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PubMed1999

Locating animals from their sounds and tomography of the atmosphere: experimental demonstration.

Calling animals are located using widely distributed receivers, and the sounds from the animals are used to map the sound speed and wind fields by means of tomography. In particular, two Red-Winged Blackbirds Agelaius phoeniceus are correctly located within a meter using recordings from five receivers spread over a 20 by 30 m region. The demonstration hinges on two new developments. First, a new algorithm for blindly estimating the impulse response of the channel is shown capable of estimating the differences in the time of first arrivals at two receivers. Since it is known that the first arrivals travel along nearly straight paths, the difference in time constrains the animal's location to a hyperboloid, and the animal is located by intersecting hyperboloids from many pairs of receivers. Second, in order to accurately find the intersection point and map the sound speed and wind fields using tomography, a nonlinear equation is solved. The new algorithm for blindly estimating the impulse response of a channel offers a new way for locating sounds and making tomographic maps of the environment without any requirement for a model for the propagation of sound such as is needed for focalization and matched field processing.

Algorithms↗
PubMed1999

Individual differences in external-ear transfer functions reduced by scaling in frequency.

This study examined inter-subject differences in the transfer functions from the free field to the human ear canal, which are commonly know as head-related transfer functions. The directional components of such transfer functions are referred here to as directional transfer functions (DTFs). The DTFs of 45 subjects varied systematically among subjects in regard to the frequencies of spectral features such as peaks and notches. Inter-subject spectral differences in DTFs were quantified between 3.7 and 12.9 kHz for sound-source directions throughout the coordinate sphere. For each pair of subjects, an optimal frequency scale factor aligned spectral features between subjects and, thus, minimized inter-subject spectral differences. Frequency scaling of DTFs reduced spectral differences by a median value of 15.5% across all pairs of subjects and by more than half in 9.5% of subject pairs. Optimal scale factors showed a median value of 1.061 and a maximum of 1.38. The optimal scale factor between any pair of subjects correlated highly with the ratios of subjects' maximum interaural delays, sizes of their external ears, and widths of their heads.

Adult↗
PubMed2002

Gated visual input to the central auditory system.

The central auditory system translates sound localization cues into a map of space guided, in part, by visual experience. In barn owls, this process takes place in the external nucleus of the inferior colliculus (ICX). However, to date, no trace of visual activity has been observed in this auditory nucleus. Here we show that strong visual responses, which are appropriate to guide auditory plasticity, appear in the ICX when inhibition is blocked in the optic tectum. Thus, visual spatial information is gated into the auditory system by an inhibitory mechanism that operates at a higher level in the brain.

Acoustic Stimulation↗
PubMed2003

Glutamatergic calcium responses in the developing lateral superior olive: receptor types and their specific activation by synaptic activity patterns.

The lateral superior olive (LSO) is a binaural auditory brain stem nucleus that plays a central role in sound localization. Survival and maturation of developing LSO neurons critically depend on intracellular calcium signaling. Here we investigated the mechanisms by which glutamatergic afferents from the cochlear nucleus increase intracellular calcium concentration in LSO neurons. Using fura-2 calcium imaging in slices prepared from neonatal mice, we found that cochlear nucleus afferents can activate all major classes of ionotropic and metabotropic glutamate receptors, each of which contributes to an increase in intracellular calcium. The specific activation of different glutamate receptor classes was dependent on response amplitudes and afferent stimulus patterns. Low-amplitude responses elicited by single stimuli were entirely mediated by calcium-impermeable AMPA/kainate receptors that activated voltage-gated calcium channels. Larger-amplitude responses elicited by either single stimuli or stimulus trains resulted in additional calcium influx through N-methyl-d-aspartate receptors. Finally, high-frequency stimulation also recruited group I and group II metabotropic glutamate receptors, both of which mobilized intracellular calcium. This calcium release in turn activated a strong influx of extracellular calcium through a membrane calcium channel that is distinct from voltage-gated calcium channels. Together, these results indicate that before hearing onset, distinct patterns of afferent activity generate qualitatively distinct types of calcium responses, which likely serve in guiding different aspects of LSO development.

Afferent Pathways↗
PubMed2003

Modulation of cochlear afferent response by the lateral olivocochlear system: activation via electrical stimulation of the inferior colliculus.

The olivocochlear (OC) efferent innervation of the mammalian inner ear consists of two subdivisions, medial (MOC) and lateral (LOC), with different peripheral terminations on outer hair cells and cochlear afferent terminals, respectively. The cochlear effects of electrically activating MOC efferents are well known, i.e., response suppression effected by reducing outer hair cells' contribution to cochlear amplification. LOC peripheral effects are unknown, because their unmyelinated axons are difficult to electrically stimulate. Here, stimulating electrodes are placed in the inferior colliculus (IC) to indirectly activate the LOC system, while recording cochlear responses bilaterally from anesthetized guinea pigs. Shocks at some IC sites produced novel cochlear effects attributable to activation of the LOC system: long-lasting (5-20 min) enhancement or suppression of cochlear neural responses (compound action potentials and round window noise), without changes in cochlear responses dominated by outer hair cells (otoacoustic emissions and cochlear microphonics). These novel effects also differed from classic MOC effects in their lack of dependence on the level and frequency of the acoustic stimulus. These effects disappeared on sectioning the entire OC bundle, but not after selective lesioning of the MOC tracts or the cochlea's autonomic innervation. We conclude that the LOC pathway comprises two functional subdivisions, capable of inducing slow increases or decreases in response magnitudes in the auditory nerve. Such a system may be useful in maintaining accurate binaural comparisons necessary for sound localization in the face of slow changes in interaural sensitivity.

Acoustic Stimulation↗
PubMed2002

Neural correlates of the precedence effect in the inferior colliculus: effect of localization cues.

The precedence effect (PE) is an auditory phenomenon involved in suppressing the perception of echoes in reverberant environments, and is thought to facilitate accurate localization of sound sources. We investigated physiological correlates of the PE in the inferior colliculus (IC) of anesthetized cats, with a focus on directional mechanisms for this phenomenon. We used a virtual space (VS) technique, where two clicks (a "lead" and a "lag") separated by a brief time delay were each filtered through head-related transfer functions (HRTFs). For nearly all neurons, the response to the lag was suppressed for short delays and recovered at long delays. In general, both the time course and the directional patterns of suppression resembled those reported in free-field studies in many respects, suggesting that our VS simulation contained the essential cues for studying PE phenomena. The relationship between the directionality of the response to the lead and that of its suppressive effect on the lag varied a great deal among IC neurons. For a majority of units, both excitation produced by the lead and suppression of the lag response were highly directional, and the two were similar to one another. For these neurons, the long-lasting inhibitory inputs thought to be responsible for suppression seem to have similar spatial tuning as the inputs that determine the excitatory response to the lead. Further, the behavior of these neurons is consistent with psychophysical observations that the PE is strongest when the lead and the lag originate from neighboring spatial locations. For other neurons, either there was no obvious relationship between the directionality of the excitatory lead response and the directionality of suppression, or the suppression was highly directional whereas the excitation was not, or vice versa. For these neurons, the excitation and the suppression produced by the lead seem to depend on different mechanisms. Manipulation of the directional cues (such as interaural time and level differences) contained in the lead revealed further dissociations between excitation and suppression. Specifically, for about one-third of the neurons, suppression depended on different directional cues than did the response to the lead, even though the directionality of suppression was similar to that of the lead response when all cues were present. This finding suggests that the inhibitory inputs causing suppression may originate in part from subcollicular auditory nuclei processing different directional cues than the inputs that determine the excitatory response to the lead. Neurons showing such dissociations may play an important role in the PE when the lead and the lag originate from very different directions.

Acoustic Stimulation↗
PubMed2004

Multiple large inputs to principal cells in the mouse medial nucleus of the trapezoid body.

The calyx of Held is a giant nerve terminal that forms a synapse directly onto the principal cells of the medial nucleus of the trapezoid body (MNTB) in the mammalian auditory brain stem. This central synapse, which is involved in sound localization, has become widely used for studying synaptic transmission. Anatomical studies of this nucleus have indicated that each principal cell is innervated by only one calyx. Here we use previously established electrophysiological criteria of excitatory postsynaptic current amplitude, kinetics, and transmitter type, as well as other characteristics commonly reported for this synapse, to examine the input properties of principal neurons. Our findings indicate that some principal cells receive more than one strong excitatory input. These inputs meet previously established electrophysiological criteria for identification as calyceal nerve terminals. Implications for the execution and analysis of experiments to avoid errors due to such multiple inputs are discussed.

Animals↗
PubMed2005

Microsecond precision of phase delay in the auditory system of the barn owl.

The auditory system encodes time with sub-millisecond accuracy. To shed new light on the basic mechanism underlying this precise temporal neuronal coding, we analyzed the neurophonic potential, a characteristic multiunit response, in the barn owl's nucleus laminaris. We report here that the relative time measure of phase delay is robust against changes in sound level, with a precision sharper than 20 micros. Absolute measures of delay, such as group delay or signal-front delay, had much greater temporal jitter, for example due to their strong dependence on sound level. Our findings support the hypothesis that phase delay underlies the sub-millisecond precision of the representation of interaural time difference needed for sound localization.

Acoustic Stimulation↗
PubMed1993

High-frequency neurons in the inferior colliculus that are sensitive to interaural delays of amplitude-modulated tones: evidence for dual binaural influences.

1. Localization of sounds has traditionally been considered to be performed by a duplex mechanism utilizing interaural temporal differences (ITDs) at low frequencies and interaural intensity differences at higher frequencies. More recently, it has been found that listeners can detect ITDs at high frequencies if the amplitude of the sound varies and an ITD is present in the envelope. Here we report the responses of neurons in the inferior colliculi of unanesthetized rabbits to ITDs of the envelopes of sinusoidally amplitude-modulated (SAM) tones. 2. Neurons were studied extracellularly with glass-coated Pt-Ir or Pt-W microelectrodes. Their sensitivity to ITDs in the envelopes of high-frequency sounds (> or = 2 kHz) was assessed using SAM tones that were presented binaurally. The tones at the two ears had the same carrier frequency but modulation frequencies that differed by 1 Hz. This caused a cyclic variation in the ITD produced by the envelope. In this "binaural SAM" stimulus, the carriers caused no ITD because they were in phase. In addition to the binaural SAM stimulus, pure tones were used to investigate responses to ipsilateral and contralateral stimulation and the nature of the interaction during binaural stimulation. 3. Neurons tended to display one of two kinds of sensitivity to ITDs. Some neurons discharged maximally at the same ITD at all modulation frequencies > 250 Hz (peak-type neurons), whereas others were maximally suppressed at the same ITD (trough-type neurons). 4. At these higher modulation frequencies (> 250 Hz), the characteristic delays that neurons exhibited tended to lie within the range that a rabbit might normally encounter (+/- 300 microseconds). The peak-type neurons favored ipsilateral delays, which correspond to sounds in the contralateral sound field. The trough-type neurons showed no such preference. 5. The preference of peak-type neurons for a particular delay was sharper than that of trough-type neurons and was comparable to that observed in neurons of the inferior colliculus that are sensitive to delays of low-frequency pure tones. 6. At lower modulation frequencies (< 150 Hz) characteristic delays often lay beyond +/- 300 microseconds. 7. Increasing the ipsilateral intensity tended to shift the preferred delay ipsilaterally at lower (< 250 Hz), but not at higher, modulation frequencies. 8. When tested with pure tones, a substantial number of peak-type neurons were found to be excited by contralateral stimulation but inhibited by ipsilateral stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1993

Hyperpolarization-activated cation current (Ih) in neurons of the medial nucleus of the trapezoid body: voltage-clamp analysis and enhancement by norepinephrine and cAMP suggest a modulatory mechanism in the auditory brain stem.

1. Principal cells in the medial nucleus of the trapezoid body (MNTB) are part of a circuit in the superior olivary complex (SOC) that processes binaural information important for sound localization. MNTB cells have two voltage-dependent currents active near rest that contribute to these cells' highly nonlinear membrane properties and shape their responses to synaptic input. One of these currents, a low-threshold, 4-aminopyridine (4-AP)-sensitive K+ current, has been studied previously under current clamp. Using the single-electrode voltage-clamp technique, we have investigated the other of these currents, a hyperpolarization-activated, mixed cation current (Ih), in brain slices of the rat SOC. 2. Ih is responsible for a prominent "sag" in the voltage response to a steady hyperpolarizing current recorded under current clamp in MNTB cells. In voltage-clamp recordings, hyperpolarizing voltage steps from the resting potential elicited a large inward current that activated and deactivated with biexponential kinetics. Activation time constants were voltage dependent, with tau 1 and tau 2 = 246 and 1620 ms at -75 mV and 107 and 560 ms at -100 mV. 3. Ih was blocked by 1-5 mM cesium and had a reversal potential of -43 mV. Steady-state activation curves derived from tail currents yielded a half-activation voltage of -75.7 mV and slope factor of 5.7 mV, corresponding to < 10% activation of Ih at rest. 4. Application of norepinephrine (15-20 microM) or 8-bromoadenosine 3',5'-cyclic monophosphate (8-Br-cAMP) (1 mM) caused a depolarizing shift in the steady-state activation curve and decreased the activation time constants. The shift in the activation curve resulted in a large increase in the activation of Ih at rest, an inward shift in the holding current, and an increase in the resting membrane conductance. In current-clamp recordings, this increase in the resting activation level of Ih resulted in membrane depolarization of 2-3 mV in the absence of 4-AP, and 5-10 mV in the presence of 4-AP, an increase in the input conductance, and a reduction in the voltage sag in response to hyperpolarizing currents. 5. The resulting change in the resting point of MNTB cells exposed to norepinephrine or 8-Br-cAMP is likely to alter the responses of these cells to synaptic input, both via the direct effect on the resting membrane conductance and by changing the activation of the low-threshold, 4-AP-sensitive potassium current.(ABSTRACT TRUNCATED AT 400 WORDS)

4-Aminopyridine↗
PubMed1994

Interaction of excitation and inhibition in processing of pure tone and amplitude-modulated stimuli in the medial superior olive of the mustached bat.

1. In mammals with good low-frequency hearing, the medial superior olive (MSO) processes interaural time or phase differences that are important cues for sound localization. Its cells receive excitatory projections from both cochlear nuclei and are thought to function as coincidence detectors. The response patterns of MSO neurons in most mammals are predominantly sustained. In contrast, the MSO in the mustached bat is a monaural nucleus containing neurons with phasic discharge patterns. These neurons receive projections from the contralateral anteroventral cochlear nucleus (AVCN) and the ipsilateral medial nucleus of the trapezoid body (MNTB). 2. To further investigate the role of the MSO in the bat, the responses of 252 single units in the MSO to pure tones and sinusoidal amplitude-modulated (SAM) stimuli were recorded. The results confirmed that the MSO in the mustached bat is tonotopically organized, with low frequencies in the dorsal part and high frequencies in the ventral part. The 61-kHz region is overrepresented. Most neurons tested (88%) were monaural and discharged only in response to contralateral stimuli. Their response could not be influenced by stimulation of the ipsilateral ear. 3. Only 11% of all MSO neurons were spontaneously active. In these neurons the spontaneous discharge rate was suppressed during the stimulus presentation. 4. The majority of cells (85%) responded with a phasic discharge pattern. About one-half (51%) responded with a level-independent phasic ON response. Other phasic response patterns included phasic OFF or phasic ON-OFF, depending on the stimulus frequency. Neurons with ON-OFF discharge patterns were most common in the 61-kHz region and absent in the high-frequency region. 5. Double tone experiments showed that at short intertone intervals the ON response to the second stimulus or the OFF response to the first stimulus was inhibited. 6. In neuropharmacological experiments, glycine applied to MSO neurons (n = 71) inhibited any tone-evoked response. In the presence of the glycine antagonist strychnine the response patterns changed from phasic to sustained (n = 35) and the neurons responded to both tones presented in double tone experiments independent of the intertone interval (n = 5). The effects of strychnine were reversible. 7. Twenty of 21 neurons tested with sinusoidally amplitude-modulated (SAM) signals exhibited low-pass or band-pass filter characteristics. Tests with SAM signals also revealed a weak temporal summation of inhibition in 13 of the 21 cells tested.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Azimuthal receptive fields are shaped by GABAergic inhibition in the inferior colliculus of the mustache bat.

1. In this study we examine the effects of GABAergic inhibition on the response properties and the constructed azimuthal receptive fields of 54 excitatory/inhibitory (EI) neurons tuned to 60 kHz in the inferior colliculus of the mustache bat. The constructed azimuthal receptive fields predict the spike counts that would be evoked by different intensities of 60-kHz sounds presented from each of 13 azimuthal locations in the frontal sound field. 2. Action potentials were recorded with a micropipette attached to a multibarrel glass electrode. Bicuculline, an antagonist specific for gamma-aminobutyric acid-A (GABAA) receptors, was iontophoretically applied through the multibarrel electrode. Both monaural and binaural response properties were initially recorded at a variety of interaural intensity disparities (IIDs) and absolute intensities, and the same response properties were subsequently assessed while GABAergic inhibition was blocked by bicuculline. Azimuthal receptive fields both before and during the application of bicuculline were constructed from response properties obtained with earphones after correcting for the directional properties of the ear and the IIDs generated by 60-kHz sounds presented from a variety of azimuthal locations. 3. Bicuculline had virtually no effect on either the monaural or binaural properties of 19 cells (35%). The constructed azimuthal receptive fields of these cells were also unaffected by bicuculline. Presumably the properties of these cells were formed in a lower nucleus, most likely the contralateral lateral superior olive (LSO), and were imposed on the collicular cell via the crossed projection from the LSO to the inferior colliculus, which is known to be excitatory. 4. In more than half of the neurons (65%) GABAergic inhibition influenced one or more features of the cell's response properties and thus its azimuthal receptive field. Some response properties were formed in the colliculus through GABAergic inhibition, whereas others appear to have been shaped initially in a lower nucleus and then further modified by GABAergic inhibition in the inferior colliculus. Moreover, a number of features of GABAergic inhibition that acted on inferior collicular cells were evoked by stimulation of the contralateral (excitatory) ear, whereas other features were influenced by stimulation of the ipsilateral (inhibitory) ear. 5. In 20 cells (37%) blocking GABAergic inhibition reduced or abolished the inhibition evoked by the ipsilateral ear. The receptive fields of cells in which the ipsilaterally evoked inhibition was reduced by bicuculline expanded further into the ipsilateral sound field than they did before bicuculline.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1996

Representation of frequency in the primary auditory field of the barn owl forebrain.

1. The primary auditory field (PAF) constitutes the first telencephalic stage of auditory information processing in the classical auditory pathway. In this study we investigated the frequency representation in the PAF of the barn owl, a species with a broad frequency range of hearing and a highly advanced auditory system. 2. Single- and multiunit sites were recorded extracellularly in ketamine-anesthetized owls. The frequency response properties of PAF sites were assessed with the use of digitally synthesized dichotic stimuli. PAF sites (n = 442) either were unresponsive to tonal stimulation (but responsive to noise stimuli), were tuned for frequency, or had multipeaked frequency response profiles. Tuned sites responded best to frequencies between 0.2 and 8.8 kHz, a range that encompasses nearly the entire hearing range of the barn owl. Most sites responding best to frequencies < 4 kHz had relatively broad frequency tuning, whereas sites responding best to higher frequencies had either broad or narrow frequency tuning. Sites with multipeaked frequency response profiles typically had two response peaks. The first peak was usually between 1 and 3 kHz and the second was usually between 5 and 8 kHz; there was no systematic relationship between the two peak frequencies. 3. In dorsoventral electrode penetrations that contained sites with tuned and/or multipeaked response profiles, a "common frequency" was identified that elicited a maximal response from all of the sites in the penetration. 4. The PAF contains a single tonotopic field. Units tuned to low frequencies are located caudomedially, whereas units tuned to high frequencies are located rostrolaterally. Compared with the frequency representation along the basilar papilla and in other auditory structures, the PAF overrepresents low frequencies (< 4 kHz) that are important for barn owl vocalizations. Conversely, high frequencies (> or = 4 kHz), which are necessary for precise sound localization, are underrepresented relative to these more peripheral auditory structures. 5. There was considerable interindividual variability both in the relative magnification of different frequency ranges and in the orientation of the tonotopic map in the brain. 6. These results suggest that the barn owl PAF, like the mammalian primary auditory cortex, is a general processor of auditory information that is involved in the analysis of both the meaning (such as species-specific vocalizations) and the location of auditory stimuli. In addition, the high degree of interindividual variability in the representation of frequency information suggests that the barn owl PAF, like the mammalian auditory cortex, is subject to modification by sensory experience.

Acoustic Stimulation↗
PubMed1997

Neuronal encoding of sound direction in the auditory midbrain of the rainbow trout.

Acoustical stimulation causes displacement of the sensory hair cells relative to the otoliths of the fish inner ear. The swimbladder, transforming the acoustical pressure component into displacement, also contributes to the displacement of the hair cells. Together, this (generally) yields elliptical displacement orbits. Alternative mechanisms of fish directional hearing are proposed by the phase model, which requires a temporal neuronal code, and by the orbit model, which requires a spike density code. We investigated whether the directional selective response of auditory neurons in the midbrain torus semicircularis (TS; homologous to the inferior colliculus) is based on spike density and/or temporal encoding. Rainbow trout were mounted on top of a vibrating table that was driven in the horizontal plane to simulate sound source direction. Rectilinear and elliptical (or circular) motion was applied at 172 Hz. Generally, responses to rectilinear and elliptical/circular stimuli (irrespective of direction of revolution) were the same. The response of auditory neurons was either directionally selective (DS units, n = 85) or not (non-DS units, n = 106). The average spontaneous discharge rate of DS units was less than that of non-DS units. Most DS units (70%) had spontaneous activities < 1 spike per second. Response latencies (mode at 18 ms) were similar for both types of units. The response of DS units is transient (19%), sustained (34%), or mixed (47%). The response of 75% of the DS units synchronized to stimulus frequency, whereas just 23% of the non-DS responses did. Synchronized responses were measured at stimulus amplitudes as low as 0.5 nm (at 172 Hz), which is much lower than for auditory neurons in the medulla of the trout, suggesting strong convergence of VIIIth nerve input. The instant of firing of 42% of the units was independent of stimulus direction (shift <15 degrees), but for the other units, a direction dependent phase shift was observed. In the medial TS spatial tuning of DS units is in the rostrocaudal direction, whereas in the lateral TS all preferred directions are present. On average, medial DS units have a broader directional selectivity range, are less often synchronized, and show a smaller shift of the instant of firing as a function of stimulus direction than lateral DS units. DS response characteristics are discussed in relation to different hypotheses. We conclude that the results are more in favor of the phase model.

Air Sacs↗
PubMed1997

Responses of neurons in the inferior colliculus to binaural masking level difference stimuli measured by rate-versus-level functions.

The psychophysical detection threshold of a low-frequency tone masked by broadband noise is reduced by < or = 15 dB by inversion of the tone in one ear (called the binaural masking level difference: BMLD). The contribution of 120 low-frequency neurons (best frequencies 168-2,090 Hz) in the inferior colliculus (ICC) of the guinea pig to binaural unmasking of 500-Hz tones masked by broadband noise was examined. We measured rate-level functions of the responses to identical signals (So) and noise (No) at the two ears (NoSo) and to identical noise but with the signal inverted at one ear (NoS pi): the noise was 7-15 dB suprathreshold. The masked threshold was estimated by the standard separation, "D". The neural BMLD was estimated as the difference between the masked thresholds for NoSo and NoS pi. The presence of So and S pi tones was indicated by discharge rate increases in 55.3% of neurons. In 36.4% of neurons, the presence of So tones was indicated by an increase in discharge rate and S pi tones by a decrease. In 6.8% of neurons, both So and S pi tones caused a decrease in discharge rate. In only 1.5% of neurons was So indicated by a decrease and S pi by an increase in discharge rate. Responses to the binaural configurations were consistent with the neuron's interaural delay sensitivities; 34.4% of neurons showing increases in discharge rate to both So and S pi tones gave positive BMLDs > or = 3 dB (S pi tones were detected at lower levels than So), whereas 37.3% gave negative BMLDs > or = 3 dB. For neurons in which So signals caused an increase in the discharge rate and S pi a decrease, 72.7% gave positive BMLDs > or = 3 dB and only 4.5% gave negative BMLDs > or = 3 dB. The results suggest that the responses of single ICC neurons are consistent with the psychophysical BMLDs for NoSo versus NoS pi at 500 Hz, and with current binaural interaction models based on coincidence detection. The neurons likely to contribute to the psychophysical BMLD are those with BFs near 500 Hz, but detection of So and S pi tones may depend on different populations of neurons.

Animals↗
PubMed1997

Sensitivity to interaural temporal disparities of low- and high-frequency neurons in the superior olivary complex. I. Heterogeneity of responses.

Interaural temporal disparities (ITDs) are a cue for localization of sounds along the azimuth. Listeners can detect ITDs in the fine structure of low-frequency sounds and also in the envelopes of high-frequency sounds. Sensitivity to ITDs originates in the main nuclei of the superior olivary complex (SOC), the medial and lateral superior olives (MSO and LSO, respectively). This sensitivity is believed to arise from bilateral excitation converging on neurons of the MSO and ipsilateral excitation converging with contralateral inhibition on neurons of the LSO. Here we investigate whether the sensitivity of neurons in the SOC to ITDs can be adequately explained by one of these two mechanisms. Single and multiple units (n = 124) were studied extracellularly in the SOC of unanesthetized rabbits. We found units that were sensitive to ITDs in the fine structure of low-frequency (<2 kHz) tones and also units that were sensitive to ITDs in the envelopes of sinusoidally amplitude-modulated high-frequency tones. For both categories there were "peak-type" units that discharged maximally at a particular ITD across frequencies or modulation frequencies. These units were consistent with an MSO-type mechanism. There were also "trough-type" units that discharged minimally at a particular ITD. These units were consistent with an LSO-type mechanism. There was a general trend for peak-type units to be located in the vicinity of the MSO and for trough-type units to be located in the vicinity of the LSO. Units of both types appeared to encode ITDs within the estimated free-field range of the rabbit (+/-300 micros). Many units had varying degrees of irregularities in their responses, which manifested themselves in one of two ways. First, for some units there was no ITD at which the response was consistently maximal or minimal across frequencies. Instead there was an ITD at which the unit consistently responded at some intermediate level. Second, a unit could display considerable jitter from frequency to frequency in the ITD at which it responded maximally or minimally. Units with irregular responses had properties that were continuous with those of other units. They therefore appeared to be variants of peak- and trough-type units. The irregular responses could be modeled by assuming additional phase-locked inputs to a neuron in the MSO or LSO. The function of irregularities may be to shift the ITD sensitivity of a neuron without requiring changes in the anatomic delays of its inputs.

Acoustic Stimulation↗
PubMed1995

Physiology of primary saccular afferents of goldfish: implications for Mauthner cell response.

Mauthner cells receive neurally coded information from the otolith organs in fishes, and it is most likely that initiation and directional characteristics of the C-start response depend on this input. In the goldfish, saccular afferents are sensitive to sound pressure (< -30 dB re: 1 dyne cm-2) in the most sensitive frequency range (200 to 800 Hz). This input arises from volume fluctuations of the swimbladder in response to the sound pressure waveform and is thus nondirectional. Primary afferents of the saccule, lagena, and utricle of the goldfish also respond with great sensitivity to acoustic particle motion (< 1 nanometer between 100 and 200 Hz). This input arises from the acceleration of the fish in a sound field and is inherently directional. Saccular afferents can be divided into two groups based on their tuning: one group is tuned at about 250 Hz, and the other tuned between 400 Hz and 1 kHz. All otolithic primary afferents phaselock to sinusoids throughout the frequency range of hearing (up to about 2 kHz). Based on physiological and behavioral studies on Mauthner cells, it appears that highly correlated binaural input to the M-cell, from the sacculi responding to sound pressure, may be required for a decision to respond but that the direction of the response is extracted from small deviations from a perfect interaural correlation arising from the directional response of otolith organs to acoustic particle motion.

Animals↗
PubMed2001

A biologically motivated solution to the cocktail party problem.

We present a new approach to the cocktail party problem that uses a cortronic artificial neural network architecture (Hecht-Nielsen, 1998) as the front end of a speech processing system. Our approach is novel in three important respects. First, our method assumes and exploits detailed knowledge of the signals we wish to attend to in the cocktail party environment. Second, our goal is to provide preprocessing in advance of a pattern recognition system rather than to separate one or more of the mixed sources explicitly. Third, the neural network model we employ is more biologically feasible than are most other approaches to the cocktail party problem. Although the focus here is on the cocktail party problem, the method presented in this study can be applied to other areas of information processing.

Artificial Intelligence↗