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T C Yin

Publications and source records attributed to T C Yin.

At least 37 records · Page 2Linked to original sources

Visuomotor interactions in responses of neurons in the middle and lateral suprasylvian cortices of the behaving cat.

We studied visuomotor processing in the middle (MS) and lateral suprasylvian (LS) cortices of the alert cat by making single cell recordings while the cat was working in a behavioral task requiring visual fixation and visually guided eye movements. We found responses with three different components: visual sensory, saccade-related motor, and fixation. Some cells exhibited purely visual responses and all of their activity during visuomotor tasks could be attributed to the sensory aspects of the task. Other cells showed no sensory response properties, but discharged in relation to the saccadic eye movements that the cat made to visual targets. A smaller number of fixation cells displayed increased discharge when the cat fixated a target light and usually only when that target was in a particular region of the visual field. These response components could be present in a variety of combinations in different cells, of which the largest proportion combined visuomotor responses and could take five general forms: simple visuomotor, saccadic enhanced, visually triggered movement (VTM), enhanced VTM, and disenhanced. Simple visuomotor responses had both a visual and saccade-related component. Saccadic enhanced responses had a visual response to the appearance of a spot in the cell's receptive field that became enhanced when the cat subsequently made a saccade to that spot. The VTM responses were synchronized better to the visual stimulus than to the saccade, but they also exhibited properties expected of motor responses. The last two classes of visuomotor responses were rare: one we termed enhanced VTM and the other disenhanced. Cells could combine different visuomotor response components or even sensory, saccade-related and fixation responses in different combinations for different directions of eye movements. Generally, the timing of the saccade-related responses occurred too late to play a role in the initiation of saccades: most (83%) saccade-related responses occurred between 40 ms before to 80 ms after the onset of the eye movement. Cells of all different types could be found in both the MS and LS areas, though in general the responses in LS were more sensory in nature while those in MS were more closely related to the eye movement. About a quarter of the cells were unresponsive during any aspect of our tasks.

Animals↗

Responses to amplitude-modulated tones in the auditory nerve of the cat.

Sinusoidally amplitude-modulated (AM) tones are frequently used in psychophysical and physiological studies, yet a comprehensive study on the coding of AM tones in the auditory nerve is lacking. AM responses of single auditory-nerve fibers of the cat are studied, systematically varying modulation depth, frequency, and sound level. Synchrony-level functions were nonmonotonic with maximum values that were inversely correlated with spontaneous rate (SR). In most fibers, envelope phase-locking showed a positive gain. Modulation transfer functions were uniformly low pass. Their corner frequency increased with characteristic frequency (CF), but changed little for CFs above 10 kHz. The highest modulation frequencies to which phase locking occurred were more than 0.8 oct lower than the highest frequencies to which phase locking to pure tones occurs. Cumulative, or unwrapped, phase increased linearly with modulation frequency: The slope was inversely related to CF, and slightly higher than group delays reported for pure tones. High SR, low CF fibers showed the poorest envelope phase locking. In some low CF fibers, phase locking increased at high levels, associated with "peak-splitting" phenomena. Changes in average rate due to modulation were small, and could be enhancement or suppression.

Acoustics↗

Projections of physiologically characterized globular bushy cell axons from the cochlear nucleus of the cat.

We made intraaxonal recordings from 30 individual globular bushy cell axons in the trapezoid body of the cat using HRP-filled glass microelectrodes. With subsequent HRP injection, we determined their axonal projection patterns. For cells with characteristic frequencies (CFs) above 3 kHz, short-tone peristimulus time histograms (PSTHs) at CF were typically primarylike at low tone intensities and primarylike with notch (PLN) or onset with low sustained activity (OL) at higher stimulus levels. Cells with CFs between 1 and 3 kHz showed the same response features with the spikes in the sustained region of the response phase-locked to the stimulus tone. Cells with CFs below 1 kHz showed phase-locked PSTHs with exceptionally high levels of synchrony compared to eighth nerve fibers with comparable CFs. This exceptional phase-locking was also noted when cells with CFs of 1-3 kHz were presented with tones below 1 kHz. Although the globular bushy cell axons were not completely filled from the soma of origin to terminal fields in the contralateral brainstem, a number of consistent anatomical features were distinguished in the population. All but one of the myelinated axons crossed the midline in the middle, large fiber component of the trapezoid body. Ipsilaterally, the axon always gave off from one to four collateral branches whose major targets were the posterior periolivary nucleus (PPO) and the lateral nucleus of the trapezoid body (LNTB). Minor termination sites for ipsilateral collateral branches were the dorsolateral periolivary nucleus (DLPO) and the lateral superior olive (LSO). Contralaterally the axon gave rise to one or two calyces of Held in the medial nucleus of the trapezoid body (MNTB). Three other major collateral branches arose from the contralateral axon and innervated a consistent set of areas. One headed caudally to innervate an area just ventromedial to the facial nucleus. Another followed the sixth nerve dorsally to innervate the dorsomedial periolivary nucleus (DMPO). A third collateral headed rostrally toward the ventral nucleus of the lateral lemniscus (VNLL), giving off occasional small sidebranches. Although each injected axon gave rise to a collateral that innervated the MNTB, it did not necessarily give rise to all three of the other collateral branches.

Action Potentials↗

Dendritic and axonal morphology of HRP-injected neurons in the inferior colliculus of the cat.

The dendritic and axonal morphology of neurons in the inferior colliculus of the cat was investigated after intracellular injection of HRP, in vivo. All injected axons gave off local collaterals, and most showed a widespread distribution and lacked a specific orientation. In contrast, the dendrites of injected neurons were distinguished by their degree of orientation and the direction of the longest axis of orientation. Dendrites showed a high, moderate, or low degree of orientation. Most highly oriented cells had their longest axis in the rostrocaudal direction with fewer in the mediolateral direction. In the central nucleus, only the rostrocaudally oriented cells correspond to the disc-shaped cells identified in Golgi preparations. Unlike most cells in our sample, the two cells that were disc-shaped had axons that were parallel to the orientation of the dendritic tree. In the dorsal cortex, rostrocaudally oriented cells also were found, but they had unoriented axons. In both the central nucleus and dorsal cortex, cells with a mediolateral axis of orientation or no specific orientation correspond to stellate cells and had axons with widespread local collaterals. These results suggest that an extensive network of local axon collaterals may contribute to neural processing within the inferior colliculus. In the central nucleus, local axons may establish connections within or across the fibrodendritic laminae. In the dorsal cortex, the local and afferent axons may form a complex reticular network. Finally, some injected cells had axons terminating locally and also entering the brachium of the inferior colliculus. This suggests that cells in the inferior colliculus may function as both interneurons and projection neurons.

Animals↗

Interaural time sensitivity in the inferior colliculus of the albino cat.

Anatomical studies of the Creel albino cat have demonstrated a pronounced atrophy of cells in the medial superior olive, a structure thought to be important for the detection of interaural time differences (ITDs). We looked for physiological abnormalities in the binaural interaction of cells in three albino cats by recording from single cells in the central nucleus of the inferior colliculus to ITDs of tones and noise. We found that the sensitivity to ITDs of tones and noise was somewhat diminished in the albino cats as compared to normally pigmented cats, though this deficit was only evident when a population of cells was examined. The range of sensitivity of individual cells for both tones and noise was the same in albinos and pigmented animals. Our anatomical measurements showed a smaller reduction in cross-sectional area of cells in the medial superior olive than that reported earlier, and the cell bodies in the medial superior olive of the albinos were less elongated than in normal cats.

Albinism↗

Interaural time sensitivity in medial superior olive of cat.

1. We studied the sensitivity of cells in the medial superior olive (MSO) of the anesthetized cat to variations in interaural phase differences (IPDs) of low-frequency tones and in interaural time differences (ITDs) of tones and broad-band noise signals. Our sample consisted of 39 cells histologically localized to the MSO. 2. All but one of the cells had characteristic frequencies less than 3 kHz, and 79% were sensitive to ITDs and IPDs. More than one-half (56%) of the cells responded to monaural stimulation of either ear, and both the binaural and monaural responses were highly phase locked. All of the cells that were sensitive to IPDs and monaurally driven by either ear responded in accord with that predicted by the coincidence model of Jeffress, as judged by comparisons of the phases at which the monaural and binaural responses occurred. The optimal IPDs were tightly clustered between 0.0 and 0.2 cycles. Most cells exhibited facilitation of the response at favorable ITDs and inhibition at unfavorable ITDs compared with the monaural responses. 3. Cells in the MSO exhibited characteristic delay, as judged by a linear relationship between the mean interaural phase and stimulating frequency. Characteristic phases were clustered near 0 indicating the most cells responded maximally when the two input tones were in phase. With the use of the binaural beat stimulus we found no differential selectivity for either the direction or speed of interaural phase changes. 4. The cells were also sensitive to ITDs of broad-band noise signals. The ITD curve in response to broad-band noise was similar to that predicted by the composite curve, which was calculated by linearly summating the tonal responses over the frequencies in the response area of the cell. Most (93%) of the peaks of the composite curves were between 0 and +400 microseconds, corresponding to locations in the contralateral sound field. Moreover, computer cross correlations of the monaural spike trains were similar to the ITD curve generated binaurally for both correlated and uncorrelated noise signals to the two ears. Thus our data suggest that the cells in the MSO behave much like cross-correlators. 5. By combining data from different animals and lcoating each cell on a standard MSO, we found evidence for a spatial map of ITDs across the anterior-posterior (A-P) axis of the MSO.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Responses of low-frequency cells in the inferior colliculus to interaural time differences of clicks: excitatory and inhibitory components.

1. We studied extracellular responses of low-frequency cells in the central nucleus of the inferior colliculus (ICC) to interaural time differences (ITDs) of clicks and compared their responses to ITDs of noise and tones. Most cells that displayed sensitivity to ITDs of clicks responded cyclically as a function of ITD with central peaks and troughs at the same ITDs as in response to noise. The positions of these peaks and troughs also matched those predicted from tonal ITD curves. Thus over the range of physiologically relevant ITDs, the binaural cells in the ICC showed similar sensitivity to ITDs of tones, noise, and clicks. 2. The transient nature of the response to a click allowed association of individual discharges with either the ipsilateral or contralateral stimulus when the binaural stimulus included a large ITD. We studied the influence of the click presented to one side on responses to the click presented to the other side. By examining responses to clicks with large ITDs, ranging from 2 to 3 up to 200 ms, we could identify both excitatory and inhibitory components in response to binaural clicks. 3. For many cells, there was evidence for a short-lasting excitation arising from one or both inputs of the binaural stimulus. Inhibitory interactions could also be demonstrated over a large range of ITDs. Long-lasting, late inhibitory components arose from both contralateral and ipsilateral inputs. In 87% of cells that were driven by the contralateral input, a late inhibitory component originating from the ipsilateral side was detected. In all cells that were driven by the ipsilateral side, a late inhibitory contralateral component was detected. This late inhibition of the excitatory response to one side by a leading stimulus to the other side could be evoked even when the leading stimulus was not effective in evoking an excitatory response. 4. Some cells also exhibited an early inhibitory component that preceded the excitation. An early contralateral inhibition was detected in 44% of cells that were driven by the ipsilateral input, whereas an early ipsilateral component was detected in 17% of cells driven by the contralateral input. 5. We confirmed hypotheses about the laterality and time course of the inhibitory and excitatory components by introducing interaural level differences (ILDs) into the binaural clicks and thus varying the strengths of the different components. 6. Inhibitory components may play a role in shaping the sensitivity of individual cells to ITDs of stimuli other than clicks; they were also apparent in responses to noise.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Temporal coding of resonances by low-frequency auditory nerve fibers: single-fiber responses and a population model.

1. We recorded responses of low-frequency auditory nerve fibers (characteristic frequency (CF) less than 3 kHz) in the cat to resonant stimuli with varied natural frequencies, damping coefficients, and sound pressure levels. Responses to resonances were synchronized to frequencies lying between the peak frequency of the stimulus spectrum and a frequency near the fiber's CF. The frequency of the dominant synchrony in the response varied systematically as a function of the stimulus parameters. 2. More lightly damped resonances, which have sharp spectral peaks, elicited synchrony closer to the peak frequency, whereas the broader peaks of more highly damped resonances elicited synchrony closer to the fiber's CF. Thus as the stimulus was varied from an undamped tone to a highly damped transient, the dominant component of the synchronized response moved from the peak frequency of the stimulus toward the CF of the fiber. The trajectory of the dominant component varied as a function of stimulus level, with higher levels resulting in synchrony biased toward the peak of the stimulus spectrum over a wider range of damping. 3. The frequency tuning and synchronization characteristics of a fiber, along with the stimulus parameters, determined the temporal properties of its response to complex stimuli. Using reverse correlation (revcor) filters to characterize the tuning and synchronization of auditory nerve fibers, we were able to predict the temporal properties of responses to resonant stimuli. 4. A parametric model was fit to measured revcor functions derived from responses of auditory nerve fibers to wideband noise. In this way, a bank of model revcor filters was developed based on our population of measured filters. 5. The filter bank was used to model the response of a population of auditory nerve fibers to resonances. Temporal patterns present in the response of a population of fibers encoded the parameters of resonant stimuli. 6. The model revcor filter bank provided a means of studying temporal response patterns of the population of fibers to other complex sounds. 7. The output of the population model is a representation of the temporal information provided by the auditory periphery to the central nervous system; thus it provides a potentially useful tool for testing hypotheses concerning the processing of temporal information by the central auditory system.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Effects of interaural time delays of noise stimuli on low-frequency cells in the cat's inferior colliculus. II. Responses to band-pass filtered noises.

1. We studied cells in the central nucleus of the inferior colliculus of the cat that were sensitive to interaural time delays (ITDs) in order to evaluate the influence of the stimulus spectrum of noise signals. Stimuli were sharply filtered low-, high-, and band-pass noise signals whose cutoff frequencies and bandwidths were systematically varied. The responses to ITDs of these noise signals were compared with responses obtained to ITDs of broadband noise and pure tones. 2. The discharge rate in response to band-pass noise as a function of ITD was usually a cyclic function with decreasing peak amplitudes at longer ITDs. The reciprocal of the mean interval between adjacent peaks indicated how rapidly the response rate varied with ITD and was termed the response frequency (RF). This RF was approximately equal to the median frequency of the stimulus spectrum filtered by the cell's sync-rate curve, which was the product of the synchronization to interaural phase and the discharge rate plotted against frequency. This suggests that the RF was determined by all the spectral components in the stimulus that fell within the frequency range in which the cell's response was synchronized. The contribution of each component was proportional to the sync-rate for that frequency. 3. The central peak of the ITD function usually fell within the physiological range of ITDs (+/- 400 microseconds). The location of this peak did not vary significantly with changes in stimulus spectrum by comparison with responses to tones of different frequency. Its shape also remained constant, except for a decrease in width when high-frequency components within the range of the sync-rate curve were added to the stimulus. A few cells responded with a minimal discharge instead of a maximal near-zero ITD, and this central minimum had similar properties as the central peak. The amplitude of the secondary peaks of the ITD function decreased as the stimulus bandwidth that overlapped the sync-rate curve broadened. 4. The sum of the ITD functions to two band-pass signals was similar to that of a broadband signal whose spectrum was composed of the sum of the band-pass spectra. 5. From these binaural responses we could make inferences about the response characteristics of the monaural inputs to binaural neurons. We then verified these predictions by studying responses of low-frequency trapezoid body fibers to band-pass noises.

Acoustic Stimulation↗

Effects of interaural time delays of noise stimuli on low-frequency cells in the cat's inferior colliculus. III. Evidence for cross-correlation.

1. We tested the coincidence, or cross-correlation, model of Jeffress, which proposes a neuronal mechanism for sensitivity to interaural time differences (ITDs) in low-frequency cells in the central nucleus of the inferior colliculus (ICC) of the cat. Different tokens of Gaussian noise stimuli were delivered to the two ears. We studied the neural responses to changes in ITDs of these stimuli and examined the manner in which the binaural cells responded to them. All of our results support the idea that the central binaural neurons perform an operation very similar to cross-correlation on the inputs arriving from each side. These inputs are transformed from the actual acoustic signal by the peripheral auditory system, and these transformations are reflected in the properties of the cross-correlations. 2. The responses to ITDs of identical broadband noise stimuli to the two ears varies cyclically as a function of ITD at a frequency close to the best frequency of the neuron. This cyclic response is a consequence of the narrowband filtering of the wideband acoustic signal by the auditory nerve fibers. To examine the effects of using stimuli to the two ears that were correlated to each other to different degrees, we generated pairs of noises. Each pair consisted of one standard noise, which was delivered to one ear, and a linear sum of two standard uncorrelated noises, which was delivered to the other ear. The responses of 34 neurons in the ICC to ITDs of noises with variable interaural coherence were examined. When partially correlated noises were delivered, there was a positive and approximately linear relationship between the degree of modulation of the response as a function of ITD and interaural coherence. The degree of modulation was measured by the synchronization coefficient, or vector strength, over one period of the ITD curve. 3. We examined the effects of altering the interaural phase relationships of the input noise stimuli. The phase of the noise stimuli was changed by digitally filtering the standard noise so that only a phase delay was imposed. The responses to ITDs with differing interaural phase relationships were then studied by delivering a phase-shifted noise to one ear and the standard noise to the other. The ITD curves in response to phase-shifted noise were shifted by about the same amount as the shift of the stimulus; the shift of the response was measured with respect to the case with identical noises to the two ears.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Effects of interaural time delays of noise stimuli on low-frequency cells in the cat's inferior colliculus. I. Responses to wideband noise.

We examined the responses of low-frequency neurons in the central nucleus of the inferior colliculus (ICC) of the cat to interaurally delayed, wideband noise stimuli. The stimuli were pseudorandom noise signals that were generated digitally with a nominal bandwidth of 60-4,000 Hz. We also compared the responses to noise with those obtained from interaural phase differences of pure tones. We studied 144 neurons with characteristic frequencies below 2.5 kHz. Eighty-five percent (85%) of these were sensitive to changes in both interaural time differences (ITDs) of noise and interaural phase differences of pure tones, only 2% were sensitive to one stimulus but not the other, and the remainder were insensitive to both stimuli. For most cells the discharge rate was modulated in an approximately cyclic fashion by changes in ITDs of the wideband noise stimuli. The maximal spike counts often occurred near zero ITD, and there was considerable variability in the nature of the cycling, though it usually disappeared for ITDs greater than +/- 4,000 microseconds. The position of the central peak was usually (65%) within the physiologically relevant range of +/- 400 microseconds, and most (80%) occurred at positive ITDs, which corresponded to delays to the ipsilateral stimulus. In general, the shapes of the responses were not affected by changes in stimulus level above threshold. As long as identical noises were delivered to both ears, the responses were not sensitive to the particular noise stimulus used. When uncorrelated noises were delivered to the two ears, there was no sensitivity to ITDs. Composite curves were computed by linear summation of the responses to ITDs of pure tones at frequencies spaced at equal intervals throughout each cell's response area. The shapes of composite curves were similar to the responses of the same cell to ITDs of wideband noise stimuli. The positions of the central peaks of these two functions were highly correlated (r = 0.91, slope = 0.97). The values of characteristic delay and characteristic phase computed from the tonal responses were found to be good indicators of the shapes of the noise delay curves. Characteristic phases (CPs) near zero were associated with noise delay curves symmetric about the central peak, CPs near 0.5 cycles with those symmetric about the trough, while CPs between 0 and 0.5 or between 0.5 and 1.0 had noise delay curves that were asymmetric with a prominent trough to the left or right, respectively, of the central peak.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Responses of neurons in the cat's superior colliculus to acoustic stimuli. I. Monaural and binaural response properties.

Using extracellular electrodes we studied acoustic responses in the superior colliculus (SC) of the barbiturate-anesthetized cat. Pure tonal stimuli were delivered through sealed and calibrated earphones and were presented either monaurally or binaurally with interaural intensity differences (IIDs) and interaural time differences (ITDs). Acoustically sensitive cells were found in the intermediate and deep layers of the SC throughout its rostrocaudal and mediolateral extent. Most cells (80%) discharged only at stimulus onset; the rest had more complex firing patterns. For 88% of our sample the mean first-spike latency measured at 20 dB above threshold ranged between 6 and 16 ms. The sharpness and threshold intensity of the frequency tuning curves varied widely. In the SC, the average characteristic frequency and threshold intensity were higher than in other auditory brain stem nuclei. Neurons whose characteristic frequency was low were never sharply tuned. The probability of response decreased when the repetition rate at which the stimuli were delivered increased. The mean stimulus interval at which spike count reached 50% of maximum was 360 ms. Most (83%) of the cells discharged only to monaural stimulation of the contralateral ear, 7% responded to tones applied to either ear and only 1% to only ipsilateral input. The remaining cells responded only to stimulation of both ears. With binaural stimuli, most neurons (80%) could be shown to receive input from both ears. Seventy percent of the binaural cells showed predominant binaural inhibition (BI), 25% binaural facilitation (BF), and 5% a more complex mixture. Because the majority of SC neurons had high characteristic frequencies, we examined their responses to IIDs. The spike count vs. IID functions of BI cells were monotonic and sigmoidal, those of BF cells were nonmonotonic and bell-shaped. The slopes and horizontal positions of the curves varied among neurons. IIDs favoring the contralateral ear were the most effective. For a given cell, increasing the mean binaural level extended the range of IIDs that evoked maximal discharge. A small number of cells was sensitive to physiologically significant interaural time differences of low-frequency tones or the envelopes of amplitude-modulated, high-frequency tones.

Animals↗

Responses of neurons in the cat's superior colliculus to acoustic stimuli. II. A model of interaural intensity sensitivity.

Most neurons in the deep and intermediate layers of the superior colliculus (SC) that respond to acoustic stimuli are sensitive to interaural intensity disparities (IIDs). We examine a model for the generation of sensitivity to IIDs that depends upon temporal coincidence of the inputs from each ear at a given binaural neuron. Because the neural response latency decreases with increasing stimulus intensity, IIDs affect the relative timing of arrival of the inputs. If this model were true, the neurons sensitive to IIDs should also respond to interaural time differences (ITDs) of isointensive stimuli, provided that the magnitude of the delays reflect the neural latency-intensity relationship. For both major classes of binaural cells in the SC, namely those that exhibit binaural inhibition (BI) and binaural facilitation (BF), our results support the model in that the detection of IIDs is largely due to their sensitivity to the temporal overlap of inputs from each ear. The shapes of the IID and ITD functions for each class are similar. The summation of inputs includes inhibitory as well as facilitatory interactions. Estimates of the durations of the subliminal excitatory events in BF cells using the model indicate that they are relatively short (1-4 ms), whereas the durations of the inhibitory processes in BI cells are much longer. The model specifies a common neuronal mechanism for comparison of interaural disparities of time and intensity and does not separate the processing of IIDs and ITDs, as the classic duplex theory suggests. The model provides a physiological explanation for certain features of the psychophysical phenomenon of time-intensity trading. It is also consistent with recent experiments that have shown that the auditory system is sensitive to behaviorally significant ITDs of high-frequency complex signals. The model applies only to the processing of transient stimuli and does not address neural sensitivity to IIDs of continuous high-frequency tones.

Animals↗

Coverage of wounds in the distal lower leg by advancement of an enlarged medial gastrocnemius skin flap.

Advancement of a medial gastrocnemius flap is a new technique in treating skin defects of the lower third of the lower leg, including that at the anteromedial aspect of the distal portion of the lower leg. Satisfactory results have been obtained in two clinical cases using this technique. In this modified technique, the size of the flap is enlarged, the origin of the medial head of the gastrocnemius is severed so that the island myocutaneous flap may be advanced 8 to 9 cm distally, and good blood supply to the flap is also maintained. In the authors' opinion, this method is a new approach that is simple, easy, safe, and reliable in treating skin defects of the lower leg.

Adult↗

Interaural time sensitivity of high-frequency neurons in the inferior colliculus.

Recent psychoacoustic experiments have shown that interaural time differences provide adequate cues for lateralizing high-frequency sounds, provided the stimuli are complex and not pure tones. We present here physiological evidence in support of these findings. Neurons of high best frequency in the cat inferior colliculus respond to interaural phase differences of amplitude modulated waveforms, and this response depends upon preservation of phase information of the modulating signal. Interaural phase differences were introduced in two ways: by interaural delays of the entire waveform and by binaural beats in which there was an interaural frequency difference in the modulating waveform. Results obtained with these two methods are similar. Our results show that high-frequency cells can respond to interaural time differences of amplitude modulated signals and that they do so by a sensitivity to interaural phase differences of the modulating waveform.

Animals↗

Binaural interaction in low-frequency neurons in inferior colliculus of the cat. IV. Comparison of monaural and binaural response properties.

We studied the monaural and binaural response properties of 82 low-frequency inferior colliculus (IC) neurons that display a clear sensitivity to changes in interaural phase. Most cells (60%) are excited by sound delivered to either ear, the remainder being excited only by stimulation of one ear; 70% of the neurons receive their stronger or sole excitatory input from the contralateral ear. A monotonic relation between spike discharge and sound pressure level (SPL) is seen in 65% of the monaural response areas, i.e., the range of stimulus frequencies and intensities effective in eliciting a response, while 30% show a nonmonotonic response pattern. In 33% of the cases there is a significant shift in the most effective frequency as a function of SPL. Most discharge patterns are classified as sustained (69%) and the remainder as onset. However, there is considerable variability within these patterns and often two types of discharges are present at different points in the same response area of a single cell. The sustained responses show a broad range of latencies, while onset patterns show a tighter distribution and shorter first spike latencies. Thus, IC neurons showing sensitivity to changes in interaural phase can differ in laterality preferences, response area characteristics, discharge patterns, and latency parameters. Given the diversity of inputs to the IC from lower brain stem structures, this heterogeneity is not surprising. For most neurons excited by stimulation to either ear, the characteristic frequencies, discharge patterns, and first spike latencies are similar, suggesting that the monaural inputs to a binaural cell are of the same type. A neuron's most effective frequencies at a particular SPL for monaural and binaural stimulation are, in general, the same. In some cases a neuron's monaural and binaural response areas can show remarkable similarities, suggesting that certain monaural features are intimately related to the binaural response. In 18% of the IC cells, phase locking to the monaural stimulating frequency is seen. When both inputs are phase locked, a simple coincidence model can predict the interaural phase or delay at which the maximal binaural discharge occurs.

Acoustic Stimulation↗

Binaural interaction in low-frequency neurons in inferior colliculus of the cat. II. Effects of changing rate and direction of interaural phase.

We used the binaural beat stimulus to study the interaural phase sensitivity of inferior colliculus (IC) neurons in the cat. The binaural beat, produced by delivering tones of slightly different frequencies to the two ears, generates continuous and graded changes in interaural phase. Over 90% of the cells that exhibit a sensitivity to changes in the interaural delay also show a sensitivity to interaural phase disparities with the binaural beat. Cells respond with a burst of impulses with each complete cycle of the beat frequency. The period histogram obtained by binning the poststimulus time histogram on the beat frequency gives a measure of the interaural phase sensitivity of the cell. In general, there is good correspondence in the shapes of the period histograms generated from binaural beats and the interaural phase curves derived from interaural delays and in the mean interaural phase angle calculated from them. The magnitude of the beat frequency determines the rate of change of interaural phase and the sign determines the direction of phase change. While most cells respond in a phase-locked manner up to beat frequencies of 10 Hz, there are some cells tht will phase lock up to 80 Hz. Beat frequency and mean interaural phase angle are linearly related for most cells. Most cells respond equally in the two directions of phase change and with different rates of change, at least up to 10 Hz. However, some IC cells exhibit marked sensitivity to the speed of phase change, either responding more vigorously at low beat frequencies or at high beat frequencies. In addition, other cells demonstrate a clear directional sensitivity. The cells that show sensitivity to the direction and speed of phase changes would be expected to demonstrate a sensitivity to moving sound sources in the free field. Changes in the mean interaural phase of the binaural beat period histograms are used to determine the effects of changes in average and interaural intensity on the phase sensitivity of the cells. The effects of both forms of intensity variation are continuously distributed. The binaural beat offers a number of advantages for studying the interaural phase sensitivity of binaural cells. The dynamic characteristics of the interaural phase can be varied so that the speed and direction of phase change are under direct control. The data can be obtained in a much more efficient manner, as the binaural beat is about 10 times faster in terms of data collection than the interaural delay.

Acoustic Stimulation↗