Noise masking of tone responses and critical ratios in single units of the mouse cochlear nerve and cochlear nucleus

Responses of single units in the cochlear nerve and cochlear nucleus to tone bursts in a background of continuous white broadband noise were recorded. Tone and noise intensities ranged from threshold to saturation levels. Masking of the tone response by the noise was demonstrated by comparing peristimulus-time histograms and spike rates recorded during the tone and between tone presentations. The response of a unit to masking was found to be predictable based upon the difference in its rate of response to the tone and to the noise when the tone was masked. Several nonlinearities of the masking process are described. The most prominent one is an increase in the difference between tone and noise levels at the threshold of masking with increasing tone levels, i.e. neural critical ratios increase with increasing tone level. On the average, the frequency dependence of single unit effective bandwidths and of critical ratio bandwidths is similar to that of mean behavioral critical ratio bands.     

Spatial organization of the reciprocal connections between the cat dorsal and anteroventral cochlear nuclei

We are studying the interconnections between the anteroventral cochlear nucleus (AVCN) and the dorsal cochlear nucleus (DCN). Biotinylated dextran was injected into the DCN, where the best frequency of responses was also recorded. Ventrotubercular neurons in AVCN were labeled, along with cochlear nerve fibers and the axons of cells in DCN. In AVCN, a central band of labeled cochlear nerve axons and large endbulbs was labeled. Bordering this band was a ‘fringe' of smaller tuberculoventral axonal endings forming pericellular nests. Most AVCN neurons projecting to DCN were stellate, elongate, or giant cells, located in the posterior division of AVCN, regardless of the DCN injection site. About 75% of the labeled AVCN cells lay within the bands of labeled cochlear nerve fibers. Another 15% were in the outer fringes on either side of these bands, while 10% were outside the bands and the fringes. These findings suggest that most AVCN neurons projecting to the DCN conform to the tonotopic map. A significant portion of the ventrotubercular neurons occupy side-bands in AVCN. Reciprocally, the tuberculoventral tract forms a robust fringe of axonal endings flanking the central bands. The neuronal and axonal bands and side-bands may underlie excitatory and inhibitory signal transformations.     

Type II cochlear ganglion cells in the chinchilla

In order to ascertain whether Type II cochlear ganglion cells project to the brain, we have studied the retrograde transport of horseradish peroxidase (HRP) from the cochlear nucleus to the spiral ganglion of the chinchilla. In this animal there exist two types of ganglion neurons, which closely correspond to those previously described in guinea pigs, cats and rats. As in the guinea pig, the majority population (Type I) consists of relatively large, myelinated neurons. The minority population (Type II, 10% of the total population) consists of small, mostly unmyelinated cells, with filamentous cytoplasm and finely grained nuclear chromatin. Type II neurons tend to be clustered toward the peripheral side of Rosenthal's canal, often in close proximity to the intraganglionic spiral bundle. By 24 h after injections of HRP into the cochlear nucleus, incubation of the cochlear ganglion in diaminobenzidine/H2O2 reveals abundant HRP label in both Type I and Type II neurons. Type II neurons, however, tend to be labelled less intensely than Type I neurons. Control experiments, consisting of spillage of HRP solution over the cochlear nucleus, were carried out to determine how much HRP might be picked up by neurons after HRP diffusion. Comparison of cochleae from injected animals and from the control animals suggests that most of the label that was found in ganglion neurons after cochlear nucleus injections represents axonally transported HRP. We conclude, at least tentatively, that Type II neurons project to the brain. The fact that less label is found in Type II neurons that in Type I neurons suggests that the former have thinner axons and/or finer terminals in the cochlear nucleus.     

Gamma-aminobutyric acidergic and glycinergic inputs shape coding of amplitude modulation in the chinchilla cochlear nucleus.

Amplitude modulation is a prominent acoustic feature of biologically relevant sounds, such as speech and animal vocalizations. Enhanced temporal coding of amplitude modulation signals is found in certain dorsal and posteroventral cochlear nucleus neurons when they are compared to auditory nerve. Although mechanisms underlying this improved temporal selectivity are not known, involvement of inhibition has been suggested. gamma-Aminobutyric acid- and glycine-mediated inhibition have been shown to shape the dorsal cochlear nucleus and posteroventral cochlear nucleus response properties to other acoustic stimuli. In the present study, responses to amplitude modulation tones were obtained from chinchilla dorsal cochlear nucleus and posteroventral cochlear nucleus neurons. The amplitude modulation carrier was set to the neuron's characteristic frequency and the modulating frequency varied from 10 Hz. Rate and temporal modulation transfer functions were compared across neurons. Bandpass temporal modulation transfer functions were observed in 74% of the neurons studied. Most cochlear nucleus neurons (90%) displayed flat or lowpass rate modulation transfer functions to amplitude modulation signals presented at 2540 dB (re: characteristic frequency threshold). The role of inhibition in shaping responses to amplitude modulation stimuli was examined using iontophoretic application of glycine or gamma-aminobutyric acidA receptor agonists and antagonists. Blockade of gamma-aminobutyric acidA or glycine receptors increased stimulus-evoked discharge rates for a majority of neurons tested. Synchronization to the envelope was reduced, particularly at low and middle modulating frequencies, with temporal modulation transfer functions becoming flattened and less bandpass in appearance. Application of glycine, gamma-aminobutyric acid or muscimol increased the modulation gain over the low- and mid-modulation frequencies and reduced the discharge rate across envelope frequencies for most neurons tested. These findings support the hypothesis that glycinergic and gamma-aminobutyric acidergic inputs onto certain dorsal cochlear nucleus and posteroventral cochlear nucleus neurons play a role in shaping responses to amplitude modulation stimuli and may be responsible for the reported preservation of amplitude modulation temporal coding in dorsal cochlear nucleus and posteroventral cochlear nucleus neurons at high stimulus intensities or in background noise.     

Rhythmic discharge properties of caudal cochlear nucleus neurons during postnatal development in cats

Action potentials recorded extracellularly from neurons within the caudal cochlear nuclei of developing cats exhibited distinctive temporal characteristics (i.e., rhythmic responses) in response to long-duration acoustic stimuli including both tone and noise bursts. Unlike the homogeneous response characteristics of auditory nerve fibers, cochlear nucleus neurons exhibited many variations in rhythmic discharge patterns. The majority of neurons within the caudal CN of kittens younger than 10 days of age responded rhythmically to long-duration acoustic stimuli, however, the percentage of neurons responding rhythmically steadily decreased thereafter, and by the end of the second postnatal week most tonically-responding neurons maintained sustained steady-state discharge rates throughout stimulation. Discharges of neurons recorded during the transitional ages (around 13 days) were rhythmic at low sensation levels and exhibited adultlike sustained patterns at higher levels. Using constant sensation level stimuli (re individual neuron thresholds), burst frequencies remained essentially constant during the period of development in which rhythmic responses were observed. Intervals separating discharge bursts decreased as stimulus intensities increased for all neurons studied during the relevant period, but were not related in an orderly way to stimulus frequency. The effects of intensity on response periodicity were not mimicked by altering the amount of neurotransmitter present at the postsynaptic cell through microiontophoresis of excitatory amino acids and their antagonists onto the surface of neurons within the caudal CN. In addition, some immature neurons which responded phasically to acoustic stimuli responded rhythmically during the simultaneous presentation of acoustic stimuli and neuroexcitatory agents (i.e., glutamate). These results suggest that the source of the rhythmicity is not intrinsic to neurons in the caudal CN. Based on these and other observations we conclude that the most probable source of response periodicity observed early in development is the domination of inner hair cell output by efferent projections of the olivocochlear bundle, the temporal discharge patterns of which are also periodic.     

The sciatic nerve of the toad Xenopus laevis as a physiological model of the human cochlear nerve

The response of single fibres of the human cochlear nerve to electrical stimulation by a cochlear implant has previously been inferred from the response of the cochlear nerve in other mammals. These experiments are hindered by stimulus artefact and the range of stimulus currents used is therefore much less than the perceptual dynamic range (from threshold to discomfort) of human subjects. We have investigated use of the sciatic nerve of the toad Xenopus laevis as a convenient physiological model of the human cochlear nerve. Use of this completely dissected nerve reduces the problems of stimulus artefact whilst maintaining the advantages of a physiological preparation. The validity of the model was assessed by measuring the refractory periods, excitation time-constant, and relative spread of single fibres using microelectrode recording. We have also investigated the response of nerve fibres to sinusoidal stimulation. Based on these measurements, we propose that the sciatic nerve may be a suitable model of the human cochlear nerve if the timescales of stimuli are decreased by a factor of about five to compensate for the slower dynamics of the sciatic nerve and if noise is added to the stimuli to compensate for the lower internal noise of sciatic nerve fibres.     

The Effect of Gaussian Noise on the Threshold,Dynamic Range,and Loudness of Analogue Cochlear Implant Stimuli

The deliberate addition of Gaussian noise to cochlear implant signals has previously been proposed to enhance the time coding of signals by the cochlear nerve. Potentially, the addition of an inaudible level of noise could also have secondary benefits: it could lower the threshold to the information-bearing signal, and by desynchronization of nerve discharges, it could increase the level at which the information-bearing signal becomes uncomfortable. Both these effects would lead to an increased dynamic range, which might be expected to enhance speech comprehension and make the choice of cochlear implant compression parameters less critical (as with a wider dynamic range, small changes in the parameters would have less effect on loudness). The hypothesized secondary effects were investigated with eight users of the Clarion cochlear implant; the stimulation was analogue and monopolar. For presentations in noise, noise at 95% of the threshold level was applied simultaneously and independently to all the electrodes. The noise was found in two-alternative forced-choice (2AFC) experiments to decrease the threshold to sinusoidal stimuli (100 Hz, 1 kHz, 5 kHz) by about 2.0 dB and increase the dynamic range by 0.7 dB. Furthermore, in 2AFC loudness balance experiments, noise was found to decrease the loudness of moderate to intense stimuli. This suggests that loudness is partially coded by the degree of phase-locking of cochlear nerve fibers. The overall gain in dynamic range was modest, and more complex noise strategies, for example, using inhibition between the noise sources, may be required to get a clinically useful benefit.     

Reverse correlation study of cochlear filtering in normal and pathological guinea pig ears

The filtering properties of single cochlear fibres have been determined in normal and kanamycin-treated guinea pigs using the reverse correlation technique. This method allows investigation of filtering over a wide dynamic range.For normal guinea pig fibres, the near threshold filter functions obtained with this, method correspond to the tone derived frequency threshold curves (FTCs). The 10 dB bandwidth of the filter functions increased monotonically with increasing noise levels above threshold. Thus with noise levels at approximately 50 dB above threshold, the 10 dB bandwidth had increased by a factor of 1.3–3. The changes in 3 dB bandwidth with increasing levels were, for some fibres, different from those of the 10 dB bandwidths.For the pathological fibres, the derived filter functions corresponded to their tone determined FTCs, and were therefore comparatively broadly tuned. Their tuning (O_10dB) approximated to those of normal fibres when the latter were measured 60 dB or more above threshold (i.e., at similar levels of stimulus), and did not increase further with increase in level.The findings in the normal guinea pig are consistent with those obtained by others in rodents, but are not consistent with those from the cat, where normal filtering is more robust to high levels of stimulus noise.     

Narrow-band analysis of compound action potentials for several stimulus conditions in the guinea pig

Compound action potentials (AP) were recorded under various stimulus conditions in 31 guinea pigs. Stimulus attenuation, decrease of inter-stimulus interval, increase of the level of a continuous wide-band noise masker, and lowering of the animal's temperature all resulted in a drop of the AP amplitude and an increase in latency. A narrow-band analysis of the compound APs makes it possible to describe these AP changes in terms of the response behaviour of small cochlear regions according to their central frequencies.The results show that intensity-dependent changes in the AP parameters can be explained on the basis of the tuning properties of the auditory nerve fibres when the effect of the rise time of the toneburst stimulus is taken into account.Shortening of the inter-stimulus interval produces a complex interaction in terms of tone-burst frequency and the region along the cochlear partition that contributes dominantly to the AP. It is concluded that response contributions from the narrow bands with a central frequency near the toneburst frequency show the most adaptation.The change in amplitude for narrow-band responses under increased masking is similar to that for stimulus attenuation. It seems, however, that the underlying masking mechanism is more comparable to the adaptation mechanism.Cooling of the animal did not affect the sharpness of tuning.In all four recording situations there seems to be a decrease in the amount of synchronization of single-fibre responses as reflected in the width of narrow-band action potentials.     

Transganglionic transport of D-aspartate from cochlear nucleus to cochlea--a quantitative autoradiographic study

This study concerns the connections of the inner and outer hair cells and the different types of ganglion cells of the mammalian cochlea with the central nervous system by making use of their putative neurotransmitters. D-[3H]Aspartate (D-ASP), a putative marker for glutamatergic neurons, was injected into the cochlear nucleus of cats and guinea pigs and the cochleas prepared for light microscopic autoradiography after varying survival times. A quantitative, statistical autoradiographic method is described. Grain counts per unit area were made for each of 14 tissue compartments in the cochlea and normalized to permit comparisons between cases. An operationally defined background labeling level was computed for each case and a statistical test for significance applied to the neuron-containing tissue compartments. With increasing survival times, significant labeling appeared successively in the cochlear nerve root, in each type of spiral ganglion cell, and in the neuron-containing tissue compartments of the organ of Corti. The findings are consistent with uptake of D-ASP and retrograde transport by cochlear nerve axons from the cochlear nucleus to the perikarya and peripheral processes of the spiral ganglion. We conclude that axons of all spiral ganglion cells project to the cochlear nucleus and that this nucleus is directly connected with both the inner and outer hair cells. Transganglionic transport of D-ASP from the cochlear nucleus is consistent with the hypothesis that the cochlear nerve axons use glutamate or aspartate as a neurotransmitter.     

Surface topography of the cochlear nuclei in humans: two- and three-dimensional analysis

We used three-dimensional reconstruction to study the cochlear nuclear complex (CN) in postmortem adult brains. Resulting data show that the largest part of the CN surface, particularly the dorsal cochlear nucleus (DCN), is fully within the lateral recess of the fourth ventricle. The surface of another subdivision, the ventral cochlear nucleus (VCN), is also almost entirely within the recess, except for a narrow zone adjacent to the caudoventral border of the nucleus. The caudal portion of the exposed zone of the VCN is in the vicinity of the rootlets of the glossopharyngeal (IX) nerve, and the ventral portion is close to the terminal part of the vestibulocochlear (VIII) nerve. The border between the intraventricular part of the CN and the extraventricular portion and also the terminal part of the VIII nerve approximately coincides with the line of attachment of the inferior medullary velum of the fourth ventricle (tenia of the choroid plexus). In the narrow strip of this ventral most part of the tenia we did not observe big blood vessels or neurons. Accordingly this could be a reasonably safe surgical route to the intraventricular surface of the CN.     

The Sharpening of cochlear frequency selectivity in the normal and abnormal cochlea

In the normal (anaesthetized) animal cochlea, the frequency threshold curves for single primary fibres are up to an order of magnitude sharper than the analogous functions derived from various reported measurements of the basilar membrane amplitude of vibration. This enhanced neural frequency selectivity is found in the same species and under conditions similar to those in which the mechanical measurements are taken. The sharpening process (at least near threshold) appears to be linear and is not dependent upon lateral inhibitory mechanisms. The variability of the neural frequency selectivity and its vulnerability to metabolic, chemical and pathological influences suggests the hypothesis that the sharpening is due to some form of ‘second filter’ subsequent to the relatively broadly tuned basilar membrane.

All fibres recorded from in the cochlear nerve in the normal cochlea show this enhanced frequency selectivity; in contrast, in pathological cochleas, all fibres, or a substantial proportion, have high-threshold, broadly tuned characteristics, approximating to those of the basilar membrane.

The frequency selectivity of normal cochlear fibres is adequate to account for the analogous psychophysical measures of hearing. It is proposed that loss of this normal frequency selectivity occurs in deafness of cochlear origin, accounting for widening of the critical band. A new hypothesis for recruitment is proposed on this basis.

Finally, invetigations of the cochlear nerve fibre frequency responses under conditions of hypoxia give grounds for the speculation that more than one mechanism is involved in the excitation of a single fibre, related to the separate functioning of the inner and outer hair cells.     

Place and Time Coding of Frequency in the Peripheral Auditory System: Some Physiological Pros and Cons

Evidence for and against classical theories of ‘place’ and ‘period’ mechanisms for the coding of frequency, and the modifications of the theories invoked to account for the pitch of ‘residue’ and other types of stimuli, are examined in the light of physiological data. These include new data on the temporal discharge patterns of cochlear nerve fibres under stimulation with two-tone complexes, harmonic and inharmonic three-tone complexes, and five-tone complexes of differing relative phase. They show, in particular, that certain arguments against ‘period’ coding of ‘residue’ pitch are invalid. The interspike intervals in the discharge patterns of cochlear fibres under these conditions are consistent with the pitches heard. On the other hand, the classical ‘period’ theory needs to be modified to take into account the normally relatively sharp frequency selectivity of cochlear fibres, and requires certain inefficiencies on the part of the central processor for pitch.

Comparison of measures of cochlear fibre frequency selectivity with analogous psychophysical data in man, including those on the ‘existence region’ of ‘residue’ pitch, suggests that ‘residue’-type stimuli judged to be tonal in quality could both: (a) be sufficiently resolved spectrally at the cochlear fibre level to serve as input to any of the current spectral ‘pattern recognition’ mechanisms proposed for the pitch extraction of complex signals, and also, (b) could generate patterns of temporal discharge reflecting enough waveform interaction between the harmonics to convey the pitch heard, because of the shape of the cochlear filters. (This conclusion might have to be qualified in the light of further physiological experiments on the ‘second effect’ of pitch shift.)

The present evidence, both psychophysical and physiological, suggests the following synthesis: musical interval recognition and relatively crude frequency discrimination can be accomplished by trained observers on signals where the frequency appears to be coded exclusively in terms of temporal information. However, the pitch quality of these signals is judged to be poor or absent. Likewise, signals, apparently coded exclusively by ‘place’ mechanisms, while having tonality, allow relatively crude frequency discrimination and judgment of musical intervals. With the possible exception of psychophysical data on the phenomenon of diplacusis, the present evidence cannot exclude the possibility that the central pitch extractor mechanism utilizes both the ‘place’ and ‘period’ cues produced by pure-tone signals (below 5 kHz) and ‘residue’-type signals, both signals evoking strong pitch and fine acuity of frequency discrimination. The degree of salience of a signal's pitch could well depend on the coherence of the two types of cue.

However, the greatest obstacle to the acceptance of ‘place’ coding mechanisms for frequency, particularly of the frequency components of a complex sound, is the restricted dynamic range of the peripheral elements of the auditory nervous system. Because of this, it is not clear how differences in the spectral energy distribution in signals at medium to high sound levels can be established in terms of patterns of mean discharge rate across the cochlear fibre array. At high sound levels, physiological evidence suggests that the discharge rates of the majority of stimulated fibres will be saturated, whereas psychophysical evidence suggests that the coding of the frequency and the relative level of even single-component signals can be carried out over a wide dynamic range in the absence of cues derived from spread of activity across the fibre array. Some new data, however, indicate that this problem may be circumvented at the cochlear nucleus level, but the coding mechanisms involved at the primary neurone level are obscure. One intriguing possibility exists that the auditory nervous system may utilize the fine temporal structure of cochlear fibre discharge patterns for the transmission of ‘place’ information.     

Degenerative alterations in the ventral cochlear nucleus of the guinea pig after impulse noise exposure. A preliminary light and electron microscopic study.

Guinea pigs were exposed to the noise of 40 shots of an alarm pistol held at a distance of about 60 cm. The ventral cochlear nuclei were studied in phase contrast and electron microscopy after both survival periods and longer periods of up to 55 days survival. Marked degeneration of primary cochlear nerve endings and of synapting secondary neurons of the posterior caudal part of the ventral cochlear nucleus (AVCN) and the octupus cell area (OCA) of the posterior ventral cochlear nucleus (PVCN) was found most distinctly after 5-55 days. As criteria of degeneration of the second neuron of the afferent auditory pathway we used: 1. The loss of the synapting nerve endings, mainly 'shrinking". 2. The formation of huge mitochondria in the second order neurons and their dendrites. 3. The phagocytosis by glial cells of nerve endings, of the second order neurons and of their dendrites. After 5 days survival time no distinct changes were found in the granular cell area of PVCN, where as all stages of degeneration could be found in OCA at this time. In the discussion of these findings it is concluded that additional studies of the morphology of the cochlear nuclei seem necessary, as these may lead to a better understanding of the pathology of hearing following heavy noise exposure.     

Electrical stimulation of the auditory nerve in deaf kittens: effects on cochlear nucleus morphology.

The present study examines the effects of long-term electrical stimulation of the auditory nerve on the morphology of neurons in the cochlear nucleus in young, sensorineural deaf animals. Kittens, systemically deafened using kanamycin and ethacrynic acid, received bilateral cochlear implants and were stimulated unilaterally for periods of up to four months. After sacrifice, cross-sectional areas of neuron somata were measured with an image-analysis system and compared using nonparametric statistics. The areas of cell somata within the anteroventral cochlear nucleus (AVCN) on the stimulated side were significantly larger than those of corresponding somata on the control, unstimulated side (P less than 0.001). However, there was no statistically significant difference among dorsal cochlear nucleus (DCN) neurons. These results indicate that long-term electrical stimulation of the auditory nerve can at least partially negate some effects of early postnatal auditory deprivation at the level of the cochlear nucleus.     

Interaction of cochlear nucleus explants with semiconductor materials

OBJECTIVE/HYPOTHESIS: Implantable hearing devices such as cochlear implants and auditory brainstem implants deliver auditory information through electrical stimulation of auditory neurons. The combination of microelectronic electrodes with auditory nerve cells may lead to further improvement of the hearing quality with these devices. Whereas several kinds of neurons are known to grow on semiconductor substrates, interactions of cochlear nucleus (CN) neurons with such materials have yet to be described. MATERIALS AND METHODS: To investigate survival and growth behavior of CN neurons on different semiconductor materials. CN explants from postnatal day 10 Sprague-Dawley rats were cultured for 96 hours in Neurobasal medium on polished and unpolished silicon wafers (p-type Si [100] and p-type Si3N4[100]) as well as plastic surface. These surfaces had been coated with poly-L-lysine and laminin. Neuronal outgrowth was examined using image analysis software after immunohistologic staining for neurofilament. Neurite length and directional changes were quantified. Additionally, neurite morphology and adhesion to the semiconductor material was evaluated by scanning electron microscopy. RESULTS: Although proper adhesion of CN explants was seen, no neurite growth could be detected on unpolished silicon wafers (Si and Si3N4). Compared with the other test conditions, polished, laminin-coated Si3N4 wafers showed best biocompatibility regarding neurite length and number per explant. CN explants developed a mean of eight neurons with an average length of 236 mum in 96 hours of culture on these wafers. CONCLUSION: The results of this study demonstrate the general possibility of CN neuron growth in culture on semiconductors in vitro. The differences in neuron length and number per explant indicate that the growth of CN neurons is influenced by the semiconductor substrate as well as extracellular matrix proteins, with laminin-coated p-type Si3N4[100] being a preferable material for future hybrid experiments on auditory-neuron semiconductor chips.     

Responses of chopper units in the ventral cochlear nucleus of the anaesthetised guinea pig to clicks-in-noise and click trains

Auditory brainstem responses (ABRs) have been measured with clicks, clicks masked by noise, click trains and pseudorandom maximum length sequences (MLS) of clicks. To investigate the neuronal populations contributing to the ABR under these stimulation conditions, we measured the extracellular responses of ventral cochlear nucleus (VCN) units in the urethane-anaesthetised guinea pig. We studied 23 chopper, 7 primary-like and 7 onset units. This report focuses on the responses from chopper units. The probability of discharge for chopper units increased with increasing click level reaching nearly 100% in many units, over a range of about 20–30 dB. Following each response to a click there was a 5–10 ms suppression of the spontaneous or noise evoked activity. As the level of the noise was increased over a range of 20–30 dB, the response to the clicks gradually decreased leading to a complete abolition of the click response at high noise levels. In a few units, low level noise produced a facilitation of the response to single clicks. In response to constant level equally spaced click trains, discharge probability increased with increasing minimum pulse interval (MPI), approaching 100% for MPIs of 4–8 ms in some units. The recovery afforded by the gaps in the MLS train often resulted in higher discharge probability for MLS than click trains with the same MPI, while response probabilities for MLS and click trains were similar when compared at equivalent average click rates. At short MPIs (0.5 and 1.0 ms), peri stimulus time histograms in response to click trains resembled those to best frequency (BF) tones and noisebursts, with chopping peaks unrelated to unit BF. VCN units show highly synchronised and reliable responses to click trains, MLS trains and clicks masked by noise. The decrease in discharge rate and increase in latency of chopper units with decreasing click level, increasing click rate and increasing masker level parallel the peak amplitude and latency changes observed in the auditory brainstem response.     

Postnatal development of auditory nerve and cochlear nucleus neuronal responses in kittens

Neurons located within the auditory periphery of kittens (i.e., primary auditory nerve fibers and neurons of the cochlear nucleus (CN) exhibit similar response properties throughout the early stages of postnatal development. Neural thresholds to acoustic stimuli are uniformly high, spontaneous and acoustically-evoked discharge rates are low, input/output slopes are shallow, and temporal discharge patterns are markedly immature. Phase-locking abilities are poor in developing mammals and all neurons exhibit broad bandpass tuning curves, with center frequencies clustering near 1.5 kHz. Throughout the first week, response thresholds, maximum discharge rates, rate-intensity slopes, dynamic ranges and other response indices remain essentially unchanged. Thereafter, between the 7th and 20th postnatal days, peripheral auditory development proceeds rapidly, such that thresholds, tuning properties, temporal discharge patterns, and input/output functions achieve maturity. The role of synaptogenesis in the development of adult response properties has been studied through microionophoresis of neuroactive molecules onto the surface of neurons in the caudal divisions of the cochlear nuclei of developing kittens. Results of preliminary experiments suggest that inhibitory postsynaptic receptor function precedes intrinsic excitatory neurotransmission. Furthermore, during the first two weeks of postnatal development in kittens, GABA microionophoresis onto immature caudal CN neurons, exhibiting sustained responses to acoustic stimuli, converts response patterns to the onset type in the majority of neurons encountered.     

Rate Representation of Tones in Noise in the Inferior Colliculus of Decerebrate Cats

Neurons in the central nucleus of the inferior colliculus (ICC) of decerebrate cats show three major response patterns when tones of different frequencies and sound-pressure levels (SPLs) are presented to the contralateral ear. The frequency response maps of type I units are uniquely defined by a narrow excitatory area at best frequency (BF: a unit's most sensitive frequency) and surrounding inhibition at higher and lower frequencies. As a result of this receptive field organization, type I units exhibit strong excitatory responses to BF tones but respond only weakly to broadband noise (BBN). These response characteristics predict that type I units are well suited to encode narrowband signals in the presence of background noise. To test this hypothesis, the dynamic range properties of ICC unit types were measured under quiet conditions and in multiple levels of continuous noise. As observed in previous studies of the auditory nerve and cochlear nucleus, type I units showed upward threshold shifts and discharge rate compression in background noise that partially degraded the dynamic range properties of neural representations at high noise levels. Although the other two unit types in the ICC showed similar trends in threshold shift and noise compression, their ability to encode auditory signals was compromised more severely in increasing noise levels. When binaural masking effects were simulated, only type I units showed an enhanced representation of spatially separated signals and maskers that was consistent with human perceptual performance in independent psychoacoustic observations. These results support the interpretation that type I units play an important role in the auditory processing of narrowband signals in background noise and suggest a physiological basis for spatial factors that govern signal detection under free-field listening conditions.     

Recovery from short-term adaptation in single neurons in the cochlear nucleus

Recovery from short-term adaptation was measured in single neurons in the cochlear nucleus using a forward masking stimulus paradigm. The response to a short-duration, low-level probe tone at a unit's characteristic frequency (CF) was measured before and after presentation of a masker tone at the unit's CF. The degree of adaptation was defined as the ratio of firing to the probe in the adapted and unadapted conditions. The level of the masker and time difference between the masker offset and probe onset ('DT') were varied. As DT increased, the response to the probe increased in most Primarylike, Primarylike-notch, and Chopper units. Recovery was approximately linear in log time for most of these units. However, approximately half the Pauser/Buildup and On units showed very different recovery patterns, ranging from no adaptation to very non-linear recovery patterns. The results suggest that little alteration in the recovery process occurs between the auditory nerve and Primarylike, Primarylike-notch, and Chopper units, but that significant changes in the recovery process occur in Pauser-Buildup and On units.