Luis Robles

Basilar-membrane responses to tones at the base of the chinchilla cochlea.

Basilar-membrane responses to single tones were measured, using laser velocimetry, at a site of the chinchilla cochlea located 3.5 mm from its basal end. Responses to low-level (< 10-20 dB SPL) characteristic-frequency (CF) tones (9-10 kHz) grow linearly with stimulus intensity and exhibit gains of 66-76 dB relative to stapes motion. At higher levels, CF responses grow monotonically at compressive rates, with input-output slopes as low as 0.2 dB/dB in the intensity range 40-80 dB. Compressive growth, which is significantly correlated with response sensitivity, is evident even at stimulus levels higher than 100 dB. Responses become rapidly linear as stimulus frequency departs from CF. As a result, at stimulus levels > 80 dB the largest responses are elicited by tones with frequency about 0.4-0.5 octave below CF. For stimulus frequencies well above CF, responses stop decreasing with increasing frequency: A plateau is reached. The compressive growth of responses to tones with frequency near CF is accompanied by intensity-dependent phase shifts. Death abolishes all nonlinearities, reduces sensitivity at CF by as much as 60-81 dB, and causes a relative phase lead at CF.

Basilar membrane mechanics at the base of the chinchilla cochlea. I. Input-output functions, tuning curves, and response phases.

Basilar membrane (BM) velocity was measured at a site 3.5 mm from the basal end of the chinchilla cochlea using the Mössbauer technique. The threshold of the compound action potential recorded at the round window in response to tone bursts was used as an indicator of the physiological state of the cochlea. The BM input-output functions display a compressive nonlinearity for frequencies around the characteristic frequency (CF, 8 to 8.75 kHz), but are linear for frequencies below 7 and above 10.5 kHz. In preparations with little surgical damage, isovelocity tuning curves at 0.1 mm/s are sharply tuned, have Q10s of about 6, minima as low as 13 dB SPL, tip-to-tail ratios (at 1 kHz) of 56 to 76 dB, and high-frequency slopes of about 300 dB/oct. These mechanical responses are as sharply tuned as frequency-threshold curves of chinchilla auditory nerve fibers with corresponding CF. There is a progressive loss of sensitivity of the mechanical response with time for the frequencies around CF, but not for frequencies on the tail of the tuning curve. In some experiments the nonlinearity was maintained for several hours, in spite of a considerable loss of sensitivity of the BM response. High-frequency plateaus were observed in both isovelocity tuning curves and phase-frequency curves.

Evidence from Mössbauer experiments for nonlinear vibration in the cochlea

The Mossbauer technique has been applied to the measurement of vibration of the basilar membrane in the squirrel monkey cochlea. Both steady‐state and transient responses have been recorded in the 7–8‐kHz locus of the cochlea. The steady‐state response indicates that the basilar membrane vibrates nonlinearly for frequencies of stimulation near or greater than the characteristic frequency. The nonlinearity can be observed at the lowest levels of stimulation, 70–80 dB SPL, for which measurements could be made. The nonlinearity extends to lower frequencies and the basilar membrane transfer function tends to broaden as SPL is increased. Rapid postmortem changes occur in the cochlea: (1) the amplitude of the transfer ratio (basilar membrane/malleus) decreases 10–15 dB over a period of several hours with a downward shift of 1.5–3 kHz in the characteristic frequency of the basilar membrane at a given location; (2) the low‐frequency slope of the transfer ratio settles to 6 dB/octave by 6 h after death; (3) the sl...

Transient response of the basilar membrane measured in squirrel monkeys using the Mössbauer effect.

Measurements of the transient response of the basilar membrane were conducted using the Mossbauer effect on 33 squirrel monkeys using an experimental preparation identical to that of Rhode (1971). The stimuli were acoustic clicks 150 μsec in duration repeated 100 000–400 000 times. The amplitude of the click was varied and the responses of the malleus and of the basilar membrane at a point in the basal turn were measured. The basilar membrane’s click response is oscillatory, with a period near that of the characteristic frequency. The first few response peaks behave almost linearly with stimulus intensity, while the later peaks exhibit a pronounced nonlinearity. This behavior is shown to be consistent with the nonlinearity reported using steady‐state measurement methods (Rhode, 1971). The transient response observed in some of the preparations was very lightly damped; however, a wide range in the damping of the responses was found in the different animals. A progressive increase in the rate of decay of th...

Selective Attention to Visual Stimuli Reduces Cochlear Sensitivity in Chinchillas

It is generally accepted that during periods of attention to specific stimuli there are changes in the neural activity of central auditory structures; however, it is controversial whether attention can modulate auditory responses at the cochlear level. Several studies performed in animals as well as in humans have attempted to find a modulation of cochlear responses during visual attention with contradictory results. Here, we have appraised cochlear sensitivity in behaving chinchillas by measuring, with a chronically implanted round-window electrode, sound-evoked auditory-nerve compound action potentials and cochlear microphonics, a measure of outer hair cell function, during selective attention to visual stimuli. Chinchillas were trained in a visual discrimination or in an auditory frequency discrimination two-choice task. We found a significant decrease of cochlear sensitivity during the period of attention to visual stimuli in the animals performing the visual discrimination task, but not in those performing the auditory task, demonstrating that this physiological effect is related to selective attention to visual stimuli rather than to an increment in arousal level. Furthermore, the magnitude of the cochlear-sensitivity reductions increased in sessions performed with shorter target-light durations (4–0.5 s), suggesting that this effect is stronger for higher attentional demands of the task. These results demonstrate that afferent auditory activity is modulated by selective attention as early as at sensory transduction, possibly through activation of olivocochlear efferent fibers.

Basilar membrane mechanics at the base of the chinchilla cochlea. II. Responses to low‐frequency tones and relationship to microphonics and spike initiation in the VIII Nerve

Low-frequency stimuli (40- to 1000-Hz tones) have been used to correlate the motion of the 8-to 9-kHz place of the chinchilla basilar membrane with the cochlear microphonics recorded at the round window and with the responses of auditory nerve fibers with appropriate characteristic frequency. At the lowest stimulus frequencies, maximum displacement of the basilar membrane toward scala tympani occurs in near synchrony with maximum rarefaction at the eardrum and maximum negativity at the round window; at higher frequencies, the mechanical and microphonic response phases progressively lag rarefaction, reaching - 240 deg at 1000 Hz. At most frequencies (40-1000 Hz) near-threshold neural responses, once corrected for neural travel-time and synaptic delays, somewhat lead (by some 40 deg) maximal scala tympani displacement and maximal negativity of the round window microphonics. The variation of sensitivity with frequency is similar for basilar membrane displacement and microphonic responses: Under open-bulla conditions, sensitivity is constant for frequencies between 100 and 1000 Hz; below 100 Hz, sensitivity decreases at rates close to 12 dB/oct toward lower frequencies. Neural response sensitivity matches BM displacement more closely than BM velocity.

Nonlinear Interactions in the Mechanical Response of the Cochlea to Two-Tone Stimuli

The reduction of the auditory response to one tone due to the simultaneous presence of a second tone is known as two-tone suppression. This nonlinear cochlear response was first described by Galambos and Davis (1944) in electrophysiological recordings from the cat auditory nerve. Since then two-tone suppression has been extensively studied in cochlear afferents of mammals (Nomoto et al., 1964; Kiang et al., 1965; Hind et al., 1967; Sachs and Kiang, 1968) and also of amphibia (Frischkopf and Goldstein, 1963; Feng et al., 1975), birds (Sachs et al., 1974) and reptiles (Holton and Weiss, 1978). Two-tone suppression seems to be demonstrable in all well-studied cochlear fibers, for suppressors at frequencies on both sides of the tuning curve (Sachs and Kiang, 1968; Sachs, 1969; Abbas and Sachs, 1976), and has also been observed in guinea pig inner hair cells (Sellick and Russell, 1979).

Saccadic modulation of cell discharges in the lateral geniculate nucleus

Abstract The effect of fast phases of vestibular nystagmus on evoked and spontaneous discharges of neurons in the lateral geniculate nucleus of the rat have been studied. In order to determine the central influence of saccadic activity, the receptive fields were stimulated in a fixed eye after sectioning the extraocular muscles. Data analysis was carried out using a LINC computer. Results show that the probability of firing of “concentric” cells decreases at about 30–40 msec to 160–200 msec after the onset of saccades, under a wide range of illumination and with different visual patterns. The effect was the opposite for “on-off” type of cells i.e. an increase in discharge frequency occurred with a maximum at about 150 msec after saccades. Directional cells were affected by saccades depending upon their preferred directions. The time-course of the effect correlates with the saccadic slow waves in the LGN (previously described), and with the psychophysical saccadic suppression phenomenon, when latency of transmission from retina to LGN is considered. A neuronal mechanism based on the selective effect on directional cells is proposed to account for the stabilization of the visual world during eye movements.

The Olivocochlear Reflex Strength and Cochlear Sensitivity are Independently Modulated by Auditory Cortex Microstimulation

In mammals, efferent projections to the cochlear receptor are constituted by olivocochlear (OC) fibers that originate in the superior olivary complex. Medial and lateral OC neurons make synapses with outer hair cells and with auditory nerve fibers, respectively. In addition to the OC system, there are also descending projections from the auditory cortex that are directed towards the thalamus, inferior colliculus, cochlear nucleus, and superior olivary complex. Olivocochlear function can be assessed by measuring a brainstem reflex mediated by auditory nerve fibers, cochlear nucleus neurons, and OC fibers. Although it is known that the OC reflex is activated by contralateral acoustic stimulation and produces a suppression of cochlear responses, the influence of cortical descending pathways in the OC reflex is largely unknown. Here, we used auditory cortex electrical microstimulation in chinchillas to study a possible cortical modulation of cochlear and auditory nerve responses to tones in the absence and presence of contralateral noise. We found that cortical microstimulation produces two different peripheral modulations: (i) changes in cochlear sensitivity evidenced by amplitude modulation of cochlear microphonics and auditory nerve compound action potentials and (ii) enhancement or suppression of the OC reflex strength as measured by auditory nerve responses, which depended on the intersubject variability of the OC reflex. Moreover, both corticofugal effects were not correlated, suggesting the presence of two functionally different efferent pathways. These results demonstrate that auditory cortex electrical microstimulation independently modulates the OC reflex strength and cochlear sensitivity.

Stronger efferent suppression of cochlear neural potentials by contralateral acoustic stimulation in awake than in anesthetized chinchilla

There are two types of sensory cells in the mammalian cochlea, inner hair cells, which make synaptic contact with auditory-nerve afferent fibers, and outer hair cells that are innervated by crossed and uncrossed medial olivocochlear (MOC) efferent fibers. Contralateral acoustic stimulation activates the uncrossed efferent MOC fibers reducing cochlear neural responses, thus modifying the input to the central auditory system. The chinchilla, among all studied mammals, displays the lowest percentage of uncrossed MOC fibers raising questions about the strength and frequency distribution of the contralateral-sound effect in this species. On the other hand, MOC effects on cochlear sensitivity have been mainly studied in anesthetized animals and since the MOC-neuron activity depends on the level of anesthesia, it is important to assess the influence of anesthesia in the strength of efferent effects. Seven adult chinchillas (Chinchilla laniger) were chronically implanted with round-window electrodes in both cochleae. We compared the effect of contralateral sound in awake and anesthetized condition. Compound action potentials (CAP) and cochlear microphonics (CM) were measured in the ipsilateral cochlea in response to tones in absence and presence of contralateral sound. Control measurements performed after middle-ear muscles section in one animal discarded any possible middle-ear reflex activation. Contralateral sound produced CAP amplitude reductions in all chinchillas, with suppression effects greater by about 1–3 dB in awake than in anesthetized animals. In contrast, CM amplitude increases of up to 1.9 dB were found in only three awake chinchillas. In both conditions the strongest efferent effects were produced by contralateral tones at frequencies equal or close to those of ipsilateral tones. Contralateral CAP suppressions for 1–6 kHz ipsilateral tones corresponded to a span of uncrossed MOC fiber innervation reaching at least the central third of the chinchilla cochlea.