Microphonic Potentials Research Articles

The history of audiology

The paper presents historical data on development of audiology as a medical speciality. It gives the chronological overview of the most significant discoveries which have contributed to the progress and constant development of the science of hearing. The insights and discoveries encompass the ancient, medieval and contemporary medical science. THE TERM "AUDIOLOGY" AND FIRST ASSOCIATIONS OF AUDIOLOGISTS: The paper reviews the origin of the term "audiology" and the time of its occurrence. The First World Congress of Audiologists was held in 1948, and the Conference of Audiologists and Phoniatrists of Yugoslavia was established in 1974. The ancient scientists and philosophers mentioned in the paper are as follows: Democritus, Hippocrates, Empedocles, Plato, Aristotle, and Galenus. Discoveries of Vesalius, Fallopio, Eustachio, Duverney, Schelhammer, Catugno and Helmholtz marked their epochs and made a great contribution to the development of the entire medical science, including audiology. A significant contribution to audiology was made by Schwartz in 1920 and Fletcher in 1926, constructors of audiometers. Fowler, Weigel and Fletcher promoted an audiogram in 1922. Wever and Bray first introduced the cochlear microphonic potentials. The first modern audiometer with a flat zero line for all pitches was constructed in 1937. In 1947, Bekesy constructed the automatic audiometer, and the theory of mobile waves was introduced in 1928. Fletcher and Steinberg promoted speech audiometries in 1929. Mendel and Goldstein described medium latency responses in 1969. The first hearing aids worn within the ear appeared in the same year. William House pioneered the cochlear implantation in adults in 1969, and the program of infant cochlear implants. Jewett described the evoked auditory potentials in 1970. James Jerger classified tympanometric curves into three tympanogram types (A, B, C). Portmann and Arran introduced transtympanic electrocochleography in 1971. Kemp introduced otoacoustic emission in 1978. finally, the paper lists the titles of available dissertations and sub-specialisations in the field of audiology defended in Novi Sad and Belgrade.

The Development of a Single Frequency Place in the Mammalian Cochlea: The Cochlear Resonance in the Mustached BatPteronotus parnellii

Cochlear microphonic potentials (CMs) were recorded from the sharply tuned, strongly resonant auditory foveae of 1- to 5-week-old mustached bats that were anesthetized with Rompun and Ketavet. The fovea processes Doppler-shifted echo responses of the constant-frequency component of echolocation calls. During development, the frequency and tuning sharpness of the cochlear resonance increases, and CM ringing persists for longer after the tone. CM is relatively insensitive at tone onset and grows linearly with increased stimulus level. During the tone, the CM is more sensitive and grows compressively with increased stimulus level and phase leads onset CM by 90 degrees for frequencies below the resonance. CM during the ringing is also sensitive and compressive and phase leads onset CM by 180 degrees below the resonance and lags it by 180 degrees above the resonance. Throughout postnatal development, CMs measured during the tone and in the ringing increase both in sensitivity and compression. The cochlear resonance appears to be attributable to interaction between two oscillators. The more broadly tuned oscillator dominates the onset response, and the narrowly tuned oscillator dominates the ringing. Early in development, mechanical coupling between the oscillators results in a relatively broadly tuned system with several frequency modes in the CM at tone onset and in the CM ringing. Beating occurs between the resonance and the stimulus response during the tone and between two components of the narrowly tuned oscillator at tone offset. At maturity, the CM has three modes for frequencies within 10 kHz of the resonance at tone onset and a single, sharply tuned mode in the ringing.

Synchronization of a Nonlinear Oscillator: Processing the Cf Component of the Echo-Response Signal in the Cochlea of the Mustached Bat

Cochlear microphonic potential (CM) was recorded from the CF2 region and the sparsely innervated zone (the mustached bat's cochlea fovea) that is specialized for analyzing the Doppler-shifted echoes of the first-harmonic (approximately 61 kHz) of the constant-frequency component of the echolocation call. Temporal analysis of the CM, which is tuned sharply to the 61 kHz cochlear resonance, revealed that at the resonance frequency, and within 1 msec of tone onset, CM is broadly tuned with linear magnitude level functions. CM measured during the ongoing tone and in the ringing after tone offset is 50 dB more sensitive, is sharply tuned, has compressive level functions, and the phase leads onset CM by 90 degrees: an indication that cochlear responses are amplified during maximum basilar membrane velocity. For high-level tones above the resonance frequency, CM appears at tone onset and after tone offset. Measurements indicate that the two oscillators responsible for the cochlear resonance, presumably the basilar and tectorial membranes, move together in phase during the ongoing tone, thereby minimizing net shear between them and hair cell excitation. For tones within 2 kHz of the cochlear resonance the frequency of CM measured within 2 msec of tone onset is not that of the stimulus but is proportional to it. For tones just below the cochlear resonance region CM frequency is a constant amount below that of the stimulus depending on CM measurement delay from tone onset. The frequency responses of the CM recorded from the cochlear fovea can be accounted for through synchronization between the nonlinear oscillators responsible for the cochlear resonance and the stimulus tone.

Loud sound-induced changes in cochlear mechanics.

To investigate the inner ear response to intense sound and the mechanisms behind temporary threshold shifts, anesthetized guinea pigs were exposed to tones at 100-112 dB SPL. Basilar membrane vibration was measured using laser velocimetry, and the cochlear microphonic potential, compound action potential of the auditory nerve, and local electric AC potentials in the organ of Corti were used as additional indicators of cochlear function. After exposure to a 12-kHz intense tone, basilar membrane vibrations in response to probe tones at the characteristic frequency of the recording location (17 kHz) were transiently reduced. This reduction recovered over the course of 50 ms in most cases. Organ of Corti AC potentials were also reduced and recovered with a time course similar to the basilar membrane. When using a probe tone at either 1 or 4 kHz, organ of Corti AC potentials were unaffected by loud sound, indicating that transducer channels remained intact. In most experiments, both the basilar membrane and the cochlear microphonic response to the 12-kHz overstimulation was constant throughout the duration of the intense stimulus, despite a large loss of cochlear sensitivity. It is concluded that the reduction of basilar membrane velocity that followed loud sound was caused by changes in cochlear amplification and that the cochlear response to intense stimulation is determined by the passive mechanical properties of the inner ear structures.

Mechanical leverage in the middle ear of the American bullfrog, Rana catesbeiana

Textbooks lump the middle ears of ‘submammalian Tetrapoda’ as being ‘one-ossicle ears’. Conventionally the anuran middle ear is depicted with a shaft-like skeletal unit connecting the tympanic membrane to the inner ear. This shaft comprises mediad a long bony columella and laterad a short cartilaginous extracolumella. But dissection of Rana catesbeiana ears showed: the extracolumella, as long as the columella, is proximally expanded in the vertical plane, forming dorsal and ventral heads. The medio-dorsal head is movably jointed to the columella, between these two there is an obtuse angle ventrad; the extracolumellar medio-ventral head is anchored by a ligament to the middle-ear cavity ceiling. When the tympanic membrane moves outwards, pulling the extracolumella, the medio-dorsal head of the extracolumella must be forced inwards, rotating on the ventral anchorage, pushing the columella towards the inner ear. The ossicular chain thus includes a mechanical lever, possessing the magnitude of the ratio length:width of the extracolumella; this is additional to the lever known from the columellar footplate, which rotates on its firm ventral attachment. These levers are confirmed physiologically, by the difference between the inner-ear sensitivity (shown by isopotential audiograms of microphonic potentials) when stimulated by a vibrator first at the tympanic membrane, then at the proximal stump of the amputated columella. Perusal of the primary literature showed that this morphology is widespread among anuran ears.

Non-Linear Aspects of Outer Hair Cell Transduction and the Temporary Threshold Shifts after Acoustic Trauma

The 200-Hz cochlear microphonic potential (CM) and the compound action potential (CAP) of the auditory nerve evoked by tone-bursts were recorded in the basal turn of the cochlea of anaesthetised guinea pigs, before and after exposure to traumatic high-frequency tones that produce a temporary threshold shift (TTS) in this cochlear region. The drop in CM and the TTS were highly correlated, suggesting that it is the disruption of the outer hair cells generating the CM that causes the TTS. The previously measured rise in endocochlear potential and drop in organ of Corti K⁺ levels suggest that the TTS is due to a temporary closure of outer hair cell mechanoelectrical transduction (MET) channels, which produces a drop in the mechanical sensitivity of the organ of Corti, due to disruption of the active process provided by outer hair cells. The time course of the onset and recovery of TTS is consistent with a kinetic folding and refolding of MET channels over a time course of hours and days. Mathematical modelling of this putative channel folding suggests that TTS recovery may be accelerated by the presentation of additional sounds during the recovery period. We present electrophysiological data (CM and CAP measurements) showing that this accelerated recovery occurs. Using two-tone complexes (phase-locked 5- and 10-kHz traumatic tones, and 10-kHz traumatic tones with 25-Hz bias tones), we also show that the mechanisms producing TTS are non-linear and asymmetric, and that the greatest ‘trauma’ occurs when the hair bundles of outer hair cells are deflected away from the basal body of these cells (i.e. in the direction normally causing hyperpolarisation of the cell membrane potential).

Cochlear microphonic potentials: a new recording technique.

A new instrumentation and a particular method for detecting and recording cochlear microphonic potentials (CMPs) are described here. The CMPs were recorded in rats by means of pure tones (4,000, 2,000, 1,000, 500, and 250 Hz) and intraepidermic electrodes; the electrocochleography technique was avoided. An experimental design that included the use of a glutamatergic agonist (kainic acid [KA]) and an aminoglycoside antibiotic (kanamycin [KANA]) was carried out to demonstrate the origin of the recorded potential. Morphological studies showed that KA selectively eliminated the afferent type I dendrites of the spiral ganglion, while the administration of KANA resulted in the absence of outer hair cells. When CMPs were recorded after KA administration, no alterations were detected. In contrast, KANA administration resulted in the absence of any selective electrophysiological activity corresponding to CMPs. All these results were compared with the recording of the compound action potential of the eighth nerve obtained by electrocochleography. These findings and the great specificity of the reproduction of the sound stimulus confirm that the CMPs can be recorded by the new equipment.

Determination of auditory thresholds in the brushtail possum ( Trichosurus vulpecula)

The auditory abilities of the brushtail possum ( Trichosurus vulpecula) have been measured by cochlear potential readings, but no behavioural determinations of their auditory abilities have been published. Six experimentally naive possums were trained to perform a two-response conditional discrimination between the presence and absence of an 880-Hz tone (at 80 dB). All six possums readily learned this task. The behavioural threshold for this tone was determined using a modified tracking procedure and found to be similar to that reported using cochlear microphonic potentials. One concern with the current method was the communal nature of the experimental environment so a further threshold determination in a sound-attenuating chamber was conducted. No substantial difference was noted between the results obtained in the two threshold determinations. The success of the current method means that a full, behavioural audiogram for the brushtail possum, which would complement the existing cochlear potential data, is now possible.

Speech evaluation of partially implantable piezoelectric middle ear implants in vivo.

Cochlear microphonic responses (CMs) were measured in a rabbit model in the intact ear (CM1) and in ears with a partially implantable piezoelectric middle ear implant (P-MEI) (CM2) to investigate the characteristics of speech transmission of the P-MEI in vivo. The spectra of pure tones, voices, and elicited CM1 and CM2 obtained from the round window before and after the implantation of a P-MEI device were calculated by using fast Fourier transform. Frequency response functions of CM1 and CM2 were used to demonstrate the functional similarity between the implanted P-MEI and the normal ossicular chain. The coherence functions between the voices and CM1 and between CM1 and CM2 were evaluated to characterize speech transmission of the P-MEI in vivo. Ten rabbit ears were used in this study. Pure tones, six Chinese vowels, and six Chinese characters were the acoustic stimuli. The CMs elicited by a list of bisyllabic words were tape-recorded and then recognized by subjects with normal hearing. Using pure tones at the same intensity of 90 dB SPL, frequency response functions of the CMs between the two states (the intact ear with normal hearing and ears with a P-MEI device at the medium volume) were calculated showing great resemblance in shape. Compared with that at 1 kHz, gain factors were 10 and 20 dB, respectively, at higher frequencies. The correlation and spectral analyses of the vocalizations, CM1 and CM2, demonstrated that the harmonics of CM1 were approximately identical as those of the voices between 0.5 and 5.0 kHz with coherence functions of about 0.7 to approximately 1 at the formants' frequencies, whereas the harmonics of CM2 between 0.5 and 2.5 kHz were enhanced with the coherences near to unity at the formants' frequencies, and others <0.5 kHz and >2.5 kHz were attenuated. The recognition score of the CMs elicited by a list of bisyllabic words was >90% using subjects with normal hearing. Data from this study suggest that cochlear microphonic potentials can be used as an important tool to evaluate objectively whether the implantation of a P-MEI device is successful and whether the quality in speech transmission of the P-MEI is satisfactory. Thus, the method would be of significance to clinically ascertain the performance of the device in vivo.

Reversible and irreversible damage to cochlear afferent neurons by kainic acid excitotoxicity.

Kainic acid (KA) selectively damages afferent synapses that innervate, in chickens, mainly tall hair cells. To better understand the nature of KA-induced excitotoxic damage to the cochlear afferent neurons, KA, at two different concentrations (0.3 or 5 mM), was injected directly into the inner ear of adult chickens. Pathologic changes in the afferent nerve ending and cell body were evaluated with light and transmission electron microscopy at various time points after KA application. The compound action potential (CAP) and cochlear microphonic (CM) potential were recorded to monitor the physiologic status of the afferent neurons and hair cells, respectively. Hair cell morphology and function were essentially normal after KA treatment. However, afferent synapses beneath tall hair cells were swollen within 30 minutes after KA at both low (KA-L) and high (KA-H) doses. In the KA-L group, the swelling disappeared within 1 day and the morphology of the postsynaptic region returned to near normal condition. In the KA-H group, by contrast, the vacant region beneath tall hair cells remained evident even 20 weeks after KA. The number of cochlear ganglion neurons in the KA-H group decreased progressively from 1 to 8-20 weeks, whereas hair cells in the basilar papilla remained morphologically intact out to 20 weeks after KA. There was no significant change in neuron number in the KA-L group. Temporal changes in the CAP amplitude paralleled the anatomic changes, although the CAP only partially recovered. These results suggest that KA induces partially reversible damage to cochlear afferent neurons with low KA concentration; above this level, KA triggers irreversible, progressive neurodegeneration.

Selective loss of inner hair cells and type-I ganglion neurons in carboplatin-treated chinchillas. Mechanisms of damage and protection.

Carboplatin preferentially destroys inner hair cells (IHCs) and type-I spiral ganglion neurons while sparing outer hair cells (OHCs). Loss of IHCs and type-I ganglion cells is associated with a significant reduction of the compound action potential (CAP). However, the cochlear microphonic (CM) potential and distortion product otoacoustic emissions (DPOAEs) remain normal, indicating that the OHCs are functionally intact. In the vestibular system, carboplatin selectively destroys type-I hair cells and their afferent neurons. Damage of type-I vestibular hair cells and their afferent terminals is associated with significant depression of nystagmus induced by cold, caloric stimulation. Histochemical studies revealed a rapid decrease in succinate dehydrogenase (SDH) staining in IHCs soon after carboplatin treatment, and staining intensity remained depressed in surviving IHCs for at least 1 month after carboplatin treatment. These results suggest that carboplatin depresses the metabolic function in surviving IHCs. Several lines of evidence suggest that free radicals may contribute to carboplatin-induced sensory cell damage. Intracochlear infusion of L-buthionine-[S,R]-sulfoximine (BSO), which depletes intracellular glutathione (GSH), increases IHC and OHC loss. Previous in vitro studies have shown that neurotrophin 4/5 (NT-4/5) promotes the survival of spiral ganglion neurons from cisplatin ototoxicity. In vivo perfusion of NT-4/5 promoted the survival of spiral ganglion neurons, but did not protect the hair cells.

Hearing impairment in patients with acoustic neuroma—analysis by electrocochleography

Objective: In patients with acoustic neuroma, the site and severity of hearing impairment are important in discussing surgical approaches. Since the effectiveness of conventional auditory psychological testing is limited, we studied objectively hearing impairment of the cochlea and the cochlear nerve due to the tumor. Methods: Electrocochleography (ECochG) was carried out in 21 patients with acoustic neuroma. Cochlear microphonic potential (CM) and action potential (AP) in ECochG evoked with clicks and short tone bursts were recorded through a transtympanic needle electrode technique. Cochlear function was studied using the detection thresholds of CM, and cochlear nerve involvement was analyzed by differences between AP and CM detection thresholds. Results: The 1 kHz CM detection threshold was elevated in 17 (81.0%) of 21 patients indicating cochlear impairment. Of seven patients with normal hearing or mild sensorineural hearing loss in pure tone audiometry, three had a slightly elevated CM detection threshold. Of five patients with pronounced pure tone levels, four showed a CM response and were thought to have mild cochlear dysfunction. Cochlear nerve impairment was confirmed in three of four patients with well-developed CM based on elevated AP detection thresholds. Three patients had CM response but no AP response, suggesting severe cochlear nerve impairment. Conclusion: Disorders of the cochlea and the cochlear nerve can be evaluated with ECochG AP and CM measurement. The findings of ECochG are thought to be important information to judge hearing prognosis, thereby enhancing its clinical utility.

Supporting Cells Contribute to Control of Hearing Sensitivity

The mammalian hearing organ, the organ of Corti, was studied in an in vitro preparation of the guinea pig temporal bone. As in vivo, the hearing organ responded with an electrical potential, the cochlear microphonic potential, when stimulated with a test tone. After exposure to intense sound, the response to the test tone was reduced. The electrical response either recovered within 10-20 min or remained permanently reduced, thus corresponding to a temporary or sustained loss of sensitivity. Using laser scanning confocal microscopy, stimulus-induced changes of the cellular structure of the hearing organ were simultaneously studied. The cells in the organ were labeled with two fluorescent probes, a membrane dye and a cytoplasm dye, showing enzymatic activity in living cells. Confocal microscopy images were collected and compared before and after intense sound exposure. The results were as follows. (1) The organ of Corti could be divided into two different structural entities in terms of their susceptibility to damage: an inner, structurally stable region comprised of the inner hair cell with its supporting cells and the inner and outer pillar cells; and an outer region that exhibited dynamic structural changes and consisted of the outer hair cells and the third Deiters' cell with its attached Hensen's cells. (2) Exposure to intense sound caused the Deiters' cells and Hensen's cells to move in toward the center of the cochlear turn. (3) This event coincided with a reduced sensitivity to the test tone (i.e., reduced cochlear microphonic potential). (4) The displacement and sensitivity loss could be reversible. It is concluded that these observations have relevance for understanding the mechanisms behind hearing loss after noise exposure and that the supporting cells take an active part in protection against trauma during high-intensity sound exposure.

Development of acetylcholine-induced responses in neonatal gerbil outer hair cells.

Cochlear outer hair cells (OHCs) are dominantly innervated by efferents, with acetylcholine (ACh) being their principal neurotransmitter. ACh activation of the cholinergic receptors on isolated OHCs induces calcium influx through the ionotropic receptors, followed by a large outward K+ current through nearby Ca2+-activated K+ channels. The outward K+ current hyperpolarizes the cell, resulting in the fast inhibitory effects of efferent action. Although the ACh receptors (AChRs) in adult OHCs have been identified and the ACh-induced current responses have been characterized, it is unclear when the ACh-induced current responses occur during development. In this study we attempt to address this question by determining the time of onset of the ACh-induced currents in neonatal gerbil OHCs, using whole cell patch-clamp techniques. Developing gerbils ranging in age from 4 to 12 days were used in these experiments, because efferent synaptogenesis and functional maturation of OHCs occur after birth. Results show that the first detectable ACh-induced current occurred at 6 days after birth (DAB) in 12% of the basal turn cells with a small outward current. The fraction of responsive cells and the size of outward currents increased as development progressed. By 11 DAB, the fraction of responsive cells and the current size were comparable with those of adult OHCs. The results indicate that the maturation of the ACh-induced response begins around 6 DAB. It appears that the development of ACh-induced responses occur during the same time period when OHCs develop motility but before the onset of auditory function, which is around 12 DAB when cochlear microphonic potentials can first be evoked with acoustic stimulation in gerbils.

The Goldman-Hodgkin-Katz equation and graphical `load-line' analysis of ionic flow through outer hair cells

While the `membrane potential' of a cell which has a homogeneous membrane and surrounding environment, and which is not pumping ions electrogenically (passing no net current through its membranes), can be estimated from the Goldman voltage equation, this equation is inappropriate for other cells. In the mammalian cochlea such problematic cells include the cells of stria vascularis and the sensory hair cells of the organ of Corti. Not only is the Goldman voltage equation inappropriate, but in asymmetric cells the concept of a single `membrane potential' is misleading: a different transmembrane voltage is required to define the electrical state of each section of the cell's heterogeneous membrane. This paper presents a graphical `load-line analysis' of currents through one such asymmetric cell, the outer hair cells of the organ of Corti. The approach is extremely useful in discussing the effects of various cochlear manipulations on the electrical potential within hair cells, even without a detailed knowledge of their membrane conductance. The paper discusses how modified Goldman-Hodgkin-Katz equations can be used to describe stretch-activated channels, voltage-controlled channels, ligand-mediated channels, and how the combination of these channels and the extracellular ionic concentrations should affect the hair cell's resting intracellular potential and resting transcellular current, its receptor current and receptor potential, and the extracellular microphonic potential around these cells. Two other issues discussed are the role of voltage-controlled channels in genetically determining membrane potential, and the insensitivity of hair cells to changes of extracellular potassium concentration under some conditions.

Tonic efferent-induced cochlear damping in roosting and echolocating mustached bats

The activity of the medial olivocochlear (MOC) efferent system in mustached bats, Pteronotus p. parnellii, was studied by monitoring changes in the mechanical properties of the cochlea. The changing properties were expressed by the decay time (DT) of cochlear microphonic potentials produced by transient-induced ringing ( Henson et al., 1995). Tape-recorded roost noise (biosonar and communication sounds) produced sudden, marked decreases in DT when presented to the contralateral ear of animals adapted to the quiet. When the animals were first removed from their roosts the DT was relatively short (1.2–1.5 ms) but this gradually lengthened by about 0.5–1.0 ms as they rested in a quiet chamber. The time required to reach a stable, quiet-adapted state after noise exposure varied with SPL and exposure time; in many experiments recovery was in the range of 90–120 min. When quiet-adapted bats were isolated and allowed to fly and echolocate for 20 min, the DTs measured within a few minutes after the end of the flight were also short and only slowly returned to longer preflight values. The administration of a single dose of gentamicin, which blocks MOC effects, greatly reduced the amount of suppression (damping) observed after periods of noise and echolocation sound exposure. We conclude that tonic MOC activity is induced by the natural vocalizations and roost noise and this activity probably regulates and protects the highly resonant cochlear partition.

Changes in cochlear mechanics during vocalization: evidence for a phasic medial efferent effect

The mustached bat, Pteronotus p. parnellii, has a finely tuned cochlea that rings at its resonant frequency in response to an acoustic tone pip. The decay time (DT) and frequency of these damped oscillations can be measured from the cochlear microphonic potential (CM) to study changes in cochlear mechanics. In this report, we describe phasic changes that occur in synchrony with communication sound vocalizations of the bat. Three animals with chronically implanted electrodes were studied. During the experiments, 1–2 ms tone pips were emitted from a speaker every 200 ms. This triggered a computer analysis of the resulting CM to determine the DT and cochlear resonance frequency (CRF) of the ringing. The time relative to vocalizations was determined by monitoring the output of a microphone placed near a bat's mouth. Similar results were obtained from all three bats tested. In a representative case, the average DT was 2.33±0.25 ms while the bat was quiet, but it decreased by 46% to 1.26±0.75 during vocalizations, which indicates a greater damping of the cochlear partition. Sometimes, DT started decreasing immediately before the bat vocalized. After the end of a vocalization, the return to baseline values varied from rapid (milliseconds) to gradual (1–2 seconds). The CRF also changed from baseline values during vocalization, although the amount and direction of change were not predictable. When gentamicin was administered to block the action of medial olivocochlear (MOC) efferents, DT reduction was still evident during vocalization but less pronounced. We conclude that phasic changes in damping occur in synchrony with vocalization, and that the MOC system plays a role in causing suppression. Since suppression can begin prior to vocalization, this may be a synkinetic effect, mediated by neural outflow to the ear in synchrony with neural outflow to the middle ear muscles and the muscles used for vocalization.

Postnatal maturation of the organ of Corti in gerbils: Morphology and physiological responses

The organ of Corti, the sensory epithelium of hearing in mammals, matures postnatally in the gerbil. Quantitative analyses of the postnatal development of the organ of Corti, including supporting cells and the basilar membrane, were carried out. The morphological study confirmed that maturation of the sensory cells proceeds with a base-to-apex gradient, with the outer hair cells appearing to mature before the inner hair cells. Maturation of the supporting cells and the basilar membrane commenced first in the middle turn. Expansion of the second row of Deiters' cells began at 6 days after birth in the middle turn, before enlargement of the pillar cell heads at 8 days postnatally. Pillar cell head enlargement continued until 20 days postnatally in the middle turn. The tunnel of Corti and spaces of Nuel appeared first in the middle turn between 8 and 10 days postnatally. The maturation of the basilar membrane involved the thickening of the central hyaline layer and a reduction in the epithelial cells on the tympanic aspect. This process continued until about 20 days after birth. The cochlear microphonic potential, whole nerve action potential, and stimulus frequency otoacoustic emissions were recorded from 12 days after birth onward and related to changes in organ of Corti morphology. The results show that changes in the accessory structures continue throughout the period of onset and development of cochlear responses between 12 and 20 days after birth, and may therefore influence the micromechanical responses of the organ of Corti to acoustic stimuli during this period.

Microphonic and DPOAE measurements suggest a micromechanical mechanism for the ‘bounce’ phenomenon following low-frequency tones

Neural auditory thresholds in the guinea pig can be temporarily improved by up to 6 dB about 2 min after the cessation of an moderately intense low-frequency tone (Kirk and Patuzzi, 1997). We have measured changes in the f 2-f 1 distortion product otoacoustic emission (DPOAE) and low-frequency microphonic potential in scala tympani before, during and after a low-frequency tone (200 Hz) to determine the cause of this so-called bounce phenomenon. In particular we have analysed the low-frequency microphonic waveform in detail to estimate changes in the maximal receptor current through the outer hair cells (OHCs), the sensitivity of the OHC forward transduction process and the change in OHC operating point on the mechano-electrical transduction transfer curve. Our results indicate that a 200 Hz tone changes the maximal current and sensitivity of the OHCs minimally, but more importantly, it transiently changes the operating point on the OHC transfer curve. In particular, the operating point changes are consistent with a movement of the OHC stereocilia away from the OHC basal body at the peak of the bounce. These changes detected using the microphonic potential are associated with changes in the level of the f 2-f 1 DPOAE that correlate well with the electrical measurements. We suggest that the shift in operating point is largely responsible for the increase in cochlear sensitivity, and is due to a disruption of the salt balance within the cochlea during the intense low-frequency tone.

Pressure-induced basilar membrane position shifts and the stimulus-evoked potentials in the low-frequency region of the guinea pig cochlea.

We have used the guinea pig isolated temporal bone preparation to investigate changes in the non-linear properties of the tone-evoked cochlear potentials during reversible step displacements of the basilar membrane towards either the scala tympani or the scala vestibuli. The position shifts were produced by changing the hydrostatic pressure in the scala tympani. The pressures involved were calculated from measurements of the fluid flow through the system, and the cochlear DC impedance calculated (1.5 x 10(11) kg m-4 s-1, n = 10). Confocal microscopic visualization of the organ of Corti showed that pressure increases in the scala tympani caused alterations of the position of the reticular lamina and stereocilia bundles. For low pressures, there was a sigmoidal relation between the DC pressure applied to the scala tympani (and thus the position shift of the organ of Corti) and the amplitude of the summating potential. The cochlear microphonic potential also showed a pronounced dependence on the applied pressure: pressure changes altered the amplitude of the fundamental as well as its harmonics. In addition, the sound pressure level at which the responses began to saturate was increased, implying a transition towards a linear behaviour. An increase of the phase lag of the cochlear microphonic potential was seen when the basilar membrane was shifted towards the scala vestibuli. We have also measured the intracochlear DC pressure using piezoresistive pressure transducers. The results are discussed in terms of changes in the non-linear properties of cochlear transduction. In addition, the implications of these results for the pathophysiology and diagnosis of Meniérè's disease are discussed.