Shigeyuki Kuwada

Scalp potentials of normal and hearing-impaired subjects in response to sinusoidally amplitude-modulated tones.

None of the current electrical audiometric procedures, alone or in combination, has yet achieved the precision of conventional audiometric testing that is used to assess hearing in verbally capable children and adults. The reason for this, in part, lies in the use of stimuli which have a wide frequency content. We have measured scalp potentials which follow the envelopes of sinusoidally amplitude-modulated tones: a frequency-specific stimulus. In normal subjects such amplitude-modulation following responses (AMFRs) appear to be generated by two sources. One source has a latency of about 30 ms, generates large responses and is only observed at modulations below 55 Hz, while the other source has a latency of 7-9 ms, generates smaller responses, and is only observed at modulations from 100-350 Hz. The latencies of these two sources are consistent with origins in the cortex and midbrain, respectively. We examined AMFRs to low frequency (50 Hz) modulations as a possible audiometric tool. In normal subjects, the amplitude of the AMFR increased as a function of intensity, decreased as a function of carrier frequency, and could be evoked across the whole audiometric range (250-8000 Hz). In hearing-impaired subjects, the AMFR amplitudes as a function of carrier frequency accurately reflected the pattern of hearing loss on a frequency-by-frequency basis. In most subjects, the threshold for evoking the AMFR was within 0-25 dB of hearing threshold. It therefore appears that the AMFR may be a potentially useful tool to assess hearing in those unable to undergo conventional audiometric testing.

A neuronal population code for sound localization

The accuracy with which listeners can locate sounds is much greater than the spatial sensitivity of single neurons. The broad spatial tuning of auditory neurons indicates that a code based on the responses of ensembles of neurons, a population code, must be used to determine the position of a sound in space. Here we show that the tuning of neurons to the most potent localization cue, the interaural time difference in low-frequency signals (<∼2 kHz; refs 4, 5), becomes sharper as the information ascends through the auditory system. We also show that this sharper tuning increases the efficiency of the population code, in the sense that fewer neurons are required to achieve a given acuity.

Intracellular Recordings in Response to Monaural and Binaural Stimulation of Neurons in the Inferior Colliculus of the Cat

The inferior colliculus (IC) is a major auditory structure that integrates synaptic inputs from ascending, descending, and intrinsic sources. Intracellular recording in situ allows direct examination of synaptic inputs to the IC in response to acoustic stimulation. Using this technique and monaural or binaural stimulation, responses in the IC that reflect input from a lower center can be distinguished from responses that reflect synaptic integration within the IC. Our results indicate that many IC neurons receive synaptic inputs from multiple sources. Few, if any, IC neurons acted as simple relay cells. Responses often displayed complex interactions between excitatory and inhibitory sources, such that different synaptic mechanisms could underlie similar response patterns. Thus, it may be an oversimplification to classify the responses of IC neurons as simply excitatory or inhibitory, as is done in many studies. In addition, inhibition and intrinsic membrane properties appeared to play key roles in creating de novo temporal response patterns in the IC.

Eosinophilic oesophagitis in patients presenting with dysphagia – a prospective analysis

Background  Eosinophilic oesophagitis (EoO) may be a common finding in adults presenting with dysphagia.

SIMULTANEOUS ANTEROGRADE LABELING OF AXONAL LAYERS FROM LATERAL SUPERIOR OLIVE AND DORSAL COCHLEAR NUCLEUS IN THE INFERIOR COLLICULUS OF CAT

The laminar organization of the central nucleus of inferior colliculus includes layers of axons that may be important in shaping the responses of neurons. Depending on their source, some layered axons are afferents that are superimposed and terminate on the same postsynaptic neurons, while other layered afferents, such as those from the ipsilateral and contralateral lateral superior olive, terminate side‐by‐side. The specific pattern of convergence may dictate which populations of axons are presynaptic to layered disc‐shaped neurons in the central nucleus.

Interaural time sensitivity of high‐frequency neurons in the inferior colliculus

Recent psychoacoustic experiments have shown that interaural time differences provide adequate cues for lateralizing high-frequency sounds, provided the stimuli are complex and not pure tones. We present here physiological evidence in support of these findings. Neurons of high best frequency in the cat inferior colliculus respond to interaural phase differences of amplitude modulated waveforms, and this response depends upon preservation of phase information of the modulating signal. Interaural phase differences were introduced in two ways: by interaural delays of the entire waveform and by binaural beats in which there was an interaural frequency difference in the modulating waveform. Results obtained with these two methods are similar. Our results show that high-frequency cells can respond to interaural time differences of amplitude modulated signals and that they do so by a sensitivity to interaural phase differences of the modulating waveform.

The frequency-following response to continuous tones in humans

Previous studies of the frequency-following response (FFR) in man suggest that it has multiple sources. Identification of these sources has been complicated by the use of tone bursts to evoke FFRs and the lack of precise methods to calculate their amplitude and latency. Tone bursts produce transient responses which confound measurements of the FFR. The use of continuous tones avoids this problem and the Fast Fourier Transform can be used to assess accurately and efficiently the presence, amplitude and phase angle of the FFR. In this study we systematically examined the frequency and intensity range over which FFRs to continuous tones could be evoked using FFRs to tone bursts for comparison. We then analyzed FFRs to continuous tones to determine the sources of this potential. FFRs to both stimuli have similar thresholds (65-90 dB SPL) and can be evoked by the same range of frequencies. Neurogenic FFRs in man occur only below 1000 Hz. The source for this potential has a latency of 8.2 +/- 0.1 ms (mean +/- SD) and is consistent with a midbrain source. At higher frequencies FFRs have a latency of less than 1 ms and are most likely cochlear microphonic. The small variation in the latency of the neurogenic FFR suggests this as a possible tool for assessing neurological disorders.

GABAA Synapses Shape Neuronal Responses to Sound Intensity in the Inferior Colliculus

Neurons in the inferior colliculus (IC) change their firing rates with sound pressure level. Some neurons maintain monotonic increases in firing rate over a wide range of sound intensities, whereas other neurons are monotonic over limited intensity ranges. We examined the conditions necessary for monotonicity in this nucleus in vitro in rat brain slices and in vivo in the unanesthetized rabbit. Our in vitro recordings indicate that concurrent activation of GABAA synapses with excitatory inputs facilitates monotonic increases in firing rate with increases in stimulus strength. In the absence of synaptic inhibition, excitatory input to IC neurons causes large depolarizations that result in firing block and nonmonotonicity. In vivo, although GABAA synapses decrease the firing rate in all IC neurons, they can have opposing effects on rate-level functions. GABAergic inputs activated by all sound intensities maintain monotonicity by keeping the postsynaptic potential below the level at which depolarization block occurs. When these inputs are blocked, firing block can occur and rate-level functions become nonmonotonic. High-threshold GABAergic inputs, in contrast, cause nonmonotonic responses by decreasing the firing rate at high intensities. Our results suggest that a dynamic regulation of the postsynaptic membrane potential by synaptic inhibition is necessary to allow neurons to respond monotonically to a wide range of sound intensities.

Transformations in processing interaural time differences between the superior olivary complex and inferior colliculus: beyond the Jeffress model

Interaural time differences (ITDs) are used to localize sounds and improve signal detection in noise. Encoding ITDs in neurons depends on specialized mechanisms for comparing inputs from the two ears. Most studies have emphasized how the responses of ITD-sensitive neurons are consistent with the tenets of the Jeffress model. The Jeffress model uses neuronal coincidence detectors that compare inputs from both sides and delay lines so that different neurons achieve coincidence at different ITDs. Although Jeffress-type models are successful at predicting sensitivity to ITDs in humans, in many respects they are a limited representation of the responses seen in neurons. In the superior olivary complex (SOC), ITD-sensitive neurons are distributed across both the medial (MSO) and lateral (LSO) superior olives. Similar response types are found in neurons sensitive to ITDs in two signal types: low-frequency sounds and envelopes of high-frequency sounds. Excitatory-excitatory interactions in the MSO are associated with peak-type responses, and excitatory-inhibitory interactions in the LSO are associated with trough-type responses. There are also neurons with responses intermediate between peak- and trough-type. In the inferior colliculus (IC), the same basic types remain, presumably due to inputs arising from the MSO and LSO. Using recordings from the SOC and IC, we describe how the response types can be described within a continuum that extends to very large values of ITD, and compare the functional organization at the two levels.

Physiological Studies of Directional Hearing

This review concerns physiological studies of the neural mechanisms of sound localization. We will describe the responses of neurons in the central auditory system to sounds presented dichotically or in the free field. The free‐field studies have sought to define the spatial receptive fields of these neurons and their topographical organization in the brain. Although the functional aspects of these cells are most effectively addressed by free‐field methods, the mechanism by which these cells accomplish this task is best studied using dichotic stimulation. Influenced by the duplex theory and human psychoacoustics, the major focus of neurophysiological studies using dichotic stimulation has been the investigation of the effects of varying interaural phase and intensity. The neuronal responses to these stimuli and their relationship to the frequency domain will be discussed with a particular emphasis on the concept of characteristic delay. In an attempt to define the underlying circuitry, we will compare the...