Search PubMed⌕ Search

Biomedical subjects

M B Sachs

Publications and source records attributed to M B Sachs.

At least 37 records · Page 2Linked to original sources

Strategies for the representation of a tone in background noise in the temporal aspects of the discharge patterns of auditory-nerve fibers.

The responses of populations of auditory-nerve fibers to both a 1.0-kHz tone, and 1.0-kHz tone in broadband noise, have been measured. Period histograms were generated from fiber spike trains and discrete Fourier transforms (DFTs) with a resolution of 125 Hz were computed from each histogram. Sample mean and sample variance statistics were generated for period histograms of response and for temporal response measures derived from discrete Fourier transforms. It is demonstrated how the statistical properties of auditory-nerve fiber response determine the strategy for the estimation and discrimination of particular stimulus components. When the tone is presented alone, the entire population of auditory-nerve fibers provides statistically reliable estimates of the 1.0-kHz tone. Upon addition of the broadband noise stimulus only those units with characteristic frequencies which are close in frequency to the 1.0-kHz stimulus provide spectral estimates which have high signal-to-noise ratios (mean-squared-to-variance ratios). Estimates of the 1.0-kHz-tone stimulus derived from auditory-nerve fibers with characteristic frequencies which are far from the 1.0-kHz stimulus are statistically unreliable. Based on the responses of the population of auditory-nerve fibers, the strategy for estimating the 1.0-kHz-tone stimulus is to derive estimates of the 1.0-kHz stimulus from the subpopulation of neurons with characteristic frequencies close to the 1.0-kHz stimulus. It is concluded that neurons which are tuned close to 1.0-kHz provide the central nervous system (CNS) with the most salient information about the 1.0-kHz stimulus in the presence of the broadband background.

Analysis of Variance↗

Effect of electrical stimulation of the crossed olivocochlear bundle on auditory nerve response to tones in noise.

The discharge rates of single auditory-nerve fibers responding to best-frequency (BF) tones of varying level presented simultaneously with fixed level broadband noise were recorded with and without electrical stimulation of the crossed olivocochlear bundle (COCB). In the absence of COCB stimulation, monotonic increases in noise level produce monotonic increases in the low-level noise-driven response rate of auditory nerve fibers. As a result of adaptation, these increases in noise-driven response rate produce monotonic decreases in saturation discharge rate. At high noise levels, these compressive effects may eliminate the differential rate response of auditory nerve fibers to BF tones. COCB stimulation can restore this differential rate response by producing large decreases in noise-driven response rate and large increases in saturation discharge rate. In backgrounds of quiet, COCB stimulation is known to shift the dynamic range of single auditory nerve fiber BF tone responses to higher stimulus levels. In the presence of background noise, COCB stimulation produces upward shift of dynamic range, which decreases with increasing noise level. At high noise levels, COCB-induced decompression of rate-level functions may occur with little or no dynamic range shift. This enables auditory nerve fibers to signal changes in tone level with changes in discharge rate at lower signal-to-noise ratios than would be possible otherwise. Broadband noise also produces upward shift of the dynamic range of single auditory nerve fiber BF tone response. Noise-induced dynamic range shift of BF tone response was measured as a function of noise level with and without COCB stimulation. COCB stimulation elevates the threshold of noise-induced dynamic range shift. This shift is thought to result from two-tone rate suppression. Increases in the threshold of noise-induced shift due to COCB stimulation therefore suggests an interaction between the mechanism of two-tone rate suppression and the mechanism by which COCB stimulation produces dynamic range shift. These interactions were further investigated by recording auditory nerve fiber rate responses to fixed-level BF excitor tones presented simultaneously with fixed-frequency variable level suppressor tones. Rate responses were recorded with and without COCB stimulation. Experimental results were quantified using a phenomenological model of two-tone rate suppression presented by Sachs and Abbas.

Acoustic Stimulation↗

Effects of noise on pure-tone thresholds in blackbirds (Agelaius phoeniceus and Molothrus ater) and pigeons (Columba livia).

Blackbirds and pigeons were trained to detect tones in quiet and in broadband noise by using positive-reinforcement techniques. In Experiment 1, thresholds in noise were obtained in blackbirds as a function of both tone frequency and noise intensity for a pulsed noise masker (noise gated on and off with tone). For blackbirds, critical ratios (the ratio of the power of the just-detectable tone in noise to the power of the noise masker) obtained in pulsed noise showed no consistent relation to tone frequency. For pigeons, on the other hand, critical ratios obtained in continuous noise increased by about 3 dB/octave across their range of hearing, being similar to known critical ratio functions for cats and humans. In Experiment 2, critical ratios in blackbirds obtained with both continuous noise and pulsed noise were compared. Blackbird critical ratios were more stable in continuous noise and averaged 4 dB lower than critical ratios in pulsed noise. The blackbird critical ratio function obtained with continuous noise was similar to the known critical ratio function of another avian species, the parakeet. Thus, small birds appear to have atypical critical ratio functions, compared with pigeons and other vertebrates.

Animals↗

Representation of a low-frequency tone in the discharge rate of populations of auditory nerve fibers.

Discharge rates for populations of single auditory nerve fibers in response to 1.5 kHz tone bursts were measured in anesthetized cats. Separate plots of average rate vs. best frequency (rate-place profiles) were made for high, medium and low spontaneous rate (SR) auditory nerve fibers. At the lowest sound levels studied (34 dB SPL), all three SR groups show a peak in the rate-place profile centered around 1.5 kHz. At the highest sound levels studied (87 dB SPL), the average rates of the high and medium SR fibers are saturated across a wide range of best frequencies, but a peak around 1.5 kHz is maintained in the rate-place representation of the low SR fibers. These results show that in addition to the temporal information present in the discharge patterns of auditory nerve fibers, a rate-place representation of a single low-frequency tone exists in the auditory nerve over a wide range of sound levels.

Acoustics↗

Representation of voice pitch in discharge patterns of auditory-nerve fibers.

Responses of populations of auditory-nerve fibers were measured for synthesized consonant-vowel stimuli. This paper explores the encoding of fundamental frequency (pitch) in these responses. Post-stimulus time (PST) histograms were computed from 25 ms segments of the spike trains. Discrete Fourier transforms with a 40 Hz resolution were computed from the histograms. Two representations of pitch are considered. The first representation is based on the pitch-related temporal properties of the speech signal. Histograms for individual units can show envelope modulations directly related to the pitch period. These modulations reflect the responses of these fibers to a number of stimulus harmonics near fiber CF. Responses of fibers near formant frequencies are dominated by a single large harmonic component, and thus show small or no pitch-related enveloped modulations. Envelope modulations are reduced in the presence of background noise. The second representation uses both temporal properties of auditory-nerve responses and cochlear place to encode the pitch-related harmonic structure of speech. As a measure of the response of the population of fibers to each harmonic of 40 Hz the magnitude of the component of the Fourier transform at that frequency was averaged across all fibers whose characteristic frequencies were within one-fourth octave of that harmonic. We call this measure the average localized synchronized rate (ALSR). The ALSR provides a good representation of stimulus spectrum, even in the presence of background noise. From the harmonic structure of the ALSR, we are able to extract the stimulus pitch frequency. The relationship of these two representations to pitch perception in both acoustic and electrical stimulation (via cochlear implants) is discussed.

Acoustic Stimulation↗

Neural coding of complex sounds: speech.

The studies that we have reviewed here demonstrate that a temporal-place code can represent fine details in the spectra of vowels and stop-consonants. These features include formant frequencies, formant-frequency transitions, and pitch. On the other hand, such a phase locking-based representation may have difficulty with certain fricative consonants. Detailed format structure of vowels is present in a rate-place code at moderate stimulus levels, but is maintained at high levels only in the small population of low-spontaneous-rate fibers. A rate-place code preserves the formant structure at high stimulus levels better for stop-consonants than for vowels. Formant structure of some fricatives may be represented in a place-rate code in cases where a temporal-place code fails. Voice pitch is well preserved in a temporal code but not in a rate-place code. From this summary we must consider the possibility that the central nervous system utilizes both rate-place and temporal-place information in processing all of the relevant features of speech. This review points out a number of issues that currently confronts us in the coding of complex stimuli. We conclude by briefly summarizing these issues. First, we must attempt to clarify the role of the low-spontaneous-rate, high-threshold fibers in the representation of speech at high-stimulus levels. A second major question that must be addressed is the role of the cochlear efferents in the peripheral encoding of speech, especially in the presence of background noise. Closely related to the role of the efferents may be the role of the middle ear muscles. Finally, if we are to seriously consider a temporal-based code for speech, we must develop quantitative hypotheses about how spectral information might be extracted from temporal patterns in the central nervous system with real neural "hardware."

Electrophysiology↗

Representation of stop consonants in the discharge patterns of auditory-nerve fibers.

The representation of the speech syllables /da/ and /ba/ in populations of auditory-nerve fibers was studied. Post-stimulus-time histograms were computed from 20-ms segments of fiber spike trains occurring in response to the stimulus. Discrete Fourier transforms with a resolution of 50 Hz were computed from each histogram. As a measure of the response of the population of fibers to each harmonic of the 50-Hz resolution frequency of the transform, the magnitude of the response to that frequency was averaged across all fibers whose characteristic frequencies were within one-fourth octave of that harmonic. We have previously called this measure the average localized synchronized rate (ALSR). Response profiles for the 20-ms segments of the stimulus were generated by plotting the ALSR versus frequency. Time-varying spectral features of the /da/ and /ba/ stimuli are well preserved by such profiles. For example, the onset spectrum and formant transitions of the consonant-vowel syllable are well represented. Furthermore, the fine structure in the speech spectrum related to the pitch of the excitation source is maintained in these ALSR plots. Average discharge rate profiles were generated in a manner similar to that for the ALSR; in this case average rate replaces Fourier transform components as response measure. Such average rate profiles can represent the transitions of at least formants two and three. However, such average rate profiles do not represent the steady-state formants or the voice pitch.

Animals↗

Auditory nerve representation of vowels in background noise.

Responses of auditory nerve fibers to steady-state vowels presented alone and in the presence of background noise were obtained from anesthetized cats. Representation of vowels based on average discharge rate and representation based primarily on phase-locked properties of responses are considered. Profiles of average discharge rate versus characteristic frequency (CF) ("rate-place" representation) can show peaks of discharge rate in the vicinity of formant frequencies when vowels are presented alone. These profiles change drastically in the presence of background noise, however. At moderate vowel and noise levels and signal/noise ratios of +9 dB, there are not peaks of rate near the second and third formant frequencies. In fact, because of two-tone suppression, rate to vowels plus noise is less than rate to noise alone for fibers with CFs above the first formant. Rate profiles measured over 5-ms intervals near stimulus onset show clear formant-related peaks at higher sound levels than do profiles measured over intervals later in the stimulus (i.e., in the steady state). However, in background noise, rate profiles at onset are similar to those in the steady state. Specifically, for fibers with CFs above the first formant, response rates to the noise are suppressed by the addition of the vowel at both vowel onset and steady state. When rate profiles are plotted for low spontaneous rate fibers, formant-related peaks appear at stimulus levels higher than those at which peaks disappear for high spontaneous fibers. In the presence of background noise, however, the low spontaneous fibers do not preserve formant peaks better than do the high spontaneous fibers. In fact, the suppression of noise-evoked rate mentioned above is greater for the low spontaneous fibers than for high. Representations that reflect phase-locked properties as well as discharge rate ("temporal-place" representations) are much less affected by background noise. We have used synchronized discharge rate averaged over fibers with CFs near (+/- 0.25 octave) a stimulus component as a measure of the population temporal response to that component. Plots of this average localized synchronized rate (ALSR) versus frequency show clear first and second formant peaks at all vowel and noise levels used. Except at the highest level (vowel at 85 dB sound pressure level (SPL), signal/noise = +9 dB), there is also a clear third formant peak. At signal-to-noise ratios where there are no second formant peaks in rate profiles, human observers are able to discriminate second formant shifts of less than 112 Hz. ALSR plots show clear second formant peaks at these signal/noise ratios.

Animals↗

Representation of whispered vowels in discharge patterns of auditory-nerve fibers.

We have previously shown that the spectra of speech sounds can be represented in the temporal patterns of discharge of populations of auditory-nerve fibers. These results were obtained using perfectly periodic stimuli, for which a temporal representation is straightforward. In order to see if our results could be generalized to nonperiodic stimuli, we have studied responses to a whispered vowel with formant frequencies typical of /epsilon/. The whispered vowel was generated by exciting a vocal tract model with noise; this signal was therefore aperiodic. Temporal patterns of responses to the vowel in populations of auditory-nerve fibers were analyzed using interval histograms. Fourier transforms of these histograms show large components at frequencies near the formant frequencies of the vowel. With these Fourier transform components as a measure of the temporal response, a temporal place representation of the response of populations of fibers preserves the spectral features of the aperiodic vowel stimulus. Profiles of average rate versus characteristic frequency for fibers with spontaneous rates greater than 1/s show little if any formant-related structure. On the other hand, such profiles for fibers with spontaneous rates less than 1/s may show peaks in the region of the formants.

Animals↗

Responses of auditory-nerve fibers to characteristic-frequency tones and low-frequency suppressors.

Responses of single auditory-nerve fibers to combinations of low-frequency (suppressor) and characteristic frequency (CF) tones were recorded. The shapes of period histograms were observed as the level of the suppressor was raised. For fibers with CFs above 5.0 kHz, at the lowest suppressor levels used, the phase of the suppressor tone which caused instantaneous rate to increase when presented alone caused a rate decrease when added to a CF tone. For suppressor frequencies of 500 and 100 Hz, as suppressor level is increased, two peaks appear in the period histograms. Further increase in level causes first one, then the other of these peaks to disappear and the histogram takes on the shape of the histogram to the suppressor alone. A similar progression of histogram shape follows a decrease in CF tone level for fixed suppressor level.

Acoustic Stimulation↗

Aspects of frequency discrimination in passerine birds and pigeons.

With the use of a positive reinforcement operant technique, pulsed-tone, frequency difference limens (DLs) were measured in two Icterid passerine species, the redwing blackbird (Agelaius phoeniceus) and the brown-headed cowbird (Molothrus ater). Direct comparisons were made with two nonpasserine control (pigeon) subjects. Blackbirds were more sensitive than pigeons to frequency changes above 1.0 kHz but less sensitive below 1.0 kHz. Measured DLs in all these birds were within the range of nonhuman mammals but higher than those of humans. The lowest delta f/f ratio for all birds occurred at 4.0 kHz, a frequency for which they exhibit minimal absolute auditory thresholds and which is a dominant component in the song of the redwing. As in humans, blackbird sensitivity at 1.0 and 2.0 kHz declined as sensation level (SL) was decreased, but at 4.0 kHz, sensitivity showed no clear dependence on SL. At 8.0 kHz, near the upper limit of blackbird hearing, sensitivity increased with a decrease in SL. This latter frequency is a dominant component in the song of the cowbird. Differences in frequency sensitivty were also found as a function of the direction of frequency shift from the standard: Upward shifts produced lower DLs at 1.0 and 2.0 kHz, and downward shifts produced lower DLs at 8.0 kHz.

Animals↗

Auditory intensity discrimination in blackbirds and pigeons.

Redwing blackbirds, brown-headed cowbirds, and pigeons were trained with operant conditioning techniques to respond to small increases in the intensity of pulsed tone trains at three frequencies: .5, 1.0, and 2.0 kHz. All three species produced similar intensity difference limens (DLs) at the frequencies tested. Intensity DLs decreased as sensation level (intensity level above absolute threshold) increased at all three frequencies, with the slopes of these sensation level functions being greatest at 2.0 kHz . The median intensity DLs at 50 dB sensation level were 3.3, 2.7, and 2.9 dB at .5, 1.0, and 2.0 kHz, respectively, averaged over the three species. Some subjects were also required to detect decreases in intensity. They produced intensity DLs two to three times larger than the DLs obtained when these same subjects were required to detect increases in intensity. Avian intensity DLs generally appear to be 1-2 dB higher than the DLs of those mammals that have been tested (rat, cat, monkey, humans).

Animals↗

Nonlinearities in auditory-nerve fiber responses to bandlimited noise.

Discharge rate was measured as a function of spectral level for noise bursts of one bandwidth and center frequency. Such rate-level functions were measured for a number of bandwidths; either the low- or high-cutoff frequencies were set at fiber characteristic frequency (CF). Rate-level functions were also measured, simultaneously, for single tones at CF. We define dynamic range as the range in descibels over which rate increases from 10% to 80% of the maximum driven rate to CF tones. When pooling data across CF in single cats, dynamic range is an increasing function of fiber threshold for CF tones and noise stimuli. Narrow bands of noise produce rate-level functions that are similar to those for CF tones. For noise bands centered above CF, rate-level functions become less steep as bandwidth is increased, and are always monotonic. For wide bands of noise centered below CF, rate-level functions can be nonmonotonic or appear to plateau at rates less than the saturation rate to CF tones. Thus, wide bands of noise centered above or below CF can produce lower discharge rates than do narrow bands at the same spectral level. This rate reduction has properties similar to those for two-tone suppression. The suppressive effects observed for bandlimited noise are most pronounced on low spontaneous units and least pronounced on high spontaneous units.

Acoustic Stimulation↗

Encoding of steady-state vowels in the auditory nerve: representation in terms of discharge rate.

Responses of large populations of auditory-nerve fibers to synthesized steady-state vowels were recorded in anesthetized cats. Driven discharge rate to vowels, normalized by dividing by saturation rate (estimated from the driven rate to CF tones 50 dB above threshold), was plotted versus fiber CF for a number of vowel levels. For the vowels /I/ and /e/, such rate profiles showed a peak in the region of the first formant and another in the region of the second and third formants, for sound levels below about 70 dB SPL. For /a/ at levels below about 40 dB SPL there are peaks in the region of the first and second formants. At higher levels these peaks disappear for all the vowels because of a combination of rate saturation and two-tone suppression. This must be qualified by saying that rate profiles plotted separately for units with spontaneous rates less than one spike per second may retain peaks at higher levels. Rate versus level functions for units with CFs above the first formant can saturate at rates less than the saturation rate to CF to-es or they can be nonmonotonic; these effects are most likely produced by the same mechanism as that involved in two-tone suppression.

Acoustic Stimulation↗

Representation of steady-state vowels in the temporal aspects of the discharge patterns of populations of auditory-nerve fibers.

This paper is concerned with the representation of the spectra of synthesized steady-state vowels in the temporal aspects of the discharges of auditory-nerve fibers. The results are based on a study of the responses of large numbers of single auditory-nerve fibers in anesthetized cats. By presenting the same set of stimuli to all the fibers encountered in each cat, we can directly estimate the population response to those stimuli. Period histograms of the responses of each unit to the vowels were constructed. The temporal response of a fiber to each harmonic component of the stimulus is taken to be the amplitude of the corresponding component in the Fourier transform of the unit's period histogram. At low sound levels, the temporal response to each stimulus component is maximal among units with CFs near the frequency of the component (i.e., near its place). Responses to formant components are larger than responses to other stimulus components. As sound level is increased, the responses to the formants, particularly the first formant, increase near their places and spread to adjacent regions, particularly toward higher CFs. Responses to nonformant components, exept for harmonics and intermodulation products of the formants (2F1,2F2,F1 + F2, etc), are suppressed; at the highest sound levels used (approximately 80 dB SPL), temporal responses occur almost exclusively at the first two or three formants and their harmonics and intermodulation products. We describe a simple calculation which combines rate, place, and temporal information to provide a good representation of the vowels' spectra, including a clear indication of at least the first two formant frequencies. This representation is stable with changes in sound level at least up to 80 dB SPL; its stability is in sharp contrast to the behavior of the representation of the vowels' spectra in terms of discharge rate which degenerates at stimulus levels within the conversational range.

Acoustic Stimulation↗