Search PubMed⌕ Search

Biomedical subjects

S Kuwada

Publications and source records attributed to S Kuwada.

33 records · Page 2Linked to original sources
PubMed1991

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↗
PubMed1989

Temporal coding of envelopes and their interaural delays in the inferior colliculus of the unanesthetized rabbit.

1. The difference in the time of arrival of a sound at the two ears can be used to locate its source along the azimuth. Traditionally, it has been thought that only the on-going interaural temporal disparities (ITDs) produced by sounds of lower frequency (approximately less than 2 kHz) could be used for this purpose. However, ongoing ITDs of low frequency are also produced by envelopes of amplitude-modulated (AM) tones. These ITDs can be detected and used to lateralize complex high-frequency sounds (1, 8, 12, 15, 22, 24, 26). Auditory neurons synchronize to the modulation envelope, but do so at progressively lower modulation frequencies at higher levels of the auditory pathway. Some neurons of the cochlear nucleus synchronize best to frequencies as high as 700 Hz, but those of the inferior colliculus (IC) exhibit their best synchrony below 200 Hz. Even though synchrony to higher modulation frequencies is reduced at higher levels of the auditory pathway, is information about ITDs retained? 2. We answered this question by extracellularly recording the responses of neurons in the IC of the unanesthetized rabbit. We used an unanesthetized preparation because anesthesia alters the responses of neurons in the IC to both monaurally presented tones and ITDs. The unanesthetized rabbit is ideal for auditory research. Recordings can be maintained for long periods, and the acoustic stimulus to each ear can be independently controlled. 3. We studied the responses of 89 units to sinusoidally AM tones presented to the contralateral ear. For each unit, we recorded the response at several modulation frequencies. The degree of phase locking to the envelope at each frequency was measured using the synchronization coefficient. Two measures were used to assess the range of modulation frequencies over which phase locking occurred. The "best AM frequency" was the frequency at which we observed the greatest phase locking. The "highest AM frequency" was the highest frequency at which significant phase locking (0.001 level) was observed. We could not assess synchrony to ipsilateral AM tones directly, because most units did not respond to ipsilateral stimulation. 4. We studied the sensitivity of 63 units to ITDs produced by the envelopes of AM tones. Sensitivity to ITDs was tested by presenting AM tones to the two ears that had the same carrier frequency, but modulation frequencies that differed by 1 Hz. Units that were sensitive to ITDs responded to this stimulus by varying their response rate cyclically at the difference frequency, i.e., 1 Hz.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1989

Monaural and binaural response properties of neurons in the inferior colliculus of the rabbit: effects of sodium pentobarbital.

1. We studied the effects of sodium pentobarbital on 22 neurons in the inferior colliculus (IC) of the rabbit. We recorded changes in the sensitivity of these neurons to monaural stimulation and to ongoing interaural time differences (ITDs). Monaural stimuli were tone bursts at or near the neuron's best frequency. The ITD was varied by delivering tones that differed by 1 Hz to the two ears, resulting in a 1-Hz binaural beat. 2. We assessed a neuron's ITD sensitivity by calculating three measures from the responses to binaural beats: composite delay, characteristic delay (CD), and characteristic phase (CP). To obtain the composite delay, we first derived period histograms by averaging, showing the response at each stimulating frequency over one period of the beat frequency. Second, the period histograms were replotted as a function of their equivalent interaural delay and then averaged together to yield the composite delay curve. Last, we calculated the composite peak or trough delay by fitting a parabola to the peak or trough of this composite curve. The composite delay curve represents the average response to all frequencies within the neuron's responsive range, and the peak reflects the interaural delay that produces the maximum response. The CD and CP were estimated from a weighted fit of a regression line to the plot of the mean interaural phase of the response versus the stimulating frequency. The slope and phase intercept of this regression line yielded estimates of CD and CP, respectively. These two quantities are thought to reflect the mechanism of ITD sensitivity, which involves the convergence of phase-locked inputs on a binaural cell. The CD estimates the difference in the time required for the two inputs to travel from either ear to this cell, whereas the CP reflects the interaural phase difference of the inputs at this cell. 3. Injections of sodium pentobarbital at subsurgical dosages (less than 25 mg/kg) almost invariably altered the neuron's response rate, response latency, response pattern, and spontaneous activity. Most of these changes were predictable and consistent with an enhancement of inhibitory influences. For example, if the earliest response was inhibitory, later excitation was usually reduced and latency increased. If the earliest response was excitatory, the level of this excitation was unaltered or slightly enhanced, and changes in latency were minimal. 4. The neuron's response pattern also changed in a predictable way. For example, a response with an inhibitory pause could either change to a response with a longer pause or to a response with an onset only.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1987

Interaural phase-sensitive units in the inferior colliculus of the unanesthetized rabbit: effects of changing frequency.

We studied the interaural phase sensitivity of 85 units in the inferior colliculus (IC) of the unanesthetized rabbit. We assessed this sensitivity at several frequencies within each unit's responsive range. The interaural phase disparity was varied by delivering tones that differed by 1 Hz to the two ears, resulting in a 1-Hz binaural beat. We analyzed each unit's response to different frequencies by calculating four measures: characteristic delay (CD), characteristic phase (CP), composite peak delay, and mean peak delay. We estimated the CD and CP from the slope and phase intercept, respectively, of the regression line fitted to a plot of the mean interaural phase against stimulating frequency. The composite peak delay was estimated from the peak of a composite delay curve. This was generated by replotting the response to changes in interaural phase, as a function of the equivalent interaural delay and averaging the resultant interaural delay curves. The composite delay curve reflects the unit's average response to interaural delays across frequencies. Last, we calculated a mean peak delay, derived by converting the mean interaural phase of the response at each frequency to an equivalent delay and then averaging these delays. Interaural phase sensitivity was observed to frequencies as high as 2,150 Hz. However, the majority of units showed such sensitivity below 1,500 Hz. For most units, the interaural delay curves measured at several frequencies coincided near the peak discharge. This result is consistent with a neural model, where excitatory inputs from each ear converge upon a binaural cell, evoking maximum discharge only when the two inputs arrive simultaneously. As a first approximation, our data fit this model, indicating that IC neurons can act like coincidence detectors or cross-correlators. The distributions of CD, composite peak delay, and mean peak delay showed that most units preferred ipsilateral stimulus delays, which in the natural situation corresponds to sounds emanating from the contralateral field. Moreover, most units preferred delays that were within the estimated physiological range of the rabbit. These results support the viewpoint that neurons in the IC participate in sound localization. The distributions of CP and CD differ substantially from those found in the IC of the anesthetized cat. These differences may reflect species differences, the effects of anesthesia, or a difference in the population of units sampled. For each unit, we assessed the linearity of the plot of mean interaural phase against frequency of stimulation using a chi 2 method. For most units the plots were significantly nonlinear.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1986

The frequency-following response to continuous tones in humans.

Previous studies of the frequency-following response (FFR) in man suggest that it has multiple sources. Identification of these sources has been complicated by the use of tone bursts to evoke FFRs and the lack of precise methods to calculate their amplitude and latency. Tone bursts produce transient responses which confound measurements of the FFR. The use of continuous tones avoids this problem and the Fast Fourier Transform can be used to assess accurately and efficiently the presence, amplitude and phase angle of the FFR. In this study we systematically examined the frequency and intensity range over which FFRs to continuous tones could be evoked using FFRs to tone bursts for comparison. We then analyzed FFRs to continuous tones to determine the sources of this potential. FFRs to both stimuli have similar thresholds (65-90 dB SPL) and can be evoked by the same range of frequencies. Neurogenic FFRs in man occur only below 1000 Hz. The source for this potential has a latency of 8.2 +/- 0.1 ms (mean +/- SD) and is consistent with a midbrain source. At higher frequencies FFRs have a latency of less than 1 ms and are most likely cochlear microphonic. The small variation in the latency of the neurogenic FFR suggests this as a possible tool for assessing neurological disorders.

Adolescent↗
PubMed1986

Scalp potentials of normal and hearing-impaired subjects in response to sinusoidally amplitude-modulated tones.

None of the current electrical audiometric procedures, alone or in combination, has yet achieved the precision of conventional audiometric testing that is used to assess hearing in verbally capable children and adults. The reason for this, in part, lies in the use of stimuli which have a wide frequency content. We have measured scalp potentials which follow the envelopes of sinusoidally amplitude-modulated tones: a frequency-specific stimulus. In normal subjects such amplitude-modulation following responses (AMFRs) appear to be generated by two sources. One source has a latency of about 30 ms, generates large responses and is only observed at modulations below 55 Hz, while the other source has a latency of 7-9 ms, generates smaller responses, and is only observed at modulations from 100-350 Hz. The latencies of these two sources are consistent with origins in the cortex and midbrain, respectively. We examined AMFRs to low frequency (50 Hz) modulations as a possible audiometric tool. In normal subjects, the amplitude of the AMFR increased as a function of intensity, decreased as a function of carrier frequency, and could be evoked across the whole audiometric range (250-8000 Hz). In hearing-impaired subjects, the AMFR amplitudes as a function of carrier frequency accurately reflected the pattern of hearing loss on a frequency-by-frequency basis. In most subjects, the threshold for evoking the AMFR was within 0-25 dB of hearing threshold. It therefore appears that the AMFR may be a potentially useful tool to assess hearing in those unable to undergo conventional audiometric testing.

Acoustic Stimulation↗
PubMed1984

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↗
PubMed1984

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↗
PubMed1983

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↗
PubMed1983

Binaural interaction in low-frequency neurons in inferior colliculus of the cat. III. Effects of changing frequency.

The effects of changing stimulus frequency on the interaural phase sensitivity of neurons in the inferior colliculus (IC) were studied in barbiturate-anesthetized cats in order to reexamine the issue of characteristic delay (CD). Since the results obtained with the interaural delay and binaural beat stimuli are similar, we used the averaged interaural delay curves and binaural beat period histograms as comparable expressions of a neuron's interaural phase sensitivity. When the averaged interaural delay curves at different frequencies are plotted on a common time axis, for some cells the resulting superimposed delay curves show peaks or troughs that coincide at some CD. For most cells, though, this method of detecting a CD by visual inspection yields ambiguous and uncertain results. Composite curves, computed from the average of all the normalized superimposed delay curves, are also not helpful for showing CD. In order to provide a more objective means of analyzing the data, we plotted the mean interaural phase versus the stimulating frequency and computed the linear regression line, using the mean square error as a measure of linearity. The slope of the regression line is the CD for the neuron, and the phase intercept is referred to as the characteristic phase (CP). Cells that display a CD at the peak discharge have a CP = 0.0 cycles, while those that show a CD at the minimum discharge have a CP = 0.5. Cells that exhibit a CP at any value other than 0.0, 0.5, or 1.0 will have a CD at some relative amplitude other than the peak or trough. For cells that exhibit a CD at the peak or trough, results of the analysis procedure using the phase-frequency plot correspond to those obtained from visual inspection. For cells that do not show a common peak or trough, the analysis procedure not only specifies the location of the CD but also provides a statistical criterion of the linearity. From this analysis about 60% of the runs were identified as satisfying the criteria for CD at the P less than 0.005 level and 71% of these CDs are between +/- 300 micros. Most CD cells do not have the CD at the peak or trough of the response. Our results differ from those found in previous studies but they are in essential agreement with the original concept put forth by Rose et al. (31). Some cels exhibit little change in the CD or CP with variations in intensity, while others display marked systematic shifts in both CD and CP. In general, the peaks and troughs of the composite curves show less variability with intensity than the CD.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1983

Binaural interaction in low-frequency neurons in inferior colliculus of the cat. I. Effects of long interaural delays, intensity, and repetition rate on interaural delay function.

Detailed, quantitative studies were made of the interaural phase sensitivity of 197 neurons with low best frequency in the inferior colliculus (IC) of the barbiturate-anesthetized cat. We analyzed the responses of single cells to interaural delays in which tone bursts were delivered to the two ears via sealed earphones and the onset of the tone to one ear with respect to the other was varied. For most (80%) cells the discharge rate is a cyclic function of interaural delay at a period corresponding to that of the stimulating frequency. The cyclic nature of the interaural delay curve indicates that these cells are sensitive to the interaural phase difference. These cells are distributed throughout the low-frequency zone of the IC, but they are less numerous in the medial and caudal zones. Cells with a wide variety of response patterns will exhibit interaural phase sensitivities at stimulating frequencies up to 3,100 Hz, although above 2,500 Hz the number of such cells decrease markedly. Using dichotic stimuli we could study the cell's sensitivity to the onset delay and interaural phase independently. The large majority of IC cells respond only to changes in interaural phase, with no sensitivity to the onset delay. However, a small number (7%) of cells exhibit a sensitivity to the onset delay as well as to the interaural phase disparity, and most of these cells show an onset response. The effects of changing the stimulus intensity equally to both ears or of changing the interaural intensity difference on the mean interaural phase were studied. While some neurons are not affected by level changes, others exhibit systematic phase shifts for both average and interaural intensity variations, and there is a continuous distribution of sensitivities between these extremes. A few cells also showed systematic changes in the shape of the interaural delay curves as a function of interaural intensity difference, especially at very long delays. These shifts can be interpreted as a form of time-intensity trading. A few cells demonstrated orderly changes in the interaural delay curve as the repetition rate of the stimulus was varied. Some of these changes are consonant with an inhibitory effect that occurs at stimulus offset. The responses of the neurons show a strong bias for stimuli that would originate from he contralateral sound field; 77% of the responses display mean interaural phase angles that are less than 0.5 of a cycle, which are delays to the ipsilateral tone.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1979

Response of cat inferior colliculus neurons to binaural beat stimuli: possible mechanisms for sound localization.

The interaural phase sensitivity of neurons was studied through the use of binaural beat stimuli. The response of most cells was phase-locked to the beat frequency, which provides a possible neural correlate to the human sensation of binaural beats. In addition, this stimulus allowed the direction and rate of interaural phase change to be varied. Some neurons in our sample responded selectively to manipulations of these two variables, which suggests a sensitivity to direction or speed of movement.

Animals↗