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PubMed1993

Cochlear nerve fiber responses to amplitude-modulated stimuli: variations with spontaneous rate and other response characteristics.

1. Single-fiber responses to sinusoidally amplitude-modulated (AM) tones were recorded from the cochlear nerves of anesthetized guinea pigs. Stimuli were presented at the fiber's characteristic frequency (CF) and covered the intensity range between the fiber's minimum rate threshold and 90-100 dB SPL in 5- or 6-dB steps. The amount of modulation in each fiber's response and the average rate of the responses were quantified. The observed response modulation was compared with the modulation to be expected on the assumption that the instantaneous discharge rates varied with intensity in the same way that the average rates did (i.e., as predicted from each fiber's average-rate vs. level function). 2. The difference between the observed and expected response modulation varied widely across fibers. In most fibers' the responses to a limited range of stimulus intensities (typically between 20 and 30 dB above the fiber's rate threshold) were modulated far more than expected on the basis of their average rates, with responses to stimuli either above or below this range differing progressively less from expectation. Little or no response modulation was observed above approximately 70 dB SPL in these fibers. Other fibers exhibited response modulation that exceeded the expected modulation by smaller amounts, but maintained this modulation to much higher sound pressure levels. 3. The discrepancy between the observed and expected responses to AM stimuli also varied with the frequency of modulation (fm) within individual fibers. The discrepancies were least pronounced at low fms (e.g., 10 Hz) but became progressively larger as fm was increased to between 50 and 320 Hz (subject to the inter-fiber variations described in 2, above). 4. The AM response characteristics varied systematically with the fiber's spontaneous rate and other response characteristics (e.g., rate threshold, CF rate vs. level function type, and rapid adaptation characteristics). In particular, the most sensitive, high spontaneous rate fibers had responses that adapted rapidly after the onset of a stimulus, and showed the greatest enhancement of AM-related information at low-to-moderate stimulus intensities. However, these fibers appeared incapable of encoding AM-related information at high intensities, since their response rates "saturated" and their AM response enhancements diminished around 30 dB above threshold. In contrast, the less sensitive (i.e., higher threshold), lower spontaneous rate fibers showed less evidence of rapid adaptation near the onsets of their response, and lesser enhancements of the modulated responses predicted from their average-rate versus level functions.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
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

Effects of ear plugging on single-unit azimuth sensitivity in cat primary auditory cortex. I. Evidence for monaural directional cues.

1. Single-unit recordings were carried out in primary auditory cortex (AI) of barbiturate-anesthetized cats. Neurons, sensitive to sound direction in the horizontal plane (azimuth), were identified by their responses to noise bursts, presented in the free field, that varied in azimuth and sound pressure level (SPL). SPLs typically varied between 0 and 80 dB and were presented at each azimuth that was tested. Each azimuth-sensitive neuron responded well to some SPLs at certain azimuths and did not respond well to any SPL at other azimuths. This report describes AI neurons that were sensitive to the azimuth of monaurally presented noise bursts. 2. Unilateral ear plugging was used to test each azimuth-sensitive neuron's response to monaural stimulation. Ear plugs, produced by injecting a plastic ear mold compound into the concha and ear canal, attenuated sound reaching the tympanic membrane by 25-70 dB. Binaural interactions were inferred by comparing responses obtained under binaural (no plug) and monaural (ear plug) conditions. 3. Of the total sample of 131 azimuth-sensitive cells whose responses to ear plugging were studied, 27 were sensitive to the azimuth of monaurally presented noise bursts. We refer to these as monaural directional (MD) cells, and this report describes their properties. The remainder of the sample consisted of cells that either required binaural stimulation for azimuth sensitivity (63/131), because they were insensitive to azimuth under unilateral ear plug conditions or responded too unreliably to permit detailed conclusions regarding the effect of ear plugging (41/131). 4. Most (25/27) MD cells received either monaural input (MD-E0) or binaural excitatory/inhibitory input (MD-EI), as inferred from ear plugging. Two MD cells showed other characteristics. The contralateral ear was excitatory for 25/27 MD cells. 5. MD-E0 cells (22%, 6/27) were monaural. They were unaffected by unilateral ear plugging, showing that they received excitatory input from one ear, and that stimulation of the other ear was without apparent effect. On the other hand, some monaural cells in AI were insensitive to the azimuth of noise bursts, showing that sensitivity to monaural directional cues is not a property of all monaural cells in AI. 6. MD-EI cells (70%, 19/27) exhibited an increase in responsiveness on the side of the plugged ear, showing that they received excitatory drive from one ear and inhibitory drive from the other. MD-EI cells remained azimuth sensitive with the inhibitory ear plugged, showing that they were sensitive to monaural directional cues at the excitatory ear.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1993

Sensitivity of auditory nerve fibers to spectral notches.

1. Listeners use direction-dependent spectral cues introduced by the torso, head, and pinnae to localize the source of a sound in space. Among the prominent direction-dependent spectral features in the free field-to-eardrum transfer function are narrow regions of low acoustic energy referred to as spectral notches. In this paper, we studied the sensitivity of single auditory nerve fibers in the barbiturate-anesthetized cat to broadband noise that had been filtered by a function whose shape approximated natural notches in the free field-to-eardrum transfer function. 2. Two experimental paradigms were employed. The first was the repeated presentation of a burst of broadband noise filtered by the simulated-notch function. Center frequency of the notch was held constant at or around the fiber characteristic frequency (CF). We refer to this as a "stationary" notch stimulus. The second paradigm was the repeated presentation of a broadband noise that was constructed from noise segments, each filtered by the simulated notch, whose CF was incremented and then decremented in a systematic way. We refer to this as a "moving" notch stimulus. Results from these two paradigms were compared with respect to notch detection. 3. Data were obtained from 161 single auditory nerve fibers having CFs ranging from 0.4 to 40 kHz. Most fibers studied had CFs > 5 kHz, and they detected the presence of the spectral notch in an intensity- and frequency-dependent manner. Each fiber responded vigorously to the presence of broadband noise. When the CF of the notch encroached on the response area of the fiber, there was a demonstrable reduction in discharge rate. The greatest reduction in discharge rate occurred when the notch was centered at the fiber's CF and when the level of the notch signal was some 15-55 dB above the fiber's noise threshold. There was close association in the frequency-intensity plane between the position of the most effective notch and the fiber's threshold tuning curve. 4. For high-spontaneous rate fibers, a moving-notch stimulus, but not a stationary one, reduced the discharge below the spontaneous rate at and in the immediate vicinity of the most effective notch frequency. This increases sensitivity to a spectral notch and suggests a mechanism by which localization ability is enhanced when there is relative motion between a sound source and the head.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1993

Virtual-space receptive fields of single auditory nerve fibers.

1. Sounds reaching the tympanic membranes are first modified by the acoustic properties of the torso, head, and external ear. For certain frequencies in the incident sound there results a complex, direction-dependent spatial distribution of sound pressure at the eardrum such that, within a sound field, localized areas of pressure maxima are flanked by areas of pressure minima. Listeners may use these spatial maxima and minima in localizing the source of a sound in space. The results presented describe how information about this spatial pressure pattern is transmitted from the cochlea to the central auditory system via single fibers of the auditory nerve. 2. Discharges of single fibers of the auditory nerve were studied in Nembutal-anesthetized cats [characteristic frequencies (CFs) ranged from 0.4 to 40 kHz]. Click stimuli were derived from sound-pressure waveforms that were generated by a loudspeaker placed at 1,800 locations around the cat's head and recorded at the tympanic membrane with miniature microphones. Recorded signals were converted to acoustic stimuli and delivered to the ear via a calibrated and sealed earphone. The full complement of signals is referred to as "virtual acoustic space," and the spatial distribution of discharges to this array of signals is referred to as a "virtual-space receptive field" (VSRF). 3. Fibers detect both pressure maxima and pressure minima in virtual acoustic space. Thus VSRFs take on complex shapes. 4. VSRFs of fibers of the same or similar CF having low spontaneous rates had the same overall pattern as those from high-spontaneous rate (HSR) fibers. For HSR fibers, the VSRF is obscured by the high background spike activity. 5. Comparison of the VSRF and isolevel contour maps of the stimulus derived at various frequencies revealed that auditory nerve fibers most accurately extract spectral information contained in the stimulus at a frequency close to or slightly higher than CF.

Animals↗
PubMed1993

Neural encoding of single-formant stimuli in the cat. I. Responses of auditory nerve fibers.

1. We have studied auditory responses to a set of speech-related narrowband sounds, single-formant stimuli (SFSs), in populations of auditory nerve fibers (ANFs). An analytic method was developed to extract the envelope of temporal discharge patterns of the ANF responses to nonsinusoidally modulated stimuli, whose spectra have multiple clusters of components. Such responses are often encountered in the auditory system when complex stimuli are used and have traditionally been studied by analyzing the fundamental component of the responses. 2. The envelope modulation in the SFSs is shown to be represented by the response patterns of ANFs. When the whole ANF population is considered, the information on modulation in stimulus envelope does not disappear at the highest sound level tested at all best frequencies (BFs) we studied (1-10 kHz). The representation is the best at medium sound levels and degrades at high sound levels. Low/medium-spontaneous rate (SR) ANFs showed greater envelope modulation in their responses at high sound levels than do high-SR ANFs. The quality of the representation at high sound levels is, on average, proportional to BF threshold of an ANF. On the basis of populations of ANFs with all SRs, the envelope modulation in the SFSs is represented over a wide range of sound levels. 3. We found that low-BF ANFs differ from high-BF ANFs in representing envelope modulation in the SFSs. For ANFs with BFs less than approximately 6 kHz, information on stimulus envelope is not only contained in spectral components near direct current but also in components at the vicinities of frequencies equal to BF and its multiples. In fact, for ANFs with BFs < 3 kHz, the contribution from spectral components centered at BF to overall response modulation is greater than that from spectral components near direct current. These findings indicate that, by using measures solely based on the fundamental component, the amount of modulation in the responses to narrowband stimuli is underestimated for low-BF ANFs. 4. Off-BF stimulation of ANFs with SFSs was found to result in increased envelope modulation in responses at high sound levels. The further away the stimulus is centered relative to unit BF, the greater the modulation it induces, provided that the stimulus is capable of exciting the unit. An SFS centered as close as 15% off unit BF can produce a significant increase in the modulation of responses at very high sound levels.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1993

Antimasking effects of the olivocochlear reflex. II. Enhancement of auditory-nerve response to masked tones.

1. The antimasking effects of olivocochlear (OC) efferent feedback were studied in anesthetized or decerebrate cats by comparing responses of single auditory-nerve fibers (ANFs) to tone bursts in continuous masking noise seen with and without addition of a moderate-level contralateral noise known to activate the OC reflex. Responses were measured as a function of tone-burst intensity, tone-burst frequency, and masker intensity and were analyzed so as to allow quantitative estimates of the detectability of the tone bursts against the noise background. 2. Addition of the contralateral OC elicitor both increased the maximum discharge rates to the masked tone bursts and decreased the rates to the ipsilateral masker. The rate increases to the tone bursts could be explained on the basis of a decrease in adaptation caused by decreasing the steady response to the masker. The result is a steepening of the rate-versus-level function for masked tone bursts and a concomitant increase in the estimated discriminability of small increments of tone-burst intensity. 3. For tone bursts at the fiber's characteristic frequency (CF), the OC effects on detection threshold for the masked tone bursts depended on masker level, with small increases in threshold for low masker levels and somewhat larger decreases in threshold for higher masker levels. For tone bursts below CF, OC effects, when present, always decreased the detection threshold. 4. The largest antimasking effects were seen for fibers with CFs between 6 and 12 kHz and for masker levels within 20 dB of the fiber's threshold to the masker. These trends appeared to hold for fibers of all spontaneous rates (SRs). 5. Enhancement of the response to unmasked tone bursts and concomitant decrease in the "spontaneous rate" was elicited by OC activation in fibers if threshold sensitivity approached -10 dB SPL. This "enhancement-in-quiet" appears to arise when an animal-generated noise produces a continuous response (in the absence of purposely applied sound) that is suppressed by OC activity. This finding raises questions as to the range of "true" spontaneous rates in the cat. 6. The results highlight two important distinctions between the effects of OC feedback in quiet versus those in noise. In quiet, the effects are predominately suppressive and are restricted to stimuli at frequencies near a fiber's CF and at intensities within its dynamic range. In continuous background noise, the OC reflex can enhance the responses to transient stimuli. Such effects are seen throughout the fiber's response area.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Monaural and binaural spectrum level cues in the ferret: acoustics and the neural representation of auditory space.

1. The role of the structures of the outer ear in producing monaural and binaural spectral cues to sound location was examined acoustically in the ferret. A probe microphone was introduced across the wall of the external auditory canal and its responses to digitally constructed wideband signals were recorded for a large number of free field locations. 2. In the intact animal the patterns of both monaural and binaural cues were asymmetrical for horizontal locations about the interaural axis. For anterior sound locations the monaural transformations demonstrated relative gains at middle and high frequencies and a location-dependent frequency notch. Changing elevation resulted in variations in the corner frequencies of these spectral features. Additionally, there was greater front-back asymmetry in the binaural spectral cues for locations in lateral space when compared with locations near the midline. 3. Surgical removal of the pinna and concha (pinnectomy) eliminated all the major front-back asymmetrical features in the horizon monaural and binaural spectral transformations as well as the elevation-dependent variations in the monaural spectra. Thus the residual transformations were ambiguous for sound locations in lateral space, resulting in "cones of confusion" centered on the interaural axis. 4. These cues were reflected in the topographic representation of auditory space in the deeper layers of the superior colliculus (SC). Previous studies have shown that spatial tuning at near-threshold sound levels is based on monaural pinna cues, whereas binaural inputs are utilized at higher levels that stimulate both ears. In the intact ferret we examined statistically the topography of the representation of sound azimuth for near-threshold and suprathreshold stimuli and the alignment of the auditory and visual representations in the SC. The distributions of auditory best positions within the SC for near- and suprathreshold stimulus levels were statistically indistinguishable, suggesting that both monaural and binaural cues are integrated in this neural representation of space. 5. Pinnectomy resulted in a large increase in the number of auditory units that responded best to two distinct locations in space. One lobe of the response was tuned appropriately in terms of the position of the unit within the SC, demonstrating that the residual acoustical cues are sufficient for the construction of a topographic representation of auditory space. However, the second region of space, thereby producing an ambiguous representation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Enhancement of neural synchronization in the anteroventral cochlear nucleus. II. Responses in the tuning curve tail.

1. Discharges of neurons in the peripheral auditory system contain information about the temporal features of acoustic stimuli. Phase-locking of neurons in the anteroventral cochlear nucleus (AVCN) is usually reported to be less robust than in auditory nerve (AN) fibers, which provide their major input. In a companion paper we reported that some cells in AVCN of the cat show enhanced phase-locking compared with the AN when stimulated at the frequency to which they are most sensitive [characteristic frequency (CF)]. We called neurons "high-sync" when they showed vector strengths (R, a measure of phase-locking) > or = 0.9. Here we report phase-locking properties to stimuli at frequencies below CF. 2. Horseradish peroxidase-filled glass micropipettes or metal microelectrodes were inserted into the trapezoid body (TB), which is the large output tract of the AVCN. Acoustically driven fibers were classified on the basis of the shape of the poststimulus time (PST) histograms to short tone bursts at CF. We then presented low-frequency tones of increasing SPL and determined the maximum R value at 500 Hz (R500) for each fiber. Using the same experimental protocol we studied phase-locking in the ANs of two animals because maximal R values at the tuning curve tail have not been reported for AN fibers. 3. Although phase-locking in AN fibers is usually assumed to be independent of CF, we found that fibers with CF > 2 kHz tended to have higher R500 values than fibers with CF < or = 2 kHz. Moreover, R500 was > or = 0.9 in 20% (42 of 196) of the fibers studied and could be as high as 0.95. This population of fibers was defined as having "high-sync tails" and consisted almost entirely of fibers with low or medium spontaneous rate. 4. High-CF TB fibers stimulated at 500 Hz showed very high phase-locking. High-sync tails (R500 > or = 0.9) were found in 41 of 70 TB fibers. For a subset of these fibers (1/3 in total: 23 of 70) phase-locking was higher than is ever observed in the AN (R500 > or = 0.95); these fibers were defined as showing synchronization "enhancement." Virtually all fibers showing synchronization enhancement had primary-like-with-notch (PLN) PST histograms. Chopper and primary-like fibers showed high-sync tails for CFs > 3 kHz. 5. Synchronization filter functions were obtained for high-CF AN fibers by determining maximum synchronization for a range of stimuli below CF.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Encoding of amplitude modulation in the cochlear nucleus of the cat.

1. Amplitude modulation (AM) is a pervasive property of acoustic communication systems. In the present study we investigate neural temporal mechanisms in the auditory nerve and cochlear nuclei of the pentobarbital sodium-anesthesized cat associated with the neural coding of 100% AM tones, both in quiet and in the presence of wideband, quasi-flat-spectrum noise. The AM carrier frequency was set to the neuron's characteristic frequency (CF) and the sound pressure level (SPL) of acoustic stimuli was varied over a wide dynamic range of intensities (< or = 40 dB). The temporal AM-encoding capability of auditory neurons was measured by computing the synchronization coefficient (SC) of the neural response to the signal's modulation and carrier frequency. The temporal modulation transfer function (tMTF) of a neuron was then computed by measuring the SC of the response to signals of variable fmod (50-2550 Hz). 2. Neurons in the cochlear nuclei synchronize on average more highly to the modulation frequency than fibers of comparable CF, threshold, and spontaneous rate in the auditory nerve. The disparity in performance is greatest at high SPLs and low signal-to-noise ratios. However, there is a significant degree of diversity in AM-encoding capability among neurons in both the cochlear nuclei and auditory nerve. Among auditory nerve fibers (ANFs), low- and medium-spontaneous-rate (SR) units (SR < 18 spike/s) phase-lock with greater precision than comparable high-SR units at any given frequency, particularly at moderate to high SPLs, consistent with previous studies. 3. The phase-locking capabilities of neurons in the cochlear nucleus are considerably more variable than in the auditory nerve. Moreover, the variability itself depends on two distinct measures of phase-locking performance. Most ANFs are capable of phase-locking to frequencies as high as 3-4 kHz. In the cochlear nucleus many unit types do not phase-lock to modulation frequencies > 1 kHz. As a result, phase-locking performance is measured on the basis of two parameters, maximum synchronization, irrespective of stimulus frequency, and the upper frequency limit for significant phase-locking. 4. Cochlear nucleus neurons may be divided into three distinct groups on the basis of maximum synchronization capability. In group 1 are the primary-like (PL) units of the anteroventral division, whose phase-locking capabilities are comparable with those of high-SR ANFs.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Influence of stimulus level on acoustic motion-direction sensitivity in barn owl midbrain neurons.

1. Many neurons in the owl's midbrain are able to discriminate by their responses the motion of an acoustic source in one direction, the preferred direction, from the motion of the source in the opposite direction, the null direction. This differential response can be understood by the action of an acoustic motion detector. However, the motion detector would work in an effective way only if the motion direction-sensitive responses of a cell were largely independent of stimulus level. 2. We investigated the level dependence of acoustic motion sensitivity in midbrain neurons with standard extracellular recording techniques. We used an apparent motion paradigm with sounds moving clockwise or counterclockwise in the horizontal plane or stationary sounds for stimulation from free-field speakers. In most cases we added a constant level of background noise to the stimuli having a variable level. Thus we could analyze the responses as a function of the signal-to-noise ratio. 3. Moving and stationary sounds yielded similar response thresholds and similar dynamic ranges. 4. When the background level was changed the response threshold changed accordingly, so that the response threshold always occurred at a constant signal-to-noise ratio independent of background level. 5. The preferred direction of all cells was independent of stimulus level and some 22 percent of the cells showed a directional bias of their response at threshold. In no cell was a change in the preferred direction observed when stimulus level was varied. Typically, however, the motion direction sensitivity, measured by a signed directionality index, was not maximal at threshold. However, the dynamic range, i.e., the range of stimulus levels over which motion direction sensitivity changed, was < or = 20 dB in most cases. Therefore these cells are indeed able to encode motion direction largely independently of stimulus level. 6. We discuss the data within the framework of the acoustic motion detector mentioned above and with respect to earlier reports on acoustic motion sensitivity in the owl and other animals.

Animals↗
PubMed1994

The roles of GABAergic and glycinergic inhibition on binaural processing in the dorsal nucleus of the lateral lemniscus of the mustache bat.

1. We studied the monaural and binaural response properties of 99 neurons in the dorsal nucleus of the lateral lemniscus (DNLL) of the mustache bat before and during the iontophoretic application of antagonists that blocked gamma-aminobutyric acid-A (GABAA) receptors (bicuculline) or glycine receptors (strychnine). All cells were driven by monaural stimulation of the contralateral ear, whereas monaural stimulation of the ipsilateral ear never evoked discharges. The binaural properties of 81 neurons were determined by holding the intensity constant at the contralateral ear and presenting a variety of intensities to the ipsilateral ear. This procedure generated interaural intensity disparity (IID) functions and allowed us to determine the effect of ipsilaterally evoked inhibition on a constant excitatory drive evoked by the contralateral ear. 2. One of the main findings is that the IID functions in the majority of DNLL neurons were not affected by application of either strychnine or bicuculline. Blocking glycinergic inhibition with strychnine had no effect on the IID functions in 75% of the cells studied. However, strychnine did change the IID functions in approximately 25% of the DNLL population. In those cells glycinergic inhibition appeared to be partially, or, in a few cases, entirely responsible for the ipsilaterally evoked spike suppression. In contrast, blocking GABAergic inhibition with bicuculline had no discernible effect on the ipsilaterally evoked spike suppression in any of the excitatory/inhibitory cells that we recorded. GABAergic inhibition, therefore, plays no role in the formation of IID functions of neurons in the DNLL. Furthermore, the results suggest that glycinergic inhibition also does not contribute to the suppression of spikes evoked by stimulation of the contralateral ear in the vast majority of DNLL neurons. 3. Although the majority of IID functions were not influenced when either GABAergic or glycinergic innervation was blocked, ipsilateral stimulation alone evoked both a glycinergic and GABAergic inhibition in most DNLL cells. These inhibitory events were demonstrated in 18 other cells by evoking discharges with the iontophoretic application of glutamate. Stimulating the ipsilateral ear alone under these conditions caused a suppression of the glutamate-evoked discharges. Furthermore, the spike suppression persisted for a period of time that was longer than the duration of the tone burst at the ipsilateral ear. 4. The application of bicuculline or strychnine had different effects on the glutamate-elicited spikes. Bicuculline reduced the duration of the inhibition, and it was always the latter portion of the inhibition that was abolished by bicuculline. In more than half of the cells studied strychnine also reduced the duration of the inhibition.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

GABA and glycine have different effects on monaural response properties in the dorsal nucleus of the lateral lemniscus of the mustache bat.

1. We studied the monaural response properties of 81 neurons in the dorsal nucleus of the lateral lemniscus (DNLL) of the mustache bat before and during the iontophoretic application of antagonists that blocked gamma-aminobutyric acid-A (GABAA) receptors (bicuculline) or glycine receptors (strychnine). The main finding is that GABAergic inhibition had substantial effects, whereas glycine had little or no effect on the activity evoked by contralateral stimulation. 2. Before the application of drugs, the monaural response properties of DNLL cells were characterized by two main features. The first was that the majority (86%) of neurons had monotonic rate-intensity functions, whereas only 14% had weakly nonmonotonic functions. The second was that most (66%) neurons displayed some form of chopping response pattern, in which there was a regular interval between discharges that was unrelated to the period of the tone burst frequency. 3. Bicuculline had two major effects on the majority of DNLL cells. It caused large increases in spike counts and changes in temporal discharge patterns. In 38 of 47 cells (81%) bicuculline changed the temporal discharge patterns into a sustained chopper pattern. In addition, the duration of the discharge train continued for a period of time longer than the duration of the tone burst in many but not all neurons. Prolonged firing of this sort was rarely seen in the predrug condition. Furthermore, in a few cells bicuculline caused a decrease in the interspike interval as well as a lengthening of the discharge train. 4. Blocking glycine, in contrast, caused either small increases in spike count or no increase at all and did not affect the temporal discharge patterns in the majority (87%) of neurons. 5. In most DNLL cells the shapes of the rate-intensity functions were virtually the same before and during the application of either antagonist. The rate-intensity functions of 91% of the cells were unaffected by bicuculline and 98% were unaffected by strychnine. 6. Blocking either GABAergic inhibition or glycinergic inhibition had no effect on discharge latency in the vast majority of DNLL cells. In a few neurons application of bicuculline or strychnine had a small influence and caused discharge latency to decrease by < or = 1 ms. 7. These results show that the excitation from stimulation of the contralateral ear evokes a sustained chopping discharge pattern in the vast majority of DNLL neurons. The sustained chopping response is changed into another discharge pattern by the GABAergic innervation that is also evoked by stimulation of the contralateral ear.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Effects of ear plugging on single-unit azimuth sensitivity in cat primary auditory cortex. II. Azimuth tuning dependent upon binaural stimulation.

1. Single-unit recordings were carried out in primary auditory cortex (AI) of barbiturate-anesthetized cats. Observations were based on a sample of 131 high-best-frequency (> 5 kHz), azimuth-sensitive neurons. These were identified by their responses to a set of noise bursts, presented in the free field, that varied in azimuth and sound-pressure level (SPL). Each azimuth-sensitive neuron responded well to some levels at certain azimuths, but did not respond well to any level at other azimuths. 2. Unilateral ear plugging was used to infer each neuron's response to monaural stimulation. Ear plugs, produced by injecting a plastic ear mold compound into the external ear, attenuated sound reaching the tympanic membrane by 25-70 dB. The azimuth tuning of a large proportion of the sample (62/131), referred to as binaural directional (BD), was completely dependent upon binaural stimulation because with one ear plugged, these cells were insensitive to azimuth (either responded well at all azimuths or failed to respond at any azimuth) or in a few cases exhibited striking changes in location of azimuth function peaks. This report describes patterns of monaural responses and binaural interactions exhibited by BD neurons and relates them to each cell's azimuth and level tuning. The response of BD cells to ear plugging is consistent with the hypothesis that they derive azimuth tuning from interaural level differences present in noise bursts. Another component of the sample consisted of monaural directional (27/131) cells that derived azimuth tuning in part or entirely from monaural spectral cues. Cells in the remaining portion of the sample (42/131) responded too unreliably to permit specific conclusions. 3. Binaural interactions were inferred by statistical comparison of a cell's responses to monaural (unilateral plug) and binaural (no plug) stimulation. A larger binaural response than either monaural response was taken as evidence for binaural facilitation. A smaller binaural than monaural response was taken as evidence for binaural inhibition. Binaural facilitation was exhibited by 65% (40/62) of the BD sample (facilitatory cells). Many of these exhibited mixed interactions, i.e., binaural facilitation occurred in response to some azimuth-level combinations, and binaural inhibition to others. Binaural inhibition in the absence of binaural facilitation occurred in 35% (22/62) of the BD sample, a majority of which were EI cells, so called because they received excitatory (E) input from one ear (excitatory ear) and inhibitory (I) input from the other (inhibitory ear). One cell that exhibited binaural inhibition received excitatory input from each ear.(ABSTRACT TRUNCATED AT 400 WORDS)

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↗
PubMed1994

Auditory tuning for spatial cues in the barn owl basal ganglia.

1. The basal ganglia are known to contribute to spatially guided behavior. In this study, we investigated the auditory response properties of neurons in the barn owl paleostriatum augmentum (PA), the homologue of the mammalian striatum. The data suggest that the barn owl PA is specialized to process spatial cues and, like the mammalian striatum, is involved in spatial behavior. 2. Single- and multiunit sites were recorded extracellularly in ketamine-anesthetized owls. Spatial receptive fields were measured with a free-field sound source, and tuning for frequency and interaural differences in timing (ITD) and level (ILD) was assessed using digitally synthesized dichotic stimuli. 3. Spatial receptive fields measured at nine multiunit sites were tuned to restricted regions of space: tuning widths at half-maximum response averaged 22 +/- 9.6 degrees (mean +/- SD) in azimuth and 54 +/- 22 degrees in elevation. 4. PA sites responded strongly to broadband sounds. When frequency tuning could be measured (n = 145/201 sites), tuning was broad, averaging 2.7 kHz at half-maximum response, and tended to be centered near the high end of the owl's audible range. The mean best frequency was 6.2 kHz. 5. All PA sites (n = 201) were selective for both ITD and ILD. ITD tuning curves typically exhibited a single, large "primary" peak and often smaller, "secondary" peaks at ITDs ipsilateral and/or contralateral to the primary peak. Three indices quantified the selectivity of PA sites for ITD. The first index, which was the percent difference between the minimum and maximum response as a function of ITD, averaged 100 +/- 29%. The second index, which represented the size of the largest secondary peak relative to that of the primary peak, averaged 49 +/- 23%. The third index, which was the width of the primary ITD peak at half-maximum response, averaged only 66 +/- 35 microseconds. 6. The majority (96%; n = 192/201) of PA sites were tuned to a single "best" value of ILD. The widths of ILD tuning curves at half-maximum response averaged 24 +/- 9 dB. 7. On average, sound level had no effect on a site's best ITD or best ILD nor did it affect ITD tuning widths. ILD tuning widths did, however, tend to increase slightly with sound level (average effect was 0.1 dB ILD/dB). 8. Most PA sites responded best to contralateral-ear leading ITDs with a majority being tuned to ITDs near 0 microsecond (corresponding to sound-source locations just contralateral to the midline).(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↗
PubMed1994

Functional organization of sound direction and sound pressure level in primary auditory cortex of the cat.

1. The functional organization of neuronal tuning to the azimuthal location and sound pressure level (SPL) of noise bursts was studied in high-frequency primary auditory cortex (AI) of barbiturate-anesthetized cats. Three data collection strategies were used to map neural responses: 1) electrode penetrations oriented normal to the cortical surface provided information on the radial organization of neurons' responses; 2) neurons' responses were examined at a few points in the middle cortical layers in multiple normal penetrations across AI to produce fine-grain maps of azimuth and level selectivity; and 3) electrode penetrations oriented tangential to the cortical surface provided information on neurons' responses along the isofrequency dimension. 2. An azimuth-level data set was obtained for each single- or multiple- (multi-) unit recording; this consisted of responses to noise bursts at five SPLs (0-80 dB in 20-dB steps) from seven azimuthal locations in the frontal hemifield (-90 to +90 degrees in 30 degrees steps; 0 degree elevation). An azimuth function was derived from these data by averaging response magnitude over all SPLs at each azimuth tested. A preferred azimuth range (PAR; range of azimuths over which the response was > or = 75% of maximum) was calculated from the azimuth function and provided a level-independent measure of azimuth selectivity. Each PAR was assigned to one of four azimuth preference categories (contralateral-, midline-, ipsilateral-preferring, or broad/multipeaked) according to its location and extent. A level function obtained from the data set (responsiveness averaged over all azimuths) was classified as monotonic if it showed a decrease of < or = 25% (relative to maximum) at the highest SPL tested (usually 80 dB), and nonmonotonic if it showed a decrease of > 25%. The percentage reduction in responsiveness, relative to maximum, at the highest level tested (termed nonmonotonic strength) and the preferred level range (PLR; range of SPLs over which responsiveness was > or = 75% of maximum) of each response was also determined. 3. Normal penetrations typically showed a predominance of one azimuth preference category and/or level function type. The majority of penetrations (26/36: 72.2%) showed statistically significant azimuth preference homogeneity, and approximately one-half (17/36: 47.2%) showed significant level function type homogeneity. Over one-third (13/36) showed significant homogeneity for both azimuth preference and level function type. 4. Mapping experiments (n = 4) sampled the azimuth and level response functions at two or more depths in closely spaced normal penetrations that covered several square millimeters of AI.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗