Central denervation hypersensitivity in the auditory system of the cat.

Data are reported for seven cats with a total of 29 electrodes permanently placed in or near the cochlear nucleus, the superior olivary complex, the nucleus of the inferior colliculus, and the medial geniculate body. Detection thresholds for pulsate electrical stimuli were measured using an operant behavioral procedure. Electrical stimulation thresholds were measured prior to and following bilateral destruction of the cochleas in all animals. In addition, four of the animals were tested using a site-of-stimulation discrimination prior to and following the cochlear lesion. Finally, hearing loss was evaluated in all cats after the completion of the experiments. Electrical stimulation thresholds showed a mean reduction of 7.9 dB throughout the brain stem auditory system fater cochlear destruction. The ability of the animals to perform the site-of-stimulation discrimination was not permanently impaired by the cochlear lesion. The data indicated the presence of increased sensitivity to electrical stimulation in most regions of the subcortical auditory system, although a lesser effect was found at the thalamic level. It was concluded that stimulation threshold provides an index relevant to the state of auditory neurons proximal to the electrode tip.     

Neuromagnetic steady-state responses to auditory stimuli

Steady-state magnetic responses to clicks presented at rates between 10 and 70 Hz have been recorded in healthy humans. The responses were highest in amplitude around 40 Hz. This amplitude enhancement is satisfactorily explained by summation of responses evoked by single clicks. The field maps suggest activation of the auditory cortex at all stimulus frequencies. Similar responses were obtained with gated noise bursts and by pauses in a series of clicks. The mean "apparent latency," determined from the phase lag at rates 30-70 Hz, was 54 ms. The physiological relevance of this quantity is shown to be questionable.     

Psychophysical evidence against the autocorrelation theory of auditory temporal processing.

Nowadays, it is widely believed that the temporal structure of the auditory nerve fibers' response to sound stimuli plays an important role in auditory perception. An influential hypothesis is that information is extracted from this temporal structure by neural operations akin to an autocorrelation algorithm. The goal of the present work was to test this hypothesis. The stimuli consisted of sequences of unipolar clicks that were high-pass filtered and mixed with low-pass noise so as to exclude spectral cues. In experiment 1, "interfering" clicks were inserted in an otherwise periodic (isochronous) click sequence. Each click belonging to the periodic sequence was followed, after a random portion of the period, by one interfering click. This disrupted the detection of temporal regularity, even when the interfering clicks were 5 dB less intense than the periodic clicks. Experiments 2-4 used click sequences that showed a single peak in their autocorrelation functions. For some sequences, this peak originated from "first-order" temporal regularities, that is from the temporal relations between consecutive clicks. For other sequences, the peak originated instead from "second-order" regularities, relative to nonconsecutive clicks. The detection of second-order regularities appeared to be much more difficult than the detection of comparable first-order regularities. Overall, these results do not tally with the current autocorrelation models of temporal processing. They suggest that the extraction of temporal information from a group of closely spaced spectral components makes no use of time intervals between nonconsecutive peaks of the amplitude envelope.     

Cochlear nucleus, inferior colliculus, and medial geniculate responses during the behavioral detection of threshold-level auditory stimuli in the rabbit

Rabbits were conditioned to respond behaviorally to auditory stimuli by pairing a white-noise conditioned stimulus (CS) with a corneal airpuff unconditioned stimulus (US). The conditioned response (CR) was movement of the nictitating membrane (NM). After the subjects were responding at better than the 90% correct level, the intensity of the auditory stimulus was reduced to behavioral threshold using a staircase procedure. Simultaneous measurements of neural unit activity and behavioral NM responses were then made in rabbits performing at behavioral threshold. After the experiment was completed neural unit responses during behavioral detection trials were compared to neural responses made during nondetection trials. Neural unit responses to a constant intensity, white-noise stimulus at behavioral threshold were well defined and essentially identical on behavioral detection and nondetection trials in the ventral cochlear nucleus, the ventrolateral division of the central nucleus of the inferior colliculus, and the ventral division of the medial geniculate body. This suggests that an auditory stimulus can be neuronally "detected" without being behaviorally detected, and that the neural "decision" to respond behaviorally is not made in these nuclei. Responses recorded from the dorsomedial division of the central nucleus of the inferior colliculus, the pericentral nucleus of the inferior colliculus, and less commonly in the medial division of the medial geniculate body were also clearly present and nearly identical during the onset of the auditory stimulus, but were sometimes consistently different for detection and nondetection conditions during the latter part of the auditory stimulus. These brain regions appear to receive both auditory and nonauditory inputs, and show responses which are more highly correlated with detection behavior.     

Intratympanic manganese administration revealed sound intensity and frequency dependent functional activity in rat auditory pathway

The cochlear plays a vital role in the sense and sensitivity of hearing; however, there is currently a lack of knowledge regarding the relationships between mechanical transduction of sound at different intensities and frequencies in the cochlear and the neurochemical processes that lead to neuronal responses in the central auditory system. In the current study, we introduced manganese-enhanced MRI (MEMRI), a convenient in vivo imaging method, for investigation of how sound, at different intensities and frequencies, is propagated from the cochlear to the central auditory system. Using MEMRI with intratympanic administration, we demonstrated differential manganese signal enhancements according to sound intensity and frequencies in the ascending auditory pathway of the rat after administration ofintratympanicMnCl₂.Compared to signal enhancement without explicit sound stimuli, auditory structures in the ascending auditory pathway showed stronger signal enhancement in rats who received sound stimuli of 10 and 40 kHz. In addition, signal enhancement with a stimulation frequency of 40 kHz was stronger than that with 10 kHz. Therefore, the results of this study seem to suggest that, in order to achieve an effective response to high sound intensity or frequency, more firing of auditory neurons, or firing of many auditory neurons together for the pooled neural activity is needed.     

The representation of periodic sounds in simulated sustained chopper units of the ventral cochlear nucleus

The nature of the neural processing underlying the extraction of pitch information from harmonic complex sounds is still unclear. Electrophysiological studies in the auditory nerve and many psychophysical and modeling studies suggest that pitch might be extracted successfully by applying a mechanism like autocorrelation to the temporal discharge patterns of auditory-nerve fibers. The current modeling study investigates the possible role of populations of sustained chopper (Chop-S) units located in the mammalian ventral cochlear nucleus (VCN) in this process. First, it is shown that computer simulations can predict responses to periodic and quasiperiodic sounds of individual Chop-S units recorded in the guinea-pig VCN. Second, it is shown that the fundamental period of a periodic or quasiperiodic sound is represented in the first-order, interspike interval statistics of a population of simulated Chop-S units. This is true across a wide range of characteristic frequencies when the chopping rate is equal to the f0 of the sound. The model was able to simulate the results of psychophysical studies involving the pitch height and pitch strength of iterated ripple noise, the dominance region of pitch, the effect of phase on pitch height and pitch strength, pitch of inharmonic stimuli, and of sinusoidally amplitude modulated noise. Simulation results indicate that changes in the interspike interval statistics of populations of Chop-S units compare well with changes in the pitch perceived by humans. It is proposed that Chop-S units in the ventral cochlear nucleus may play an important role in pitch extraction: They can convert a purely temporal pitch code as observed in the auditory nerve into a temporal place code of pitch in populations of cochlear-nucleus, Chop-S with different characteristic frequencies, and chopping rates. Thus, populations of cochlear-nucleus Chop-S units, together with their target units presumably located in the inferior colliculus, may serve to establish a stable rate-place code of pitch at the level of the auditory cortex.     

Electrically evoked whole-nerve action potentials: parametric data from the cat

In a companion paper [Brown et al., J. Acoust. Soc. Am. 88, 1385-1391 (1990)], a method for recording the electrically evoked whole-nerve action potential in human cochlear implant users was reported. The procedure for recording the response requires that two biphasic current pulses, a "masker" and a "probe," be presented at a rate and level sufficient to drive the auditory nerve into a refractory state. The present study was designed to assess the sensitivity of that recording technique to variations in stimulation parameters. The experiments described in this paper demonstrate that: (1) the EAP as recorded in the cat is triphasic and is defined by two negative peaks occurring at latencies of approximately 0.26 and 0.82 ms; (2) EAP amplitude is independent of the level of the masker stimulus for current levels equal to or greater than the current level of the probe stimulus; and (3) the time course of recovery of the EAP from the refractory state is stable over a range of both probe and masker current levels.     

Neural time and movement time in choice of whistle or pulse burst responses to different auditory stimuli by dolphins

Echolocating dolphins emit trains of clicks and receive echoes from ocean targets. They often emit each successive ranging click about 20 ms after arrival of the target echo. In echolocation, decisions must be made about the target--fish or fowl, predator or food. In the first test of dolphin auditory decision speed, three bottlenose dolphins (Tursiops truncatus) chose whistle or pulse burst responses to different auditory stimuli randomly presented without warning in rapid succession under computer control. The animals were trained to hold pressure catheters in the nasal cavity so that pressure increases required for sound production could be used to split response time (RT) into neural time and movement time. Mean RT in the youngest and fastest dolphin ranged from 175 to 213 ms when responding to tones and from 213 to 275 ms responding to pulse trains. The fastest neural times and movement times were around 60 ms. The results suggest that echolocating dolphins tune to a rhythm so that succeeding pulses in a train are produced about 20 ms over target round-trip travel time. The dolphin nervous system has evolved for rapid processing of acoustic stimuli to accommodate for the more rapid sound speed in water compared to air.     

Modeling the cochlear nucleus: a site for monaural echo suppression?

Echo suppression plays an important role in identifying and localizing auditory objects. One can distinguish between binaural and monaural echo suppression, although the former is the one commonly referred to. Based on biological findings we introduce and analyze a mathematical model for a neural implementation of monaural echo suppression in the cochlear nucleus. The model's behavior has been verified by analytical calculations as well as by numerical simulations for several types of input signal. It shows that in the perception of a pair of clicks the leading click suppresses the lagging one and that suppression is maximal for an interclick interval of 2-3 ms. Similarly, ongoing stimuli will be affected by the suppression mechanism primarily a couple of milliseconds after onset, resulting in a reduced perception of a sound shortly after its start. Both effects match experimental data.     

Neural network models of sound localization based on directional filtering by the pinna.

Three-layer neural-network functions were developed to transform spectral representations of pinna-filtered stimuli at the input to a space-mapped representation of sound-source direction at the output. The inputs are modeled after transfer functions of the external ear of the cat; the output is modeled on the spatial sensitivity of superior colliculus neurons. Network solutions are obtained by backpropagation and by a method that enforces uniform task distribution in the hidden layer of the model. Solutions are characterized using bandlimited inputs to study the relative strength of potential sound localization cues in various frequency regions. This analysis suggests that the frequency region containing the first spectral notch (5-18 kHz) provides the best localization cues. Response properties of model neurons were studied using input patterns modeled after auditory nerve response profiles to pure tones at various frequencies and sound levels. The response properties of hidden layer model neurons resemble cochlear nucleus types III and IV and their composites. Neurons in both hidden and output layers show the properties of spectral notch detectors. Although neural networks have limitations as models of real neural systems, the results illustrate how they can provide insight into the computation of complex transformations in the nervous system.     

Postnatal functional development of the dorsal and posteroventral cochlear nuclei of the cat

The postnatal development of firing patterns and response areas was determined for single neurons in the dorsal (DCN) and posteroventral ( PVCN ) cochlear nuclei of the kitten. Extracellular, single-unit responses to pure-tone stimulation were recorded in ketamine and sodium pentobarbital anesthetized kittens between the ages of 5 and 52 days. Within the first two weeks of postnatal life threshold is high, first-spike latency is long, and maximal discharge rate is low as compared to older kittens and adult cats. Prior to the end of the second postnatal week the tone-evoked temporal discharge patterns that characterize neurons of the DCN and PVCN in the adult cat are routinely recorded. These patterns, which appear within the first 50 ms of tonal stimulation, include the so-called " primarylike ," "chopper," " pauser ," "buildup," and "onset" types and their variants. In animals younger than about 10-12 days of age, the driven activity that occurs later than about 50 ms after stimulus onset often is not sustained, but breaks up during the stimulus into bursts that are separated by intervals of about 100-150 ms. Also within the first two weeks of postnatal life, many of the response-area properties of DCN and PVCN neurons are similar to those recorded in adult cats. The excitation and inhibition found within the so-called type II/III, type IV, and type V response areas of the adult occur in this early postnatal period. We conclude that many of the cellular mechanisms that underlie the temporal firing patterns and the organization of the response areas of DCN and PVCN neurons are active in the growing, differentiating cochlear nuclei and that the emergence of these mechanisms does not depend on afferent activity generated in the cochlear and auditory nerve by the animal's acoustic environment. Furthermore, if temporal firing patterns and response-area profiles remain relatively constant over the life span of the animal, then so must the spatial and temporal relationships of the inputs that produce and maintain them as these neurons, and the circuits of which they are a part, grow in size and complexity.     

Comparison of electrophonic and auditory-nerve electroneural responses

Electrophonic and auditory-nerve electroneural responses were recorded from the inferior colliculus of the cat. The electrophonic response appeared at a latency 1.0-1.5 ms later than the electroneural response, due to the time requirements for cochlear transduction. The electrophonic response also demonstrated very slow growth of response amplitude with increasing stimulus current as compared to the electroneural response. Aminoglycoside perfusion of the cochlea eliminated the electrophonic component from the evoked response record and left the electroneural component relatively unchanged, indicating that the electrophonic is an acoustic stimulus that requires an intact auditory end organ for transduction.     

Human brain-stem auditory evoked responses obtained by cross correlation to trains of clicks, noise bursts, and tone bursts

Brain-stem auditory evoked responses (BAERs) were obtained in eight normal-hearing young adults. Stimuli included clicks, noise bursts, and tone bursts. Tone bursts included carrier frequencies of 1, 2, 4, and 8 kHz. All stimuli were presented at 60 dB nHL. BAERs were obtained by presenting stimuli in pseudorandom trains, called maximum length sequences (MLSs). BAERs were recovered by cross correlating the responses with a recovery sequence. MLS-BAERs were obtained with minimum pulse intervals (MPIs) of 6, 4, and 2 ms. Conventional BAERs were also obtained for stimuli presented at a rate of 30 Hz. BAERs were obtained for all stimuli, for both the conventional averaging technique and for the cross-correlation technique. BAERs were observed for MPIs as short as 2 ms for all stimuli. Wave V was the only peak consistently identifiable for these stimuli. For all stimuli, wave V latency increased and wave V amplitude decreased with decreasing MPI. This is the first demonstration of the use of maximum length sequences combined with cross correlation to obtain BAERs to noise burst and tone burst stimuli.     

Investigation of the relationship among three common measures of precedence: fusion, localization dominance, and discrimination suppression

Listeners have a remarkable ability to localize and identify sound sources in reverberant environments. The term "precedence effect" (PE; also known as the "Haas effect," "law of the first wavefront," and "echo suppression") refers to a group of auditory phenomena that is thought to be related to this ability. Traditionally, three measures have been used to quantify the PE: (1) Fusion: at short delays (1-5 ms for clicks) the lead and lag perceptually fuse into one auditory event; (2) Localization dominance: the perceived location of the leading source dominates that of the lagging source; and (3) Discrimination suppression: at short delays, changes in the location or interaural parameters of the lag are difficult to discriminate compared with changes in characteristics of the lead. Little is known about the relation among these aspects of the PE, since they are rarely studied in the same listeners. In the present study, extensive measurements of these phenomena were made for six normal-hearing listeners using 1-ms noise bursts. The results suggest that, for clicks, fusion lasts 1-5 ms; by 5 ms most listeners hear two sounds on a majority of trials. However, localization dominance and discrimination suppression remain potent for delays of 10 ms or longer. Results are consistent with a simple model in which information from the lead and lag interacts perceptually and in which the strength of this interaction decreases with spatiotemporal separation of the lead and lag. At short delays, lead and lag both contribute to spatial perception, but the lead dominates (to the extent that only one position is ever heard). At the longest delays tested, two distinct sounds are perceived (as measured in a fusion task), but they are not always heard at independent spatial locations (as measured in a localization dominance task). These results suggest that directional cues from the lag are not necessarily salient for all conditions in which the lag is subjectively heard as a separate event.     

Isolating mechanisms that influence measures of the precedence effect: theoretical predictions and behavioral tests

This study tests how peripheral auditory processing and spectral dominance impact lateralization of precedence effect (PE) stimuli consisting of a pair of leading and lagging clicks. Predictions from a model whose parameters were set from established physiological results were tested with specific behavioral experiments. To generate predictions, an auditory nerve model drove a binaural, cross correlation computation whose outputs were summed across frequency using weightings derived from past physiological studies. The model predicted that lateralization (1) depends on stimulus center frequency and the inter-stimulus delay (ISD) between leading and lagging clicks for narrowband clicks and (2) changes differently with lead click level for different ISDs. Behaviorally, subjects lateralized narrowband and wideband click pairs whose stimulus parameters were chosen based on modeling results to test how peripheral processing and frequency dominance contribute to lateralization of PE stimuli. Behavioral results (including unique measures with the lead attenuated relative to the lag) suggest that peripheral interactions between leading and lagging clicks on the basilar membrane and strong weighting of cues around 750 Hz influence lateralization of paired clicks with short ISDs. When combined with auditory nerve adaptation, which emphasizes onset information, lateralization of PE click pairs with a short ISD can be well predicted.     

The effect of an additional reflection in a precedence effect experiment

Studies on the precedence effect typically utilize a two-source paradigm, which is not realistic relative to real world situations where multiple reflections exist. A step closer to multiple-reflection situations was studied using a three-source paradigm. Discrimination of interaural time differences (ITDs) was measured for one-, two-, and three-source stimuli, using clicks presented over headphones. The ITD was varied in either the first, second, or the third source. The inter-source intervals ranged from 0-130 ms. A perceptual weighting model was extended to incorporate the three-source stimuli and used to interpret the data. The effect of adding a third source could mostly, but not entirely, be understood by the interaction of effects observed in the precedence effect with two sources. Specifically, for delays between 1 and 8 ms, the ITD information of prior sources was typically weighted more heavily than subsequent sources. For delays greater than 8 ms, subsequent sources were typically weighted slightly more heavily than prior sources. However, there were specific conditions that showed a more complex interaction between the sources. These findings suggest that the two-source paradigm provides a strong basis for understanding how the auditory system processes reflections in spatial hearing tasks.     

A neural circuit transforming temporal periodicity information into a rate-based representation in the mammalian auditory system

Periodic amplitude modulations (AMs) of an acoustic stimulus are presumed to be encoded in temporal activity patterns of neurons in the cochlear nucleus. Physiological recordings indicate that this temporal AM code is transformed into a rate-based periodicity code along the ascending auditory pathway. The present study suggests a neural circuit for the transformation from the temporal to the rate-based code. Due to the neural connectivity of the circuit, bandpass shaped rate modulation transfer functions are obtained that correspond to recorded functions of inferior colliculus (IC) neurons. In contrast to previous modeling studies, the present circuit does not employ a continuously changing temporal parameter to obtain different best modulation frequencies (BMFs) of the IC bandpass units. Instead, different BMFs are yielded from varying the number of input units projecting onto different bandpass units. In order to investigate the compatibility of the neural circuit with a linear modulation filterbank analysis as proposed in psychophysical studies, complex stimuli such as tones modulated by the sum of two sinusoids, narrowband noise, and iterated rippled noise were processed by the model. The model accounts for the encoding of AM depth over a large dynamic range and for modulation frequency selective processing of complex sounds.     

Some features of sensing by cochlear neurons of low-frequency signals

Some features of sensing by human cochlear neurons of the infrasound vibrations generated by a change in frequency and volume of the audible range are discussed. It has been shown that auditory neurons respond to a volume envelope that selects corresponding infrasonic frequencies for their further processing. This mechanism is possible if the ear operates nonlinearly performing detection with further time averaging of no less than over 40–50 ms. If the frequency of the sound code coincides with any rhythm of the brain, resonance may occur, increasing the amplitude of the corresponding rhythm.     

Breakdown of echo suppression in the precedence effect

A new phenomenon is reported in which a change in spatial location of the leading sound source disrupts the normal echo suppression of the precedence effect. Click trains were presented through two loudspeakers, one leading the other by a few milliseconds. When leading and lagging signals were switched, listeners heard clicks momentarily for as long as several seconds from both loudspeakers before echo suppression was reestablished.     

Electrically evoked whole-nerve action potentials: data from human cochlear implant users

This study describes a method for recording the electrically evoked, whole-nerve action potential (EAP) in users of the Ineraid cochlear implant. The method is an adaptation of one originally used by Charlet de Sauvage et al. [J. Acoust. Soc. Am. 73, 615-627 (1983)] in guinea pigs. The response, recorded from 11 subjects, consists of a single negative peak that occurs with a latency of approximately 0.4 ms. EAP input/output functions are steeply sloping and monotonic. Response amplitudes ranging up to 160 micro V have been recorded. Slope of the EAP input/output function correlates modestly (approximately 0.6-0.69) with results of tests measuring word recognition skills. The refractory properties of the auditory nerve were also assessed. Differences across subjects were found in the rate of recovery from the refractory state. These findings imply that there may be difference across subjects in the accuracy with which rapid temporal cues can be coded at the level of the auditory nerve. Reasonably strong correlations (approximately 0.74-0.85) have been found between the magnitude of the slope of these recovery curves and performance on tests of word recognition.