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John C Middlebrooks

Publications and source records attributed to John C Middlebrooks.

7 recordsLinked to original sources

Spatial sensitivity in field PAF of cat auditory cortex.

We compared the spatial tuning properties of neurons in two fields [primary auditory cortex (A1) and posterior auditory field (PAF)] of cat auditory cortex. Broadband noise bursts of 80-ms duration were presented from loudspeakers throughout 360 degrees in the horizontal plane (azimuth) or 260 degrees in the vertical median plane (elevation). Sound levels varied from 20 to 40 dB above units' thresholds. We recorded neural spike activity simultaneously from 16 sites in field PAF and/or A1 of alpha-chloralose-anesthetized cats. We assessed spatial sensitivity by examining the dependence of spike count and response latency on stimulus location. In addition, we used an artificial neural network (ANN) to assess the information about stimulus location carried by spike patterns of single units and of ensembles of 2-32 units. The results indicate increased spatial sensitivity, more uniform distributions of preferred locations, and greater tolerance to changes in stimulus intensity among PAF units relative to A1 units. Compared to A1 units, PAF units responded at significantly longer latencies, and latencies varied more strongly with stimulus location. ANN analysis revealed significantly greater information transmission by spike patterns of PAF than A1 units, primarily reflecting the information transmitted by latency variation in PAF. Finally, information rates grew more rapidly with the number of units included in neural ensembles for PAF than A1. The latter finding suggests more accurate population coding of space in PAF, made possible by a more diverse population of neural response types.

Acoustic Stimulation↗

Listener weighting of cues for lateral angle: the duplex theory of sound localization revisited.

The virtual auditory space technique was used to quantify the relative strengths of interaural time difference (ITD), interaural level difference (ILD), and spectral cues in determining the perceived lateral angle of wideband, low-pass, and high-pass noise bursts. Listeners reported the apparent locations of virtual targets that were presented over headphones and filtered with listeners' own directional transfer functions. The stimuli were manipulated by delaying or attenuating the signal to one ear (by up to 600 micros or 20 dB) or by altering the spectral cues at one or both ears. Listener weighting of the manipulated cues was determined by examining the resulting localization response biases. In accordance with the Duplex Theory defined for pure-tones, listeners gave high weight to ITD and low weight to ILD for low-pass stimuli, and high weight to ILD for high-pass stimuli. Most (but not all) listeners gave low weight to ITD for high-pass stimuli. This weight could be increased by amplitude-modulating the stimuli or reduced by lengthening stimulus onsets. For wideband stimuli, the ITD weight was greater than or equal to that given to ILD. Manipulations of monaural spectral cues and the interaural level spectrum had little influence on lateral angle judgements.

Cues↗

Auditory cortical images of cochlear-implant stimuli: coding of stimulus channel and current level.

This study quantified the accuracy with which populations of neurons in the auditory cortex can represent aspects of electrical cochlear stimuli presented through a cochlear implant. We tested the accuracy of coding of the place of stimulation (i.e., identification of the active stimulation channel) and of the stimulus current level. Physiological data came from the companion study, which recorded spike activity of neurons simultaneously from 16 sites along the tonotopic axis of the guinea pig's auditory cortex. In that study, cochlear electrical stimuli were presented to acutely deafened animals through a 6-electrode animal version of the 22-electrode Nucleus banded electrode array (Cochlear). Cochlear electrode configurations consisted of monopolar (MP), bipolar (BP + N) with N inactive electrodes between the active and return electrodes (0 < or = N < or = 3), tripolar (TP) with one active electrode and two flanking return electrodes, and common ground (CG) with one active electrode and as many as five return electrodes. In the present analysis, an artificial neural network was trained to recognize spatiotemporal patterns of cortical activity in response to single presentations of particular stimuli and, thereby, to identify those stimuli. The accuracy of pair-wise discrimination of stimulation channels or of current levels was represented by the discrimination index, d', where d' = 1 was taken as threshold. In many cases, the threshold for discrimination of place of cochlear stimulation was < 0.75 mm, and the threshold for discrimination of current levels was < 1 dB. Cochlear electrode configurations varied in the accuracy with which they signaled to the auditory cortex the place of cochlear stimulation. The BP + N and TP configurations provided considerably greater sensitivity to place of stimulation than did the MP configuration. The TP configuration maintained accurate signaling of place of stimulation up to the highest current levels, whereas sensitivity was degraded at high current levels in BP + N configurations. Electrode configurations also varied in the dynamic range over which they signaled stimulus current level. Dynamic ranges were widest for the BP + 0 configuration and narrowest for the TP configuration. That is, the configuration that showed the most accurate signaling of cochlear place of stimulation (TP) showed the most restricted dynamic range for signaling of current level. These results suggest that the choice of the optimal electrode configuration for use by human cochlear-prosthesis users would depend on the particular demands of the speech-processing strategy that is to be employed.

Action Potentials↗

Auditory cortical images of cochlear-implant stimuli: dependence on electrode configuration.

This study examines patterns of auditory cortical activity elicited by single-pulse cochlear implant stimuli that vary in electrode configuration, cochlear place of stimulation, and stimulus level. Recordings were made from the primary auditory cortex (area A1) of ketamine-anesthetized guinea pigs. The spatiotemporal pattern of neural spike activity was measured simultaneously across 16 cortical locations spanning approximately 2-3 octaves of the tonotopic axis. Such a pattern, averaged over 40 presentations of any particular stimulus, was defined as the "cortical image" of that stimulus. Acutely deafened guinea pigs were implanted with a 6-electrode animal version of the 22-electrode Nucleus banded electrode array (Cochlear). Cochlear electrode configurations consisted of monopolar (MP), bipolar (BP + N) with N inactive electrodes between the active and return electrodes (0 < or = N < or = 4), tripolar (TP) with one active electrode and two flanking return electrodes, and common ground (CG) with one active electrode and as many as five return electrodes. Cortical images typically showed a focus of maximum spike probability and minimum latency. Spike probabilities tended to decrease, and latencies tended to increase, with increasing cortical distance from that focus. Cortical images of TP stimuli were the most spatially compact, followed by BP + N images, and then MP images, which were the broadest. Images of CG stimuli were rather variable across animals and stimulus channels. The locations of cortical images shifted systematically from caudal to rostral as the cochlear place of stimulation changed from basal to apical. At the most sensitive cortical site for each condition, the dynamic ranges over which spike rates increased with increased current level were restricted to about 1-2 dB, regardless of configuration. Dynamic ranges tended to increase with increasing cortical distance from the most sensitive site. Electrode configurations that produced compact cortical images (e.g., TP and BP + 0) showed the greatest range of thresholds within each cortical image and the largest dynamic range at cortical sites removed from the most sensitive site.

Action Potentials↗

Cortical representation of auditory space: information-bearing features of spike patterns.

Previous studies have demonstrated that the spike patterns of cortical neurons vary systematically as a function of sound-source location such that the response of a single neuron can signal the location of a sound source throughout 360 degrees of azimuth. The present study examined specific features of spike patterns that might transmit information related to sound-source location. Analysis was based on responses of well-isolated single units recorded from cortical area A2 in alpha-chloralose-anesthetized cats. Stimuli were 80-ms noise bursts presented from loudspeakers in the horizontal plane; source azimuths ranged through 360 degrees in 20 degrees steps. Spike patterns were averaged across samples of eight trials. A competitive artificial neural network (ANN) identified sound-source locations by recognizing spike patterns; the ANN was trained using the learning vector quantization learning rule. The information about stimulus location that was transmitted by spike patterns was computed from joint stimulus-response probability matrices. Spike patterns were manipulated in various ways to isolate particular features. Full-spike patterns, which contained all spike-count information and spike timing with 100-micros precision, transmitted the most stimulus-related information. Transmitted information was sensitive to disruption of spike timing on a scale of more than approximately 4 ms and was reduced by an average of approximately 35% when spike-timing information was obliterated entirely. In a condition in which all but the first spike in each pattern were eliminated, transmitted information decreased by an average of only approximately 11%. In many cases, that condition showed essentially no loss of transmitted information. Three unidimensional features were extracted from spike patterns. Of those features, spike latency transmitted approximately 60% more information than that transmitted either by spike count or by a measure of latency dispersion. Information transmission by spike patterns recorded on single trials was substantially reduced compared with the information transmitted by averages of eight trials. In a comparison of averaged and nonaveraged responses, however, the information transmitted by latencies was reduced by only approximately 29%, whereas information transmitted by spike counts was reduced by 79%. Spike counts clearly are sensitive to sound-source location and could transmit information about sound-source locations. Nevertheless, the present results demonstrate that the timing of the first poststimulus spike carries a substantial amount, probably the majority, of the location-related information present in spike patterns. The results indicate that any complete model of the cortical representation of auditory space must incorporate the temporal characteristics of neuronal response patterns.

Action Potentials↗

Cortical neurons that localize sounds.

Efforts to locate a cortical map of auditory space generally have proven unsuccessful. At moderate sound levels, cortical neurons generally show large or unbounded spatial receptive fields. Within those large receptive fields, however, changes in sound location result in systematic changes in the temporal firing patterns such that single-neuron firing patterns can signal the locations of sound sources throughout as much as 360 degrees of auditory space. Neurons in the cat's auditory cortex show accurate localization of broad-band sounds, which human listeners localize accurately. Conversely, in response to filtered sounds that produce spatial illusions in human listeners, neurons signal systematically incorrect locations that can be predicted by a model that also predicts the listeners' illusory reports. These results from the cat's auditory cortex, as well as more limited results from nonhuman primates, suggest a model in which the location of any particular sound source is represented in a distributed fashion within individual auditory cortical areas and among multiple cortical areas.

Animals↗