Units In Dorsal Cochlear Nucleus Research Articles

Decreasing dorsal cochlear nucleus activity ameliorates noise-induced tinnitus perception in mice

BackgroundThe dorsal cochlear nucleus (DCN) is a region known to integrate somatosensory and auditory inputs and is identified as a potential key structure in the generation of phantom sound perception, especially noise-induced tinnitus. Yet, how altered homeostatic plasticity of the DCN induces and maintains the sensation of tinnitus is not clear. Here, we chemogenetically decrease activity of a subgroup of DCN neurons, Ca2+/Calmodulin kinase 2 α (CaMKII α)-positive DCN neurons, using Gi-coupled human M4 Designer Receptors Exclusively Activated by Designer Drugs (hM4Di DREADDs), to investigate their role in noise-induced tinnitus.ResultsMice were exposed to loud noise (9–11kHz, 90dBSPL, 1h, followed by 2h of silence), and auditory brainstem responses (ABRs) and gap prepulse inhibition of acoustic startle (GPIAS) were recorded 2 days before and 2 weeks after noise exposure to identify animals with a significantly decreased inhibition of startle, indicating tinnitus but without permanent hearing loss. Neuronal activity of CaMKII α+ neurons expressing hM4Di in the DCN was lowered by administration of clozapine-N-oxide (CNO). We found that acutely decreasing firing rate of CaMKII α+ DCN units decrease tinnitus-like responses (p = 3e −3, n = 11 mice), compared to the control group that showed no improvement in GPIAS (control virus; CaMKII α-YFP + CNO, p = 0.696, n = 7 mice). Extracellular recordings confirmed CNO to decrease unit firing frequency of CaMKII α-hM4Di+ mice and alter best frequency and tuning width of response to sound. However, these effects were not seen if CNO had been previously administered during the noise exposure (n = 6 experimental and 6 control mice).ConclusionWe found that lowering DCN activity in mice displaying tinnitus-related behavior reduces tinnitus, but lowering DCN activity during noise exposure does not prevent noise-induced tinnitus. Our results suggest that CaMKII α-positive cells in the DCN are not crucial for tinnitus induction but play a significant role in maintaining tinnitus perception in mice.

Single neuron recordings in dorsal cochlear nucleus (DCN) of awake gerbil

The auditory responses of neurons in the dorsal cochlear nucleus (DCN) are known to be sensitive to anesthesia, and consequently many studies have used an unanesthetized, decerebrate preparation. Decerebration, however, severs multiple descending pathways to the DCN and is traumatic to the brain, and so sensory responses may be influenced. It was possible, by combining sterile surgery, animal conditioning, and mild restraint to allow recordings in awake gerbils ( Merionus unguiculatus) . Response maps (RMs), post-stimulus time histograms (PSTHs), and responses to notch noise stimuli were recorded in awake gerbils. Some units’ responses were compared to those from previous experiments in anesthetized and decerebrate gerbil preparations. All RM types observed in decerebrate gerbils were also observed in the awake gerbil, with the notable exception of type IV units. Type III units were still the most commonly recorded units. No significant changes were observed in either RM shape or notch noise responses. Although most awake DCN units had spontaneous activity (typically <20 spikes/s), these rates were lower than those in decerebrate gerbils. Type III units’ spontaneous activity rates were significantly different from those in the decerebrate prep ( P < 0.05). DCN neurons may be more inhibited in awake gerbils. Although type III units were most sensitive to spectral notches, different unit types seem to respond slightly differently to notch noise. This study indicates that responses of DCN units in awake and decerebrate gerbils are different, though the shapes of RMs are the same.

Multisensory integration in the dorsal cochlear nucleus: unit responses to acoustic and trigeminal ganglion stimulation

A necessary requirement for multisensory integration is the convergence of pathways from different senses. The dorsal cochlear nucleus (DCN) receives auditory input directly via the VIIIth nerve and somatosensory input indirectly from the Vth nerve via granule cells. Multisensory integration may occur in DCN cells that receive both trigeminal and auditory nerve input, such as the fusiform cell. We investigated trigeminal system influences on guinea pig DCN cells by stimulating the trigeminal ganglion while recording spontaneous and sound-driven activity from DCN neurons. A bipolar stimulating electrode was placed into the trigeminal ganglion of anesthetized guinea pigs using stereotaxic co-ordinates. Electrical stimuli were applied as bipolar pulses (100 micros per phase) with amplitudes ranging from 10 to 100 microA. Responses from DCN units were obtained using a 16-channel, four-shank electrode. Current pulses were presented alone or preceding 100- or 200-ms broadband noise (BBN) bursts. Thirty percent of DCN units showed either excitatory, inhibitory or excitatory-inhibitory responses to trigeminal ganglion stimulation. When paired with BBN stimulation, trigeminal stimulation suppressed or facilitated the firing rate in response to BBN in 78% of units, reflecting multisensory integration. Pulses preceding the acoustic stimuli by as much as 95 ms were able to alter responses to BBN. Bimodal suppression may play a role in attenuating body-generated sounds, such as vocalization or respiration, whereas bimodal enhancement may serve to direct attention in low signal-to-noise environments.

Intrinsic oscillations in spike trains indicate non-renewal statistics due to convergence of inputs in dorsal cochlear nucleus neurons

The occurrence of intrinsic oscillations (IOs) in a unit’s discharge is reflected by a prominent peak in the power spectrum (i.e., Fourier transform of the autocorrelation function) of spike trains obtained from single-unit discharge, at a frequency independent of stimulus spectral characteristics. IOs have been reported by researchers in the dorsal cochlear nucleus (DCN) of both the cat and the Mongolian gerbil. It has been hypothesized that IOs are related to inter-spike interval (ISI) regularity (e.g., [Hear. Res. 58 (1992) 153]). This hypothesis is tested in this paper. Responses to multiple presentations of 50–300 ms duration tone bursts, at and near the unit’s best frequency (BF) at 20–60 dB re threshold were recorded from DCN units of barbiturate-anesthetized (30 units), as well as decerebrate (53 units) Mongolian gerbils. IOs in the recordings were then compared with the IOs in simulations of spiking-neuron models. The models were selected because: (1) their ISI regularity characteristics follow those of experimental data and (2) their IO properties are completely determined by their ISI regularity. Such comparison reveals that Ghoshal’s hypothesis fails for a fraction of the units. These results suggest a re-evaluation of the purported relationship between IOs, ISI regularity, and SAM response. Alternate hypotheses are proposed here using computational models that are based on convergence of multiple neural inputs onto the unit under study. These models produce non-renewal statistics that resemble those of the experimental data, as is evident from IO-based analysis.

Proprioceptive Information from the Pinna Provides Somatosensory Input to Cat Dorsal Cochlear Nucleus

The dorsal cochlear nucleus (DCN) is a second-order auditory structure that also receives nonauditory information, including somatosensory inputs from the dorsal column and spinal trigeminal nuclei. Here we investigate the peripheral sources of the somatosensory inputs to DCN. Electrical stimulation was applied to cervical nerves C1-C8, branches of C2, branches of the trigeminal nerve, and hindlimb nerves. The largest evoked potentials in the DCN were produced by C2 stimulation and by stimulation of its branches that innervate the pinna. Electrical stimulation of C2 produced a pattern of inhibition and excitation of DCN principal cells comparable with that seen in previous studies with stimulation of the primary somatosensory nuclei, suggesting that the same pathway was activated. Because C2 contains both proprioceptive and cutaneous fibers, we applied peripheral somatosensory stimulation to identify the effective somatosensory modalities. Only stimuli that activate pinna muscle receptors, such as stretch or vibration of the muscles connected to the pinna, were effective in driving DCN units, whereas cutaneous stimuli such as light touch, brushing of hairs, and stretching of skin were ineffective. These results suggest that the largest somatosensory inputs to the DCN originate from muscle receptors associated with the pinna. They support the hypothesis that a role of the DCN in hearing is to coordinate pinna orientation to sounds or to support correction for the effects of pinna orientation on sound-localization cues.

Temporal encoding and transmitting of amplitude and frequency modulations in dorsal cochlear nucleus

Amplitude- and frequency-modulated (AM and FM, respectively) tones have been considered as simplified models of natural sounds. The responses of auditory neurons can phase-lock to the modulation frequency (fm). The encoding and transmitting of such modulation phase-locking are interesting since there is no any fm physical peak in spectrum. In the present study, we approached these issues by recording the phase-locked responses of the dorsal cochlear nucleus (DCN) units in guinea pigs to different AM and FM tones. For AM noise tones without the spectral cues of fm, the unit's discharges still phase-locked to the envelope cycles, but it was generally weaker than to sinusoidal AM (SAM) tones. At 50% modulation depth (dm), the mean modulation gains of Pauser/Buildup (P/B) units (n=7) to AM noise tones was −0.61 dB whereas they had a 6.48 dB mean to SAM tones. Similar to the case of AM tones, phase-locking to sinusoidal FM (SFM) tones represented the time courses of frequency changes, and it could be separated and changeable corresponding to the frequency increasing and decreasing. There were differences between the phase-locking to SAM and SFM tones in an identical unit. Both ON and type I/III units tended to have stronger phase-locking to the SFM tones than to the SAM tones. The phase-locking to the possible demodulated fm components was further examined with different carrier frequencies (fc) and pure tones. The DCN units showed poor or no responses to modulation tones out of their response areas even in the low characteristic frequency (CF) units, but the low-CF units had clear phase-locking to pure tones at the similar fm ranges. The pure-tone phase-locking had a band-pass shape different from the low-pass shape of the auditory nerve fibers. These data suggest that the modulation phase-locking in the DCN units may be based on the temporal modulation cues and transmitted in the carrier place. The temporal integration of modulation information over the unit's response area as an across-frequency temporal processing model was discussed for modulation enhancement in the CN units. © Elsevier Science B.V. All rights reserved.

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

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

Why do cats need a dorsal cochlear nucleus?

The dorsal cochlear nucleus (DCN), one of the three major divisions of the cochlear nucleus (CN), has a complex internal structure, multiple inputs (some of them non-auditory), and multiple output pathways. Response properties of DCN units are accordingly complex. The principal cells of the DCN have type IV response characteristics, characterized by relatively high levels of spontaneous activity and inhibition by high level best frequency (BF) tones. We showed previously that type IV units are inhibited by two separate inhibitory mechanisms, one of them sensitive to narrow band stimuli and the other to wide band stimuli. One result of the wide band inhibition of type IV units is their sensitivity to spectral notches in the region of their BF - stimuli with such notches inhibit type IV units. The source of the narrow band inhibition is an interneuron in the DCN which has type II response characteristics - it does not have spontaneous activity and is strongly activated by BF tones. The neurons giving rise to type II responses are presumably vertical cells, which also project to other divisions of the CN. From anatomical studies, it is known that type IV units are also inhibited by a third system, which carries non-auditory information; movements of the pinna inhibit type IV units through this system. We hypothesize that type IV units signal important events to the auditory system by being inhibited. Such events are either auditory, e.g. spectral maxima and minima, or non-auditory, such as the somatosensory inputs from the pinnae. We hypothesize that the projection of type II units to the ventral cochlear nucleus (VCN) plays a role in reducing the effects of spectral notches introduced by the pinnae in the core auditory pathway. We conclude that although the DCN lies close to the auditory periphery, it already performs sophisticated tasks of auditory processing.

Processing of modulation frequency in the dorsal cochlear nucleus of the guinea pig: sinusoidal frequency-modulated tones

Frequency- and amplitude-modulated (FM and AM. respectively) tones are important information-bearing elements in voice sounds and can also be produced by the spatial movement of sound sources. Zhao and Liang (1995) recently reported the response features of dorsal cochlear nucleus (DCN) neurons to AM tones. In the present study, the responses of the guinea pig DCN neurons to sinusoidal FM (SFM) tones were examined. Discharges of the DCN units to the SFM tones phase-locked to the stimulus modulation frequencies ( f m). The phase-locked discharge patterns existed over broad ranges of modulation parameters and at stimulus levels as high as 95 dB SPL or modulation depths ( d m) as low as 2%. Robust phase-locking to the f m was observed in samples of all DCN unit types studied. The means of best f m ( Bf m and upper limit f m ( ULf m) of all recorded units were 510 Hz and 940 Hz, respectively. Pauser/Buildup (P/B) units had mean maximum synchronization index ( SI max) of 0.57. ON units had the highest Bf m with the mean of 646 Hz and subtype ON-S showed the highest mean of Sl max at 0.63. Phase-locking to the ƒ m was independent of discharge rates and existed even when th discharge rates were reduced to the background spontaneous rate (SR). A few units showed stronger synchronous responses to the square and triangular FM stimuli instead of the SFM tones. The relationship between the modulated responses and the unit's response area were further examined. The f, phase-locking occurred to modulation bands (or frequency ranges) within the response area, with the modulation bands as narrow as ± 160 Hz in the central inhibitory areas of the type IV units. As the width of the modulation band changed within a unit's response area, the phases of the f m, phase-locked responses of P/B units linearly changed while for Onset units, the change was lesser. The P/B and Onset units had a π phase shift and a π/2 phase change, respectively, as carrier frequencies ( f cs) passed through characteristic frequencies (CF) and the excitatory/inhibitory response boundaries. The phase-locked responses to the f ms were dependent on the SR but were independent of the CF. Low-SR ( > 2 spikes/s) units had higher synchronization of responses to the f m, than the high-SR ( > 2 spikes/s) units ( SI max = 0.64 and 0.42, respectively). These results suggest that the temporal characteristics of the fm is effectively represented in the responses of DCN units to the SFM tones. Such temporal encoding behavior can play an important role in the processing of the complex sounds in the auditory system. These results also have implications for a possible role for the DCN is in identifying the spatial movement of a sound source.

Processing of modulation frequency in the dorsal cochlear nucleus of the guinea pig: Amplitude modulated tones

The modulation frequency (Fm), particularly high Fm (> 200 Hz), in amplitude modulated (AM) tones can elicit the perception of the periodicity pitch (Langner, 1992). In this study, single unit responses to the Fms of the sinusoidal AM tones were investigated at 50 to 90 dB SPL. The recordings were made from the dorsal cochlear nucleus (DCN) of neuroleptic anesthetized guinea pigs with an intact cerebellum. The DCN units show a good capability of phase-locking to Fm at 400–1200 Hz. On-S-type II and Pauser/Buildup (P/B) units have a high modulation gain (7.2–8.3 dB). P/B units can preserve the high modulation gain (5–9 dB) up to 90 dB SPL. The modulation gain exponentially increases with decreasing modulation depth (Dm) and the phase-locking is detectable even at the Dm as low as 2–5%. The ‘central skipping’ of the phase-locking peak has been found at deep Dms in a few cases. The synchronization is independent of the discharge rate and can remain high even when the responses to AM tones are inhibited below the spontaneous activity. Such encoding behaviors over the unit's response area show that the Fm phase-locking is strong near or at its characteristic frequency (CF). The synchronization index (SI) versus carrier frequency (Fc) curve is similar to the inverse shape of tuning curve but more narrowly tuned than the iso-intensity function of pure tones at moderate to high intensity levels. The phase-locking is related to the unit's spontaneous rate (SR). The average modulation gain of the lower SR (≤ 2 spikes/s) units is 5 dB higher than that of the higher SR (> 2 spikes/s) units (8.16 and 2.92 dB, respectively) at 70 dB SPL. These results suggest that AM information is temporally encoded over broad ranges of modulation parameters in the DCN and is conveyed by the Fc channel. Such a timing mechanism can play an important role in processing of complex sounds under normal acoustic conditions.

Spontaneous and sound-evoked discharge characteristics of complex-spiking neurons in the dorsal cochlear nucleus of the unanesthetized decerebrate cat.

1. We examined the spontaneous and sound-evoked discharge characteristics of 20 complex-spiking units recorded in the dorsal cochlear nucleus (DCN) of 15 unanesthetized, decerebrate cats. 2. The extracellularly recorded complex spikes consisted of bursts of two to five action potentials whose size gradually decreased during the burst. Complex spikes were observed both in the spontaneous and sound-evoked activity of the units in our sample. 3. The spontaneous rates (SRs) of DCN complex-spiking units ranged from 0 to 30 spikes/s. Spontaneous activity consisted of complex and simple (i.e., the common single neuronal action potential) spikes. Comparison of the SR distributions of the DCN complex-spiking units with that of a total sample of 194 DCN units (from 9 cats) suggests that the complex-spiking units tended to be in the lower half of the DCN SR distribution. 4. Sound-evoked discharges could consist of both complex and simple spikes. On the basis of their sound-driven responses, we divided the DCN complex-spiking units into two groups. The majority (15 of 20, 75%) were weakly driven by pure tones and inhibited by broadband noise. They tended to have broad response areas. Their response latencies to pure tone and noise stimuli were relatively long (10-20 ms). The recording depths of these units tended to be superficial (i.e., 10 of 15 units were located within 400 microns of the dorsal surface of the DCN). A minority (5 of 20, 25%) of the complex-spiking units were strongly driven by pure tone and broadband noise stimuli. These units had more clearly defined excitatory regions of response areas than the weakly driven units. Their response latencies to pure tone and noise stimuli were short (< 10 ms). The recording depths of these units tended to be deeper (i.e., 4 of 5 units were located at 400-700 microns) than those of the weakly driven units. 5. Intracellular recording and labeling studies of in vitro DCN slice preparations have correlated complex spikes with the superficially located cartwheel cells. Given the complex spikes of the units, many of which were located superficially, we suggest that our sample, particularly the weakly driven group of neurons, corresponds to the cartwheel cells. 6. Cartwheel cells are putative inhibitory interneurons whose axons primarily contact on the main projection neurons of DCN, the fusiform cells. The present finding of sound-evoked discharges by the superficially located complex-spiking units suggests that cartwheel cells should play a role in modifying the sound-evoked responses of the fusiform cells.

Neural modeling of the dorsal cochlear nucleus: cross-correlation analysis of short-duration tone-burst responses.

A conceptual model of a portion of dorsal cochlear nucleus (DCN) neural circuitry has emerged over the past two decades. This model suggests that the response properties of the DCN's major projection neurons, called type IV units, are due, in part, to the behavior of local circuit inhibitory interneurons called type II units (Young and Brownell 1976). Cross-correlation studies of simultaneously recorded pairs of DCN units in decerebrate cat derived from 50-s best frequency (BF) stimuli are consistent with and have extended this conceptual model (Voigt and Young 1980, 1985, 1988, 1990). Interestingly, Gochin et al. (1989) found no signs of inhibition in the anesthetized rat DCN in cross-correlograms derived from 55-ms short-duration BF tone bursts. This seemingly contradictory result has motivated this study. Computer simulations were run using our network model of the intrinsic DCN neural circuitry. This model has previously been shown to reproduce the major features of both type II and type IV rate-level curves and the inhibitory trough (IT) observed in cross-correlograms derived from long-duration stimuli (Voigt and Davis 1994). The goal was to study the stimulus-duration-dependent strength of ITs in the cross-correlograms derived from short-duration BF tone-burst stimuli. The results suggest that ITs may not be detectable when the stimulus duration is 50 ms but may be detectable when the stimulus duration is 200 ms or greater. Furthermore, when the ITs are detected in cross-correlograms derived from 200-ms data sets, the strength of the IT, as measured by effectiveness, is comparable to the strength of ITs measured when the stimulus duration is 50 s.

Dorsal cochlear nucleus single neurons can enhance temporal processing capabilities in background noise.

Envelope temporal fluctuations are critical for effective processing of biologically relevant sounds including speech, animal vocalizations, sound-source location and pitch. Amplitude modulation (AM) of sound envelopes can be encoded in quiet with high fidelity by some auditory neurons, including those of the cochlear nucleus. From both neurophysiological and clinical perspectives, it is important to understand the effects of background noise on the processing of AM. To further this goal, single-unit recordings were made from dorsal cochlear nucleus (DCN) units in urethane-anesthetized chinchillas. All units of this study were classified as pauser/buildup or On-s units according to PSTH response patterns, first spike latencies, and shape of best-frequency (BF) rate-intensity functions. BF pure-tone and AM (10-500 Hz) tone bursts were presented at several sound levels, in quiet and in the presence of a continuous wideband masker. The following was found: (1) DCN units can enhance their AM coding relative to quiet in the presence of loud noise (+14 or +19 dB S/N) and at high signal levels (e.g. 75 dB SPL); (2) for the sample of units of the present study, this is usually achieved by lowering the average firing rate and increasing the synchronous (fundamental frequency) response; (3) for some units, the AM coding stays the same or declines in the background noise. The nature of these findings suggests that part of a DCN unit's abilities to preserve or enhance AM coding with masking noise results from peripheral operating range shifts, whereas part comes from intrinsic circuitry (inhibitory inputs) or cellular mechanisms (dendritic filtering of sound temporal features) within the DCN.

Spectral and temporal response patterns of single units in the chinchilla dorsal cochlear nucleus

Spectral and temporal response patterns to pure-tone stimuli were collected from single units in the dorsal cochlear nucleus of anesthetized chinchillas. The spectral response profiles were divisible into groups based on the balance of excitation and inhibition. Temporal responses were characterized in chloralose-anesthetized animals by collecting PST-histograms. There appeared to be no simple one-to-one relationship between a unit's spectral and its temporal response pattern. Excitatory spectral responses were generally sharply tuned areas resembling those of auditory nerve fibers. However, unlike the latter, the majority of these had chopper or pauser/buildup temporal responses. Inhibitory spectral responses were of two distinct types: one included lateral inhibitory areas flanking the tuned excitatory areas which occasionally invaded the latter creating a nonmonotonic excitatory response at the unit's characteristic frequency. The other included sharply tuned inhibitory areas. The characteristic frequencies of these units were found to be in close correspondence with those of sharply tuned excitatory units from the same penetration suggesting that these inhibitory units were tonotopically mapped in the same register as tuned excitatory units. The spectral response patterns were studied with three types of anesthesia: ketamine/xylazine, dial/urethane, and chloralose. In each of these groups the patterns were similar. However, the proportions of units showing inhibition was strongly dependent on the choice of anesthetic agent with chloralose yielding the highest proportions (59%) and ketamine/xylazine yielding the lowest (29%).

Inhibitory connections between AVCN and DCN: Evidence from lidocaine injection in AVCN

Principal cells of the dorsal cochlear nucleus (DCN) receive a variety of inhibitory influences, including some that are associated with the anteroventral cochlear nucleus (AVCN). Electrical stimulation in the caudal part of the AVCN produces long-lasting inhibition of DCN units [(1973) Exp. Brain Res. 17, 4287ndash;442], including unit type IV, which is associated with principal cells. This inhibition may be mediated either by orthodromic activation of AVCN axons that terminate in DCN or by antidromic stimulation of DCN inhibitory interneurons whose axons send collaterals to the AVCN. These two possibilities cannot be distinguished with electrical stimulation alone. In this study, microinjections of lidocaine were made in the AVCN. These injections should block AVCN to DCN axons without directly affecting the activity, within the DCN, of DCN interneurons. Two effects of lidocaine injections were observed. In some cases, all activity of DCN cells was abolished by lidocaine. These cases involved either spread of lidocaine to the recording site or into the auditory nerve. In other cases, the inhibitory responses of DCN type IV units were weakened by lidocaine without changing the units' thresholds or spontaneous activity. The latter result demonstrates that there are inhibitory pathways originating in AVCN or passing through AVCN and terminating in the DCN.

Acoustic response properties of units in dorsal cochlear nucleus of unanesthetized, decerebrate gerbil

Single unit recordings were made in the dorsal cochlear nucleus (DCN) of unanesthetized, decerebrate Mongolian gerbils. Access to the gerbil DCN was obtained using a newly described approach [Frisina et al., Hearing Res. 6, 259 (1982)]. Single units, recorded with 3M NaCl filled micropipette electrodes (20 Mohm typical), were classified into three types (II, III, or IV) based on their responses to best frequency (BF) tone bursts and their rates of spontaneous discharges. Type II units have little or no spontaneous activity and are excited by BF tones at all levels above threshold. Type III units are also excited by BF tones at all levels above threshold, but have spontaneous activity rates that exceed 2.5 spikes/s. Type IV units are excited over a narrow range of sound levels near BF threshold, but are strongly inhibited at higher levels. Thus all unit types found in the unanesthetized, decerebrate cat DCN [Young and Brownell, J. Neurophysiol. 39, 282 (1976); Young and Voigt, Hearing Res. 6, 153 (1982)] are also found in gerbil. [Work supported by NIH.]

Response properties of type II and type III units in dorsal cochlear nucleus

Type II and type III units in the dorsal cochlear nucleus (DCN) of unanesthetized (decerebrate) cats are those with excitatory responses to best frequency (BF) tones at all levels above threshold. They are distinguished from type IV units which give predominantly inhibitory responses to tones. Type II and type III units are distinguished from one another by their rates of spontaneous activity. Type II units have low rates of spontaneous activity (below 2.5 spikes/s), whereas type HI units are spontaneously active at rates up to 95 spikes/s. In this paper we show that segregation of units according to spontaneous activity produces a segregation of several other properties as well. A typical type II unit responds vigorously to BF tones (median maximum rate of 242 spikes/s), has a slightly nonmonotonic rate versus level function, and responds weakly or not at all to broadband noise. These units tend to be located in the deep layer of the DCN. Type III units tend to respond to BF tones at moderate rates (median maximum driven rate of 124 spikes/s), have monotonie or nonmonotonic rate versus level functions, and respond to noise at rates comparable to their tone rates. They are distributed somewhat more uniformly in all DCN layers, but a majority were found in the fusiform cell and molecular layers. Most evidence indicates that type II units are recorded from inhibitory interneurons in the DCN.

Cochlear nucleus unit responses to pure tones in the unanesthetized rabbit

Seventy-six units were studied in the cochlear nucleus of unanesthetized, paralyzed rabbits. Most were located in the dorsal cochlear nucleus. Fifty-eight units were characterized according to their response area by the scheme of Evans and Nelson (1973, Exp. Brain Res., 17, 402–427): 22.5% showed purely excitatory response areas (type I II ), 72.5% showed mixed excitatory and inhibitory response areas (types III and IV), and 5% showed purely inhibitory response areas (type V). Ten units had virtually no spontaneous activity so were unclassifiable. When characterized according to the scheme of Pfeiffer (1966, Exp. Brain Res., 1, 220–235), 41% of the units were primary-like, 24% were choppers, 20% were pausers, 3% were buildup, and 1% were on units. There was no overall correlation between the two methods of unit classification, but type I II units showed a high number of chopper patterns. Dorsal cochlear nucleus units in the unanesthetized rabbit showed response characteristics very similar to those which were reported in the decerebrate cat. However, there was a greater prevalence of primary-like, purely excitatory units in the rabbit dorsal cochlear nucleus. This may have reflected a species difference or have been due to technical considerations. It appears that the rabbit cochlear nucleus has very similar physiologic, as well as anatomic, characteristics as does that of the cat.

Cochlear Nucleus Intracellular Study of Responses to Steady-State and Frequency-Modulated Stimuli

We have studied a total of 164 cochlear nucleus cells and 22 eighth nerve fibers in 46 pentobarbital anesthetized cats. Tuning curves, peristimulus time histograms, and responses to triangular frequency-modulated sweeps were made. Eighth nerve fibers have a characteristic PSTH characterized by gradual adaptation. Responses to FM signals were symmetrical over the intensity and sweep rates tested although firing was enhanced with faster modulation rates. The same type of PSTH and FM responses were seen in many cells in the ventral cochlear nucleus and only occasionally in the dorsal cochlear nucleus (DCN). Many units in the dorsal cochlear nucleus show, instead, complex response patterns to both steady state and FM signals. Some DCN units which have a symmetrical response to FM signals at high intensities show asymmetry or directionality as signal strength is lowered. Units with only an onset response to tone burst or showing nonmonotonic intensity functions were likely to show asymmetry or directionality in their FM response. Both the discharge data and the membrane potential shifts suggest that the ventral cochlear nucleus acts as a relay station transmitting information coded at the periphery. The dorsal cochlear nucleus, however, seems to have an active role in extracting and recoding information. [Supported by NINDS and Giannini Foundation.]