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T F Weiss

Publications and source records attributed to T F Weiss.

At least 19 recordsLinked to original sources

Equilibrium behavior of an isotropic polyelectrolyte gel model of the tectorial membrane: effect of pH.

An isotropic polyelectrolyte gel model (Weiss and Freeman, 1996a) of the tectorial membrane (TM) was extended to incorporate the effect of pH. The effect of pH is analyzed in order to interpret measurements of the effect of pH on the dimensions of the TM (Freeman et al., 1996a). The pH dependence of the model results from the binding of hydrogen ions to TM macromolecules--to both neutral sites with basic pK and negatively charged sites with acidic pK. Parameters of the model can be based on estimates of the concentration in the TM of collagen and glycosaminoglycans (GAGs), two identified constituents of the TM that account for approximately 40% of its dry weight. The resulting model shows swelling responses at both high and low pH that are qualitatively similar to the measurement on the TM but that differ quantitatively. Alternatively, parameters of the model can be chosen in an ad hoc fashion to closely fit the measurements of TM swelling as a function of pH. Taken together these results suggest that either estimates of the collagen and GAG content of the TM are in error or constituents of the TM, which have not yet been identified, contain appreciable pH-dependent fixed charge and contribute to the swelling behavior of the TM.

Binding Sites↗

The osmotic response of the isolated, unfixed mouse tectorial membrane to isosmotic solutions: effect of Na+, K+, and Ca2+ concentration.

Changes in the size, shape, and structure of the isolated tectorial membrane (TM) of the mouse were measured in response to isosmotic changes in the ionic composition of the bathing solution. Substitution of artificial perilymph (AP) for artificial endolymph (AE) caused a small (approximately 1%) shrinkage of the TM's thickness. This substitution alters not only the predominate cation (from K+ to Na+) but also the Ca2+ concentration (from 20 mumol/l to 2 mmol/l). When the predominate cation was changed from K+ to Na+, while holding Ca2+ concentration constant, results depended on Ca2+ concentration: there was a small (approximately 1%) swelling for 20 mumol/l Ca2+, larger (approximately 14%) swelling for lower (< 7 mumol/l) concentrations of Ca2+, and little response for 2 mmol/l Ca2+ or for solutions containing the Ca2+ chelator EGTA. Addition of Ca2+ while holding the predominate cation constant caused shrinkage of the TM; both removal of Ca2+ and addition of the Ca2+ chelator EGTA caused swelling. Swelling responses were largely reversible if the magnitude of the swelling was small. Responses greater than a few percent were only partially reversible and caused long-lasting changes. Changes in ionic composition of the bath affected not only the thickness of the TM but also its other dimensions. Solution changes that increase TM thickness tend to cause radial shearing motions of the surfaces of the TM, which are accompanied by small decreases in width. Little change in length was observed. Although the responses were non-isotropic, increases in thickness were highly correlated with increases in volume. Swelling of the TM was also accompanied by a reduction in prominence of its radially oriented fibrillar structure. These results for the isolated TM of the mouse are qualitatively similar to those obtained previously for the isolated chick TM (Freeman et al., 1994) but different from those obtained for the in vitro mouse TM (Kronester-Frei, 1979a).

Acoustic Stimulation↗

The osmotic response of the isolated tectorial membrane of the chick to isosmotic solutions: effect of Na+, K+, and Ca2+ concentration.

Changes in the size, shape, and structure of the isolated tectorial membrane of the chick were measured in response to isosmotic changes in the ionic composition of the perfusion solution. Substitution of artificial perilymph (AP) for artificial endolymph (AE) caused a small (approximately 15%), slow (time constants tau approximately 12 min) shrinkage of the thickness of the tectorial membrane that was largely reversed on return to AE. Substitution of AP for AE alters not only the predominate cation (from K+ to Na+) but also the Ca2+ concentration (from < 7 mumol/l to 2 mmol/l). Additional experiments were performed to separate effects of each of these changes. When a high-Na+, low-Ca2+ solution was substituted for a high-K+, low-Ca2+ solution (AE), the tectorial membrane swelled significantly, often to more than twice its original thickness (the largest swelling was 337%), with a slow time course (tau approximately 23 min). Addition of the Ca2+ to either high-K+ or high-Na+ solutions caused rapid shrinkage of the tectorial membrane (tau approximately 2-3 min). Addition of the Ca2+ chelator EGTA caused rapid swelling (tau approximately 4 min). Large osmotic responses were only partially reversible and caused long-lasting changes. For example, long-duration solution changes that produced large, rapid osmotic responses early in an experiment tended to produce smaller and slower responses later in the experiment. In contrast, the small osmotic responses to short-duration solution changes were repeatable for tens of hours. Changes in ionic composition of the bath affected not only the thickness of the tectorial membrane but also its other dimensions. Responses were not generally isotropic; both the size and shape of the tectorial membrane generally changed. Consistent changes in microstructure accompanied the osmotic changes.

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Effect of lymph composition on an in vitro preparation of the alligator lizard cochlea.

The effects of different artificial lymphs on the cochlear duct of the alligator lizard were studied in an in vitro preparation. The duct was dissected and cemented to the glass floor of a chamber that had been filled with an artificial lymph. The vestibular membrane was removed and latex beads (1-5 microns in diameter) were allowed to settle on the endolymphatic surface of the duct. During perfusion with an artificial lymph solution, the positions of beads were measured and video images of the duct were obtained. Artificial lymphs were isosmotic and included artificial endolymph (AE), artificial perilymph (AP), Leibovitz's L-15 culture medium, an AE solution whose calcium concentration was the same as that of AP, and AE and AP solutions in which gluconate was substituted for chloride ions. Results obtained in AE were consistently different from those in other lymphs. The displacements of beads, the projected area of the papilla, the occurrence of blebs, and direct observation of cells in the duct all indicated that the tissue swelled in AE (with or without 2 mmol/l Ca) but showed no consistent shrinking or swelling in any of the other artificial lymphs. Thus for the solutions we used, the presence of both potassium and chloride was required to elicit the swelling response to isosmotic artificial lymphs. There were some regional differences in the swelling response: the swelling of the endolymphatic surface of the tissue in a direction orthogonal to the basilar membrane surface was smaller on the free-standing region of the basilar papilla than either on the tectorial membrane or on the hyaline epithelial cells. The preparation was osmotically stable in AP and in both AE and AP solutions in which gluconate was substituted for chloride ions. After exposure to these solutions for as much as 300 min, the preparation showed no gross signs of deterioration visible with the light microscope, and continued to exhibit a highly specific osmotic response to the composition of the bathing medium.

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Dependence of discharge rate on sound pressure level in cochlear nerve fibers of the alligator lizard: implications for cochlear mechanisms.

1. Rate-level functions for individual cochlear nerve fibers of the alligator lizard, Gerrhonotus multicarinatus, were generated by measuring a fiber's driven discharge rate (the difference between the average discharge rates in the presence and absence of a tone burst) as a function of sound pressure level. 2. When plotted in double logarithmic coordinates, the rate-level function approaches a straight line at low sound pressure levels and saturates at high levels. Thus the rate-level function is a saturating power function of sound pressure. We developed an algorithm to estimate the exponent of the straight-line portion of the function. When tested on simulated data with known parameters, the algorithm provided unbiased estimates of the exponent. 3. Nerve fibers innervating two distinct regions of the alligator lizard's auditory organ, the free-standing region and the tectorial region, have differing rate-level functions. 4. The mean exponent estimate of the rate-level functions of fibers innervating the free-standing region is approximately 2 at all frequencies. For stimulus frequencies at the characteristic frequency (CF), the mean value was 2.1 +/- 0.10 (SE, n = 131). For stimulus frequencies above and below CF, the mean exponent estimates were 2.1 +/- 0.13 (n = 49) and 2.1 +/- 0.11 (n = 34), respectively. A value of 2 is expected for a broad class of nonlinear systems. 5. The mean exponent estimates of the rate-level functions of fibers innervating the tectorial region were 3.0 +/- 0.30 (n = 32) for stimulus frequencies at CF, 2.5 +/- 0.33 (n = 3) for stimulus frequencies below CF, and 1.0 +/- 0.21 (n = 16) for stimulus frequencies above CF. Both the deviation from square-law behavior at CF and the frequency dependence of the exponent imply that nonlinear processing in the tectorial region differs intrinsically from that in the free-standing region. 6. For free-standing fibers, the saturation rate of the rate-level function (the maximum driven rate) is independent of stimulus frequency. This suggests that, in the free-standing region, 1) the alternating (AC) component of the receptor potential makes no significant contribution to the average rate of discharge and 2) neural saturation results from a process that occurs after the narrow-band frequency-selective process(es). 7. In tectorial fibers, the saturation rate is a bandpass function of sound frequency, with a broad peak between 150 and 300 Hz. This function appears to be common to all tectorial fibers.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Mechanisms that degrade timing information in the cochlea.

Action potentials of cochlear nerve fibers are synchronized to the temporal variations of sounds, but this synchronization is attenuated for high-frequency sounds. In cochleas from a number of vertebrates, the frequency dependence of synchronization can be represented as a lowpass filter process whose order is at least three (Weiss and Rose, 1988a); i.e. at least three first-order kinetic processes may be responsible for the loss of synchronization. In this paper we assess the extent to which calcium processes, that are essential for chemical transmission at the hair-cell neuron junction, contribute to this attenuation of synchronization. We analyze a model of calcium processes in hair cells (Lewis, 1985; Hudspeth and Lewis, 1988a) for sinusoidal receptor potentials. We show that: (1) the relation between the receptor potential and the calcium current, which is nonlinear, acts approximately as a first-order lowpass filter whose cut-off frequency decreases with increasing receptor potential magnitude; (2) the relation between calcium current and calcium-concentration is a first-order, lowpass filter with constant cutoff frequency. These two calcium processes plus the lowpass-filter process resulting from the electrical resistance and capacitance of the hair-cell membrane - which limits the rate at which the receptor potential can change (Weiss and Rose, 1988b) - can account for much, although perhaps not for all, of the loss of synchronization of cochlear nerve fibers.

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Superposition of hydrodynamic forces on a hair bundle.

Vertebrates sense sound, orientation, and motion by means of bundles of microscopic sensory hairs that protrude from the surfaces of receptor (hair) cells. To determine the effects of the sensory epithelium, tectorial structures, and fluids on the motions of hair bundles, we examine a class of mathematical models in which hair-cell organs are represented as a system of rigid mechanical structures surrounded by fluid. The epithelium and tectorial structures are represented by rigid basal and tectorial plates, respectively; the hair bundle by a rigid body hinged to the basal plate. When the displacements of these structures are small, the equations of motion for the fluid are predominately linear. Therefore, both the fluid velocity and the force of fluid origin on the body can be expressed as a sum of components; each component results from motion of a single structure while all others are stationary. This analysis leads to a network description of the motion of the rigid body in which hydrodynamic forces are segregated from mechanical forces. The separation of hydrodynamics and mechanics not only clarifies the effects of fluids on motion but also minimizes the number of hydrodynamic computations needed to analyze models of hair-bundle motion.

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Hydrodynamic forces on hair bundles at low frequencies.

We have analyzed a model for the motion of hair bundles of hair cells at low frequencies. In the model, hair-cell organs are represented as a system of rigid mechanical structures surrounded by fluid. A rigid body, that represents a hair bundle, is hinged to a vibrating plate that represents the sensory epithelium. These structures are surmounted by a second vibrating plate that represents a tectorial structure. The analysis shows that both viscous and inertial properties of the fluid are important even at asymptotically low frequencies. The relative importance of these properties depends critically on the presence and mode of motion of the tectorial plate. As a result, the angular displacement of the body at low frequencies can be proportional to basal plate displacement, velocity, acceleration, or to no simple integral of its motion; the functional relation depends upon the disposition of the tectorial plate.

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Hydrodynamic forces on hair bundles at high frequencies.

We have analyzed a model for the motion of hair bundles of hair cells at high frequencies. In the model, hair-cell organs are represented as a system of rigid mechanical structures surrounded by fluid. A rigid body, that represents a hair bundle, is hinged to a vibrating plate that represents the sensory epithelium. These structures are surmounted by a second vibrating plate that represents a tectorial structure. The analysis shows that at high frequencies, fluid forces cause the rigid body to move as though it were attached to the plates with a system of levers. As a result, the angular displacement of the rigid body is proportional to the displacements of the plates even when there are no mechanical attachments of the body to the tectorial plate. This result is independent of both the size and the shape of the rigid body and independent of the presence and proximity of the tectorial plate, although the constant of proportionality depends upon these factors. Therefore, the mechanical stimulation of hair cells may be particularly simple at high frequencies where the structural differences in hair bundles and tectorial attachments--that have been shown to be important at low frequencies--play a less important role.

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Hydrodynamic analysis of a two-dimensional model for micromechanical resonance of free-standing hair bundles.

To investigate the role of inner ear fluids and structures on mechanical stimulation of the hair bundles of hair cells, we analyzed a two-dimensional structure that consists of: a rectangular flap (which represents a hair bundle) attached to a flat basal plate (which represents the surface of the epithelium that contains the hair cells) with a spring-loaded hinge (that represents the compliant attachment of a hair bundle to the hair cell body) and surrounded by a viscous fluid (that represents endolymph). We computed the fluid velocity as well as the forces on and motion of the flap in response to sinusoidal vibration of the plate by numerical integration of the hydrodynamic equations, and--at asymptotically low and high frequencies--by analytic methods. The results suggest that: (1) the surface of the sensory epithelium, from which hair bundles project into fluid, plays an important part in the production of fluid forces on hair bundles; (2) both fluid inertia and viscosity play a key role in hair bundle mechanics; (3) passive mechanical resonances are likely to contribute to both frequency selectivity and frequency-to-place coding in the inner ear.

Algorithms↗

The role of fluid inertia in mechanical stimulation of hair cells.

Hair cells, which are the receptor cells of hearing and equilibrium in vertebrates, produce electrical responses when their hair bundles are displaced by sensory stimuli. This paper summarizes the results of a theoretical study of the fluid mechanics of hair-bundle motion. The principal conclusion is that fluid inertia, which has not been included in previous studies, plays a critical role in the mechanics of hair bundles and, hence, in the processes of sensory reception in hair-cell organs.

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Frequency dependence of synchronization of cochlear nerve fibers in the alligator lizard: evidence for a cochlear origin of timing and non-timing neural pathways.

The dependence of synchronization of spike discharges on tone frequency was measured in cochlear nerve fibers of anesthetized alligator lizards at 21 degrees C. Synchronization measures were based on the fundamental component of a Fourier analysis of the instantaneous discharge rate in response to tone bursts. Measurements were obtained from fibers innervating hair cells in both the region of the cochlea that contains a tectorial membrane (tectorial fibers) and the region where hair-cell stereocilia are free-standing in scala media (free-standing fibers). Both rate and synchronization tuning-curves were measured automatically as a function of tone frequency. For tectorial fibers, the shapes of synchronization tuning-curves are roughly similar to the shapes of rate tuning-curves: the characteristic frequencies (CF's) of both curves are approximately equal. For free-standing fibers, the shape of synchronization tuning-curves differ markedly from those of rate tuning-curves. The CF's of synchronization and rate tuning-curves differ - the ranges are 0.2-0.6 kHz and 1-4 kHz, respectively - and the two CF's are uncorrelated. Synchronization filter-functions, which are contours of synchronization index at constant average discharge rate, were measured as a function of tone frequency for both tectorial and free-standing fibers. These synchronization filter-functions have the shapes of lowpass filters. For the populations of tectorial fibers and of free-standing fibers taken separately, these functions are independent of CF. The corner frequency of these functions is 0.50 +/- 0.038 kHz for tectorial fibers and 0.37 +/- 0.037 kHz for free-standing fibers. We conclude that these populations are characterized by different synchronization filters. For free-standing fibers, synchronization filter-functions measured at average driven discharge rates of about 20 and 40 spikes/s do not differ appreciably, and the high-frequency slope is -80 to -115 dB/decade. The results show that tectorial fibers encode timing information for low-level stimuli, whereas free-standing fibers do not. It is proposed that in the alligator lizard, neural pathways that encode timing information originate in the tectorial region and those that encode non-timing information originate in the free-standing region.

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Stages of degradation of timing information in the cochlea: a comparison of hair-cell and nerve-fiber responses in the alligator lizard.

Responses to clicks and tone bursts of hair cells and nerve fibers in the free-standing region of the alligator lizard cochlea were compared. The objective was to determine the extent to which the hair-cell processes that produce the receptor potential are also responsible for the attenuation of the synchronized responses of nerve fibers. The AC component of the receptor potential of these hair cells has a high-frequency attenuation of 20 dB/decade [Holton and Weiss (1983) J. Physiol. 345, 205-240], whereas the synchronized response of cochlear neurons is attenuated at a rate of least 80 dB/decade [Rose and Weiss (1988) Hear. Res. 33, 151-166]. Therefore, the processes that link the receptor potential to the nerve discharge act as a lowpass filter with a high-frequency attenuation of at least 60 dB/decade. This could be obtained from a cascade of at least three first-order lowpass filter processes.

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A comparison of synchronization filters in different auditory receptor organs.

Measurements of the frequency dependence of synchronization of cochlear nerve fibers obtained in different auditory receptor organs are compared. These synchronization filter-functions are lowpass filter-functions and differ primarily in corner frequencies which we estimate to be (in kHz): 2.5 (cat), 1.1 (guinea pig), 0.48 (alligator-lizard tectorial fibers), 0.42 (tree frog), and 0.34 (alligator-lizard free-standing fibers). Some of this variation in corner frequency can be explained by temperature-dependent lowpass-filter mechanisms with a temperature factor of 2.6-3.3 for a change in temperature of 10 degrees C. However, factors in addition to temperature must be involved in producing the differences in corner frequency between cat and guinea pig fibers and between tectorial and free-standing fibers in the alligator lizard.

Acoustic Stimulation↗

A model for signal transmission in an ear having hair cells with free-standing stereocilia. I. Empirical basis for model structure.

No adequate theory for the signal-transmission properties of the peripheral auditory system exists for any vertebrate ear. Because the mammalian ear seems to pose conceptual and technical problems that complicate the development of an adequate theory, it is worthwhile to investigate simpler ears. The ear of the alligator lizard is simpler than mammalian ears in several respects: the motion of the basilar membrane is approximately independent of longitudinal position and is approximately linearly related to the sound pressure at the tympanic membrane; in a large region of the cochlea the hair cells have free-standing stereocilia that are not in contact with a tectorial membrane; the receptor potential of these hair cells is related to the sound pressure at the tympanic membrane in a relatively simple manner; the cochlear-nerve fiber responses from this region do not exhibit two-tone rate suppression. Also, the relative accessibility of this ear has enabled measurement of several response variables: tympanic-membrane volume velocity, extracolumella velocity, basilar-membrane velocity, hair-cell stereociliary displacement, hair-cell receptor potentials, and cochlear-nerve-fiber discharges. A model is developed to represent these results in terms of underlying anatomical structures and physiological mechanisms.(ABSTRACT TRUNCATED AT 250 WORDS)

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A model for signal transmission in an ear having hair cells with free-standing stereocilia. II. Macromechanical stage.

A model of the signal-processing properties of the macromechanical system of the alligator-lizard middle and inner ear is developed. The model is based on measurements (as a function of tone frequency) of ossicular and basilar-membrane motion and of acoustic input-admittance at the tympanic membrane before and after alterations of middle-ear and inner-ear structures. From the structure of the ear, we formulate an equivalent electric network consisting of six admittance blocks and two transformers. The admittance blocks represent the mechanical properties of particular structures, e.g. the tympanic membrane (TM), the ossicle, the TM-ossicular joint, the vestibule, the basilar membrane and the helicotrema. The transformers represent the conversions from acoustical to mechanical variables performed by the TM-ossicular system and by the columella footplate. The block admittances and transformer ratios are inferred from the measurements, and each admittance is approximated by a network with a few simple elements. The resulting model fits the experimental results satisfactorily, allows correlations between specific structures and response behavior, and predicts the macromechanical transfer function. This transfer function acts as the first stage of a model of the alligator-lizard auditory periphery (Weiss et al., Weiss and Leong, 1985, Hearing Res. 20, 131-138, 157-174, 175-195). In the model the frequency dependence of basilar-membrane motion (1) is determined primarily by the tympanic membrane and extracolumella in the middle frequencies and (2) is affected by the helicotrema at low frequencies and the TM-ossicular joint at high frequencies.

Acoustics↗

A model for signal transmission in an ear having hair cells with free-standing stereocilia. III. Micromechanical stage.

Measurements have shown that the sound-induced motion of free-standing stereocilia of hair cells in the alligator lizard cochlea exhibits tonotopically organized frequency selectivity that is correlated with the geometry of the stereociliary tuft. We propose a model in which basilar-membrane motion causes vibration of the receptor organ which drags the stereocilia back and forth through the endolymph. The stereociliary tuft is represented as a rigid rod attached to the cuticular plate by a compliant hinge. Viscous and inertial forces exerted by the endolymph on the rod are computed approximately. A transfer function, H mu(f), is derived that relates rod angular displacement to basilar-membrane velocity. H mu(f) has low- and high-frequency slopes of 10 and -20 dB/decade, respectively. The resonant frequency of H mu(f) depends on the dimensions of the rod because this frequency is inversely proportional to the square root of the product of the moment of inertia of the rod, which depends on rod dimensions, and the compliance of the hinge, which does not. In most respects, measurements of frequency selectivity and tonotopic organization of hair cells and cochlear neurons in the alligator lizard, can be accounted for by an input transfer function, HI(f) = Hm(f)H mu(f)Ha(f), where Hm(f) is the macromechanical transfer function that relates sound pressure at the tympanic membrane to basilar-membrane velocity (Rosowski et al., 1985, Hearing Res. 20, 139-155), and Ha(f) is a first-order lowpass filter. Mechanisms that could produce the additional lowpass filter process are discussed.

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A model for signal transmission in an ear having hair cells with free-standing stereocilia. IV. Mechanoelectric transduction stage.

A model is described for mechanoelectric transduction in hair cells with free-standing stereocilia in the alligator lizard cochlea. The model relates the angular displacement of the stereocilia to the receptor potential in the absence of the hair cell's membrane capacitance whose effect is considered elsewhere (Leong and Weiss, 1985, in preparation). The model consists of two parts: a population of membrane ionic channels and an electric network that relates the channel conductance to the equivalent resistance of the hair cell. The membrane ionic channels tend to open when the stereociliary tuft is displaced toward the kinocilium (or basal body) and tend to close when the tuft is oppositely displaced. The fraction of channels that is open for a given tuft displacement is governed by Boltzmann statistics and the energies of open and closed configurations of the channels are separated by a single energy barrier whose height depends on the angular displacement of the stereociliary tuft. The resulting channel conductance is a hyperbolic-tangent type function of the angular displacement of the stereociliary tuft. The channel conductance is coupled to the rest of the hair cell by an equivalent electric network containing constant resistance and a capacitance. The Thévenin equivalent resistance change across the basolateral membrane of the hair cell, called the transducer function, is also a hyperbolic-tangent type function of angular displacement. The parameters of the channel conductance and the values of resistances in the hair-cell electric model determine the scale factors and the location of the operating point of this hyperbolic-tangent type function. The hyperbolic-tangent type function is a specific example of a class of monotonically decreasing and saturating, or sigmoidal, transducer functions. The spectral properties of sigmoidal transducer functions are examined for sinusoidal angular displacements of amplitude theta. General results are obtained for arbitrary sigmoidal transducer functions; particular results are obtained for the hyperbolic-tangent type function. General conclusions concerning spectral components of the resistance change include: all spectral components are independent of the frequency of the angular displacement; the constant or DC component can be positive or negative; the fundamental component is 180 degrees out of phase with the angular displacement, i.e. the resistance decreases when the stereocilia are displaced towards the kinocilium; for small values of theta, the magnitude of the nth harmonic tends to grow as theta n for n greater than 0; the zeroth harmonic or DC component grows as theta 2.(ABSTRACT TRUNCATED AT 400 WORDS)

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