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.     

Developmental changes of frequency representation in the rat cochlea.

The place-frequency map of the developing rat cochlea was measured by iontophoretic HRP-injections into the ventral cochlear nucleus at electrophysiologically characterized positions. Distribution of retrograde HRP transport in cochlear spiral ganglion cells was analysed by means of a three dimensional reconstruction of the cochlea. Cochlear place-frequency maps were derived in rats of two ages groups: 13 to 22 days, and 36/37 day old animals. These maps were compared with the place-frequency map of adult rats (Müller, 1991). No systematic difference in the place-frequency map between 36/37 day old and adult rats was observed. In animals of the younger age group the place-frequency map (for frequencies above 4 kHz) was shifted towards lower frequencies for a given place along the basilar membrane. The morphological and physiological basis for this frequency shift during development is discussed.     

Frequency representation and spiral ganglion cell density in the cochlea of the gerbil Pachyuromys duprasi.

The tonotopic map of the cochlea in the gerbil Pachyuromys duprasi was analysed by local iontophoretic HRP-application into physiologically defined regions of the cochlear nucleus and mapping of subsequent HRP transport patterns in cochlear spiral ganglion cells. Furthermore the spiral ganglion cell density along the cochlear duct was determined. The cochlear tonotopic map was established in the frequency range between 0.6 and 17.5 kHz. These frequencies corresponded to locations between 86 and 3% basilar membrane length (0% = cochlear base). It was found that the slope of the place-frequency map varied with frequency, the maximum slope being found between 1 and 4 kHz. This frequency range corresponds to the frequency range of highest auditory sensitivity as determined from cochlear microphonic recordings (Plassmann et al., 1987). The density of spiral ganglion cells also varied along the cochlear duct. A pronounced maximum (1927 cells/mm) was located at around 70% basilar membrane length, compared to values of 800 cell per mm near the cochlear apex and base. This region of high ganglion cell density also corresponds to the frequency range of highest auditory sensitivity.     

Postnatal development of the organ of Corti in cats: a light microscopic morphometric study

This study quantitatively characterizes the development of the major morphological features of the organ of Corti during the first 2 weeks postnatal, the period when the cat auditory system makes the transition from being essentially non-functional to having nearly adult-like responses. Four groups of kittens (n=3) were studied at one day postnatal (P1), P5, P10, P15, and compared to adults. Measurements were made of the organ of Corti at 3 cochlear locations: 20%, 60% and 85% of basilar membrane length from the base – cochlear locations which in the adult correspond to best frequencies of ≈20 kHz, 2 kHz and 500 Hz, respectively. In addition, measurements of basilar membrane length and opening of the tunnel of Corti were made in 20 cochlear specimens from kittens aged P0–P6. Results indicate that: (i) at P0 the basilar membrane has attained adult length, and the tunnel of Corti is open over approximately the basal one-half of the cochlea; (ii) the initial opening of the tunnel of Corti occurs at a site about 4 mm from the cochlear base (best frequency of ≈25 kHz in the adult cochlea); (iii) the thickness of the tympanic cell layer decreases markedly at the basal 20-kHz location; (iv) the areas of the tunnel of Corti and space of Nuel and the angulation of the inner hair cells (IHC) relative to the basilar membrane all show marked postnatal increases at both the middle and apical locations; (v) IHC are nearly adult-like in length and shape at birth, whereas the OHC (at 2-kHz and 500-Hz locations) undergo marked postnatal changes; (vi) disappearance of the marginal pillars and maturation of the supporting cells are not yet complete by P15.     

Basilar membrane tonotopicity in the hook region of the cat cochlea.

Middle-ear to basilar membrane (BM) velocity transfer functions are reported for seven locations in the hook region of a single cat cochlea. These transfer functions were recorded at high sound pressure levels in a linearized, or passive cochlea, and resemble those reported previously by Wilson and Evans (1983). They demonstrate longitudinal tonotopicity with a gradient of approximately 3.6 mm/octave. When allowances are made for the nonlinear mechanisms previously demonstrated in active hook region preparations (Cooper and Rhode, 1992), the data are also consistent with the tonotopic map derived from the intracellular dye-filling studies of Liberman (1982).     

Measurement of guinea pig basilar membrane using computer-aided three-dimensional reconstruction system

Cochleas are known to have the ability to analyze a frequency widely, and this ability seems to be owed mostly to the basilar membrane (BM) configuration. However, the relationship between the cochlear frequency-position map and the BM configuration is not clear. Therefore, in this paper, the internal structures of a guinea pig cochlea, especially the BM configuration, were reconstructed and measured using a computer-aided three-dimensional (3-D) reconstruction system. Then, an attempt was made to examine the influence of the BM configuration on the cochlear frequency-position map. The measurement results indicate that the width of the BM increased and its thickness decreased with an increase in the distance from the basal turn towards the apical turn. Theoretical consideration reveals that the wide frequency-position of the cochlea is achieved by not only the BM configuration change along the length of the cochlea but also the change of the Young's modulus of the BM along the length of the cochlea.     

Basilar Membrane Vibrations Near the Round Window of the Gerbil Cochlea

Using a laser velocimeter, responses to tones were measured at a basilar membrane site located about 1.2 mm from the extreme basal end of the gerbil cochlea. In two exceptional cochleae in which function was only moderately disrupted by surgical preparations, basilar membrane responses had characteristic frequencies (CFs) of 34–37 kHz and exhibited a CF-specific compressive nonlinearity: Sensitivity near the CF decreased systematically and the response peaks shifted toward lower frequencies with increasing stimulus level. Response phases also changed with increases in stimulus level, exhibiting small relative lags and leads at frequencies just lower and higher than CF, respectively. Basilar membrane responses to low-level CF tones exceeded the magnitude of stapes vibrations by 54–56 dB. Response phases led stapes vibrations by about 90° at low stimulus frequencies; at higher frequencies, basilar membrane responses increasingly lagged stapes vibration, accumulating 1.5 periods of phase lag at CF. Postmortem, nonlinearities were abolished and responses peaked at ~0.5 octave below CF, with phases which lagged and led in vivo responses at frequencies lower and higher than CF, respectively. In conclusion, basilar membrane responses near the round window of the gerbil cochlea closely resemble those for other basal cochlear sites in gerbil and other species.     

Postnatal development of the organ of Corti in the wild house mouse, laboratory mouse, and their hybrid

Length of the basilar membrane, number and distribution of cochlear receptors, and the width of the triad of outer hair cells were analyzed in the course of the postnatal development and in adult individuals in wild and laboratory house mice and in hybrids of these species. While in newborn animals the triad of outer hair cells was wide at the base and narrow at the apex, the opposite was true for adult animals. The parameter decreased at the base and increased at the apex during postnatal development. The center of differentiation of (the reticular lamina of) the organ of Corti was localized at 40-50% of the basilar membrane length from the base and corresponded to the region with the maximum density of inner hair cells. The reticular lamina in the apical half of the cochlea matured earlier than in the basal half. Distribution of receptors did not change after birth. The shortest basilar membrane and the slowest rate of maturation were found in wild mice. Hybrids had the longest basilar membrane and the highest rate of maturation. These facts are considered an effect of heterosis.     

Predicting the effect of post-implant cochlear fibrosis on residual hearing

Intracochlear scarring is a well-described sequela of cochlear implantation. We developed a mathematical model of passive cochlear mechanics to predict the impact that this might have upon residual acoustical hearing after implantation. The cochlea was modeled using lumped impedance terms for scala vestibuli (SV), scala tympani (ST), and the cochlear partition (CP). The damping of ST and CP was increased in the basal one half of the cochlea to simulate the effect of scar tissue. We found that increasing the damping of the ST predominantly reduced basilar membrane vibrations in the apex of the cochlea while increasing the damping of the CP predominantly reduced basilar membrane vibrations in the base of the cochlea. As long as intracochlear scarring continues to occur with cochlear implantation, there will be limitations on hearing preservation. Newer surgical techniques and electrode technologies that do not result in as much scar tissue formation will permit improved hearing preservation.     

应用CT三维重建对正常人群耳蜗方位和盘曲模式的研究

目的：运用CT三维重建等影像手段对正常人群耳蜗进行形态学研究,为指导内耳畸形的诊断及人工耳蜗植入等内耳手术的术前评估提供理论依据。方法：收集100例（200耳）排除内耳解剖异常的患者作为研究对象,依年龄分为5组,利用计算机工作站的3D立体重建和2D多平面重建技术对其耳蜗基底部的长和宽、底周内最长直线距离、耳蜗的高度、第一周和第二周之间的角度及正中矢状面与底周间夹角等结构进行系统测量,统计分析以上各结构指标在不同年龄组、性别、左右耳之间的差异。结果：耳蜗基底部的长为（8．56±0．52）mm、宽为（6．63±0．56）mm、底周内最长直线距离为（7．33±0．56）mm、耳蜗的高度为（3．76±0．28）mm、第一周和第二周之间的角度为（15．824±2．78）°。以上所有指标在不同年龄、性别及左右耳间差异无统计学意义（P〉0．05）。正中矢状面与耳蜗底周间角度在各年龄组间差异有统计学意义,该角度随着年龄的增长而缩小（P〈0．05）。第一和第二周之间的角度在各年龄组内存在变异,提示耳蜗的盘曲模式在个体间差异明显。结论：3D和2D多平面重建技术能准确评价内耳形态结构,测定国人耳蜗各主要结构的正常值,可为临床诊断及人工耳蜗等内耳手术治疗提供重要的参考依据。     

蜗内直流电对耳蜗基底膜振动的影响

目的探讨外加直流电流后对耳蜗基底膜振动的影响.方法在豚鼠耳蜗底回距圆窗龛缘2.4mm处开一直径约0.4mm小孔,作为测量活体基底膜的振动速度测试窗.在测试窗上、下缘的鼓阶、前庭阶各开一小孔,将铂-铱电极置入鼓阶、前庭阶作为跨蜗管的电刺激电极.用激光多普勒干涉测速仪观察直流电流对纯音诱发的基底膜振动速度的影响.结果当外加电流前庭阶极性为正,鼓阶极性为负时,可以看到基底膜振动速度显著增大,给相反极性电流时,基底膜振动速度减小.结论生理状态下的正内淋巴电位是耳蜗将声音能量转变为神经冲动的必要条件.适当提高外毛细胞顶端正电位,有助于提高耳蜗放大器的增益.外加负电位则严重影响耳蜗放大器的增益.     

The interaction of noise and aspirin in the chick basilar papilla. Noise and aspirin toxicity

The possible synergism between noise and aspirin for causing cochlear damage was examined histologically. Six chicks fed aspirin for five days and five chicks fed a normal diet only were paired and placed in sound chambers. They were exposed to a 1500-Hz tone at 115 dB sound pressure level for eight hours. The mean serum salicylate level just before noise exposure was 24 mg/dL (1.74 mmol/L). Ten days later they were killed, and the temporal bones were processed. Hair cell counts were made at 100-microns intervals throughout the length of the basilar papilla (cochlea). The noise produced a discrete cochlear lesion centered about 30% of the distance from the base to apex. The addition of aspirin did not significantly alter the extent or location of this lesion. One aspirin-fed chick had a unilateral middle ear effusion, and a striking shift in the center of damage toward the apex was noted in this cochlea.     

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.     

蜗内直流电对耳蜗基底膜振动的影响

目的：探讨外加直流电流后对耳蜗基底膜振动的影响。方法：在豚鼠耳蜗底回距圆窗龛缘2．4mm处开一直径约0．4mm小孔，作为测量活体基底膜的振动速度测试窗，在测试窗上，下缘的鼓阶，前庭阶各开一小孔，将铂－铱电极置入鼓阶，前庭阶作为跨蜗管的电刺激电极，用激光多普勒干涉测速仪观察直流电流对纯音诱发的基底膜振动速度的影响。结果：当外加电流前庭阶极性为正，鼓阶极性为负时，可以看到基底膜振动速度显著增大，给相反极性电流时，基底膜振动速度减小。结论：生理状态下的正内淋巴电位是耳蜗将声音能量转变为神经冲动的必要条件，适当提高外毛细胞顶端正电位，有助于提高耳蜗放大器的增益，外加负电位则严重影响耳蜗放大器的增益。     

Recent developments in cochlear physiology

Recent findings in cochlear physiology have caused many of our long held ideas about how sound is analyzed by the ear to be reevaluated. This article describes changes which have occurred in three classical ideas of cochlear transduction: (1) There is a gradient of frequency representation along the cochlea with high frequencies being represented at the base and lower frequencies represented progressively toward the apex. It is now known that the specific frequency which is represented at a given location along the cochlea is not invariant but changes systematically during the normal development of hearing. (2) The place code and frequency tuning along the cochlea are due to the conventional traveling wave of von Békésy and basilar membrane mechanics. Experiments in nonmammalian vertebrates which lack a traveling wave have shown that other mechanisms, including the mechanical resonance of hair cell stereocilia, may contribute to tonotopic organization and frequency tuning. It is possible that hair cell stereocilia also contribute to frequency representation and tuning in the mammalian cochlea. (3) The vibration of the basilar membrane to sound is determined by its passive mechanical properties. It is now known that the response of the basilar membrane, and that of the cochlear partition as a whole, is influenced by physiological processes which utilize metabolic energy. The active processes are likely expressed through the motile activity of outer hair cells.     

The cochleogram of the guinea pig

The cochleogram is an important tool to relate properties of the cochlea (e.g. hair cell loss, damaged hair cells) to their position in the cochlear turns, to calculate the average hair cell density, and to measure the length of the whole cochlea. In this work different methods of plotting cochleograms are compared. We suggest that a sector-wise division of the cochlea for counting a cochleogram has advantages over line diagrams that provide a higher spatial resolution but might lead to misinterpretations of the degree of missing hair cells. The scanning electron microscopic analysis of 171 guinea pig cochleas revealed a mean basilar membrane length of 16.4 ± 1.4 mm (mean ± standard deviation) with sector lengths of 6.9, 4.2, 3.2, and 1.9 mm, thus adding relevant information to the morphology of the guinea pig cochlea.     

Backward Propagation of Otoacoustic Emissions

IntroductionsSounds impinging the eardrum are transmitted viamiddle ear ossicles to the oval window. Stapes vibra-tion creates pressure difference between the scala tym-pani and the scala vestibuli. This pressure differencecauses a movement of cochlear partition and adjacentcochlear fluids. Basilar membrane vibrations result indeflection of hair cell stereocilia, which gate ion chan-nels on their tips. This mechanical-to-electrical trans-duction process converts mechanical vibrations intoelec…     

A short history of hearing research. IV: Physiology]

In 1863, Hensen concluded from measurements of the width of the basilar membrane that tones of high and low pitch were represented at the base and apex of the cochlea, respectively. According to his calculations on the tonotopic representation of sound stimuli in the cochlea Helmholtz proposed additional resonators that would transmit the amplified signal to the afferent nerve endings. He speculated that the pillar cells of the tunnel of Corti or strands of the basilar membrane might be these proposed resonators. The resonance theory was contradicted by Wien in 1905. However, further experiments by Held and Kleinknecht in 1927 and by Békésy in 1928 demonstrated that Helmholtz's ideas on the tonotopic dispersion of the vibration of the basilar membrane were correct. Békésy measured the vibration of the cochlear partition in human and animal cadavers and discovered the travelling-wave of the basilar membrane. At the turn of the century Ter Kuile noted that the vibration of the cochlear partition caused a deflection of the sensory hairs of the hair cells, the auditory receptor cells. Wever and Bray described in 1930 stimulus-evoked electrical currents near the cochlea with a wave form similar to that of the original sound stimulus. It was Adrian who later coined the term "cochlear microphonics" for this phenomenon. According to calculations of Gold (1948) and others active mechanical amplification would be required for such a sharp tuning in the cochlea. The first to measure action potentials of the afferent auditory nerve was Tasaki (1954).(ABSTRACT TRUNCATED AT 250 WORDS)     

Development of the mouse cochlea database (MCD)

The mouse cochlea database (MCD) provides an interactive, image database of the mouse cochlea for learning its anatomy and data mining of its resources. The MCD website is hosted on a centrally maintained, high-speed server at the following URL: http://mousecochlea.umn.edu. The MCD contains two types of image resources, serial 2D image stacks and 3D reconstructions of cochlear structures. Complete image stacks of the cochlea from two different mouse strains were obtained using orthogonal plane fluorescence optical microscopy (OPFOS). 2D images of the cochlea are presented on the MCD website as: viewable images within a stack, 2D atlas of the cochlea, orthogonal sections, and direct volume renderings combined with isosurface reconstructions. In order to assess cochlear structures quantitatively, "true" cross-sections of the scala media along the length of the basilar membrane were generated by virtual resectioning of a cochlea orthogonal to a cochlear structure, such as the centroid of the basilar membrane or the scala media. 3D images are presented on the MCD website as: direct volume renderings, movies, interactive QuickTime VRs, flythrough, and isosurface 3D reconstructions of different cochlear structures. 3D computer models can also be used for solid model fabrication by rapid prototyping and models from different cochleas can be combined to produce an average 3D model. The MCD is the first comprehensive image resource on the mouse cochlea and is a new paradigm for understanding the anatomy of the cochlea, and establishing morphometric parameters of cochlear structures in normal and mutant mice.