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Projections from auditory cortex to the cochlear nucleus in rats: synapses on granule cell dendrites.

Previous work has demonstrated that layer V pyramidal cells of primary auditory cortex project directly to the cochlear nucleus. The postsynaptic targets of these centrifugal projections, however, are not known. For the present study, biotinylated dextran amine, an anterograde tracer, was injected into the auditory cortex of rats, and labeled terminals were examined with light and electron microscopy. Labeled corticobulbar axons and terminals in the cochlear nucleus are found almost exclusively in the granule cell domain, and the terminals appear as boutons (1-2 microns in diameter) or as small mossy fiber endings (2-5 microns in diameter). These cortical endings contain round synaptic vesicles and form asymmetric synapses on hairy dendritic profiles, from which thin (0.1 micron in diameter), nonsynaptic "hairs" protrude deep into the labeled endings. These postsynaptic dendrites, which are typical of granule cells, surround and receive synapses from large, unlabeled mossy fiber endings containing round synaptic vesicles and are also postsynaptic to unlabeled axon terminals containing pleomorphic synaptic vesicles. No labeled fibers were observed synapsing on profiles that did not fit the characteristics of granule cell dendrites. We describe a circuit in the auditory system by which ascending information in the cochlear nucleus can be modified directly by descending cortical influences.

Animals↗

Neurons in the cochlear nucleus investigated with tone and noise stimuli.

1. Responses of cochlear nucleus neurons to stationary and amplitude modulated noise stimulation are investigated and compared with responses to tonal stimuli. 2. Cross-correlation functions, computed from responses to stationary noise stimulation, showing a clear oscillation could be most easily obtained from low CF fibres presumed to be auditory nerve fibres and low CF cochlear nucleus neurons showing only activation response and a primarylike temporal pattern of response to tone bursts. This reflects good quality of phase locking in these neurons. 3. The CCF reflects strongly the frequency selectivity of the neuron as revealed in its response area but not the temporal pattern of response to tone bursts. 4. Responses to noise bursts are correlated with the responses to tone bursts of many different frequencies in both their sign (i.e. activation or suppression) and their temporal pattern. 5. The concept of two independently operating mechanisms, one depending on the fine time structure of the stimulus (the carrier) and the other on its amplitude, and determining respectively the fine time structure of the response pattern and its magnitude, is introduced. Experimental data are presented which lend support to the adequacy of the description in the majority of cases and reveal its shortcomings in others.

Animals↗

Stimulation-dependent development of neuronal cytoplasm in mouse cochlear nucleus.

The extent of neuronal development in the mouse ventral cochlear nucleus was examined in normal and developmentally auditory deprived mice. Mice were unilaterally deprived on postnatal day three by external auditory meatus removal and sacrificed with controls on day 45, which is after the developmental period. Light microscopic morphometry of neuronal nucleus and cytoplasm areas demonstrated that the normal spherical and globular cells were larger in their low frequency regions than in their high frequency regions. This size difference occurred mainly in the cytoplasm. Developmental deprivation reduced neuronal cytoplasm areas similarly in the high and low frequency regions of both neuronal types, but had no effect on the size of the neuronal nucleus. It was interpreted that cytoplasmic metabolic events are dependent on developmental levels of stimulation and high and low frequency regions normally are differentially stimulated. Furthermore, these stimulation-dependent cytoplasmic events are impaired by developmental hypostimulation, which retards neuronal metabolism and growth. No changes were observed in the cochlear nucleus contralateral to the deprived side, suggesting that compensatory changes, such as hypertrophy, did not occur.

Aging↗

Role of the dorsal cochlear nucleus in the sound localization behavior of cats.

The role of the dorsal cochlear nucleus (DCN) in directional hearing was evaluated by measuring sound localization behaviors before and after cats received lesions of the dorsal and intermediate acoustic striae (DAS/IAS). These lesions are presumed to disrupt spectral processing in the DCN without affecting binaural time and level difference cues that exit the cochlear nucleus via the ventral acoustic stria. Prior to DAS/IAS lesions, cats made accurate head orientation responses toward sound sources in the frontal sound field. After a unilateral DAS/IAS lesion, subjects showed increased errors in the azimuth and elevation of their responses; in addition, the final orientation of head movements tended to be more variable. Largest deficits in response elevation were observed in the hemifield that was ipsilateral to the lesion. When a second lesion was placed in the opposite DAS/IAS, increased orientation errors were observed throughout the frontal field. Nonetheless, bilaterally lesioned cats showed normal discrimination of changes in sound source location when tested with a spatial acuity task. These findings support previous interpretations that the DCN contributes to sound orientation behavior, and further suggest that the identification of absolute sound source locations and the discrimination between spatial locations involve independent auditory processing mechanisms.

Animals↗

Differentiation of the giant and fusiform cells in the dorsal cochlear nucleus of the hamster.

Two categories of large neurons--fusiform cells and giant cells--can be distinguished in the dorsal cochlear nucleus of the hamster. In the adult, these neurons are located in separate laminae in the nucleus and have distinct dendritic morphology. However, the two cell types are not distinguishable in the newborn hamster. At birth the large cells in the dorsal cochlear nucleus are clustered into one group and are alike morphologically. On postnatal day 5, laminae are still not apparent, but the neurons have begun to acquire their adult shapes. By day 15 laminae have formed, and the cells appear mature with the one exception that the apical dendrites of the fusiform cells have not acquired the spines which will cover their surface in the adult. The appearance of laminae coincides with the growth of axons and dendrites into a interstitial zone between the layers of cell bodies. Dendritic growth occurs during the time of axonal ingrowth and establishment of contacts between the axons and dendrites. The growth of the apical dendrites of fusiform cells, which are not contacted by these fibers, lags behind. These results demonstrate that afferent ingrowth and the differentiation of dendrites in the dorsal cochlear nucleus are temporally related. The synchronous development may serve to ensure a specific synaptic arrangement between the axons and their target dendrites.

Animals↗

The commissural pathway and cochlear nucleus bushy neurons: an in vivo intracellular investigation.

A direct commissural connection formed between cochlear nuclei allows information from the contralateral ear to rapidly influence the processing of the ascending auditory signal. Among the neuronal groups proposed to both receive, and contribute to, commissural input is the bushy cell population in the ventral cochlear nucleus (VCN). In this in vivo electrophysiological study we examine the intracellular recordings of bushy neurons during electrical stimulation of the contralateral cochlear nucleus (CN) for evidence of both their contribution to, and input from commissural projections. Activation of the commissural pathway revealed short-latency fast hyperpolarisation in 19.5% of the 41 bushy neurons examined. The hyperpolarising potentials were small in amplitude, displayed a highly variable time course between neurons, and in some cases were eliminated with injection of depolarising current. There was no indication of antidromic activity, or short-latency excitatory potentials. These results suggest that i) bushy neurons do not contribute projections to the commissural connection, and ii) a small portion of bushy neurons are hyperpolarised following commissural stimulation.

Animals↗

Glycine immunoreactive projections from the dorsal to the anteroventral cochlear nucleus.

The aim of the present study was to investigate whether projections from the dorsal cochlear nucleus (DCN) to the anteroventral cochlear nucleus (AVCN) use either of two inhibitory transmitters, glycine or GABA. Retrograde HRP labeling of DCN-to-AVCN projection neurons was combined with postembedding immunocytochemistry in the DCN of guinea pigs. Following injections of HRP in the anterior or posterior divisions of AVCN, large numbers of neurons were labeled in the DCN. All of these were located in the deep layer, except for a few granule cells. Nearly all (96%) of the projection neurons were immunoreactive for glycine and most had dendritic and somatic morphologies corresponding to those of elongate neurons (so-called 'corn' cells); only a few resembled small stellate neurons. Few (3%) retrogradely labeled neurons were immunoreactive for GABA. The results suggest that projections from the deep DCN to the AVCN are formed primarily by glycinergic elongate neurons. These projections could have a substantial inhibitory influence on the output of neurons in the AVCN.

Animals↗

Spectral integration by type II interneurons in dorsal cochlear nucleus.

The type II unit is a prominent inhibitory interneuron in the dorsal cochlear nucleus (DCN), most likely recorded from vertical cells. Type II units are characterized by low rates of spontaneous activity, weak responses to broadband noise, and vigorous, narrowly tuned responses to tones. The weak responses of type II units to broadband stimuli are unusual for neurons in the lower auditory system and suggest that these units receive strong inhibitory inputs, most likely from onset-C neurons of the ventral cochlear nucleus. The question of the definition of type II units is considered here; the characteristics listed in the preceding text define a homogeneous type II group, but the boundary between this group and other low spontaneous rate neurons in DCN (type I/III units) is not yet clear. Type II units in decerebrate cats were studied using a two-tone paradigm to map inhibitory responses to tones and using noisebands of varying width to study the inhibitory processes evoked by broadband stimuli. Iontophoresis of bicuculline and strychnine and comparisons of two-tone responses between type II units and auditory nerve fibers were used to differentiate inhibitory processes occurring near the cell from two-tone suppression in the cochlea. For type II units, a significant inhibitory region is always seen with two-tone stimuli; the bandwidth of this region corresponds roughly to the previously reported excitatory bandwidth of onset-C neurons. Bandwidth widening experiments with noisebands show a monotonic decline in response as the bandwidth increases; these data are interpreted as revealing strong inhibitory inputs with properties more like onset-C neurons than any other response type in the lower auditory system. Consistent with these properties, iontophoresis of inhibitory antagonists produces a large increase in discharge rate to broadband noise, making tone and noise responses nearly equal.

Acoustic Stimulation↗

Effects of intensity of repetitive acoustic stimuli on neural adaptation in the ventral cochlear nucleus of the rat.

To study neural adaptation as a function of stimulus intensity, auditory near-field evoked potentials were recorded from the ventral cochlear nucleus in awake Long Evans rats. Responses to 250-ms trains of repetitive clicks (pulse rates ranging from 100 to 1000 pulses per second) were collected at stimulus intensities of 5, 10, 30, 50 and 70 dB SPL. The amplitude of the first negative (N1) component of the average evoked potentials to individual pulses in the train was measured by using a subtraction method. The N1 responses were normalized with respect to the highest cochlear nucleus potential observed in the train, and then plotted as a function of click position in the train. As expected, the general trend of the curves was an exponential decay reaching a plateau more or less rapidly as a function of both intensity and rate of stimulation. Fitting these curves with exponential decay equations revealed that the rapid time constant decreased for increasing stimulus intensities whereas the short-term time constant is relatively independent of intensity. The amount of adaptation (expressed as the ratio of the plateau to the first peak amplitude) was substantially less prominent at low intensities (5-10 dB SPL) and low rates (100-200 pulses per second) than at higher intensities and high rates. These results indicate that adaptation patterns obtained in the ventral cochlear nucleus by using near-field evoked potentials exhibit properties comparable to those already present at the level of the auditory nerve.

Acoustic Stimulation↗

Frequency extent of two-tone facilitation in onset units in the ventral cochlear nucleus.

1. The frequency threshold curves (FTCs) of 91 single units in the cochlear nucleus of the anesthetized guinea pig were measured using a conventional single-tone paradigm and a two-tone paradigm designed to elucidate the frequency extent of two-tone facilitation in onset units (On). Units were classified according to existing classification schemes into primary-like (n = 3), chopper (n = 23), and three onset groups: OnI (n = 12), OnC (n = 29), and OnL (n = 24). Histological reconstructions show onset units to be widely distributed within the ventral cochlear nucleus in a manner generally consistent with its tonotopic organization. 2. The FTCs of onset units differed in their minimum thresholds, the steepness of their high- and low-frequency cutoffs, and their sharpness of tuning as quantified by the quality factor at 10 dB (Q10dB) above best frequency (BF) threshold values. There was considerable overlap in the sharpness of tuning between onset units and auditory nerve fibers, as indicated by the distribution of Q10dB values in the octave around 10 kHz: onset units had Q10dB values of 3.56 +/- 1.38 (SD), compared with 6.3 +/- 2.48 for auditory nerve fibers. The tuning of chopper units was similar to that of auditory nerve fibers (5.52 +/- 1.46). 3. Seventy-five percent of onset units showed some degree of facilitation (a threshold reduction) when their FTCs were measured in the presence of BF tones 4 dB below BF threshold. The frequency extent of such facilitation was variable, with a maximum of 6 octaves around the BF. In extreme cases facilitation could be measured when the BF tone was as low as 30 dB below BF threshold. 4. In 17% of onset units, suppressive effects were evident, as shown by noncontiguous frequency regions of facilitation. These suppressive effects might be a reflection either of suppression in the auditory nerve input or of a direct inhibitory input to the onset units. The strength of this effect suggests that inhibition is a likely explanation, consistent with the finding in previous morphological studies of profuse synapses with pleomorphic vesicles on multipolar cells. 5. FTCs of chopper and primary-like units measured in the presence of BF tones showed little facilitation. The facilitation that was observed in chopper units was confined to a narrow region around BF and disappeared when the facilitatory tone was lowered to 4 dB below BF threshold. 6. These data support the hypothesis that onset units, but not chopper or primary-like units, receive excitatory inputs from auditory nerve fibers with a wide range of BFs. However, the frequency range of facilitation and the magnitude of the threshold facilitation varied from unit to unit, suggesting that the off-BF inputs from auditory nerve fibers are not evenly distributed or equally effective in all units.

Acoustic Stimulation↗

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.

Action Potentials↗

Tonotopic projection from the dorsal to the anteroventral cochlear nucleus of mice.

To understand how auditory information is processed in the cochlear nuclei, it is crucial to know what circuitry exists and how it functions. In slice preparations, horseradish peroxidase (HRP) injections into the anteroventral cochlear nucleus (AVCN) reveal two circuits: a connection between the dorsal cochlear nucleus (DCN) and AVCN and a local circuit confined to the AVCN. Extracellular injection in the AVCN labels a band of cells in the DCN. The labeled cells in the DCN lie within a band of auditory nerve fiber terminals that are labeled by the same injection, showing that the connection from the DCN to the AVCN is frequency specific. The injections into the AVCN also labeled a cluster of neurons in the AVCN dorsal to the injection site. These cells may be interneurons that relay information from areas encoding higher frequencies to areas encoding lower frequencies within the AVCN. In the parasagittal plane, the AVCN is organized along two orthogonal axes that are indicated with HRP labeling of fibers and cell bodies. The tonotopic axis runs approximately dorsoventrally; the isofrequency axis runs approximately rostrocaudally. The axons of labeled DCN neurons and the cluster lie along the tonotopic axis, whereas the labeled auditory nerve fibers define the isofrequency axis. Where they cross is where HRP is taken up by the fibers. The area of uptake is small and lies in the middle of the darkly stained injection site.

Animals↗

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.

Animals↗

Dynamic properties of excitation and two-tone inhibition in the cochlear nucleus studied using amplitude-modulated tones.

The dynamic properties of excitation and two-tone inhibition in the cochlear nucleus were studied from extracellularly recorded unit responses to two simultaneously presented tones. One tone was presented at the unit's characteristic frequency, CF, the other at the unit's best inhibitory frequency, BIF. One or both of the tones were amplitude-modulated with pseudorandom noise. The system under study is in general nonlinear, but can be considered to function as a linear system for small changes in sound intensity around a certain operating point. The dynamic properties are likely to be different at different operating points. A suitable method for the study of dynamic properties of such a system employs tones that are amplitude-modulated with pseudorandom noise. In the present study, the dynamic properties were assessed by cross-correlating the unit discharge rate with the modulation. This was accomplished by computing the cross-covariance function between a period of noise and a period histogram of the discharges, the histogram being locked to the periodicity of the pseudorandom noise. In this way, it has been shown in previous works (Moller, 1973, 1974b), that the cross-covariance function is a valid approximation of the system's impulse response function at a certain sound intensity, provided the modulation is kept at a low value. In the present study the computed cross-covariance function is thus an approximation of the change in discharge rate of the cochlear nucleus units in response to a brief increase in stimulus intensity. As the response of the system under the given circumstances is approximately that of a linear system, the integrated cross-covariance is an approximation of the system's step response function, i.e the change in discharge rate that resulte from a hypothetical step increase in stimulus intensity. The results of the present study can be summarized as follows: 1. The impulse and step response functions computed from the responses to the modulated inhibitory tone of the great majority of units from which recording was made were found to be virtual mirror images of those obtained when the excitatory tone was modulated, the inhibitory response being somewhat smaller in amplitude than the excitatory. 2. When both tones were modulated simultaneously, the step response function was approximately the algebraic sum of the two responses obtained when the tones were modulated singly, further indicating that the system functions as a linear system when the stimulus amplitude is varied slightly around a certain operating point. 3. The shape of the cross-covariance functions is similar for all three stimulus situations, but varies with stimulus intensity and is different in different units. 4. The implication of the results is that the inhibition studied may either originate from the inhibition (suppression) seen in primary fibers or it may be the result of a true neural inhibition in the cochlear nucleus that occurs without any interneurons.

Animals↗

Axonal and non-axonal immunolocalization of cytosolic aspartate aminotransferase (cAATase), GABA and glutamic acid decarboxylase (GAD) in the rat cochlear nucleus.

The localization of both cAATase activity, by histoenzymological method, and the immunoreactivity against cAATase, were investigated in the cochlear nucleus of rats. The immunohistochemical determination of cAATase was carried out using the PAP method, with an antiserum obtained from rabbits immunized with porcine cAATase. It was also studied both the immunolocalization of GABA-like and GAD-like substances. Our observations, with light microscope, revealed weak cAATase activity in the small neurons, and a more intense one in the fibers surrounding neuronal bodies. The large neurons presented a very weakly activity within their neuronal bodies and dendrites, but it was strongly found in granulations that surround the perikaryon and dendrites. cAATase immunoreactivity presents the same distribution as the enzymological activity. In the same way, we have investigated the pattern of distribution of both GABA- and GAD-like substances. Immunolocalization of these substances was similar to that found for cAATase. In the control sections incubated with Gostatin (0.05 mM), cAATase activity was absent. The immunoreactivity was also negative in every immunohistochemical control sections. These facts suggest that aspartate could intervene as a co-neurotransmitter or neuromodulator in the rat cochlear nucleus, and that axonic endings could contain cAATase, GABA and GAD. It was also found immunoreactivity against cAATase, GABA and GAD, in neuronal bodies, dendrites and glial processes, in close association with capillary wall. These observations have led us to suggest the possible co-localization and co-release of both GABA and aspartate from synaptic and non-synaptic sites in the cochlear nucleus.

Animals↗

Aspartate aminotransferase and glutaminase activities in rat olfactory bulb and cochlear nucleus; comparisons with retina and with concentrations of substrate and product amino acids.

The quantitative distributions of aspartate aminotransferase and glutaminase were mapped in subregions of olfactory bulb and cochlear nucleus of rat, and were compared with similar data for retina and with the distributions of their substrate and product amino acids aspartate, glutamate, and glutamine. The distributions of both enzymes paralleled that of aspartate in the olfactory bulb and that of glutamate in the cochlear nucleus. In retina (excluding inner segments), there were similarities between aspartate aminotransferase and both glutamate and aspartate distributions. The distribution of gamma-aminobutyrate (GABA) was similar to those of both enzymes in olfactory bulb, to aspartate aminotransferase in cochlear nucleus, and to glutaminase in retina (excluding inner segments). The results are consistent with significant involvement of aspartate aminotransferase, especially the cytosolic isoenzyme, and glutaminase in accumulation of the neurotransmitter amino acids glutamate, aspartate, and GABA, although with preferential accumulation of different amino acids in different brain regions.

Amino Acids↗

Quantitative analysis of spiral ganglion projections to the cat cochlear nucleus.

A quantitative examination of the tonotopic organization of primary afferent projections to the cochlear nucleus (CN) in adult cats was conducted by using focal extracellular injections of Neurobiotin (NB) into the spiral ganglion of the basal cochlea. One to three injections separated by intervals of at least 2 mm were positioned along the basal one-third of the cochlea. Each injection produced discrete projection laminae that appeared as parallel horizontal sheets of labeled axons terminals distributed sequentially dorsally to ventrally across each major CN subdivision: the anteroventral, posteroventral, and dorsal cochlear nucleus, (AVCN, PVCN, and DCN, respectively). The length (rostrocaudal dimension), width (mediolateral dimension), thickness (dorsoventral dimension), and relative placement of 18 "frequency-band" laminae were measured in 10 adult cochlear nuclei. The average AVCN projection thickness was approximately twice that of the PVCN and DCN projections. In double injection cases, the center-to-center separation between AVCN laminae was also approximately twice that in the PVCN and equal to that in the DCN. Lamina thickness did not differ significantly as a function of frequency representation. However, in both width and length, mid-frequency laminae were up to two times larger than high-frequency laminae. Thus, the results indicate that DCN projections are the most discrete (i.e., are the thinnest and have the least overlap between adjacent frequency projections), whereas the AVCN projections are the largest but are as discrete as PVCN projections. In addition, high-frequency projections are smaller and more discrete than mild-frequency projections, which are larger and have greater overlap with adjacent frequency projections.

Animals↗

Topography of spiral ganglion projections to cochlear nucleus during postnatal development in cats.

A fundamenntal organizational principle of the central auditory system is that virtually all areas are tonotopically organized. However, we know very little about the timing or mechanisms that are responsible for the development of this organization. When cats are born, their auditory nervous systems are extremely immature, and their hearing thresholds are very high. Until postnatal days 7-10 (P7-10), cats have behavioral and physiological thresholds which are near or above the pain threshold for adults and also have poor frequency selectivity. Physiological thresholds for auditory nerve fibers and cochlear nucleus neurons are typically above 100-120 dB SPL (sound pressure level re 20 microPa). Three weeks later (at approximately P31), the sensitivity and frequency discrimination (tuning) of these neurons approximate adult values. This study examines the development of the tonotopic projections from the spiral ganglion to the cochlear nucleus during the period in cat development in which the auditory system undergoes the transition from being essentially nonfunctional to having adult-like function. With the animals heavily anesthetized, the cochleas were surgically exposed in kittens ranging in age from P6 to P45. Focal injections of Neurobiotin (NB) were made into Rosenthal's canal, labeling a small cluster of cells in the spiral ganglion of each cochlea. The projections of these labeled cells were visualized as frequency-specific bands of labeled axons and terminals in all major subdivisions of the cochlear nucleus. The thickness of these bands (i.e., the dimension of the bands orthogonal to the isofrequency representation and across the frequency gradient) were measured and compared to similar projections in adults. As in adult cats, the thickness of the bands varied only slightly with the location of the injection site (frequency representation) over a range of 1-7 mm from the cochlear base (45-13 kHz). Moreover, band thickness did not vary significantly with age. These data indicate that the tonotopic organization of spiral ganglion projections to the cochlear nucleus is as precise in kittens as young as P6 as it is in adults.

Afferent Pathways↗