The amplitude and the phase of vibration of the basilar membrane and the bony limbus of the cochlea were measured in living squirrel monkeys using the Mössbauer technique. In the middle ear, the vibration of the malleus (and occasionally the incus) was measured. The Mössbauer technique makes possible the measurement of very small velocities, e.g., 0.2 mm/sec. This sensitivity permits measurement of the motion of the malleus at sound-pressure levels (SPLs) of 90 to 110 dB and measurement of the motion of the basilar membrane at 70 to 120 dB SPL, depending on the frequency. The basilar membrane vibrates nonlinearly for frequencies which produce the largest deflections at the spot on the basilar membrane under observation. The ratio of the displacement of the basilar membrane to that of the malleus was observed to have the following characteristics: (1) As the frequency is increased from a low value, its amplitude increases at 6 dB/oct until just below the maximum ratio where the slope increases to about 24 dB/oct; (2) the maximum ratio was about 24 dB for the SPLs used; (3) for frequencies above that producing the maximum ratio, the drop-off rate was approximately 100 dB/oct; (4) the amplitude ratio did not drop off indefinitely but tended to level off; (5) the motion of the basilar membrane differs from the motion of the malleus by 90° at very low frequencies; (6) for frequencies below that producing the maximum ratio, the phase differences between the motion of the basilar membrane and that of the malleus is a linear function of frequency; (7) near the frequency corresponding to the maximum ratio, the phase difference decreases at a faster rate; and (8) the phase difference approaches a constant value (7π 8π or 9π) for high frequencies. Anatomical constraints allowed only a small portion of the basal turn to be studied (6.5–7.5 kHz produced maximum deflection of the basilar membrane in this region).

Physiological response properties of neurons in the ventral cochlear nucleus have a variety of features that are substantially different from the stereotypical auditory nerve responses that serve as the principal source of activation for these neurons. These emergent features are the result of the varying distribution of auditory nerve inputs on the soma and dendrites of the various cell types within the nucleus; the intrinsic membrane characteristics of the various cell types causing different responses to the same input in different cell types; and secondary excitatory and inhibitory inputs to different cell types. Well-isolated units were recorded with high-impedance glass microelectrodes, both intracellularly and extracellularly. Units were characterized by their temporal response to short tones, rate vs. intensity relation, and response areas. The principal response patterns were onset, chopper, and primary-like. Onset units are characterized by a well-timed first spike in response to tones at the characteristic frequency. For frequencies less than 1 kHz, onset units can entrain to the stimulus frequency with greater precision than their auditory nerve inputs. This implies that onset units receive converging inputs from a number of auditory nerve fibers. Onset units are divided into three subcategories, OC, OL, and OI. OC units have extraordinarily wide dynamic ranges and low-frequency selectivity. Some are capable of sustaining firing rates of 800 spikes/s at high intensities. They have the smallest standard deviation and coefficient of variation of the first spike latency of any cells in the cochlear nuclei. OC units are candidates for encoding intensity. OI and OL units differ from OC units in that they have dynamic ranges and frequency selectivity ranges much like those of auditory nerve fibers. They differ from one another in their steady-state firing rates; OI units fire mainly at the onset of a tone. OI units also differ from OL units in that they prefer frequency sweeps in the low to high direction. Primary-like-with-notch (PLN) units also respond to tones with a well-timed first spike. They differ from onset cells in that the onset peak is not always as precise as the spontaneous rate is higher. A comparison of spontaneous firing rate and saturation firing rate of PLN units with auditory nerve fibers suggest that PLN units receive one to four auditory nerve fiber inputs. Chopper units fire in a sustained regular manner when they are excited by sound.(ABSTRACT TRUNCATED AT 400 WORDS)

We distinguish two types of large multipolar cells designated sustained (CS) and onset (OC) choppers in the anterior posteroventral cochlear nucleus (A-PVCN)/nerve root region on the basis of certain anatomical and physiological features. CS axons head into the trapezoid body, while OC axons use the intermediate acoustic stria of Held. At the electron microscopic (EM) level, collateral terminals of OC axons contain pleomorphic vesicles; CS terminals contain small round vesicles. CS dendritic trees tend to be distributed in a stellate fashion while OC dendritic trees tend to be elongated. At the EM level the sustained chopper somata are sparsely innervated while the proximal dendritic tree receives considerably more input. The OC somata are highly innervated and this heavy innervation continues out onto the proximal dendrites. Distally the dendritic innervation falls off considerably for both categories. Physiologically, members of the OC population have wider dynamic ranges at the characteristic frequency (CF), wider response areas that are typically not flanked by inhibitory sidebands, and responses to short tones that do not show the same form of regularity expressed by sustained choppers. Intracellularly the sustained choppers exhibit sustained depolarization to short tones for the duration of the stimulus with resultant regular spiking at a rate that is stimulus level dependent. The response to swept tone shows this same level-dependent regularity. In response to tones, the OC cells also show a sustained depolarization whose amplitude is stimulus-level dependent but whose range is much greater and whose onset is initiated more abruptly. Although the onset component of the OC spike output is reliably initiated by these levels of depolarization, regular firing to the sustained depolarization is not initiated at levels of depolarization that would surely generate regular firing in sustained choppers. This regularity is also absent in the swept tone response despite marked levels of excitation.

To determine the correspondence between anatomical and physiological cell types in the ventral cochlear nucleus of the cat, intracellular injections of horseradish peroxidase were made into cells whose extracellular and intracellular responses to sound had been studied. Three identified cells responded to a short tone burst at their characteristic frequencies with an onset pattern. This pattern is characterized by a strong response to the onset of the stimulus. One was an octopus cell. The second cell, located in the octopus-cell area, was a giant cell with a few somatic spines and thin tapering dendrites; the intracellular record revealed that even in the absence of sound it received continuous synaptic input, while tones at characteristic frequency produced a sustained depolarization. A third cell, which had an onset response at low intensities and a chopper response at high intensities, was a stellate cell located in the intermediate acoustic stria with dendrites oriented parallel to the fiber tract. This cell had an unusually broad dynamic range in response to changes in intensity. Two cells with transient chopper response patterns were stellate cells in the posteroventral cochlear nucleus with many branched and beaded dendrites. Three cells with more sustained chopper response patterns were stellate cells in the anteroventral cochlear nucleus with fewer, less-branched, smooth dendrites. Two cells with primarylike responses to tones were bushy cells with numerous short, thin, highly branched dendrites in the posterior division of the anteroventral cochlear nucleus. Intracellular responses to tones at the characteristic frequency consisted of large brief depolarizations, which were not sustained. Another cell, which responded to tones in a phase-locked fashion, was also located in the anteroventral cochlear nucleus. It was a small, stellate cell with relatively few, smooth dendrites. The labeled cells largely support previous attempts at physiological-morphological correlations: (1) bushy cells exhibit primarylike pattern; (2) stellate cells exhibit chopper patterns; and (3) octopus cells exhibit an onset pattern. It was also demonstrated that more than one cell type can exhibit a particular response pattern.

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

Measurements from the 1-4-mm basal region of the chinchilla cochlea indicate the basilar membrane in the hook region (12-18 kHz) vibrates essentially as it does more apically, in the 5-9-kHz region. That is, a compressive nonlinearity in the region of the characteristic frequency, amplitude-dependent phase changes, and a gain relative to stapes motion that can attain nearly 10,000 at low levels. The displacement at threshold for auditory-nerve fibers in this region (20 dB SPL) was approximately 2 nm. Measurements were made at several locations in individual animals in the longitudinal and radial directions. The results indicate that there is little variability in the phase of motion radially and no indication of higher-order modes of vibration. The data from the longitudinal studies indicate that there is a shift in the location of the maximum with increasing stimulus levels toward the base. The cochlear amplifier extends over a 2-3-mm region around the location of the characteristic frequency.

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Mechanical responses to one- and two-tone acoustic stimuli were recorded from the cochlear partition in the apical turn of the chinchilla cochlea, the basal turn of the guinea pig cochlea, and the hook region of the guinea pig cochlea. The most sensitive or "best" frequencies (BFs) for the sites studied were approximately 500 Hz, 17 kHz, and 30 kHz, respectively. Responses to the cubic difference tone (CDT), 2F1 - F2 (where F1 and F2 are the frequencies of the primary stimuli), were characterized at each site. Responses to the quadratic difference tone (QDT), F2 - F1, were also characterized in the apical turn preparations (QDT responses were too small to measure in the basal cochlea). The observed responses to BF QDTs and CDTs and to BF CDTs at each site appeared similar in many ways; the relative magnitudes of the responses were highest at low-to-moderate sound pressure levels (SPLs), for example, and the absolute magnitudes grew nonmonotonically with increases in the level of either primary (L1 or L2) alone. The peak effective levels of the CDT and QDT responses were also similar, at around -20 dB re L1 and/or L2. In other respects, however, the responses to CDTs and QDTs and to BF CDTs at each site behaved quite differently. At low-to-moderate SPLs, for example, most CDT phase leads decreased with increases in either L1 or L2, whereas most QDT phase leads increased with increasing L1 and varied little with L2. Most CDT responses also varied monotonically with equal-level primaries (i.e., when L1 = L2), whereas most QDT responses varied nonmonotonically. Different responses also varied in different ways when F1 and F2 were varied. Apical turn QDT responses were observed over a very wide F1/F2 range (F1 = 1-12 kHz), but were usually largest for stimuli <2-4 kHz. Apical turn CDT levels decreased (at rates of approximately 40-80 dB/octave) only when the frequency ratio F2/F1 increased beyond approximately 1.4-1.5. In the basal turn and hook regions, the CDT levels depended nonmonotonically on F2/F1 with the eventual rates of decrease being approximately 200 dB/octave. Optimal frequency ratios for the CDT increased from (F2 < 1.1F1) to (F2 approximately 1.2F1) with increasing SPL in the basal turn, but were stable at around F2/F1 approximately 1.05 in the hook region. CDT phase leads tended to increase with increasing F2/F1 in all three regions of the cochlea, particularly at low-to-moderate SPLs. These findings are discussed in relation to previous studies of cochlear mechanics, physiology, and psychophysics.