Biomedical subjects

E D Young

Publications and source records attributed to E D Young.

At least 37 records · Page 2Linked to original sources

PubMed1997

Linear and nonlinear spectral integration in type IV neurons of the dorsal cochlear nucleus. I. Regions of linear interaction.

The principal neurons of the dorsal cochlear nucleus have complex response properties, many of which are classified as type IV. These units integrate energy in the acoustic signal in a nonlinear fashion; for example, at high sound levels the response to a noise of narrow bandwidth and to a band-reject filtered noise with a spectral notch of the same bandwidth may both be inhibitory. However, the sum of these two stimuli, which is broadband noise (BBN), generally gives an excitatory response. In other situations, linear interactions among stimulus components are observed. In this paper, three regimes of approximate linearity were identified. First, best-frequency (BF) tones and equal-energy narrow noisebands centered at BF evoke almost the same response, which is consistent with a stage of linear filtering followed by a nonlinearity that generates the rate responses of the neuron. Second, for sounds close to threshold (10-15 dB re threshold), energy over the full bandwidth of the unit is integrated linearly. Within this regime, responses to the narrow noiseband and the spectral notch mentioned above do sum to equal the response to BBN. Finally, two noisebands centered at different frequencies, such that their sum is a notch in a broad band of noise, sum linearly at low sound levels; the degree of linearity improves as the separation between the noisebands increases. The results are interpreted in terms of a model of type IV response generation containing two inhibitory interneurons: type II units, which are active for narrowband stimuli, including tones, and the wideband inhibitor, which is active for broadband stimuli. In most cases, the onset of nonlinearity occurs for stimuli that significantly activate the type II inhibitory interneuron.

Acoustic Stimulation↗

PubMed1997

Linear and nonlinear spectral integration in type IV neurons of the dorsal cochlear nucleus. II. Predicting responses with the use of nonlinear models.

Two nonlinear modeling methods were used to characterize the input/output relationships of type IV units, which are one principal cell type in the dorsal cochlear nucleus (DCN). In both cases, the goal was to derive predictive models, i.e., models that could predict the responses to other stimuli. In one method, frequency integration was estimated from response maps derived from single tones and simultaneous pairs of tones presented over a range of frequencies. This model combined linear integration of energy across frequency and nonlinear interactions of energy at different frequencies. The model was used to predict responses to noisebands with varying width and center frequency. In almost all cases, predictions using two-tone interactions were better than linear predictions based on single-tone responses only. In about half the cases, reasonable quantitative fits were achieved. The fits were best for noisebands with narrow bandwidth and low sound levels. In the second nonlinear method, the spectrotemporal receptive field (STRF) was derived from responses to broadband stimuli. The STRF could account for some qualitative features of the responses to broad noisebands and spectral notches embedded in broad noisebands. Quantitatively, however, the STRFs failed to predict the responses of type IV units even to simple broadband noise stimuli. For narrowband stimuli, the STRF failed to predict even qualitative features (such as excitatory and inhibitory frequency bands). The responses of DCN type IV units presumably result from interactions of two inhibitory sources, a strong one that is preferentially activated by narrowband stimuli and a weaker one that is preferentially activated by broadband stimuli. The results presented here suggest that the STRF measures effects related to the broadband inhibition, whereas two-tone interactions measure mostly effects related to narrowband inhibition. This explains why models based on two-tone interactions predict the responses to narrow noisebands much better then models based on STRFs. It is concluded that a minimal stimulus set for characterizing type IV units must contain both broadband and narrowband stimuli, because each stimulus class by itself activates only partially the integration mechanisms that shape the responses of type IV units. Similar conclusions are expected to hold in other parts of the auditory system: when characterizing a complex auditory unit, it is necessary to use a range of stimuli to ensure that all integration mechanisms are activated.

Acoustic Stimulation↗

PubMed1996

Effects of pinna position on head-related transfer functions in the cat.

To measure the effects of the pinna position on spectral sound localization cues, the head-related transfer function (HRTF) from the free-field to a point in the ear canal was measured for anesthetized cats with their pinnae in three positions: the relaxed, anesthetized position; pulled forward into an approximation of the "alert cat" posture; and pulled back. The general features of HRTFs are not changed by moving the pinna, although the mapping of particular HRTF features onto directions in space is changed. As an approximation, the pinna behaves like a fixed-shaped sound collector, so that HRTFs shift with the pinna when it moves; however, pinna movement changes some quantitative details of HRTFs beyond what is predicted by this approximation. When viewed as directional gain, pinna movements serve to optimize listening conditions. However, when considering sound localization, pinna movements lead to ambiguities regarding source location. If pinna position is not incorporated into the computation, and spectral cues alone are used for localization, the ambiguity is about 60 degrees in azimuth and 30 degrees in elevation. Pinna movements produce similar azimuthal ambiguity in interaural level differences. Interaural time difference cues could be used to reduce the ambiguity in azimuth, but a knowledge of pinna position seems to be necessary to resolve ambiguities in elevation.

Animals↗

PubMed1996

Effects of somatosensory and parallel-fiber stimulation on neurons in dorsal cochlear nucleus.

1. Single units and evoked potentials were recorded in the dorsal cochlear nucleus (DCN) of paralyzed decerebrate cats in response to electrical stimulation at two sites: 1) in the somatosensory dorsal column nuclei (together called MSN below for medullary somatosensory nuclei), which activates mossy-fiber inputs to granule cells in superficial DCN, and 2) on the free surface of the DCN, which activates granule cell axons (parallel fibers) directly. The goal was to evaluate hypotheses about synaptic interactions in the cerebellum-like circuitry of the superficial DCN. A four-pulse facilitation paradigm was used (50-ms interpulse interval); this allows identification of three components of the responses of DCN principal cells (type IV units) to these stimuli. The latencies of the response components were compared with the latency of the evoked potential in DCN, which signals the arrival of the parallel fiber volley at the recording site. 2. The first component is a short-latency inhibitory response; this component is seen only with MSN stimulation and is seen almost exclusively in units also showing the second component, the transient excitatory response. The short-latency inhibitory component precedes the evoked potential. No satisfactory explanation for the short-latency component can be given at present; it most likely reflects a fast-conducting inhibitory input that arrives at the type IV unit before the slowly conducting parallel fibers. 3. The second component is a transient excitatory response; this component is seen with both MSN and parallel fiber stimulation; it is weak and appears to be masked easily by the inhibitory response components. The excitatory component occurs at the same latency as the evoked potential and probably reflects direct excitation of principal cells by granule cell axons. The excitatory component is seen in about half the type IV units for both stimulating sites. With MSN stimulation, the lack of excitation in some units suggests a heterogeneity of cochlear granule cells, with some carrying somatosensory information and some not carrying this information; with parallel fiber stimulation, excitation probably requires the stimulating and recording electrodes to be lined up on the same "beam" of parallel fibers. 4. The third component is a long-lasting inhibitory response that is observed in virtually all type IV units with both MSN and parallel-fiber stimulation; its latency is longer than the evoked potential. Evidence suggests that it is produced by inhibitory input from cartwheel cells. The appearance of this inhibitory component in almost all type IV units can be accounted for by the considerable spread of cartwheel-cell axons in the direction perpendicular to the parallel fibers. 5. The evoked potential and all three components of the unit response vary systematically in size over the four pulses of the electrical stimulus. These results can be accounted for by two phenomena: 1) a facilitation of the granule cell synapses on all cell types that produces a steadily growing response through the four pulses, resembles presynaptic facilitation, and is seen with both MSN and parallel-fiber stimulation; and 2) a strong reduction in the granule cell response between the first and second pulse for MSN stimulation only. This reduction probably occurs presynaptically in the glomerulus or in the granule cell itself and could reflect inhibitory inputs. 6. The response components described above are seen in type IV units recorded in both the fusiform-cell and deep layers of the DCN; this suggests that both pyramidal and giant cells are activated similarly. The simplest interpretation is that both principal cell types are activated by the cerebellum-like circuitry in superficial DCN. Alternatively, because giant cells appear to make limited contact with the granule-cell circuits of superficial DCN, this finding may suggest the existence of currently undescribed granule cell circuits in deep DCN that are si

Animals↗

PubMed1996

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.

Animals↗

PubMed1995

Auditory-nerve encoding of pinna-based spectral cues: rate representation of high-frequency stimuli.

The elevation of sound sources and their front-back position is encoded in spectral cues produced by direction-dependent filtering in the pinna. Auditory-nerve (AN) fiber population recordings were used to analyze the neural representation of the acoustic features which carry this information. The most prominent pinna-produced spectral features occur at frequencies greater than 5 kHz, so this information must be encoded in AN discharge rates and not in measures of phase locking. However, profiles of discharge rate versus fiber best frequency (BF) reveal a poor representation of the spectra of the stimuli, primarily because of fiber-to-fiber variation in rate. The variation is not controlled by rate normalization, but a clear representation of the ratio of the magnitude spectra of two stimuli is seen when responses are plotted as the difference between the rates in response to the two stimuli. This results suggests that precise information about stimulus spectrum is present in discharge rate, which could be revealed in rate profiles constructed with suitable normalization. When binaural stimuli are presented, a weak inhibitory effect, due to the olivocochlear bundle or the middle ear muscle reflex, is observed. The rate changes are small and are not correlated with the spectrum of the contralateral ear stimulus.

Animals↗

PubMed1995

Somatosensory effects on neurons in dorsal cochlear nucleus.

1. Single units and evoked potentials were recorded in dorsal cochlear nucleus (DCN) in response to electrical stimulation of the somatosensory dorsal column and spinal trigeminal nuclei (together called MSN for medullary somatosensory nuclei) and for tactile somatosensory stimuli. Recordings were from paralyzed decerebrate cats. 2. DCN principal cells (type IV units) were strongly inhibited by electrical stimulation (single 50-microA bipolar pulse) in MSN or by somatosensory stimulation. Units recorded in the fusiform cell and deep layers of DCN were inhibited, suggesting that the inhibition affects both types of principal cells (i.e., both fusiform and giant cells). 3. Interneurons (type II units) that inhibit principal cells were only weakly inhibited by electrical stimulation and were never excited, demonstrating that the inhibitory effect on principal cells does not pass through the type II circuit. In the vicinity of the DCN/PVCN (posteroventral cochlear nucleus) boundary, units were encountered that were excited by electrical stimulation in MSN; some of these neurons responded to sound, and some did not. Their response properties are consistent with the hypothesis that they are deep-layer inhibitory interneurons conveying somatosensory information to the DCN. 4. Analysis of the evoked potentials produced by electrical stimulation in MSN suggests that the somatosensory inputs activate the granule cell system of the DCN molecular layer. A model based on previous work by Klee and Rall was used to show that the distribution of evoked potentials in DCN can be explained as resulting from radial currents produced in the DCN molecular and fusiform-cell layers by synchronous activation of granule cells inputs to fusiform and cartwheel cells. Current-source density analysis of the evoked potentials is consistent with this model. Thus molecular layer interneurons (cartwheel and stellate cells) are a second possible source of inhibition to principal cells. 5. With lower stimulus levels (20 microA) and pulse-pair stimuli (50- to 100-ms interstimulus interval), three components of the inhibitory response can be recognized in both fusiform cell layer and deep layer type IV units: a short-latency inhibition that begins before the start of the evoked potential; a longer-latency inhibition whose timing corresponds to the evoked potential; and an excitatory component that occurs on the rising phase of the evoked potential. The excitatory component is usually overwhelmed by the inhibitory components and could be derived from granule cell inputs; the long-latency inhibitory component could be derived from cartwheel cells or the hypothesized deep-layer inhibitory interneurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1994

Comparative analysis of spectro-temporal receptive fields, reverse correlation functions, and frequency tuning curves of auditory-nerve fibers.

The tuning properties of single auditory-nerve fibers (ANFs) are characterized with spectro-temporal receptive fields (STRFs), reverse correlation functions (revcors), and frequency tuning curves (FTCs). Measures of tuning and latency from the STRFs and revcors are largely comparable to the traditional measures of tuning from FTCs and measures of latency from peristimulus time histograms (PSTHs), but several important differences are found. As is well known, revcors can only characterize low (< 6 kHz) best frequency (BF) units, whereas STRFs are able to characterize all units studied (BFs ranging from 0.26-23 kHz), except for a few very low-BF examples. Whereas tuning bandwidth derived from revcor exceeds that measured from FTCs at all BFs and increases with sound level. STRF bandwidth is comparable to FTC bandwidth, except at low BFs, and is stable with sound level. The STRF may reflect nonlinear properties of auditory-nerve fibers such as refractoriness and two-tone suppression that are absent in the FTC and revcor characterizations. The principal drawback of the STRF is its narrow dynamic range.

Animals↗

PubMed1994

The electrotonic structure of regular-spiking neurons in the ventral cochlear nucleus may determine their response properties.

1. Intracellular recordings were obtained from neurons in parasagittal brain slices of the guinea pig ventral cochlear nucleus (VCN). The principal neurons of the VCN can be parceled into two categories. Regular-spiking (Type I) neurons have a linear current-voltage (I-V) relationship over a large range of intracellularly injected currents and fire tonically in response to suprathreshold depolarizing currents. Phasically spiking (Type II) neurons have a nonlinear I-V relationship and fire only phasically at the onset of a depolarizing current or offset of a hyperpolarizing current. Regular-spiking neurons have been shown to be of the stellate morphological type, whereas phasically spiking neurons have been shown to be bushy cells. 2. The electrotonic structure of regular-spiking neurons was studied by applying previously developed modeling techniques based on the somatic shunt model. In these techniques, physiological data are used to determine the set of parameters best describing the neuron. As predicted from previous theoretical investigations, the use of an anatomic constraint (somatic surface area) reduces the variance in estimates of model parameters, especially for the dendritic membrane time constant tau D. 3. Model representations of regular-spiking cells fall into two categories: those with (passive) somatic membrane properties that are nearly identical to those of the dendrite (8/15 cases), and those with a significant amount of somatic shunt (7/15). Estimates of tau D (mean = 7.7 ms) are lower than those often described in the literature. We argue that this low value of tau D may be related to the need of neurons in the auditory brainstem to operate at high firing rates and/or to encode audio-frequency temporal fluctuations. 4. Dendritic transfer functions were calculated as functions of synaptic location using somatic shunt representations of regular-spiking neurons. These transfer functions allow us to predict that mid-range auditory frequencies (approximately 1 kHz) are greatly attenuated, even for synapses near the soma. Thus it is suggested that the electrotonic architecture of regular-spiking cells creates sufficient low-pass filtering of synaptic inputs to reduce the synchronization of firing of these neurons to mid-frequency auditory stimuli.

Animals↗

PubMed1994

Two separate inhibitory mechanisms shape the responses of dorsal cochlear nucleus type IV units to narrowband and wideband stimuli.

1. The principal cells of the dorsal cochlear nucleus (DCN) are mostly inhibited by best frequency (BF) tones but are mostly excited by broadband noise (BBN), producing the so-called type IV response characteristic. The narrowband inhibitory responses can be explained by the inhibitory influence of interneurons with type II response characteristics. However, it is not clear that all the details of the type IV responses can be accounted for by this neural circuit. In particular, many type IV units are inhibited by band-reject noise (notch noise); type II units tend to be only weakly excited by these stimuli, if at all. In this paper we study the relationships between the narrowband, inhibitory and the wideband, excitatory regimens of the type IV responses and present the case for the existence of a second inhibitory source in DCN, called the wideband inhibitor (WBI) below. 2. Type IV units were studied using pure tones, noise bands arithmetically centered on BF, notch noise centered on BF, and BBN. We measured the rate-level function (response rate as function of stimulus level) for each stimulus. This paper is based on the responses of 28 type IV units. 3. Evidence for low-threshold inhibitory input to type IV units is derived from analysis of rate-level functions at sound levels just above threshold. Notch noise stimuli of the appropriate notch width produce inhibition at threshold in this regime. When BBN is presented, this inhibition appears to summate with excitation produced by energy in the band of noise centered on BF, resulting in BBN rate-level functions with decreased slope and maximum firing rate. A range of slopes and maximal firing rates is observed, but these variables are strongly correlated and they are negatively correlated with the strength of the inhibition produced by notch noise; this result supports the conclusion that a single inhibitory source is responsible for these effects. 4. By contrast, there is a weak (nonsignificant) positive correlation between the strength of the inhibitory effect of notch noise and the slope/maximal firing rate in response to narrowband stimuli, including BF tones. The contrast between this positive nonsignificant correlation and the significant negative correlation mentioned above suggests that more than one inhibitory effect operates: specifically, the type II input is responsible for inhibition by narrowband stimuli and a different inhibitory source, the WBI, produces inhibition by notch stimuli. 5. Several lines of evidence are given to show that type II units cannot produce the inhibition seen with notch noise stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1993

Discharge-rate dependence of refractory behavior of cat auditory-nerve fibers.

A descriptive model for auditory-nerve (AN) refractory periods is described. The model assumes that interspike intervals consist of a constant-length absolute refractory period (ARP), followed by random-length relative refractory period (RRP) and a random-length waiting time to the next spike. Both the RRP and waiting time are exponentially distributed. This model fits AN hazard functions sufficiently well to provide estimates of the ARP and RRP durations for each fiber. The ARP is found to be constant, independent of discharge rate, with mean value between 0.56 and 0.86 ms in data from 7 experiments. The RRP decreases in duration as discharge rate increases; RRP mean length is less than 2 ms in most cases. There is an additional, slow component of recovery, lasting 20-40 ms, which is not modeled. The variation in RRP with discharge rate is shown to be capable of accounting for the deviation of AN regularity from that predicted for a Poisson process. Finally, properties of peaks seen in hazard functions just at the end of the ARP are described; these peaks are not included in the model, but are shown to be especially prominent in low and medium spontaneous rate fibers.

Acoustic Stimulation↗

PubMed1993

Convergence of auditory nerve fibers onto bushy cells in the ventral cochlear nucleus: implications of a computational model.

1. Convergence of auditory nerve (AN) fibers onto bushy cells of the ventral cochlear nucleus (VCN) was investigated with a model that describes the electrical membrane properties of these cells. The model consists of a single compartment, representing the soma, and includes three voltage-sensitive ion channels (fast sodium, delayed-rectifier-like potassium, and low-threshold potassium). These three channels have characteristics derived from voltage clamp data of VCN bushy cells. The model also contains a leakage channel, membrane capacitance, and synaptic inputs. The model accurately reproduces the membrane rectification seen in current clamp studies of bushy cells, as well as their unique current clamp responses. 2. In this study, the number and synaptic strength of excitatory AN inputs to the model were varied to investigate the relationship between input convergence parameters and response characteristics. From 1 to 20 excitatory synaptic inputs were applied through channels in parallel with the voltage-gated channels. Each synapse was driven by an independent AN spike train; spike arrivals produced brief (approximately 0.5 ms) conductance increases. The number (NS) and conductance (AE) of these inputs were systematically varied. The input spike trains were generated as a renewal point process that accurately models characteristics of AN fibers (refractoriness, adaptation, onset latency, irregularity of discharge, and phase locking). Adaptation characteristics of both high and low spontaneous rate (SR) AN fibers were simulated. 3. As NS and AE vary over the ranges 1-20 and 3-80 nS, respectively, the full range of response types seen in VCN bushy cells are produced by the model, with AN inputs typical of high-SR AN fibers. These include primarylike (PL), primarylike-with-notch (Pri-N), and onset-L (On-L). In addition, Onset responses, whose association with bushy cells in uncertain, and "dip" responses, which are not seen in the VCN, are produced. Dip responses occur with large NS and/or AE, and are due to depolarization block. When the AN inputs have the adaptation characteristics of low-SR AN fibers, PL--but not Pri-N or On-L responses--are produced. This suggests that neurons showing Pri-N and On-L responses must receive high-SR AN inputs. 4. The regularity of discharge of the model is compared with that of VCN bushy cells, using a measure derived from the mean and standard deviation of interspike intervals. Regularity is an important constraint on the model because the regularity of VCN bushy cells is the same as that of their AN inputs.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1992

Neural organization and responses to complex stimuli in the dorsal cochlear nucleus.

The dorsal division of the cochlear nucleus (DCN) is the most complex of its subdivisions in terms of both anatomical organization and physiological response types. Hypotheses about the functional role of the DCN in hearing are as yet primitive, in part because the organizational complexity of the DCN has made development of a comprehensive and predictive model of its input-output processing difficult. The responses of DCN cells to complex stimuli, especially filtered noise, are interesting because they demonstrate properties that cannot be predicted, without further assumptions, from responses to narrow band stimuli, such as tones. In this paper, we discuss the functional organization of the DCN, i.e. the morphological organization of synaptic connections within the nucleus and the nature of synaptic interactions between its cells. We then discuss the responses of DCN principal cells to filtered noise stimuli that model the spectral sound localization cues produced by the pinna. These data imply that the DCN plays a role in interpreting sound localization cues; supporting evidence for such a role is discussed.

Acoustic Stimulation↗

PubMed1992

The parameter identification problem for the somatic shunt model.

The somatic shunt model, a generalized version of the Rall equivalent cylinder model, is used commonly to describe the passive electrotonic properties of neurons. Procedures for determining the parameters of the somatic shunt model that best describe a given neuron typically rely on the response of the cell to a small step of hyperpolarizing current injected by an intrasomatic recording electrode. In this study it is shown that the problem of estimating model parameters for the somatic shunt model using physiological data is ill-posed, in that very small errors in measured data can lead to large and unpredictable errors in parameter estimates. If the somatic shunt is assumed to be a real property of the intact neuron, the effects of these errors are not severe when predicting EPSP waveshapes resulting from synaptic input at a given location. However, if the somatic shunt is assumed to be a consequence of a leakage pathway around the recording electrode, and a correction for the shunt is applied, then the instability of the inverse problem can introduce large errors in estimates of EPSP waveshape as a function of synaptic location in the intact cell. Morphological constraints can be used to improve the accuracy of the inversion procedure in terms of both parameter estimates and predicted EPSP responses.

Mathematics↗

PubMed1992

Pinna-based spectral cues for sound localization in cat.

The directional dependence of the transfer function from free field plane waves to a point near the tympanic membrane (TM) was measured in anesthetized domestic cats. A probe tube microphone was placed approximately 3 mm from the TM from beneath the head in order to keep the pinna intact. Transfer functions were computed as the ratio of the spectrum of a click recorded near the TM to the spectrum of the click in freefield. We analyze the transfer functions in three frequency ranges: low frequencies (less than 5 kHz) where interaural level differences vary smoothly with azimuth; midfrequencies (5-18 kHz) where a prominent spectral notch is observed; and high frequencies (greater than 18 kHz) where the transfer functions vary greatly with source location. Because no two source directions produce the same transfer function, the spectrum of a broadband sound at the TM could serve as a sound localization cue for both elevation and azimuth. In particular, we show that source direction is uniquely determined, for source directions in front of the cat, from the frequencies of the midfrequency spectral notches in the two ears. The validity of the transfer functions as measures of the acoustic input to the auditory system is considered in terms of models of sound propagation in the ear canal.

Acoustic Stimulation↗

PubMed1992

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

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

Auditory Perception↗

PubMed1991

Organization of dorsal cochlear nucleus type IV unit response maps and their relationship to activation by bandlimited noise.

1. Response maps of 49 type IV neurons in cat dorsal cochlear nucleus (DCN) were studied by moving a tone in small steps along the frequency dimension and along the intensity dimension. Type IV responses are recorded from DCN principal cells. Data were collected from 38 units with best frequencies (BFs) from 2.16 to 50.3 kHz with the use of electrode penetrations along the long (strial) axis of the DCN; an additional 11 units from a previous study were analyzed. A stereotypical type IV response map is defined as consisting of two excitatory and two inhibitory regions. Type IV units from both the pyramidal cell layer (probably pyramidal cells) and the deep layer (probably giant cells) show the same types of response maps. 2. Two of the regions, one excitatory and one inhibitory, are seen in all type IV units. These regions are a low-threshold excitatory region at best frequency (BFER) and an inhibitory area at higher levels, usually centered below BF but extending upward in frequency to include BF (central inhibitory area, or CIA). The high resolution of the response maps in this paper allows us to show that type IV units fall into two groups on the basis of whether their CIAs are narrow with well-defined borders (35 units) or broad with poorly defined borders (14 units). 3. Two additional features of type IV response maps can be defined, most consistently in units with well-defined CIAs. These features are an excitatory region along the high-frequency edge of the CIA (upper excitatory region, UER) and an upper inhibitory sideband (UIS). The BFER and UER are continuous in many units, but in some cases their continuity is broken by the CIA. It seems likely that the BFER and UER represent a single excitatory input to type IV units and are revealed because the tuning curve of the stronger inhibitory inputs that produce the CIA has thresholds greater than and BFs lower than the excitatory inputs. 4. The CIA is probably produced by inhibitory inputs from DCN type II neurons. The bandwidths of type IV CIAs are about 1-3 times larger (at 40 dB above threshold) than the excitatory bandwidths of DCN type II units, suggesting a convergence of the equivalent in tuning of about two type II units onto each type IV unit. The BF of the CIA is below the excitatory BF of the type IV unit in most cases.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

PubMed1990

Cross-correlation analysis of inhibitory interactions in dorsal cochlear nucleus.

1. Cross-correlation analysis was used to study the organization of inhibitory connections between type II or type III units and type IV principal cells in cat dorsal cochlear nucleus (DCN). Pairs of units were isolated using two microelectrodes so that information about the distance over which connections are made could be analyzed. Data were obtained from 51 pairs consisting of a type II and a type IV unit and from 22 pairs consisting of a type III and a type IV unit. The analyses in this paper concentrate on type II-type IV pairs. 2. Inhibitory troughs (ITs) are observed in the cross-correlograms of type II-type IV pairs (21/51 cases). An IT is a transient decrease in discharge probability in the postsynaptic (type IV) unit immediately after spikes in the presynaptic unit (type II). The average latency to the start of ITs is 0.73 ms, and the troughs are asymmetric with a faster leading phase. Small excitatory peaks accompany the ITs in type II units, but these are probably secondary effects associated with the IT. ITs are consistent with a monosynaptic, inhibitory connection between type II and type IV units. A variety of evidence suggests that type II responses are recorded from vertical cells, an interneuron in the deep layer of the DCN that may be glycinergic. 3. The cross-correlograms of type III-type IV pairs are more complex and variable than those of type II-type IV pairs--ITs are seen in 4/22 cases, and peaks of correlation that are symmetrically located around the origin (central mound or CM) are seen in 4/22 cases; two cases have both an IT and a CM. CMs result from shared sources of input. Whereas type II-type IV correlogram features change primarily in amplitude as stimulus conditions change, correlogram features in some type III-type IV pairs change qualitatively with stimulus conditions; correlograms are flat for some stimuli and show ITs or CMs or mixtures of the two for others. This variability suggests that the circuitry associated with type III-type IV pairs is more complex than a monosynaptic connection, and further analysis of type III-type IV pairs was not done. 4. The strength of inhibition for an IT is measured as the area under the IT (effectiveness) and as effectiveness divided by the postsynaptic discharge rate (association index).(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗