A physiological frequency-position map of the chinchilla cochlea.

Accumulating evidence indicates that mammalian cochlear frequency-position maps (location of maximum vibration of the basilar membrane as a function of frequency) depend on the physiological condition of the inner ear. Cochlear damage desensitizes the ear, after the damage the original location of maximum vibration is tuned to a lower sound frequency. This suggests that frequency-position maps, derived from such desensitized ears, are shifted to lower frequencies, corresponding to a shift of the basilar membrane vibration pattern towards the base for a given stimulus frequency. To test this hypothesis, we re-mapped the cochlear frequency-position map in the chinchilla. We collected frequency-position data from chinchillas in normal physiological condition ("physiological map") and compared these to data previously established from sound overexposed ears ("anatomical map"). The characteristic frequency (CF) of neurons in the cochlear nucleus was determined. Horse-radish peroxidase (HRP) or biocytin (BCT) were injected iontophoretically to trace auditory nerve fibers towards their innervation site in the organ of Corti. The relationship between distance from the base (d, percent) and frequency (f, kHz) was described best by a simple exponential function: d = 61.2 - 42.2 x log(f). The slope of the function was 2.55 mm/octave. Compared to the "anatomical map", the "physiological map" was shifted by about 0.3 octaves to higher frequencies corresponding to a shift of the basilar membrane vibration pattern of 0.8 mm towards the apex for a given stimulus frequency. Our findings affirm that frequency-position maps in the mammalian cochlea depend on the condition of the inner ear. Damage-induced desensitization in mammalian inner ears results in similar shifts of CF (about 0.5 octaves) but different shifts of the maximum of the vibration pattern towards the base at given frequencies, dependent on the mapping constant of the species, longer basilar membranes showing a larger basal shift. Furthermore, the results substantiate the notion that "crowding" at lower frequencies appears to be a specialization rather than a general feature.

Reduction of inhibition in the inferior colliculus after inner hair cell loss.

Effects of carboplatin-induced partial loss of inner hair cells on rate-level functions of neurons in the central nucleus of the inferior colliculus of the same chinchillas before and 1-3 months after carboplatin treatment were examined. Partial inner hair cell loss caused only a small elevation of response thresholds but induced a significant increase in the proportion of monotonic and a significant decrease in the proportion of non-monotonic rate-level functions at the characteristic frequency of 210 multiunits. This indicates that inhibition in the inferior colliculus is reduced after sensory deafferentation. We conclude that some of the functional changes reported in the auditory cortex after peripheral deafferentation are already realized at the level of the inferior colliculus.

Inhibitory and excitatory response areas of neurons in the central nucleus of the inferior colliculus in unanesthetized chinchillas.

In unanesthetized chinchillas, we determined excitatory and inhibitory response regions of neurons in the central nucleus of the inferior colliculus (ICc). The responses of 250 multiunits and 47 single units in the ICc to one- and two-tone stimuli were measured by extracellular recordings. The one-tone excitatory response area of ICc neurons from awake chinchillas was classified as either narrow with a steep high-frequency slope >140 dB/oct (type 1), broad with a high-frequency slope <140 dB/oct (type 2), or complex with a negative high-frequency slope (type 3). One-tone inhibition was prominent only in units with a high spontaneous firing rate. As revealed with two-tone stimuli, inhibition in the ICc of awake chinchillas and its relation to excitatory response regions was different from what is reported in anesthetized animals. The two-tone inhibitory responses were classified as follows: (1) inhibitory regions of equal strength on both sides of the characteristic frequency; (2) asymmetrical inhibitory regions, more prominent at the high-frequency side of the characteristic frequency; (3) strong inhibitory regions overlying most of the one-tone excitatory response region; (4) inhibitory response regions lying only within the one-tone excitatory response region; and (5) neurons without clear two-tone inhibition. One-tone and two-tone inhibitory regions of the same unit were markedly different in 66% of the units with a high spontaneous rate. The neural response to frequencies within the inhibitory regions often was an onset response followed by inhibition. Excitatory and inhibitory response properties were similar over considerable penetration distances (600-1,000 microm) in a particular dorso-ventral recording track.

Shift in the cochlear place-frequency map after noise damage in the mouse.

The cochlear place-frequency map, determined from noise-damaged mice, is shifted to lower frequencies by up to one octave compared with the map determined from normal hearing mice. To test the hypothesis that the shift results from damage to the cochlear amplifier, we measured frequency tuning curves from the same neurons before and after noise exposure. Noise damage resulted in loss of tuning and elevation of thresholds. The neuronal characteristic frequency shifted by 0.6-1.2 octaves, dependent on frequency. The shift in characteristic frequency was used to calculate a shifted place-frequency map. We conclude that desensitization of areas in the inner ear after noise exposure can explain the shift of the map after noise damage relative to the normal physiological map.

A physiological place-frequency map of the cochlea in the CBA/J mouse.

Genetically manipulated mice have gained a prominent role in in vivo research on development and function of the auditory system. A prerequisite for the interpretation of normal and abnormal structural and functional features of the inner ear is the exact knowledge of the cochlear place-frequency map. Using a stereotaxic approach to the projection site of the auditory nerve fibers in the cochlear nucleus, we succeeded in labelling physiologically characterized auditory nerve afferents and determined their peripheral innervation site in the cochlea. From the neuronal characteristic frequency (CF) and the innervation site in the organ of Corti a place-frequency map was established for characteristic frequencies between 7.2 and 61.8 kHz, corresponding to locations between 90% and 10% basilar membrane length (base = 0%, apex = 100%, mean length measured under the inner hair cells 5.13 mm). The relation between normalized distance from the base (d) and frequency (kHz) can be described by a simple logarithmic function: d(%) = 156.5-82.5 x log(f), with a slope of 1.25 mm/octave of frequency. The present map, recorded under physiological conditions, differs from earlier maps determined with different methods. The simple logarithmic place-frequency relation found in the mouse indicates that mice are acoustic generalists rather than specialists.