameters in the modeling of the VSRF. The optimization equations required to solve this problem are presented. Three descriptive classes of VSRF (contralateral, frontal, and ipsilateral) approximations are investigated, together with an examination of the residual error after parameter optimization. The use of the analytic receptive field model in computational models of population coding of sound direction is discussed, together with the importance of quantifying receptive field gradients. Because spatial hearing is by its very nature three dimensional or, more precisely, two dimensional (directional) on the sphere, we find that spatial receptive field models are best developed on the sphere.

Acoustic Stimulation↗

The structure of spatial receptive fields of neurons in primary auditory cortex of the cat.

Transient broad-band stimuli that mimic in their spectrum and time waveform sounds arriving from a speaker in free space were delivered to the tympanic membranes of barbiturized cats via sealed and calibrated earphones. The full array of such signals constitutes a virtual acoustic space (VAS). The extra-cellular response to a single stimulus at each VAS direction, consisting of one or a few precisely time-locked spikes, was recorded from neurons in primary auditory cortex. Effective sound directions form a virtual space receptive field (VSRF). Near threshold, most VSRFs were confined to one quadrant of acoustic space and were located on or near the acoustic axis. Generally, VSRFs expanded monotonically with increases in stimulus intensity, with some occupying essentially all of the acoustic space. The VSRF was not homogeneous with respect to spike timing or firing strength. Typically, onset latency varied by as much as 4-5 msec across the VSRF. A substantial proportion of recorded cells exhibited a gradient of first-spike latency within the VSRF. Shortest latencies occupied a core of the VSRF, on or near the acoustic axis, with longer latency being represented progressively at directions more distant from the core. Remaining cells had VSRFs that exhibited no such gradient. The distribution of firing probability was mapped in those experiments in which multiple trials were carried out at each direction. For some cells there was a positive correlation between latency and firing probability.

Acoustic Stimulation↗

Simulation of free-field sound sources and its application to studies of cortical mechanisms of sound localization in the cat.

We synthesized a set of signals (clicks) for earphone delivery whose waveforms and amplitude spectra, measured at the eardrum, mimic those of sounds arriving from a free-field source. The complete stimulus set represents 1816 sound-source directions, which together surround the head to form a 'virtual acoustic space' for the cat. Virtual-space stimuli were delivered via calibrated earphones sealed into the external meatus in cats under barbiturate anesthesia. Neurons recorded in AI cortex exhibited sensitivity to the direction of sound in virtual acoustic space. The aggregation of effective sound directions formed a virtual space receptive field (VSRF). At 20 dB above minimal threshold, VSRFs fell into one of several categories based on spatial dimension and location. Most VSRFs were confined to either the contralateral (59%) or ipsilateral (10%) sound hemifield. Seven percent spanned the frontal quadrants and 16% were omnidirectional. Eight percent fit into no clear category and were termed 'complex'. The size, shape, and location of VSRFs remained stable over many hours of recording. The results are in essential agreement with free-field studies. VSRFs were found to be shaped by excitatory and inhibitory interactions of activity arriving from the two ears. Some cortical neurons were found to preserve the spectral information in the free-field sound which was generated by the acoustical properties of the head and pinna, filtered by the cochlea and transmitted by auditory nerve fibers.

Acoustic Stimulation↗

An insert earphone system for delivery of spectrally shaped signals for physiological studies.

Acoustic signals arriving at the eardrum in free-space carry directionally dependent temporal and spectral information resulting from the acoustical effects of the body, head, and external ear as well as from differences in the length of the sound path to each ear. Through analysis of the responses of single auditory neurons, the acoustical and neural mechanisms by which sounds in free-space are localized are being studied. The approach involves simulation of free-field signals at the two eardrums of a cat via earphones and a study of the neuronal responses to such a virtual acoustic space. This approach makes it possible to manipulate different stimulus parameters independently in order to examine their role in determining the spatial characteristics of neuronal response. This report describes an insert earphone system designed for the delivery of such simulated signals which are broadband transients having complex spectra that mimic the acoustic transfer function of the external ear for frequency components up to 30 kHz or more.

Acoustic Stimulation↗

Direction-dependent spectral properties of cat external ear: new data and cross-species comparisons.

Free-field to eardrum transfer functions were measured in anesthetized cats inside an anechoic chamber. Direction-dependent transformations were determined by measurement of sound-pressure levels using a small probe tube microphone surgically implanted in a ventral position near the tympanic membrane. Loudspeaker and probe microphone characteristics were eliminated by subtraction of the signal recorded in the free field with no animal present. Complexities of the transfer function, which include the presence of prominent spectral notches in the 8- to 18-kHz frequency region, are due primarily to the acoustical properties of the pinna. Differential amplification of frequency components within the broadband stimulus occurs as a function of source direction. Spectral features vary systematically with changes in both elevation (EL) and azimuth (AZ). The contrast between a notch and its shoulders is enhanced in the interaural spectral records. Spectral data from single source locations and spatial data for single frequencies at many locations are presented and comparisons with other species are drawn. It is suggested that spectral features in the 8- to 18-kHz region provide some of the necessary spectral information for sound localization and that the contrast in spectral energy between the frequencies at the notch and its shoulders is a potential directional cue.

Acoustic Stimulation↗

Interaural time differences: implications regarding the neurophysiology of sound localization.

Interaural time differences (ITDs) were measured from 400--7000 Hz on cats in order to provide quantitative data for use in physiological/behavioral studies on sound localization. ITDs derived from clicks and the initial portion of tone bursts showed a pronounced roughness and frequency dependence. This frequency dependence is most evident at higher angles of incidence and indicates that a single ITD will not always represent a single position on the azimuth. Controls demonstrate that most of the roughness in these functions was due to reflections off the surface supporting the animal and that the measured ITDs corresponded to predictions made by steady-state theory. Measurements made with and without the pinnae in position indicate that they have relatively little effect on these ITD functions, particularly for frequencies below 2500 Hz and for small angles of incidence. In spite of acoustic limitations exemplified by the roughness and frequency dependence of these functions, ITDs generated by sound sources situated close to the midline provide reliable localization cues that are much better than those derived from sources well out on the azimuth. Finally, it is noted that another ITD, the group ITD, can be ascribed to an acoustic signal. Calculations based on the measured steady-state ITDs show differences between the group and steady-state ITDs over a given range of frequencies. Differences between the group and steady-state ITD can be significant, and it is argued that: (1) The group ITD can provide a localization cue to the auditory system that is distinct from the steady-state ITD; and (2) it is possible these group ITDs are used by the nervous system to localize sound sources in realistic situations.

Acoustic Stimulation↗