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Biomedical subjects

R D Rabbitt

Publications and source records attributed to R D Rabbitt.

At least 19 recordsLinked to original sources

Evidence of piezoelectric resonance in isolated outer hair cells.

Our results demonstrate high-frequency electrical resonances in outer hair cells (OHCs) exhibiting features analogous to classical piezoelectric transducers. The fundamental (first) resonance frequency averaged f(n) approximately 13 kHz (Q approximately 1.7). Higher-order resonances were also observed. To obtain these results, OHCs were positioned in a custom microchamber and subjected to stimulating electric fields along the axis of the cell (1-100 kHz, 4-16 mV/80 microm). Electrodes embedded in the side walls of the microchamber were used in a voltage-divider configuration to estimate the electrical admittance of the top portion of the cell-loaded chamber (containing the electromotile lateral wall) relative to the lower portion (containing the basal plasma membrane). This ratio exhibited resonance-like electrical tuning. Resonance was also detected independently using a secondary 1-MHz radio-frequency interrogation signal applied transversely across the cell diameter. The radio-frequency interrogation revealed changes in the transverse electric impedance modulated by the axial stimulus. Modulation of the transverse electric impedance was particularly pronounced near the resonant frequencies. OHCs used in our study were isolated from the apical region of the guinea pig cochlea, a region that responds exclusively to low-frequency acoustic stimuli. In this sense, electrical resonances we observed in vitro were at least an order of magnitude higher (ultrasonic) than the best physiological frequency of the same OHCs under acoustic stimuli in vivo. These resonance data further support the piezoelectric theory of OHC function, and implicate piezoelectricity in the broad-band electromechanical behavior of OHCs underlying mammalian cochlear function.

Animals↗

Hair-cell versus afferent adaptation in the semicircular canals.

The time course and extent of adaptation in semicircular canal hair cells was compared to adaptation in primary afferent neurons for physiological stimuli in vivo to study the origins of the neural code transmitted to the brain. The oyster toadfish, Opsanus tau, was used as the experimental model. Afferent firing-rate adaptation followed a double-exponential time course in response to step cupula displacements. The dominant adaptation time constant varied considerably among afferent fibers and spanned six orders of magnitude for the population ( approximately 1 ms to >1,000 s). For sinusoidal stimuli (0.1-20 Hz), the rapidly adapting afferents exhibited a 90 degrees phase lead and frequency-dependent gain, whereas slowly adapting afferents exhibited a flat gain and no phase lead. Hair-cell voltage and current modulations were similar to the slowly adapting afferents and exhibited a relatively flat gain with very little phase lead over the physiological bandwidth and dynamic range tested. Semicircular canal microphonics also showed responses consistent with the slowly adapting subset of afferents and with hair cells. The relatively broad diversity of afferent adaptation time constants and frequency-dependent discharge modulations relative to hair-cell voltage implicate a subsequent site of adaptation that plays a major role in further shaping the temporal characteristics of semicircular canal afferent neural signals.

Adaptation, Physiological↗

Ultrastructural observations of efferent terminals in the crista ampullaris of the toadfish, Opsanus tau.

The present study was conducted to visualize the ultrastructural features of vestibular efferent boutons in the oyster toadfish, Opsanus tau. The crista ampullaris of the horizontal semicircular canal was processed for and examined by routine transmission electron microscopy. The results demonstrate that such boutons vary in size and shape, and contain a heterogeneous population of lucent vesicles with scattered dense core vesicles. Efferent contacts with hair cells are characterized by local vesicle accumulations in the presynaptic terminal and a subsynaptic cistern in the postsynaptic region of the hair cell. Serial efferent to hair cell to afferent synaptic arrangements are common, particularly in the central portion of the crista. However, direct contacts between efferent terminals and afferent neurites were not observed in our specimens. The existence of serial synaptic contacts, often with a row of vesicles in the efferent boutons lining the efferent-afferent membrane apposition, suggests that the efferent influence on the crista may involve both synaptic and nonsynaptic, secretory mechanisms. Further, it is suggested that differences in more subtle aspects of synaptic architecture and/or transmitter and receptor localization and interaction may render the efferent innervation of the peripheral crista less effective in influencing sensory processing.

Animals↗

Ultrastructural observations of efferent terminals in the crista Ampullaris of the toadfish, opsanus tau.

The present study was conducted to visualize the ultrastructural features of vestibular efferent boutons in the oyster toadfish, Opsanus tau. The crista ampullaris of the horizontal semicircular canal was processed for and examined by routine transmission electron microscopy. The results demonstrate that such boutons vary in size and shape, and contain a heterogeneous population of lucent vesicles with scattered dense core vesicles. Efferent contacts with hair cells are characterized by local vesicle accumulations in the presynaptic terminal and a subsynaptic cistern in the postsynaptic region of the hair cell. Serial efferent to hair cell to afferent synaptic arrangements are common, particularly in the central portion of the crista. However, direct contacts between efferent terminals and afferent neurites were not observed in our specimens. The existence of serial synaptic contacts, often with a row of vesicles in the efferent boutons lining the efferent-afferent membrane apposition, suggests that the efferent influence on the crista may involve both synaptic and nonsynaptic, secretory mechanisms. Further, it is suggested that differences in more subtle aspects of synaptic architecture and/or transmitter and receptor localization and interaction may render the efferent innervation of the peripheral crista less effective in influencing sensory processing.

Animals↗

Relationship between inner-ear fluid pressure and semicircular canal afferent nerve discharge.

The present study was designed to determine (1) the transcupular fluid pressure (deltaP) generated across the semicircular canal cupula in response to sinusoidal head rotation, (2) the translabyrinthine dilational pressure (P0) generated across the membranous labyrinth in response to an increase in endolymph fluid volume (hydrops), (3) afferent nerve discharge patterns generated by these distinct pressure stimuli and, (4) threshold values of deltaP and P0 required to elicit afferent neural responses. The experimental model was the oyster toadfish, Opsanus tau. Micromechanical indentation of the horizontal canal (HC) duct and utricular vestibule was used to simulate sinusoidal head rotation and fluid volume injection. Single-unit neural spike trains and endolymph pressure within the ampulla, on both sides of the cupula, were recorded simultaneously. deltaP averaged 0.013 Pa per 1 degrees/s of sinusoidal angular head velocity and P0 averaged 0.2 Pa per 1 nL of endolymph volume injection. The most responsive afferents had a threshold sensitivity to deltaP of 10(-3) Pa and to P0 of 5 x 10(-2) Pa based on a discharge modulation criterion of 1 impulse/s per cycle for 2 Hz pressure stimuli. Neural sensitivity to AP was expected on the basis of transverse cupular and hair bundle deflections. Analysis of mechanics of the end organ, neuronal projections into the crista, and individual neural firing patterns indicates that P0 sensitivity resulted from pressure-induced distension of the ampulla that led to a nonuniform cupular deformation pattern and hair bundle deflections. This explanation is consistent with predictions of a finite element model of the end organ. Results have implications regarding the role of deltaP in angular motion transduction and the role of P0 under transient hydropic conditions.

Action Potentials↗

Acoustic intensity, impedance and reflection coefficient in the human ear canal.

The sound power per unit cross-sectional area was determined in human ear canals using a new method based on measuring the pressure distribution (P) along the length of variable cross-section acoustic waveguides. The technique provides the pressure/power reflection coefficients (R/R) as well as the acoustic intensity of the nonplanar incident wave (I+, the acoustic input to the ear) and the nonplanar outgoing wave (I-, the acoustic output of the ear). Results were compared to the classical acoustic impedance (Z) and associated plane-wave power reflection coefficient (R(Z)). Performance of the method was investigated theoretically using horn equation simulations and evaluated experimentally using pressure data recorded in nonuniform waveguides. The method was applied in normal-hearing young adults to determine ear-canal position- and frequency-dependence of I(+/-), R, and R(Z) using random phase broadband stimuli (1-15 kHz; approximately 75 dB SPL). Reflection coefficient (R) measurements at two different locations within individual human ear canals exhibited a position dependence averaging deltaR approximately 0.1 (over 6 mm distance)--a difference consistent with predictions of inviscid acoustics in nonuniform waveguides. Since this position dependence was relatively small, an "optimized" position-independent reflection coefficient was defined to facilitate practical application and intersubject comparisons.

Acoustic Impedance Tests↗

Physiology of the semicircular canals after surgical plugging.

Inactivation of individual semicircular canals by surgical occlusion (plugging) of the slender duct has been used in basic studies to elucidate the role of individual canal inputs to vestibular-mediated control systems and in clinical applications to treat certain vestibular disorders. The procedure has been shown to be highly effective in blocking sensitivity of individual canals, at least for moderate angular motion stimuli. Effectiveness does not extend to stimuli involving high accelerations where a residual response persists even after complete occlusion of the duct. The residual can be quite large at high-stimulus frequencies where sensitivity to angular motion approaches that of patent canals. The overall physiological effect of canal plugging is reported here in terms of the frequency-dependent attenuation in gain and phase shift of primary afferents. Plug-canal responses are quantitatively described in terms of biomechanics of the deformable labyrinth.

Animals↗

How endolymph pressure modulates semicircular canal primary afferent discharge.

Histological observation of endolymphatic hydrops in subjects suffering from Meniere's disease/syndrome and the presence of vestibular symptoms in experimentally induced endolymphatic hydrops have led to considerable debate regarding the potential role of inner-ear pressure. Could a condition influencing endolymphatic volume regulation lead to transient changes in translabyrinthine pressure sufficiently large to alter primary afferent discharge rates? To investigate this question we built a highly sensitive laser-based pressure sensor and recorded endolymphatic pressure modulations within the horizontal semicircular canal ampulla in response to mechanical stimuli for simple head rotation and controlled volume injection. The most sensitive primary afferents responded to changes in endolymph pressures as low as 0.005 Pa (re: perilymph)--a pressure considerably lower than that observed in most experimentally induced hydropic conditions. This threshold pressure was generated by a approximately 25 picoliter volume injection stimulus. Semicircular canal afferent responses to endolymphatic pressure can be explained on the basis of pressure-induced hydrops, which causes dilation of the ampulla, deformation of the cupula, hair bundle movement, and afferent discharge. Primary afferent responses to maintained stimuli were transient in nature and recovered to pre-stimulus background discharge rates following a period of adaptation. Results demonstrate that the semicircular canals are indeed sensitive to small changes in translabyrinthine pressure in addition to head-movement-related changes in transcupular pressure.

Animals↗

Directional coding of three-dimensional movements by the vestibular semicircular canals.

A morphologically descriptive mathematical model was developed to study the role of labyrinthine geometry in determining sensitivities of each semicircular canal to angular motion stimuli in three-dimensional (3D) space. For this, equations describing viscous flow of the endolymph and poro-elastic response of the cupulae were coupled together and solved within a 3D reconstructed geometry. Results predict the existence of prime rotational directions about which the labyrinth resolves 3D angular movements into separate vectorial components. The components are predicted to be transmitted to the brain separately, one coded by each canal nerve. Prime directions predicted by the model are non-orthogonal, distinct from the anatomical canal planes, and distinct from the directions of rotation which elicit maximal responses of individual canal nerves. They occur for each canal along the intersection of the two null planes defined by its sister canals. Hence, rotation about a prime direction excites only one canal nerve. This contrasts the situation for rotations about anatomical canal planes, or about maximal response directions, where the model predicts activation of multiple canal nerves. The prime directions are sensitive to labyrinthine morphology and, hence, are predicted to vary between species and, to a lesser extent, vary between individual animals. Prime directions were estimated in the present work using a mathematical model, but could be determined experimentally based on the directional sensitivities of individual canal nerves. The model also predicts the existence of dominant eigenmodes and time constants associated with rotation in each of the prime directions. Results may have implications regarding the central representation of angular head movements in space as well as the neuronal mappings between three-canal afferent inputs and motor outputs.

Head Movements↗

Influence of surgical plugging on horizontal semicircular canal mechanics and afferent response dynamics.

Mechanical occlusion of one or more of the semicircular canals is a surgical procedure performed clinically to treat certain vestibular disorders and used experimentally to assess individual contributions of separate canals and/or otoliths to vestibular neural pathways. The present experiments were designed to determine if semicircular canal afferent nerve modulation to angular head acceleration is blocked by occlusion of the endolymphatic duct, and if not, what mechanism(s) might account for a persistent afferent response. The perilymphatic space was opened to gain acute access to the horizontal canal (HC) in the oyster toadfish, Opsanus tau. Firing rate responses of HC afferents to sinusoidal whole-body rotation were recorded in the unoccluded control condition, during the process of duct occlusion, and in the plugged condition. The results show that complete occlusion of the duct did not block horizontal canal sensitivity; individual afferents often exhibited a robust firing rate modulation in response to whole-body rotation in the plugged condition. At high stimulus frequencies (about >8 Hz) the average sensitivity (afferent gain; spikes/s per degrees /s of head velocity) in the plugged condition was nearly equal to that observed for unoccluded controls in the same animals. At low stimulus frequencies (about <0.1 Hz), the average sensitivity in the plugged condition was attenuated by more than two orders of magnitude relative to unoccluded controls. The peak afferent firing rate for sinusoidal stimuli was phase advanced approximately 90 degrees in plugged canals relative to their control counterparts for stimulus frequencies approximately 0.1-2 Hz. Data indicate that afferents normally sensitive to angular velocity in the control condition became sensitive to angular acceleration in the plugged condition, whereas afferents sensitive to angular acceleration in the control condition became sensitive to the derivative of acceleration or angular jerk in the plugged condition. At higher frequencies (>8 Hz), the phase of afferents in the plugged condition became nearly equal, on average, to that observed in controls. A three-dimensional biomechanical model of the HC was developed to interpret the residual response in the plugged condition. Labyrinthine fluids were modeled as incompressible and Newtonian; the membranous duct, osseous canal and temporal bone were modeled as visco-elastic materials. The predicted attenuation and phase shift in cupular responses were in close agreement with the observed changes in afferent response dynamics after canal plugging. The model attributes the response of plugged canals to labyrinthine fluid pressure gradients that lead to membranous duct deformation, a spatial redistribution of labyrinthine fluids and cupular displacement. Validity of the model was established through its ability to predict: the relationship between plugged canal responses and unoccluded controls (present study), the relationship between afferent responses recorded during mechanical indentation of the membranous duct and physiological head rotation, the magnitude and phase of endolymphatic pressure generated during HC duct indentation, and previous model results for cupular gain and phase in the rigid-duct case. The same model was adjusted to conform to the morphology of the squirrel monkey and of the human to investigate the possible influence of canal plugging in primates. Membranous duct stiffness and perilymphatic cavity stiffness were identified as the most salient model parameters. Simulations indicate that canal plugging may be the most effective in relatively small species having small labyrinths, stiff round windows, and stiff bony perilymphatic enclosures.

Afferent Pathways↗

Three-dimensional reconstruction of the membranous vestibular labyrinth in the toadfish, Opsanus tau.

Membranous vestibular labyrinths from the oyster toadfish, Opsanus tau, were fixed, dissected from the animal, stained, and embedded in rectangular blocks of clear histological resin. Photomicrographs of complete embedded labyrinths were taken from six orthogonal directions and used to construct three-dimensional (3D) geometrical models of the semicircular canals, ampullae, utricular vestibule and common crus. Membraneous ducts and ampullae were modeled using a set of cross-sectional elliptical curves laced together to generate curved tubular models of each structure. The ensemble of these curved tubes was used to generate a complete 3D reconstruction of the outside surface of the membranous labyrinth. When viewed from six orthogonal directions, reconstructions closely matched the embedded tissue. Dimensions of the reconstruction and histological sections were compared to measurements of fresh tissue taken from the same animals prior to fixation and used to correct the reconstructions for tissue shrinkage. Results provide estimates of the endolymphatic volumes, local cross-sectional areas and elliptical eccentricities as well as 3D orientations of the geometric canal planes relative to the skull. Ten micrometer histological sections of the material were also prepared to measure wall thickness in various regions of the labyrinth.

Animals↗

Determinants of semicircular canal afferent response dynamics in fish.

Present results separate the relative contributions of semicircular canal biomechanics from hair cell/afferent biophysics in determining the amplitude and phase of afferent responses to sinusoidal motion of the head. Separation was achieved by combining electrical polarization of the endolymph with mechanical indentation of the canal limb to modulate the instantaneous firing rate of horizontal semicircular canal afferents. The electrical stimulus drives hair cell transduction currents via modulation of the Nernst-Planck potential, whereas the mechanical stimulus mimics head rotation and modulates the open probability of the transduction channels. Responses for electrical polarization therefore reflect post-transduction-current (PTC) mechanisms, and responses for mechanical stimulation include the additional influence of canal mechanics. Linear transfer functions defining individual afferent response dynamics were obtained for low levels of each stimuli and are reported in Bode form providing gain (spikes/s per micron or mV) and phase (deg re: peak stim) over the frequency range from 0.02 to 40 Hz. Combined results for electrical and mechanical stimuli distinguish the component of sensory signal processing carried out by canal mechanics from that carried out by the hair cell/afferent complexes. Individual afferents were categorized according to their response to the mechanical stimuli as low-gain velocity (LG), high-gain velocity (HG) or acceleration (A) sensitive, groups as originally defined by Boyle and Highstein to describe interafferent diversity present within the population. In contrast to the results for mechanical stimuli, all afferent groups exhibit nearly equal increases in gain and phase for increasing frequencies of electrical stimulation. Comparison of individual afferent responses for the two stimuli leads to the conclusion that the LG, HG, and A groups are distinguished primarily by diversity in the mechanical activation of associated hair cells and not by PTC mechanisms. Even though PTC processing does not contribute significantly to determining these groups, it is the primary determinant underlying high-frequency gain and phase enhancements observed in the population average. Comparison of mechanical and electrical responses also reveals the mechanical lower-corner responsible for phase enhancements and gain decreases in all afferents at low frequencies of mechanical stimulation (< 0.05 Hz). Results imply that LG afferents encode angular head velocity by canceling a phase lag and gain attenuation due to the mechanics with a phase lead and gain enhancement due to PTC mechanisms above approximately 0.2 Hz. In contrast, A group afferents encode angular head acceleration by combining high-frequency phase leads and gain enhancements present in both the mechanics and PTC mechanisms across the physiological frequency spectrum. HG afferents fall between these two extremes, and, other than the influence of the mechanical lower-corner, their response primarily reflects PTC processing.

Action Potentials↗

Determinants of semicircular canal afferent response dynamics in the toadfish, Opsanus tau.

1. Present results determine the relative contributions of the biomechanical and the posttransduction-current (PTC) mechanisms to the sensory process carried out by the horizontal semicircular canal (HC) in the oyster toadfish, Opsanus tau. The role of each element was estimated using in vivo measurements of hair cell receptor potentials and afferent responses elicited by electrical stimuli and mechanical HC indentation. Individual afferent response dynamics are defined here using first-harmonic transfer functions presented in the form of response gain and phase for sinusoidal stimuli from approximately 0.02-30 Hz. Comparison of the response dynamics for the two types of stimuli distinguishes the mechanical and the PTC transfer functions leading to the neural response. The results show that both mechanisms contribute significantly to the overall signal processing performed by the semicircular canals. 2. Endolymphatic polarization and HC indentation. Modulation of the endolymphatic potential by current injection induces a differential voltage across the apical face of the hair cells that drives the transduction current directly via the Nernst-Planck potential. Results show that the electrical impedance of the apical tight junctions is much larger than the basal impedance to ground in O. tau, such that leakage current to the basolateral space is negligible and the voltage-sensitive basolateral currents remain fully functional during polarization of the endolymph (in the frequency range tested). Extracellular afferent responses to endolymphatic polarization were combined with responses to HC indentation to separate the relative contributions of the mechanical and the PTC mechanisms to the overall afferent response dynamics. Data show that more than one-half of the overall signal processing, as defined by the first-harmonic transfer function, persists even when canal mechanics is bypassed. 3. Hair-cell receptor potential modulation during HC indentation. Sharp microelectrodes were used to record the modulation of hair-cell receptor potentials (intracellular voltages) in vivo during physiological levels of sinusoidal HC indentation. Receptor potentials exhibit modulations dominated by the first harmonic and centered about the resting potential. The average gain of the receptor-potential modulation for HC indentation is approximately 0.88 mV/microns indent, corresponding to a value of 0.22 mV/deg/s head velocity, centered near zero phase over the range tested from 0.1-10 Hz. The present receptor potential data fall well short of spanning the full range of gain and phase present in the afferent population. Rather, intracellular hair-cell responses are consistent with the frequency-dependent mechanical activation of the transduction current as determined above. 4. Origins of individual afferent responses. The population of afferent responses forms a continuous distribution that is discussed here in terms of three groups as defined by Boyle and Highstein: velocity-sensitive low gain (LG) afferents, velocity/acceleration-sensitive high gain (HG) afferents, and acceleration-sensitive (A) afferents. The response dynamics of individual afferents were found to be determined by a mix of biomechanical and biophysical factors that vary systematically between these afferent groups. All afferents show low-frequency phase advancement and gain decrease during HC indentation associated with the mechanical lower-corner frequency and high-frequency phase and gain enhancements associated with the PTC processing. In highly phase-advanced afferents (A type), the mechanical response is additive with the PTC processing to achieve broad-band acceleration sensitive neural responses.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Mechanical indentation of the vestibular labyrinth and its relationship to head rotation in the toadfish, Opsanus tau.

1. In the present study we examine the response of the semicircular canal of the toadfish (Opsanus tau) to head rotation and to mechanical indentation of the membranous labyrinth. The relationship between the two stimuli is described by a new elastohydrodynamic model that delineates the three-dimensional (3-D) spatiotemporal distribution of endolymph pressure and flow. In vivo electrophysiological recordings of primary afferents supplying the horizontal canal (HC) were employed to validate the model predictions. Data were collected from 213 afferents in 18 fish during independent head rotation. HC indentation, utricle (U) indentation, and paired stimuli. To quantify the afferent response and the relationship between the applied sinusoidal stimuli, the magnitude (gain) and temporal relationship (phase) of the first harmonic of modulation were calculated and compared with theoretical predictions. 2. A mathematical based extensively on the 3-D morphology of a toadfish labyrinth and the physical properties of endolymph is presented to describe the relationship between head rotation and mechanical indentation. All model parameters specifying labyrinthine morphology and physical properties of endolymph are known; the model contains no free parameters. New results are independent of the structural properties of the cupula. The analysis employs an asymptotic solution of the Navier-Stokes equations in the three toroidal ducts that includes the 3-D fluid-structure interaction taking place within the enlarged ampulla. The solution addresses the differential pressure (delta P) acting across the cupula and the dilatational pressure acting on both sides of the cupula. The analysis quantifies the hydrodynamics of the HC for mechanical indentations of the long and slender portion of the canal duct (HC indentation) and the U (U indentation). Results specifically relate the indentation stimuli to head rotation. Linear commutations of HC indentation, U indentation, and rotation stimuli are analyzed by matching delta P acting across the cupula for the three stimulus modalities. 3. HC afferents show a linear correspondence between HC indentation, U indentation, and rotation stimuli. Specific experimental results for sinusoidal stimuli at frequencies < 2 Hz show 1) +/- 1 micron-HC indentation commutates with +/- 4 degrees/s rotation, 2) + 1-micron HC indentation commutates with -/+ 15-microns U indentation, and 3) -/+ 15-microns U indentation commutates with +/- 4 degrees/s rotation. These results were obtained by adjusting the relative amplitude and phase of two stimuli presented simultaneously to achieve destructive interaction that minimizes the afferent modulation (balanced). Equivalent results were obtained using afferent responses to the stimuli applied independently.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

3D brain mapping using a deformable neuroanatomy.

This paper presents two different mathematical methods that can be used separately or in conjunction to accommodate shape variabilities between normal human neuroanatomies. Both methods use a digitized textbook to represent the complex structure of a typical normal neuroanatomy. Probabilistic transformations on the textbook coordinate system are defined to accommodate shape differences between the textbook and images of other normal neuroanatomies. The transformations are constrained to be consistent with the physical properties of deformable elastic solids in the first method and those of viscous fluids in the second. Results presented in this paper demonstrate how a single deformable textbook can be used to accommodate normal shape variability.

Algorithms↗

Sensory transduction of head velocity and acceleration in the toadfish horizontal semicircular canal.

1. Sinusoidal mechanical indentation of the long-and-slender limb of the horizontal semicircular canal and/or utricle was used to produce adequate stimulation of the labyrinth. Indentation of the canal increased, while indentation of the utricle decreased the afferent discharge rate. This follows because indentation of the canal and utricle produce oppositely directed mechanical stimuli as defined by endolymph flow, transcupular pressure, and cupular deflection. Simultaneous in-phase indentations of both the canal and utricle, with amplitudes adjusted to produce equal (but opposite) magnitudes of afferent response modulation, generate destructive interaction that minimizes the afferent modulation, whereas sinusoidal indentation 180 degrees out-of-phase generates constructive interaction that maximizes the afferent modulation. This observation correlates directly with analysis of the labyrinthine elasto-hydrodynamics which predicts that balanced in-phase indentations minimize macromechanical endolymph flow through the ampullary cross section and maximize the dilatational pressure within the ampulla acting equally on both sides of the cupula and across the labyrinthine wall. 2. Two groups of afferents are identified according to their response to balanced sinusoidal indentation of the canal limb and the utricle. In one group there is complete destructive interaction and the afferent response can be effectively nulled by adjusting the relative amplitude and phase of the two stimuli. In the second group a residual afferent response remains that cannot be nulled. The residual is described in the model as unit-specific sensitivity to dilatational pressure acting equally on both sides of the cupula.

Afferent Pathways↗

Ear canal cross-sectional pressure distributions: mathematical analysis and computation.

Cross-sectional pressure distributions, natural acoustic modes, and associated cutoff frequencies are determined for real ear-canal geometries using an asymptotic theory in combination with a numerical method. The technique is particularly well suited to obtain the higher modes, which are trapped near both ends of the ear canal. Results detail the influence of the canal geometry and frequency on the spatial distribution of the pressure. Adult ear-canal geometries are determined near the concha from ear-mold sections using a light microscope interfaced to a video-data-acquisition system. Computed results compare favorably to the exact solutions for circular and square acoustic waveguides. The cutoff frequency of the two adult ear canals studied averaged 20% less than the cutoff frequency of a circular tube of identical cross-sectional area. Inserting a probe microphone into the canal decreases the rate of decay of circumferential nonplanar modes while increasing the rate of decay of radial modes. Relative to the pressure beyond the tube, insertion increases the plane-wave component of the pressure around the tube by a multiplicative factor approximately equal to the square root of the original area divided by the occluded area. Eccentric placement of the probe tube has a relatively small influence on the cutoff frequency. The transition of the pressure distribution at the entrance to a simple plane wave in the core region of the canal is calculated and shown graphically for the actual geometry of two adult subjects.

Computer Simulation↗

A hierarchy of examples illustrating the acoustic coupling of the eardrum.

Basic principles underlying the acoustic coupling of the eardrum are illustrated in the form of a hierarchy of examples ranging from a simple piston coupled to a semi-infinite acoustic duct, to a flexible partition coupled to a variable cross-section duct, and to a closed cavity. The hierarchy illuminates some of the limitations of various simplified elements commonly used to model the middle ear and demonstrates the necessity of treating the acoustics and the eardrum as an integrated subsystem. Results show that the tympanic cavity and the secondary middle-ear air chambers contribute fundamental features to the acoustic coupling of the ear. The nature of the acoustic coupling limits the passive energy absorption and transmission properties of the eardrum. The magnitude and frequency dependence of the energy dissipation within the ultrastructure of the partition, due to bending and transverse deflection, is discussed in analogy to possible dissipation mechanisms within the eardrum itself. Examples are provided for several simple systems reproducing some of the gross anatomical characteristics of the cat eardrum.

Acoustic Stimulation↗