Anatomical localization of electrophysiological recording sites by co-ordinate transformation.

A method for estimating the anatomical locations of the units recorded in electrophysiological mapping experiments is described. A total of three locations must be marked by dye injections or electrolytic lesions and identified in tissue sections. From those locations, equations are derived to translate, scale, and rotate the three-dimensional co-ordinates of the recording sites, so that they are correct for a second, three-dimensional co-ordinate system based on the anatomy of the mapped structure. There is no limit to the number of recording sites that can be localized. This differs from methods that require a dye injection or lesion to be made at the exact location at which a particular unit was recorded. The accuracy of the transformed co-ordinates is limited by the accuracy with which the co-ordinates can be measured: in test measurements and in the experiments for which this algorithm was developed, the computed co-ordinates were typically accurate to within 100 microns or less.

Effects of interaural time differences on the responses of chinchilla inferior colliculus neurons to consonant-vowel syllables.

The responses of 100 inferior colliculus neurons to syllables differing in voice onset time (VOT) presented binaurally were studied. As in a previous study of monaural responses (Chen et al., 1996), the responses consisted of 1-3 response 'components', referred to as release responses, VOT responses or vowel responses. The discharge rate of all response components could vary cyclically with the interaural time difference (ITD). The maximal rate often occurred at an ITD around +0.2 ms (contralateral ear leading). Response frequencies (RF) based on the periodicity of the delay curves varied with the characteristic frequency (CF) and VOT. RF also varied across response components. Overall, RF was correlated with the 'most effective frequency', the spectral component with the highest amplitude, relative to the tuning curve. VOT response latency for a given syllable could change by a few ms with ITD, but those changes were small, relative to the range of latencies observed over the entire range of VOTs. Changes in ITD produced large changes in the overall shape of the peristimulus time histogram. There was no relation between the histogram shape and perceptual consonant categories.

Responses of primary auditory fibers to combined noise and tonal stimuli.

Responses to tonal stimuli, with and without added noise of different bandwidths, were obtained from anesthetized cat auditory-nerve fibers using glass micropipettes. When low-pass noise with a cut-off frequency at least one octave below best (or characteristic) frequency was used, every fiber tested at high enough intensities showed a suppression of the tonal response. This suppression did not cause a general reduction of neural responsiveness to all sounds, but rather took the general form of a frequency-specific reduction in the effective intensity of the tonal stimuli. The suppression mechanism(s) involved thus adjust the sensitivity of these fibers to cover higher intensity ranges in the presence of noise. The frequency of the most severely affected tones was always at or near best frequency, in confirmation of previous work (Abbas, P.J. and Sachs, M.B. (1976): J. Acoust. Soc. Am. 59, 112-122; Kiang, N.Y.-S. and Moxon, E.C. (1974): J. Acoust. Soc. Am. 55, 620-630). The suppresson is a direct but highly nonlinear function of the intensity and bandwidth of the noise. The effects on tonal response of wide-band noise were more variable, sometimes causing suppression similar to that induced by the low-pass noise and sometimes causing only 'strong-signal capture' effects. A model of noise-induced suppression has been developed whereby each sound produces both an excitatory effect, sharply tuned at best frequency, and a suppressive effect, which also has its lowest threshold at best frequency but is more broadly tuned.

Auditory-nerve fiber responses to frequency-modulated tones.

Discharge patterns of cat auditory-nerve fibers were obtained in response to frequency-modulated (FM) tones. The rate and direction of frequency change and the sound-pressure level of the sweep tones were systematically varied, and aspects of the discharge patterns were compared to aspects of the discharge patterns elicited by pure tones. Increases in SPL broaden the frequency range over which the fiber responds, as is the case with pure-tone stimuli. Increases in the rate of frequency change have little effect on frequency selectivity for the rates tested. In general, the pure-tone response area is a good predictor of the response area to FM. Although approximately equal numbers of spikes are elicited by ascending and descending sweeps, the discharge patterns differ slightly; for each direction of frequency change, the FM response area is shifted in the direction of the earliest-occurring frequencies. Most of this shift can be accounted for by neural adaptation. This asymmetry is small, relative to those observed in the central nervous system.

Correlates of tone-on-tone masked thresholds in the chinchilla auditory nerve.

In an attempt to determine neural correlates of tone-on-tone masking, discharge patterns of chinchilla auditory-nerve fibers were obtained in response to a set of two-tone stimuli for which behavioral masking had been previously measured (Long, G.L. and Miller, J.D. (1981): Hearing Res. 4, 279-285). The lowest masked thresholds in a sample of fibers were quantitatively similar to the chinchilla's behavioral masked thresholds. In addition, the neural data were in qualitative agreement with other previously-described characteristics of tone-on-tone masking, such as the contribution of cochlear distortion products and the upward spread of masking. It thus appears that the limitations imposed by peripheral frequency analysis determine the tone-on-tone masking pattern.

Monaural response properties of single neurons in the chinchilla inferior colliculus.

The responses of 274 inferior colliculus (IC) central nucleus neurons from 20 chinchillas were studied. Characteristic frequency (CF) increased as the IC was traversed in the dorsal-ventral direction. Most units had little or no spontaneous activity, with a mean threshold for response of about 30 dB SPL across all units. Tuning curve width varied between units, with a significant increase in Q20, with increasing CF. Peri-stimulus time histogram (PSTH) types were similar to those reported for cat inferior colliculus units. Transient, sustained, pauser, and buildup types were observed, with transient responses predominating. Response area (RA) types were also similar to those of cat IC units, with most units displaying stable best frequencies across a range of stimulus intensity levels. For a few units, excitatory RA regions were surrounded by inhibitory sidebands. Nonmonotonic discharge rate vs. stimulus intensity level functions were common in all CF ranges and for all PSTH and RA types. Mean first spike latencies, however, differed across PSTH groups, owing to the temporal definitions of these PSTH shapes. Latencies of sustained units were significantly longer than those of transient units, and buildup PSTHs showed significantly longer latencies than any other group.

Electrophysiological responses of cochlear root neurons.

Cochlear root neurons (CRNs) are second-order neurons interspersed among the fibers of the cochlear nerve in certain rodents. They project, among other nuclei, mainly to the pontine reticular nucleus, and participate in the acoustic startle response (ASR), a short-latency motor reflex initiated by sudden intense sounds. The sound-evoked activity of CRNs has not previously been described. Here we describe extracellular responses of CRNs located in the infranuclear portion of the cochlear nerve root. CRNs exhibited secure responses to tone bursts, with first-spike latencies of approximately 2.2 ms. The characteristic frequencies of the recorded CRNs were about 30 kHz, and the best-characterized CRN had a threshold of 10 dB sound pressure level and sharpness of tuning similar to that of cochlear nerve fibers. The peristimulus time histograms were primary-like with notch. The observed response properties were consistent with the suggestion that CRNs provide the short-latency acoustic input to the reticular formation that leads to an ASR.

Acute spiral ganglion lesions change the tuning and tonotopic organization of cat inferior colliculus neurons.

Many studies have reported plastic changes in central auditory frequency organization after chronic cochlear lesions. These studies employed mechanical, acoustic or drug-induced disruptions of restricted regions of the organ of Corti that permanently alter its tuning and sensitivity and require an extended recovery period before central effects can be measured. In this study, mechanical lesions were made to 1 mm sectors of the spiral ganglion (SG). These lesions remove a restricted portion of the cochlear output, but leave the organ of Corti and basilar membrane intact. Multiunit mapping assessed the pre- and post-lesion tonotopic organization of the inferior colliculus (IC). Immediately after SG lesions, IC neurons previously tuned to the lesion frequencies became less sensitive to those frequencies but more sensitive to lesion edge frequencies, resulting in a shift in their characteristic frequencies (CFs). Notches in the excitatory response areas at frequencies corresponding to the lesion frequencies and expansion of spatial tuning curves were also observed. CFs of neurons tuned to unlesioned frequencies were unchanged. These results suggest that 'plastic' changes similar to those observed after long survival times in previous studies require little or no experience and occur within minutes to hours following the lesion.

Frequency organization and cellular lamination in the medial geniculate body of the rabbit.

Cellular laminae within the tonotopically organized ventral division of the medial geniculate body (MGV) of the cat have been proposed as the anatomical substrate for physiologically defined isofrequency contours. In most species, the laminae are not visible with routine Nissl stains, but are defined by the dendritic fields of principal cells and the terminal arbors of afferents arising from the inferior colliculus. In the present study, we have used the rabbit to directly examine the relationship between the laminar and tonotopic organization of the MGV. Best frequency maps of the MGV in anesthetized adult New Zealand white rabbits were generated from cluster responses recorded at 30-100 microm intervals to randomly presented tone bursts. Parallel vertical penetrations, roughly perpendicular to the laminae, revealed a low-to-high frequency gradient within the MGV. Non-laminated regions of the ventral division, generally found at the rostral or caudal poles, did not demonstrate a systematic frequency gradient. In contrast to a predicted smooth gradient, best frequencies shifted in discrete steps across the axis of the laminae. A similar step-wise frequency gradient has been shown in the central nucleus of the inferior colliculus of the cat. It is proposed that the central laminated core of the MGV represents an efficient architecture for creating narrow frequency filters involved in fine spectral analysis.

Auditory nerve fiber representation of cues to voicing in syllable-final stop consonants.

Acoustic cues to the identity of consonants such as /d/ and /t/ vary according to contextual factors such as the position of the consonant within a syllable. However, investigations of the neural coding of consonants have almost always used stimuli in which the consonant occurs in the syllable-initial position. The present experiments examined the peripheral neural representation of spectral and temporal cues that can distinguish between stop consonants /d/ and /t/ in syllable-final position. Stimulus sets consisting of the syllables /hid/, /hit/, /hud/, and /hut/ were recorded by three different talkers. During the consonant closure interval, the spectrum of /d/ was characterized by the presence of a low-frequency "voice bar." The closure interval for the voiceless consonant /t/ was longer and lacked a voice bar. Most neurons' responses were characterized by discharge rate decreases at the beginning of the closure interval and by rate increases that marked the release of the consonant closure. Exceptions were seen in the responses of neurons with characteristic frequencies (CFs) below approximately 0.7 kHz to syllables ending in /d/. These neurons responded to the voice bar with discharge rates that could approach the rates elicited by the vowel. The latencies of prominent discharge rate changes were measured for all neurons and used to compute the length of the "encoded closure interval." The encoded interval was clearly longer for syllables ending in /t/ than in /d/. The encoded interval increased with CF for both consonants but more rapidly for /t/. Differences in the encoded closure interval were small for syllables with different vowels or syllables produced by different talkers.

Average discharge rate representation of voice onset time in the chinchilla auditory nerve.

Responses of chinchilla auditory-nerve fibers to synthesized stop consonants differing in voice onset time (VOT) were obtained. The syllables, heard as /ga/-/ka/ or /da/-/ta/, were similar to those previously used by others in psychophysical experiments with human and with chinchilla subjects. Average discharge rates of neurons tuned to the frequency region near the first formant generally increased at the onset of voicing, for VOTs longer than 20 ms. These rate increases were closely related to spectral amplitude changes associated with the onset of voicing and with the activation of the first formant; as a result, they provided accurate information about VOT. Neurons tuned to frequency regions near the second and third formants did not encode VOT in their average discharge rates. Modulations in the average rates of these neurons reflected spectral variations that were independent of VOT. The results are compared to other measurements of the peripheral encoding of speech sounds and to psychophysical observations suggesting that syllables with large variations in VOT are heard as belonging to one of only two phonemic categories.

Neural mechanisms of tone-on-tone masking: patterns of discharge rate and discharge synchrony related to rates of spontaneous discharge in the chinchilla auditory nerve.

Responses of chinchilla auditory nerve fibers to brief probe tones in the presence of a fixed tonal masker were obtained. The stimulus conditions were analogous to those that have been used in many psychophysical experiments. The relation between previously described response properties of auditory nerve fibers and features of psychophysical tone-on-tone masking was examined. In psychophysical studies, a fixed narrowband masker produces a characteristic pattern of masked thresholds, which becomes broad and asymmetrical at high masker levels. In the present experiment 1, a 5,000-Hz masker was presented at 30, 50, and 70 dB SPL. Masked thresholds based on the average rate of response to probe tones were estimated for single auditory nerve fibers. The lowest of these masked thresholds formed a pattern similar to the psychophysical masking pattern, becoming broader and more asymmetrical as the masker was increased to 70 dB SPL. The masked thresholds of fibers with low and medium rates of spontaneous discharge (SR) were as low as or lower than the masked thresholds of fibers with high SRs. In certain frequency regions, masked thresholds based on responses to cochlear distortion products were lower than the masked thresholds of any fiber responding to the probe tone; this result is also similar to previous psychophysical observations. In experiment 2, responses of chinchilla auditory nerve fibers to probe tones in the presence of a masker at 1,000 Hz and 50 dB SPL were studied. Probe tone thresholds in the presence of this masker have been measured psychophysically in chinchillas. Thus the relation between behavioral and neural masked thresholds in the same species could be examined. Masked thresholds were estimated from average discharge rate responses and also from discharge synchrony. Good quantitative agreement was observed between the probe tone levels at which changes in average discharge rate were observed and the chinchilla's behavioral masked thresholds. For fibers matched for characteristic frequency, the masked thresholds based on average discharge rate of high-SR fibers tended to be elevated compared with the thresholds of medium-SR fibers. Changes in discharge rate synchronized to the probe tone occurred at levels lower than the chinchilla's behavioral masked thresholds. If discharge synchrony can be used for detection, the code would appear to be based on the relative synchrony to the probe tone and to the masking tone. Low synchrony masked thresholds were obtained from fibers with all SRs.(ABSTRACT TRUNCATED AT 400 WORDS)

Synchronized discharge rate representation of voice-onset time in the chinchilla auditory nerve.

Responses of chinchilla auditory nerve fibers to synthesized stop consonant syllables differing in voice-onset time (VOT) were obtained. The syllables, heard as /ga/-/ka/ or /da/-/ta/, were similar to those previously used by others in psychophysical experiments with human and chinchilla subjects. Synchronized discharge rates of neurons tuned to frequencies near the first formant increased at the onset of voicing for VOTs longer than 20 ms. Stimulus components near the formant or the neuron's characteristic frequency accounted for the increase. In these neurons, synchronized response changes were closely related to the same neuron's average discharge rates [D. G. Sinex and L. P. McDonald, J. Acoust. Soc. Am. 83, 1817-1827 (1988)]. Neurons tuned to frequency regions near the second and third formants usually responded to components near the second formant prior to the onset of voicing. These neurons' synchronized discharges could be captured by the first formant at the onset of voicing or with a latency of 50-60 ms, whichever was later. Since these neurons' average rate responses were unaffected by the onset of voicing, the latency of the synchronized response did provide as much additional neural cue to VOT. Overall, however, discharge synchrony did not provide as much information about VOT as was provided by the best average rate responses. The results are compared to other measurements of the peripheral encoding of speech sounds and to aspects of VOT perception.

Responses of auditory-nerve fibers to consonant-vowel syllables.

The discharge patterns elicited by a set of synthesized consonant-vowel (CV) syllables were studied in the auditory nerve of the cat. The syllables, heard as /ba/, /da/, or /ga/, included a 25-, 50-, or 75-ms formant transition followed by a segment of steady-state vowel. The data were analyzed in terms of average discharge rate and in terms of the synchrony of discharges with respect to various spectral components of the stimuli. The results differ slightly from those of previous reports of the responses to vowels [Sachs and Young, J. Acoust. Soc. Am. 66, 470-479 (1979); Young and Sachs, J. Acoust. Soc. Am. 66, 1381-1403 (1979)], in that average discharge rates appear to provide more information about the spectra of formant transitions than they do about the spectra of steady-state vowels. This difference reflects changes in the spectrum of the syllable produced by movements of the formants. The synchrony of discharges, however, may provide more detailed information about the spectra of CVs than does average discharge rate. Each fiber's response at a particular peristimulus time may be characterized by the "dominant response component," the largest peak in the Fourier transform of the period histogram. The trajectories of the first three formants can be inferred from changes in the "dominant components" in a sample of fibers.

Comparison of click responses of primary auditory fibers with minimum-phase predictions.

Minimum-phase impulse responses were constructed from the frequency threshold-response curves of primary auditory fibers in the anesthetized cat. These impulse responses had many of the same characteristics as the experimental click responses. The two types of responses had similar inter-peak intervals as well as similar (+/- 1.5 ms) latencies to the principal mode and similar (+/- 1.0 ms) intervals from response onset to the principal mode.

Neural responses to the onset of voicing are unrelated to other measures of temporal resolution.

Voice onset time (VOT) is a temporal cue that can distinguish consonants such as /d/ from /t/. It has previously been shown that neurons' responses to the onset of voicing are strongly dependent on their static spectral sensitivity. This study examined the relation between temporal resolution, determined from responses to sinusoidally amplitude-modulated (SAM) tones, and responses to syllables with different VOTs. Responses to syllables and SAM tones were obtained from low-frequency neurons in the inferior colliculus (IC) of the chinchilla. VOT and modulation period varied from 10 to 70 ms in 10-ms steps, and discharge rates elicited by stimuli whose amplitude envelopes were modulated over the same temporal interval were compared. Neurons that respond preferentially to syllables with particular VOTs might be expected to respond best to the SAM tones with comparable modulation periods. However, no consistent agreement between responses to VOT syllables and to SAM tones was obtained. These results confirm the previous suggestion that IC neurons' selectivity for VOT is determined by spectral rather than temporal sensitivity.

Responses of primary auditory fibers to brief tone bursts.

Responses of primary auditory fibers to short triangularly modulated bursts of tone were obtained in the anesthetized cat. Based on discharge rate alone, the characteristics of the "response areas" obtained with these tone bursts were found to depend on the best frequency of the fiber. For a fiber with low best frequency (below 1 kHz), tones of greater than 10-ms duration had to be presented in order for the frequency resolution of the neuron to be as good as it was for long tones. For fibers with high best frequencies (above 10 kHz), tones of 2 or even 1 ms caused responses that were nearly as frequency selective as those obtained with long tones. A linear minimum-phase model based on the steady-state frequency selectivity of the fibers has been developed and shows generally comparable responses, but with some interesting exceptions. Synchronization of discharges to the waveform of the low-frequency tone bursts was measured and also shown to be generally compatible with the minimum-phase model. Trapezoidally modulated tone bursts were also used.

Auditory-nerve fiber representation of temporal cues to voicing in word-medial stop consonants.

The length of the interval between the onset of consonant closure and the onset of voicing in a following vowel is a temporal cue that may distinguish between consonants /d/ and /t/ in word-medial environments; this interval has been called the "consonant duration" [V. W. Zue and M. Laferriere, J. Acoust. Soc. Am. 66, 1039-1050 (1979); S. Davis and W. V. Summers, J. Phon. 17, 339-353 (1989)]. The representation of this cue in the discharge patterns of chinchilla auditory-nerve fibers was measured. The two-syllable utterances /ida/, /ita/, /uda/, and /uta/ were recorded by one male and one female talker. The onset of consonant closure produced discharge rate decreases in nearly all neurons. Either the release of closure or the onset of voicing for the second vowel could elicit an increase in discharge rate. The latencies of these discharge rate changes varied across populations of neurons. A neural measure of consonant duration was extracted from the pattern of latencies. The "encoded duration" was longer for utterances with a medial /t/ than for utterances with a medial /d/. For each utterance the encoded duration increased with increases in characteristic frequency. The variability of the encoded duration measure was small enough to preserve the distinction between utterances with different word-medial consonants. The variability of the encoded duration was large, relative to the acoustic differences between utterances that included the same medial consonant. This pattern of variability could contribute to the formation of perceptual categories by reducing the audibility of within-category acoustic differences.

Comparison of the responses of auditory nerve fibers to consonant-vowel syllables with predictions from linear models.

The responses of cat auditory-nerve fibers to synthesized consonant-vowel syllables were compared with predictions from linear models based on individual fibers' threshold tuning curves. Comparisons with the linear predictions provided information about the specific effects of peripheral nonlinearities on the representation of speech sounds. Spectral peaks, such as the formants of vowels, were more prominently represented in synchronized discharge patterns than in the linear predictions. Suppression of responses to other spectral peaks and to stimulus components between spectral peaks accounted for the differences. While profiles of fibers' synchronized responses were usually dominated by a single formant, predicted linear responses often included broad responses having two or more formants as well as components near the fibers' characteristic frequencies. In contrast, when no stimulus peak fell within a fiber's response area, the agreement between the neural response and the linear prediction was quite good. The results suggest that one role for peripheral nonlinearities in the auditory system may be to enhance the neural representation of spectral features such as formants.