Herbert Voigt

IEEE Engineering in Medicine and Biology Society

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Stimulus dependencies of the gerbil brain‐stem auditory‐evoked response (BAER). I: Effects of click level, rate, and polarity

Three experiments evaluating the effects of various stimulus manipulations on the click-evoked gerbil brain-stem auditory-evoked response (BAER) are reported. In experiment 1, click polarity and level were covaried. With increasing click level, there is a parallel decrease in the latency of the first five BAER peaks (i-v) and an increase in BAER peak amplitudes. Mean wave i amplitude was greater for rarefaction than condensation clicks at high click levels; mean wave v amplitude was greater for condensation clicks at higher click levels. Experiment 2 covaried click rate and polarity. The latency of the BAER peaks increased with increasing click repetition rate. This rate-dependent latency increase was greater for the later BAER peaks, resulting in an increase in the i-v interval with increasing click rate. As rate increased, the amplitudes of waves i and v decreased monotonically, whereas the amplitudes of waves ii-iv were largely uninfluenced by click rate. As in experiment 1, mean wave i amplitude was greater for rarefaction clicks, whereas mean wave v amplitude was greater for condensation clicks. The magnitude of these polarity dependencies on waves i and v amplitude decreased with increasing click rate. Experiment 3 evaluated the effects of click polarity on BAERs to high-intensity (100 dB pSPL) clicks presented at a rate of 10 Hz. In eight of ten gerbils evaluated, wave i amplitude was greater to rarefaction clicks, and, in all ten animals, wave v amplitude was greater to condensation clicks. The effects of click level and rate on BAER peak amplitudes, latencies, and interwave intervals are reminiscent of stimulus dependencies reported for the human BAER. The effects of click polarity on the amplitudes of waves i and v of the gerbil BAER have also been reported for the human BAER.

Wideband inhibition of dorsal cochlear nucleus type IV units in cat : a computational model

AbstractA computational model of a portion of dorsal cochlear nucleus neural circuitry was used to investigate relationships between connectivity and response properties of type IV units. The model in this study consists of four neural populations. The pattern of convergence from one population to another and the strengths of those connections are the most important model parameters. Lumped parameter electrical circuit models represent individual cells. Interconnections are achieved by activating variable conductances in post-synaptic cells according to spike activity in pre-synaptic cells. Auditory nerve fibers are incorporated as a bank of logarithmically spaced gammatone filters that drive compartmental models of inner hair cell function. While it might be possible to configure the model without wideband inhibition to simulate type IV unit notch noise responses, the resulting parameters would likely be physiologically implausible. The model with wideband inhibition, however, shows the appropriate notch noise behavior. A wide variety of simulated rate versus cutoff-frequency plots are achieved varying three model parameters. The model was fit to physiological data by finding values of these three parameters that minimize the sum of squared errors. The results show that wideband inhibition can quantitatively account for the responses of type IV units to notch noise.

Brainstem Auditory-Evoked Response in the Rat Normative Studies, with Observations Concerning the Effects of Ossicular Disruption

Six young adult Sprague-Dawley rats were unilaterally cochleotomized, Brain-stem auditory-evoked responses (BAERs) to clicks and to 1-, 2-, 4-, 8- and 16-kHz tone bursts were obtained. In addition, response thresholds were estimated before and after ossicular disruption in the noncochleotomized ear of 4 animals. With increasing tone burst frequency, there was a decrease in BAER peak latencies as well as a decrease in threshold. With increasing click and tone burst intensity, there was a decrease in peak latencies and an increase in peak amplitudes. BAER peak latency/intensity functions to click stimuli ranged from -.013 to -.018 ms/dB. With increasing tone burst frequency there was a decrease in the slope of the latency/intensity function. Following ossicular disruption, BAER thresholds to clicks were elevated by an average of 49 dB. Threshold shifts to tone burst stimuli were smallest for 1- and 2-kHz tone bursts (35-36 dB) and increased with increasing frequency up to a maximum of 65 dB for 16-kHz tone bursts.

Modeling inhibition of type II units in the dorsal cochlear nucleus.

Abstract. Type II units in the dorsal cochlear nucleus (DCN) are characterized by vigorous but nonmonotonic responses to best frequency tones as a function of sound pressure level, and relatively weak responses to noise. A model of DCN neural circuitry was used to explore two hypothetical mechanisms by which neurons may be endowed with type II unit response properties. Both mechanisms assume that type II units receive excitatory input from auditory nerve (AN) fibers and inhibitory input from an unspecified class of cochlear nucleus interneurons that also receive excitatory AN input. The first mechanism, a lateral inhibition (LI) model, supposes that type II units receive inhibitory input from a number of narrowly tuned interneurons whose best frequencies (BFs) flank the BF of the type II unit. Tonal stimuli near BF result in only weak inhibitory input, but broadband stimuli recruit enough lateral inhibitors to greatly weaken the type II unit response. The second mechanism, a wideband inhibition (WBI) model, supposes that type II units receive inhibitory input from interneurons that are broadly tuned so that they respond more vigorously to broadband stimuli than to tones. Physiological and anatomical evidence points to the possible existence of such a class of neurons in the cochlear nucleus. The model extends an earlier computer model of an iso-frequency DCN patch to multiple frequency slices and adds a population of interneurons to provide the inhibition to model type II units (called I2-cells). The results show that both mechanisms accurately simulate responses of type II units to tones and noise. An experimental paradigm for distinguishing the two mechanisms is proposed.

Computational model of response maps in the dorsal cochlear nucleus

The neurons in the mammalian (gerbil, cat) dorsal cochlear nucleus (DCN) have responses to tones and noise that have been used to classify them into unit types. These types (I–V) are based on excitatory and inhibitory responses to tones organized into plots called response maps (RMs). Type I units show purely excitatory responses, while type V units are primarily inhibited. A computational model of the neural circuitry of the mammalian DCN, based on the MacGregor neuromime, was used to investigate RMs of the principal cells (P-cells) that represent the fusiform and giant cells. In gerbils, fusiform cells have been shown to have primarily type III unit response properties; however, fusiform cells in the cat DCN are thought to have type IV unit response properties. The DCN model is based on a previous computational model of the cat (Hancock and Voigt Ann Biomed Eng 27: 73–87, 1999) and gerbil (Zheng and Voigt Ann Biomed Eng 34: 697–708, 2006) DCN. The basic model for both species is architecturally the same, and to get either type III unit RMs or type IV unit RMs, connection parameters were adjusted. Interestingly, regardless of the RM type, these units in gerbils and cats show spectral notch sensitivity and are thought to play a role in sound localization in the median plane. In this study, further parameter adjustments were made to systematically explore their effect on P-cell RMs. Significantly, type I, type III, type III-i, type IV, type IV-T and type V unit RMs can be created for the modeled P-cells. Thus major RMs observed in the cat and gerbil DCN are recreated by the model. These results suggest that RMs of individual DCN projection neurons are the result of specific assortment of excitatory and inhibitory inputs to that neuron and that subtle differences in the complement of inputs can result in different RM types. Modulation of the efficacy of certain synapses suggests that RM type may change dynamically.

Response map properties of units in the dorsal cochlear nucleus of barbiturate-anesthetized gerbil (Meriones unguiculatus)

The response map scheme introduced by Evans and Nelson (1973) and modified by others, including Davis et al. (1996) for use with gerbils, has been used primarily for classifying units recorded in the cochlear nucleus of unanesthetized decerebrate preparations. Units lacking spontaneous activity (SpAc) have been classified as either type I/III or type II units based on the relative strength of their responses to broad-band noise compared to their responses to best-frequency (BF) tones. The relative noise index (rho), a ratio of these responses after SpAc is subtracted out, provides a convenient measure of this relative strength. In this paper, responses of 320 units recorded in the dorsal cochlear nucleus (DCN) of barbiturate-anesthetized gerbils to short-duration BF tones and broad-band noise were recorded. Since 87.5% of these units lacked SpAc, their response maps resembled those of type II and type I/III units. Units were characterized by rho and the normalized slope (m) of a best line fit to the BF rate versus level plot starting from the sound level corresponding to the first inflection point of the rate curve (typically its maximum value or the start of its sloping saturation). The distributions of rho and m values do not form distinct clusters as they do for units in the decerebrate preparation. Thus, the criteria developed for classifying DCN units in the decerebrate preparation do not appear appropriate for units in the barbiturate-anesthetized preparation. Deposits of horseradish peroxidase were used to locate 52 units. Most of the low SpAc units, 56% with poor noise responses (5/9) and nearly 70% with strong noise responses (25/36), and nearly all of the high SpAc units (6/7), were located either within or below the fusiform cell layer.

A statistically based method to generate response maps objectively

One scheme to classify the physiological response properties of single units in the cochlear nucleus is based on the average discharge rate of the unit and is reflected in the distribution of excitatory and inhibitory regions in a frequency-level map (response map) that spans the units receptive area (e.g., Evans and Nelson, 1973; Young and Brownell, 1976; Young and Voigt, 1982; Shofner and Young, 1985, Spirou and Young, 1991). Typically, discharge rate versus level curves are acquired at many frequencies and the investigator determines that a unit is excited or inhibited at a given level if the driven rate is above or below a spontaneous rate estimate by a specified criterion (for example, 20%). The investigator then encloses regions of excitation and inhibition where responses over adjacent frequencies and levels are consistent. In the present report, we describe an objective 3-step computer-based method to generate response maps: raw driven and spontaneous rate estimates are smoothed with a low-pass spatial filter; a unit is said to be excited or inhibited at a given level if the filtered driven rate is above or below the mean filtered spontaneous rate for that frequency by a specified criterion (percentage or statistical); and resultant response maps are median spatial filtered to eliminate spurious regions. The results shown here demonstrate that use of a statistical criterion provides a more reliable detection of excitation and inhibition than a 20% criterion, particularly when the variance of the rate estimates is high. Further, the statistically based method permits unit classification based on response map data that are more rapidly acquired with shorter duration stimuli (32 vs. 200 ms). Although this method is applied to units recorded in the dorsal cochlear nucleus, the technique may be applicable to studies of receptive fields and their plasticity in other systems.

Type III units in the gerbil dorsal cochlear nucleus may be spectral notch detectors.

AbstractBroadband sounds originating in the median plane are thought to be localized by neural processing of spectral notches introduced by the filtering action of the pinnae. Previous studies (Nelken, I., and E. D. Young. J. Neurophysiol. 71:2446–2462, 1994; Spirou, G. A., and E. D. Young. ibid. 66: 1750–1768, 1991) suggested that type IV units in decerebrate cat dorsal cochlear nucleus (DCN) are functional detectors of these spectral notches. Intracellular marking studies by Ding et al. (Ding, J., T. E. Benson, and H. F. Voigt. J. Neurophysiol. 82:3434–3457, 1999) have shown that type III units in gerbil arise from the DCNs principal output neurons, which are thought to have type IV unit properties in cat. A relative paucity of type IV units in the decerebrate gerbil (Davis, K. A., J. Ding, T. E. Benson, and H. F. Voigt. J. Neurophysiol. 75: 1411–1431, 1996) has motivated this study of spectral notch sensitivity in the gerbil DCN. Responses to notch noise stimuli were recorded from 15 gerbil type III units to investigate whether these units may function as spectral notch detectors. For narrow notch noise stimuli, all 15 units showed excitatory responses. For progressively wider notches, the discharge rate of 13/15 units became inhibited. As the maximum limits of notch width were approached, 11/15 units showed some degree of recovery from this inhibition. This response pattern in gerbil type III units possesses the salient features of notch noise responses in cat type IV units and implicates type III units in gerbilline spectral notch detection processes.

Comments on ‘‘Stimulus dependencies of the gerbil brain‐stem auditory‐evoked response (BAER). I: Effects of click level, rate and polarity’’ [J. Acoust. Soc. Am. 85, 2514–2525 (1989)]

The purpose of this comment is to propose a peak‐labeling nomenclature for the gerbil brain‐stem auditory‐evoked response (BAER) which is different than that used by Burkard and Voigt [J. Acoust. Soc. Am. 85, 2514–2525 (1989)]. This proposed nomenclature is a compromise between the use of five primary peaks (i–v) used by Burkard and Voigt (herein called the ‘‘Boston group’’) and the use of three primary peaks (I–III) proposed by Mills and colleagues (herein called the ‘‘Charleston group’’). This proposed standardization recognizes four peaks occurring within 4 to 5 ms of stimulus onset, labeled i through iv. This standardization will allow easier comparisons of gerbil BAER data obtained across laboratories.