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PubMed1986

Analysis of cochlear mechanics.

A large number of experimental results on basilar-membrane vibration, cochlear microphonics, hair-cell receptor potentials, and spike rates in auditory nerve afferents are brought together to arrive at a comprehensive concept of cochlear mechanics, including hair-cell stimulation. Beginning with basilar-membrane tuning curves, we note that some of their most detailed determinations reveal a small notch and a secondary maximum above the best frequency in addition to sharp tuning. These features tend to become more prominent as the sharpness of tuning decreases. They cannot be accounted for on the assumption that the cochlear partition represents a simple second-order system consisting of distributed, elastically suspended mass. A higher order system is required. The cross-sectional structure of the partition suggests a fifth-order system made up of two sets of distributed resonators, one consisting essentially of the distributed mass of the organ of Corti supported by the stiffness of the basilar membrane, the other of the tectorial-membrane mass and its elastic attachment to the spiral limbus. The stiff stereocilia of the outer hair cells appear to serve as the main elastic coupling between the two resonator sets. Interaction of the two resonator sets is brought into evidence particularly clearly by weakening the coupling between the tectorial membrane and the organ of Corti. This can be achieved by manipulating the tectorial membrane with a microelectrode without affecting the endolymphatic potential. The partial decoupling leads to a transformation of a unimodal CM transfer function into a bimodal one. Except for the stiffness of the tectorial-membrane attachment to the limbus, the masses and stiffnesses involved in the two coupled resonator systems can be estimated independently on the basis of available measurements. Their application to an approximate computer model of the cochlea produced a cochlear frequency map consistent with experimental findings. The computer model, whose elements are in one-to-one correspondence with the gross elements of the cochlear partition, reproduces approximately the fundamental amplitude and phase characteristics of the measured basilar-membrane vibration. In particular, it reproduces the notch and the secondary maximum located above the best frequency. Our current computer model is linear and does not reproduce the known cochlear distortion products. Nevertheless, variation of those of its stiffness and resistance elements that correspond to the hair-cell stereocilia has allowed us to reproduce typical changes in basilar-membrane vibration, which accompany changes in sound intensity or cochlear deterioration.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1992

Electron-microscopic localization of type II, IX, and V collagen in the organ of Corti of the gerbil.

The presence of types II, IX and V collagen was probed in the organ of Corti of the adult gerbil cochlea by use of immunocytochemistry at the light- and electron-microscopic levels. Type II collagen is found in the connective tissues of the osseous spiral lamina and spiral limbus. In the region of the sensory hair cells it is present in the tectorial membrane and antibodies bind to the thick unbranched radial fibers. Type IX collagen co-localizes with type II collagen in the tectorial membrane, where antibodies bind to the thick unbranched radial fibers. Type V collagen is present in the connective tissue of the spiral limbus, the osseous spiral lamina, the eighth nerve, and the tectorial membrane. In the tectorial membrane, the staining with antibodies to type V collagen is more diffuse than that seen for types II and IX collagen and antibodies to type V bind to the thin, highly branched fibers in which the thick fibers are embedded. The results indicate that collagens characteristic of cartilage are localized in the organ of Corti. Within the tectorial membrane, types II and IX collagen form heterotypic thick fibers embedded in a reticular network of type V collagen fibers. These collagens form a highly structured matrix which contributes to the rigidity of the tectorial membrane and allow it to withstand the physical stresses associated with transmission of the stimuli necessary for sensory transduction.

Animals↗

PubMed1986

Ionic environment of cochlear hair cells.

The scala media of the adult cochlea in mammals comprises a morphologically closed compartment sealed with tight junctions of the intermediate to tight types. The unique ionic composition of endolymph is maintained by the stria vascularis through active reabsorption of sodium and active secretion of potassium against ionic gradients. The subtectorial space is only a partially closed compartment which communicates with the endolymph via holes in the tectorial membrane at its outer insertion to the organ of Corti. Hardesty's membrane divides the subtectorial space into two compartments: one facing the surfaces of inner hair cells and one facing the surfaces of outer hair cells. In the study of comparative anatomy, hair cells, e.g. in the lizard, basilar papilla are of two types: those covered with a tectorial membrane and those being free-standing lacking the tectorial membrane. The ionic environment of the hair cell surface seems to be the same, independent of whether covered with a tectorial membrane or not. The tectorial membrane itself is semipermeable to ions in the endolymphatic space. Only the surface structures of the hair cell with the sensory hairs facing the subtectorial space are exposed to the high concentration of potassium, whereas the remaining parts of the hair cell are surrounded by a fluid having a more normal extracellular type of ionic composition (cortilymph/perilymph). During embryonic development the ionic composition of endolymph develops in parallel with the morphologic maturation of the stria vascularis. A completely mature composition of endolymph is reached before any electrophysiological potentials in the cochlea can be elicited. The sensory hair surface of hair cells has reached a mature morphology prior to the maturation of endolymph. In several species the tectorial membrane is morphologically only partially mature when the increase of the potassium concentration of endolymph starts. Drugs primarily affecting the stria vascularis causing a transient change of the ionic composition of endolymph result in a transient dysfunction of inner ear potentials. If the ionic changes persist for longer time, morphological changes can occur in both the stria vascularis and the hair cells of the organ of Corti. Whether such changes are primarily caused by the ototoxic drug itself or by changes in the ionic composition of endolymph has to be explored further.

Animals↗

PubMed2000

Ultrastructural changes in the spiral limbus associated with carboplatin-induced ablation of inner hair cells.

Four months after the selective ablation of inner hair cells by carboplatin, the interdental cell epithelium exhibited dilated intercellular spaces and cytosolic vacuoles not seen in controls. In addition, the wide, often electron-lucent phalanges observed in the interdental cells of the normal chinchilla collapsed into a dense stratum that projected enlarged polypoid profiles into the limbal zone of the tectorial membrane. Carboplatin treatment also resulted in the restructuring of the tectorial membrane overlying the limbus. Changes in this membrane included a variable accumulation of the basal matrix, the rearrangement of intermediate lucent spaces, and the disappearance of a superimposed filamentous mesh. These three strata are, under normal conditions, apparently involved in events underlying tectorial membrane renewal. The post-carboplatin changes in the interdental cells and tectorial membrane occurred exclusively in the proposed medial pathway for K+ diffusion from inner hair cells and presumably resulted from a reduced flow of ions and fluid secondary to the ablation of these cells.

Animals↗

PubMedDate not supplied

Functional recovery in the avian ear after hair cell regeneration.

Trauma to the inner ear in birds, due to acoustic overstimulation or ototoxic aminoglycosides, can lead to hair cell loss which is followed by regeneration of new hair cells. These processes are paralleled by hearing loss followed by significant functional recovery. After acoustic trauma, functional recovery is rapid and nearly complete. The early and major part of functional recovery after sound trauma occurs before regenerated hair cells become functional. Even very intense sound trauma causes loss of only a proportion of the hair cell population, mainly so-called short hair cells residing on the abneural mobile part of the avian basilar membrane. Uncoupling of the tectorial membrane from the hair cells during sound overexposure may serve as a protection mechanism. The rapid functional recovery after sound trauma appears not to be associated with regeneration of the lost hair cells, but with repair processes involving the surviving hair cells. Small residual functional deficits after recovery are most likely associated with the missing upper fibrous layer of the tectorial membrane which fails to regenerate after sound trauma. After aminoglycoside trauma, functional recovery is slower and parallels the structural regeneration more closely. Aminoglycosides cause damage to both types of hair cells, starting at the basal (high frequency) part of the basilar papilla. However, functional hearing loss and recovery also occur at lower frequencies, associated with areas of the papilla where hair cells survive. Functional recovery in these low frequency areas is complete, whereas functional recovery in high frequency areas with complete hair cell loss is incomplete, despite regeneration of the hair cells. Permanent residual functional deficits remain. This indicates that in low frequency regions functional recovery after aminoglycosides involves repair of nonlethal injury to hair cells and/or hair cell-neural synapses. In the high frequency regions functional recovery involves regenerated hair cells. The permanent functional deficits after the regeneration process in these areas are most likely associated with functional deficits in the regenerated hair cells or shortcomings in the synaptic reconnections of nerve fibers with the regenerated hair cells. In conclusion, the avian inner ear appears to be much more resistant to trauma than the mammalian ear and possesses a considerable capacity for functional recovery based on repair processes along with its capacity to regenerate hair cells. The functional recovery in areas with regenerated hair cells is considerable but incomplete.

Aminoglycosides↗

PubMedDate not supplied

Observations of the organ of Corti under in vivo-like conditions.

The organ of Corti of the rat has been investigated within minutes after death in different fluids bathing the organ of Corti. Static compliance of the tectorial membrane across the membrane was measured by microsprings, and the response of the membrane to a vibrating micro-electrode was studied. In addition, fluid motion about the organ caused by the placement of a vibrating micro-electrode on the tectorial membrane was observed. The static compliance of the tectorial membrane was unaffected by the bathing solutions. Dynamic characteristics of the membrane could be completed altered by the addition of aldehyde solutions. As the cochleas used by von Békésy (1960) were preserved in formaldehyde solutions, it is concluded that his observations regarding the dynamic characteristics of the tectorial membrane were in part artifact. It is suggested that a proper description of the mechanisms of the organ of Corti must include a tectorial membrane whose limbal attachment allows for both rotation and displacement. Such a description is presented. A streaming of fluid was observed directed from Hensen's cells toward the cochlea wall. Possible mechanisms for the generation of this streaming are discussed.

Animals↗

PubMed2000

Spectrum of occipitoatlantoaxial injury in young children.

OBJECT: Injuries of the occipitoatlantoaxial (Oc-C2) region are the predominant form of cervical injury in children younger than 10 years of age. Magnetic resonance (MR) imaging can be used to visualize directly the traumatic ligamentous and soft-tissue abnormalities of the Oc-C2 region. A retrospective review was undertaken to examine the spectrum of pediatric Oc-C2 injuries seen on MR imaging, their correlation with plain x-ray film and computerized tomography findings, and their clinical course. METHODS: Seventy-one consecutive children younger than 10 years of age underwent cervical MR imaging for evaluation of traumatic injury. Magnetic resonance imaging was used to document abnormalities in 23 children; 20 of these injuries involved the Oc-C2 region. Abnormalities in the Oc-C2 region included disruptions of the musculature, apical ligament, atlantooccipital joint(s), tectorial membrane, and spinal cord. A spectrum of injury with progressive involvement of these structures was seen, ranging from isolated muscular injury to the multiple soft-tissue and ligamentous disruptions with craniocervical dislocation. Involvement of the tectorial membrane was the critical threshold in the transition from stable to unstable injury. Analysis of plain x-ray films revealed that a novel interspinous C1-2:C2-3 ratio criteria of greater than or equal to 2.5 was predictive of tectorial membrane abnormalities on MR imaging, with 87% sensitivity and 100% specificity. In patients with tectorial membrane abnormalities who underwent immobilization alone, interim platybasia was demonstrated on follow-up MR images. Conclusions. A progressive spectrum of distinct Oc-C2 injuries can occur in young children; the tectorial membrane is a critical stabilizing ligamentous structure in the Oc-C2 complex; and tectorial membrane abnormalities may be identified by a C1-2:C2-3 ratio of greater than or equal to 2.5.

Atlanto-Axial Joint↗

PubMed1996

Active control of waves in a cochlear model with subpartitions.

Multiscale asymptotic methods developed previously to study macromechanical wave propagation in cochlear models are generalized here to include active control of a cochlear partition having three subpartitions, the basilar membrane, the reticular lamina, and the tectorial membrane. Activation of outer hair cells by stereocilia displacement and/or by lateral wall stretching result in a frequency-dependent force acting between the reticular lamina and basilar membrane. Wavelength-dependent fluid loads are estimated by using the unsteady Stokes' equations, except in the narrow gap between the tectorial membrane and reticular lamina, where lubrication theory is appropriate. The local wavenumber and subpartition amplitude ratios are determined from the zeroth order equations of motion. A solvability relation for the first order equations of motion determines the subpartition amplitudes. The main findings are as follows: The reticular lamina and tectorial membrane move in unison with essentially no squeezing of the gap; an active force level consistent with measurements on isolated outer hair cells can provide a 35-dB amplification and sharpening of subpartition waveforms by delaying dissipation and allowing a greater structural resonance to occur before the wave is cut off; however, previously postulated activity mechanisms for single partition models cannot achieve sharp enough tuning in subpartitioned models.

Animals↗

PubMed2002

Extracellular matrices associated with the apical surfaces of sensory epithelia in the inner ear: molecular and structural diversity.

The ultrastructure and molecular composition of the extracellular matrices that are associated with the apical surfaces of the mechanosensory epithelia in the mouse inner ear are compared. A progressive increase in molecular and structural organization is observed, with the cupula being the simplest, the otoconial membrane exhibiting an intermediate degree of complexity, and the tectorial membrane being the most elaborate of the three matrices. These differences may reflect changes that occurred in the acellular membranes of the inner ear as a mammalian hearing organ arose during evolution from a simple equilibrium receptor. A comparison of the molecular composition of the acellular membranes in the chick inner ear suggests the auditory epithelium and the striolar region of the maculae are homologous, indicating the basilar papilla may have evolved from the striolar region of an otolithic organ. A comparison of the tectorial membranes in the chick cochlear duct and the mouse cochlea reveals differences in the structure of the noncollagenous matrix in the two species that may result from differences in the stochiometry of alpha- and beta-tectorin and/or differences in the post-translational modification of alpha-tectorin. This comparison also indicates that the appearance of collagen in the mammalian tectorial membrane may have been a major step in the evolution of an electromechanically tuned vertebrate hearing organ that operates over an extended frequency range.

Acoustic Maculae↗

PubMedDate not supplied

Elemental composition of individual cells and tissues in the cochlea.

Localization of elements at the cellular and sub-cellular levels was performed with the energy dispersive X-ray microanalysis technique, using shock-frozen, freeze-dried and araldite-embedded mouse (CBA/CBA) cochleae sectioned dry. Anatomical identification occurred in the STEM (scanning transmission electron microscopy) mode. In inner hair cell stereocilia the K/Na ratio was 70:1 but only 20:1 in the cytoplasm. In outer hair cell cytoplasm the K/Na ratio was 11:1 while the ratio in stereocilia was similar to that in inner hair cells. Ca was identified in stereocilia and the upper part of the cytoplasm of both outer and inner hair cells. The elemental composition in the subtectorial space is endolymph-like and that in the inner tunnel of the organ of Corti is similar to extracellular fluid. Considerable regional differences in elemental composition occur in the tectorial membrane with regard to P, K and Ca. The highest concentration of Ca occurs in the basal part of the tectorial membrane towards the sensory hairs. The highest concentration of K occurs in the basal and outer parts whereas the middle part of the tectorial membrane contains low levels of both K and Ca. The elemental composition changes in two main directions: 1) from the limbal (growing) region to the tip of the tectorial membrane, and, 2) from upper to lower surfaces. The three cell types of the stria vascularis differ considerably in elemental composition. The highest concentration of K occurs in marginal cells. The basal cells contain more K than do the intermediate cells. A significantly higher concentration of Ca, Cl and Na occurs in marginal cell cytoplasm than in any other cell type in the stria vascularis.

Animals↗

PubMed1990

Lectin staining of saccharides in the normal and hypothyroid developing organ of Corti.

Lectin staining has been used to detect mono- and oligosaccharides in normal and hypothyroid developing organs of Corti in the rat. Eight developmental stages were studied (1, 5, 8, 10, 15, 20, 50 and 60 days after birth). Congenital hypothyroidism was induced by oral administration of propylthyouracil to pregnant rats. Labelling of the tectorial membrane with 3 lectins, Ulex europaeus agglutinin-I (UEA-I), Lens culinaris agglutinin (LCA) and Ricinus communis agglutinin-I (RCA-I) showed no significant differences between normal and hypothyroid animals. Staining with peanut agglutinin (PNA) showed that the hypothyroid adult tectorial membrane (but not the normal one) possesses the disaccharide galactose + N-acetyl galactosamine. Phaseolus vulgaris agglutinin-L (PHA-L) labels the whole tectorial membrane in both groups of animals, but the staining is more intense in the hypothyroid one for a narrow band of oligosaccharide located just between the tectorial membrane and the underlying organ of Kölliker. Both soybean agglutinin (SBA) and succinylated wheat germ agglutinin (WGA) stain the tectorial membrane as well as the cytoplasm of the cells constituting the inner portion of the organ of Kölliker; this latter feature disappears in the normal animals about the 8th postnatal day, but it is abnormally preserved until the 60th postnatal day in the hypothyroid ones. In the adult hypothyroid animals, 3 of the lectins (LCA, PHA-L and WGA) stain extracellular conglomerates located under the synaptic pole of the outer hair cells.

Animals↗

PubMed1991

Calcium-binding sites in the inner ear after pure-tone stimulation.

Five guinea pigs were exposed to an interrupted 90 dB SPL pure tone of 3.2 kHz for a total application time of 5 min. Following sound application all animals were decapitated and the cochleae were removed. After that, calcium-binding sites were located by the potassium pyroantimonate precipitation method. Another three animals served as control animals and did not receive the sound treatment. Findings confirmed previous studies showing the spatial arrangements of precipitate rich regions in the inner ear's two acellular structures (the basilar membrane and tectorial membrane) and the two cellular structures (the inner hair cells and Huschke's teeth). By using semiquantitative evaluation we found a relative diminution of precipitable calcium in inner hair cells and in Huschke's teeth of sound-exposed animals when compared to untreated animals.

Acoustic Stimulation↗

PubMed2002

Modeling 3-D deformation of outer hair cells and their production of the active force in the cochlea.

We analyze the deformation of the outer hair cell and its production of active force under physiological conditions. The active force has two components. One results from the strain caused by loading in the organ of Corti in the cochlea and depends on the level of the acoustic signal; the other is related to the intrinsic active properties of the cell membrane. We demonstrate our approach by considering, as a basic model of an outer hair cell in the organ of Corti, a cylindrical shell that is filled with an incompressible fluid and located between two planes that move relative to each other. These planes represent the basilar membrane and tectorial membrane complexes. We show that the deformed state of the cell has a 3-D nature, including bending and twisting components. This is different from the experimental conditions in which the active force is usually measured. We estimate the active force as a function of the relative position of the planes, angle of the cell's inclination, and the cell length.

Acoustic Stimulation↗

PubMed2003

Noise in magnetic resonance imaging: no risk for sensorineural function but increased amplitude variability of otoacoustic emissions.

OBJECTIVES: Objectives were to perform exact measurements of the noise exposure in a magnetic resonance imager and to investigate the effects of magnetic resonance imaging (MRI) noise on hearing sensitivity, which are still controversial, in a large number of patients. STUDY DESIGN: Prospective trial. METHODS: Acoustic noise during seven different MRI sequences was measured using a custom-made microphone containing no ferromagnetic parts. In 244 ears of 126 patients, pure-tone audiometry was performed once before and once after MRI noise, and distortion product otoacoustic emissions were measured once before and three times after MRI. RESULTS: The sound pressure level (SPL) at the patient's ear (with consideration of the sound-damping effect of the head support) ranged from 79.5 to 86.5 dB (A), depending on the MRI sequence, with brief sound pressure peaks up to 120 dB SPL. No significant incidence of temporary threshold shift and no reduction of mean distortion product otoacoustic emission amplitude were apparent. However, a significant increase in distortion product otoacoustic emission amplitude variability after noise exposure with equal distribution of increased and decreased amplitudes was observed. This variability showed a maximum at 15 minutes after noise, as demonstrated by continuous measurements. CONCLUSIONS: First, MRI noise does not impose a risk to hearing function under the measurement condition of a sound-damping head support or ear protectors. Second, a subtle effect is demonstrated by increased distortion product otoacoustic emission amplitude variability. Third, the increased otoacoustic emission amplitude variability is an audiometric parameter that is extremely sensitive to effects of acoustic stimulation, indicating more discrete changes in cochlear activity than pure-tone audiometry or otoacoustic emission amplitude reduction. A shift of the operation point (OP) of the outer hair cell (OHC) between basilar membrane and tectorial membrane is suggested as underlying cause.

Audiometry, Pure-Tone↗

PubMed1991

Hearing impairment in mice with the cmd/cmd (cartilage matrix deficiency) mutant gene.

Mice homozygous for the autosomal recessive gene cartilage matrix deficiency (cmd) are afflicted with lesions involving cartilaginous tissue which give rise to, among other things, marked hearing loss as evidenced by auditory evoked potentials. Ultrastructural studies of the inner ear reveal that while inner hair cells are normal in shape and content, the outer hair cells have disappeared and there is some debris in Nuel's space. The pillar cells are normal as are the stria vascularis, basilar membrane, and tectorial membrane. We conclude that the cmd gene, thought to be vital in the regulation of proteoglycan synthesis, is responsible for the hearing impairment and structural anomalies of the cochlea seen in these mutant, homozygous mice.

Animals↗

PubMed1987

Neural tuning in the granite spiny lizard.

Scanning electron microscopy was used to examine the basilar papilla of the granite spiny lizard. The papilla contains three distinct hair cell populations: an apical and a basal population with free-standing cilia, and a central population with a tectorial membrane. In the free-standing populations, stereocilium length decreases towards the ends of the papilla. Ciliary tuft morphology differs in the free-standing and the tectorial membrane populations, except that several of the free-standing hair cells with the shortest stereocilia have a tuft morphology like the hair cells in the tectorial membrane population. On the basis of single-fiber physiology, auditory nerve fibers can be divided into a low characteristic frequency (CF) and a high CF population. Mappings of the tonotopic organization of the nerve demonstrated two groups of high CF fibers that correspond to the two free-standing hair cell populations. The low CF fibers are associated with the tectorial membrane hair cell population. Fiber CF correlated with hair cell cilium length, not position on basilar membrane, for hair cells with free-standing cilia. Tonotopic organization of high CF fibers could be predicted reasonably well from the histogram of fiber CFs.

Animals↗

PubMed1998

Hair cell regeneration and recovery of function in the avian auditory system.

Chickens were exposed to an intense pure tone that destroyed the hair cells and tectorial membrane in a crescent shaped patch along the abneural edge of the basilar papilla. During the following weeks, when the hair cells and tectorial membrane were regenerating, psychophysical and electrophysiological measures were obtained to assess the time course and degree of recovery. Immediately after the exposure, the behavioral thresholds were elevated 30-40 dB and auditory temporal integration was greatly reduced; however, both measures fully recovered by 28 days post-exposure. In addition, tone-on-tone masking patterns recovered to normal. Immediately after the exposure, the thresholds of single cochlear ganglion neurons were elevated more than 30 dB, tuning curves were broader than normal, two-tone rate suppression (TTRS) boundary slopes were shallower than normal and spontaneous activity was reduced. Threshold and spontaneous discharge rate fully recovered after the exposure. Tuning and TTRS also recovered significantly in most neurons; however, some units with characteristic frequencies (CFs) near the exposure frequency showed abnormal tuning and TTRS suppression. The regeneration of the hair cells and lower honeycomb layer of the tectorial membrane is associated with considerable recovery of function; however, the incomplete recovery of tuning and TTRS in some neurons may be linked to the incomplete regeneration of the tectorial membrane.

Acoustic Stimulation↗

PubMed2000

Development of the gerbil inner ear observed in the hemicochlea.

A frequency-dependent change in hearing sensitivity occurs during maturation in the basal gerbil cochlea. This change takes place during the first week after the onset of hearing. It has been argued that the mass of a given cochlear segment decreases during development and thus increases the best frequency. Changes in mass during cochlear maturation have been estimated previously by measuring the changes in cochlear dimensions. Fixed, dehydrated, embedded, or sputter-coated tissues were used in such work. However, dehydration of the tissue, a part of most histological techniques, results in severe distortion of some aspects of cochlear morphology. The present experiments, using a novel preparation, the hemicochlea, show that hydrated structures, such as the tectorial membrane and the basilar membrane hyaline matrix, are up to 100% larger than estimated previous studies. Therefore, the hemicochlea was used to study the development of cochlear morphology in the gerbil between the day of birth and postnatal day 19. We used no protocols that would have resulted in severe distortion of cochlear elements. Consequently, a detailed study of cochlear morphology yields several measures that differ from previously published data. Our experiments confirm growth patterns of the cochlea that include a period of remarkably rapid change between postnatal day 6 and 8. The accelerated growth starts in the middle of the cochlea and progresses toward the base and the apex. In particular, the increase in height of Deiters' cells dominated the change, "pushing" the tectorial membrane toward scala vestibuli. This resulted in a shape change of the tectorial membrane and the organ of Corti. The tectorial membrane was properly extended above the outer hair cells by postnatal day 12. This time coincides with the onset of hearing. The basilar membrane hyaline matrix increased in thickness, whereas the multilayered tympanic cover layer cells decreased to a single band of cells by postnatal day 19. Before and after the period of rapid growth, the observed gross morphological changes are rather small. It is unlikely that dimensional changes of cochlear structures between postnatal days 12 and 19 contribute significantly in the remapping of the frequency-place code in the base of the cochlea. Instead, structural changes affecting the stiffness of the cochlear partition might be responsible for the shift in best frequency.

Animals↗