Nature's fastest motors are the cochlear outer hair cells (OHCs). These sensory cells use a m... more Nature's fastest motors are the cochlear outer hair cells (OHCs). These sensory cells use a membrane protein, Slc26a5 (prestin), to generate mechanical force at high frequencies, which is essential for explaining the exquisite hearing sensitivity of mammalian ears. Previous studies suggest that Slc26a5 continuously diffuses within the membrane, but how can a freely moving motor protein effectively convey forces critical for hearing? To provide direct evidence in OHCs for freely moving Slc26a5 molecules, we created a knockin mouse where Slc26a5 is fused with YFP. These mice and four other strains expressing fluorescently labeled membrane proteins were used to examine their lateral diffusion in the OHC lateral wall. All five proteins showed minimal diffusion, but did move after pharmacological disruption of membrane-associated structures with a cholesterol-depleting agent and salicylate. Thus, our results demonstrate that OHC lateral wall structure constrains the mobility of plasm...
... Some biologically important lipids, particularly those that support membrane curvature, can e... more ... Some biologically important lipids, particularly those that support membrane curvature, can enter an inverted hexagonal phase and form lipidic particles that resemble the particles found in the OHC PM (see Mouritsen and Kinnunen 1996 for review). ...
Quantitative confocal microscopy methods are used to measure events along the crypt-to-surface ax... more Quantitative confocal microscopy methods are used to measure events along the crypt-to-surface axis in living mouse colonic mucosa. Experiments visualize carboxyseminaphthorhodofluor-1 (SNARF-1; a pH-sensitive fluorescent dye) in the extracellular fluid to measure extracellular pH within an intact epithelium. Lucifer yellow (LY; a pH-insensitive dye) is used to control for fidelity of the optical path. Light scatter from colonic tissue caused SNARF-1 or LY fluorescence to decrease 3% per micrometer focal distance into tissue at both 640- and 580-nm emission wavelengths. However, dual emission ratios of LY fluorescence were constant as a function of focal distance into tissue or in the presence of short-chain fatty acids (SCFA). SCFA, known to cause changes in extracellular pH, cause maximal changes in crypt luminal pH at 10 microns from the crypt base. Maximal changes of pH in lamina propria occur higher along the crypt-to-surface axis than maximal changes in luminal pH. Lateral int...
1. Voltage-dependent properties of isolated guinea pig outer hair cells (OHCs) were investigated ... more 1. Voltage-dependent properties of isolated guinea pig outer hair cells (OHCs) were investigated using whole-cell recording. An inward current was detected in approximately 10% of the cells. This inward current was identified as belonging to the voltage-activated sodium current family on the basis of its high sensitivity to tetrodotoxin and the effect of substitution of impermeant ions. Although this is the first report of a sodium current in the mammalian cochlea, it differs from the classical neuronal sodium current by having a variable magnitude from cell to cell and an inactivation that is shifted to hyperpolarized potentials. The sensory processing role of hair cells in general and outer hair cells in particular could be disrupted by the presence of a regenerative voltage-dependent current. The functional role of the OHC sodium channels is puzzling, particularly as they may be silent in vivo.
Annual International Conference of the IEEE Engineering in Medicine and Biology - Proceedings, 2007
The electromotility of cochlear outer hair cells (OHCs) is a required process for normal hearing,... more The electromotility of cochlear outer hair cells (OHCs) is a required process for normal hearing, and involves a membrane-based mechanism in which the transmembrane protein, prestin, plays a central role. We have investigated the contribution of prestin to the mechanics and electromechanics of the cell membrane using membrane tethers formed from human embryonic kidney cells. Our results suggest that prestin appears to change membrane tension and amplify electrically-evoked force generation, while a single point mutation of alanine to tryptophan in prestin reduces electrically-evoked force generation without affecting the membrane tension. We propose that prestin and membrane work in synergy to produce the electrical and mechanical changes that are required during OHC electromotility. I. INTRODUCTION N the mammalian auditory system, the cochlear outer hair cells (OHCs), located within the organ of Corti, exhibit rapid electrically-induced movements known as electromotility [1, 2], a process required for the sensitivity and frequency-resolving capabilities during normal hearing. Recently, a transmembrane protein, prestin, was discovered in the OHCs of the mammalian cochlea [3]. Prestin (SLC26A5) is a member of the solute carrier family 26 (SLC26) of proteins, which are structurally related and have been shown to transport one or more anions including sulfate, chloride, bicarbonate, and iodide [4]. When prestin is expressed in non-auditory mammalian cells it endows the transfected cells with electromotility [3], and voltagedependent non-linear capacitance (NLC) [3], which serves as a reliable "signature" of electromotility [5]. Atomic force microscopy experiments have demonstrated that upon electrical stimulation of rat prestin-transfected human embryonic kidney (HEK) cells, mechanical forces up to a stimulus frequency of 20 kHz are generated [6]. Since electromotility involves the coupling between electrical and Manuscript received April 16, 2007.
Pflügers Archiv - European Journal of Physiology, 2011
Changing the concentration of cholesterol in the plasma membrane of isolated outer hair cells mod... more Changing the concentration of cholesterol in the plasma membrane of isolated outer hair cells modulates electromotility and prestin-associated charge movement, suggesting that a similar manipulation would alter cochlear mechanics. We examined cochlear function before and after depletion of membrane cholesterol with methyl-βcyclodextrin (MβCD) in an excised guinea pig temporal bone preparation. The mechanical response of the cochlear partition to acoustic and/or electrical stimulation was monitored using laser interferometry and time-resolved confocal microscopy. The electromechanical response in untreated preparations was asymmetric with greater displacements in response to positive currents. Exposure to MβCD increased the magnitude and asymmetry of the response, without changing the frequency tuning of sound-evoked mechanical responses or cochlear microphonic potentials. Sodium salicylate reversibly blocked the enhanced electromechanical response in cholesterol depleted preparations. The increase of sound-evoked vibrations during positive current injection was enhanced following MβCD in some preparations. Imaging was used to assess cellular integrity which remained unchanged after several hours of exposure to MβCD in several preparations. The enhanced electromechanical response reflects an increase in outer hair cell electromotility and may reveal features of cholesterol distribution and trafficking in outer hair cells.
The apical surface of vertebrate inner ear sensory cells is characterized by a bundle of giant mi... more The apical surface of vertebrate inner ear sensory cells is characterized by a bundle of giant microvilli commonly known as stereocilia. Stereocilia bend about a neck-like thinning near their base and more than three decades of research has established that the direction and magnitude of sideways bundle deflection is the basis of the mechanoelectrical signalling that initiates sound perception. Aside
The effect of decreasing membrane cholesterol on the mechanical response of the cochlea to acoust... more The effect of decreasing membrane cholesterol on the mechanical response of the cochlea to acoustic and∕or electrical stimulation was monitored using laser interferometry. In contrast to pharmacological interventions that typically decrease cochlear electromechanics, reducing membrane cholesterol increased the response. The electromechanical response in untreated preparations was asymmetric with greater displacements in response to positive currents and cholesterol depletion increased the
The mechanism responsible for electromotility of outer hair cells in the ear is unknown but is th... more The mechanism responsible for electromotility of outer hair cells in the ear is unknown but is thought to reside within the plasma membrane. Lipid lateral diffusion in the outer hair cell plasma membrane is a sigmoidal function of transmembrane potential and bathing media osmolality. Cell depolarization or hyposmotic challenge shorten the cell and reduce membrane fluidity by half. Changing the membrane tension with amphipathic drugs results in similar reductions. These dynamic changes in membrane fluidity represent the modulation of membrane tension by lipid-protein interactions. The voltage dependence may be associated with the force-generating motors that contribute to the exquisite sensitivity of mammalian hearing.
Tethers are thin membrane tubes that can be formed when relatively small and localized forces are... more Tethers are thin membrane tubes that can be formed when relatively small and localized forces are applied to cellular membranes and lipid bilayers. Tether pulling experiments have been used to better understand the fine membrane properties. These include the interaction between the plasma membrane and the underlying cytoskeleton, which is an important factor affecting membrane mechanics. We use a computational method aimed at the interpretation and design of tether pulling experiments in cells with a strong membranecytoskeleton attachment. In our model, we take into account the detailed information in the topology of bonds connecting the plasma membrane and the cytoskeleton. We compute the force-dependent piecewise membrane deflection and bending as well as modes of stored energy in three major regions of the system: body of the tether, membrane-cytoskeleton attachment zone, and the transition zone between the two. We apply our method to three cells: cochlear outer hair cells ͑OHCs͒, human embryonic kidney ͑HEK͒ cells, and Chinese hamster ovary ͑CHO͒ cells. OHCs have a special system of pillars connecting the membrane and the cytoskeleton, and HEK and CHO cells have the membrane-cytoskeleton adhesion arrangement via bonds ͑e.g., PIP2͒, which is common to many other cells. We also present a validation of our model by using experimental data on CHO and HEK cells. The proposed method can be an effective tool in the analyses of experiments to probe the properties of cellular membranes.
The outer hair cell can be divided into three domains: the apex, the base, and the lateral wall. ... more The outer hair cell can be divided into three domains: the apex, the base, and the lateral wall. With the use of filipin, a polyene fluorescent antibiotic that binds to cholesterol, we found under fluorescence microscopy that the lateral wall membranes were less intensely stained than the apical and basal membranes. This difference in filipin fluorescence between the lateral walls and the ends diminished when cells were incubated in water-soluble cholesterol before staining, suggesting that exogenous cholesterol enters the lateral wall. Under confocal microscopy, we studied the incorporation pattern of a fluorescent cholesterol analogue, NBD-cholesterol. NBD-cholesterol did not stain the apical membranes whereas it intensely labeled the lateral wall. The micropipette aspiration technique was used to assess the effect of cholesterol on lateral wall stiffness. The lateral wall stiffness parameter of cells treated with water-soluble cholesterol ( n= 23) was significantly higher than th...
Hearing relies on mechanical stimulation of stereocilia bundles on the sensory cells of the inner... more Hearing relies on mechanical stimulation of stereocilia bundles on the sensory cells of the inner ear. When sound hits the ear, these stereocilia pivot about a neck-like taper near their base. More than three decades of research have established that sideways deflection of stereocilia is essential for converting mechanical stimuli into electrical signals. Here we show that mammalian outer hair cell stereocilia not only move sideways but also change length during sound stimulation. Currents that enter stereocilia through mechanically sensitive ion channels control the magnitude of both length changes and bundle deflections in a reciprocal manner: the smaller the length change, the larger is the bundle deflection. Thus, the transduction current is important for maintaining the resting mechanical properties of stereocilia. Hair cell stimulation is most effective when bundles are in a state that ensures minimal length change.
The Journal of the Acoustical Society of America, 1983
We have developed a method for measuring current density within the fluid spaces of the cochlea a... more We have developed a method for measuring current density within the fluid spaces of the cochlea and report the existence of stimulus evoked radial currents in scala tympani of the guinea pig cochlea. The spatial distribution of electrical potentials in scala tympani was measured along a radial path parallel to the basilar membrane. Click evoked potentials were recorded at successive points separated by a fixed increment as the electrode was either advanced from the spiral ligament or withdrawn from a position near the modiolus. Potential differences were found to exist between recording points and gradients were calculated from the evoked potential measurements. Evoked potential gradients are observed at the same position along the path of the electrode both on advancing and on withdrawing the electrode. The largest potential gradients are located beneath the organ of Corti. Condensation and rarefaction clicks produce radial currents in opposite directions at a given location along the electrode's path. The magnitude and spatial distribution of radial currents is a function of stimulus intensity. Potential gradients of small magnitude are observed at locations other than below the organ of Corti in some penetrations. Control experiments suggest the smaller gradients are artifactual and may result from displacement of the spiral ligament by the recording electrode. The locations, magnitude, and direction of intracochlear ionic flow relate directly to the mechano-electrical transduction process in the organ of Corti.
The Journal of the Acoustical Society of America, 1995
Brownell et al. [Science 227, 194-196 (1985)] observed that an isolated, cylindrically shaped coc... more Brownell et al. [Science 227, 194-196 (1985)] observed that an isolated, cylindrically shaped cochlear outer hair cell can change its length when an electric field is applied. In their experiments, the cell was fixed at one end, and located between two electrodes which lie on the cell axis but were positioned far from the cell. Kachar et al. [Nature 322, 365-368 (1986)] had suggested that the cell's electrically evoked elongation could be caused by pressure gradients resulting from electro-osmosis of the intracellular fluid. A mathematical model is developed which predicts the length change that would result from electro-osmotically generated pressure gradients inside the cell. Estimated parameter values are included to demonstrate that the pressures generated by electro-osmosis inside the cell would result in elongations that are at least two orders of magnitude below the experimentally measured values.
Nature's fastest motors are the cochlear outer hair cells (OHCs). These sensory cells use a m... more Nature's fastest motors are the cochlear outer hair cells (OHCs). These sensory cells use a membrane protein, Slc26a5 (prestin), to generate mechanical force at high frequencies, which is essential for explaining the exquisite hearing sensitivity of mammalian ears. Previous studies suggest that Slc26a5 continuously diffuses within the membrane, but how can a freely moving motor protein effectively convey forces critical for hearing? To provide direct evidence in OHCs for freely moving Slc26a5 molecules, we created a knockin mouse where Slc26a5 is fused with YFP. These mice and four other strains expressing fluorescently labeled membrane proteins were used to examine their lateral diffusion in the OHC lateral wall. All five proteins showed minimal diffusion, but did move after pharmacological disruption of membrane-associated structures with a cholesterol-depleting agent and salicylate. Thus, our results demonstrate that OHC lateral wall structure constrains the mobility of plasm...
... Some biologically important lipids, particularly those that support membrane curvature, can e... more ... Some biologically important lipids, particularly those that support membrane curvature, can enter an inverted hexagonal phase and form lipidic particles that resemble the particles found in the OHC PM (see Mouritsen and Kinnunen 1996 for review). ...
Quantitative confocal microscopy methods are used to measure events along the crypt-to-surface ax... more Quantitative confocal microscopy methods are used to measure events along the crypt-to-surface axis in living mouse colonic mucosa. Experiments visualize carboxyseminaphthorhodofluor-1 (SNARF-1; a pH-sensitive fluorescent dye) in the extracellular fluid to measure extracellular pH within an intact epithelium. Lucifer yellow (LY; a pH-insensitive dye) is used to control for fidelity of the optical path. Light scatter from colonic tissue caused SNARF-1 or LY fluorescence to decrease 3% per micrometer focal distance into tissue at both 640- and 580-nm emission wavelengths. However, dual emission ratios of LY fluorescence were constant as a function of focal distance into tissue or in the presence of short-chain fatty acids (SCFA). SCFA, known to cause changes in extracellular pH, cause maximal changes in crypt luminal pH at 10 microns from the crypt base. Maximal changes of pH in lamina propria occur higher along the crypt-to-surface axis than maximal changes in luminal pH. Lateral int...
1. Voltage-dependent properties of isolated guinea pig outer hair cells (OHCs) were investigated ... more 1. Voltage-dependent properties of isolated guinea pig outer hair cells (OHCs) were investigated using whole-cell recording. An inward current was detected in approximately 10% of the cells. This inward current was identified as belonging to the voltage-activated sodium current family on the basis of its high sensitivity to tetrodotoxin and the effect of substitution of impermeant ions. Although this is the first report of a sodium current in the mammalian cochlea, it differs from the classical neuronal sodium current by having a variable magnitude from cell to cell and an inactivation that is shifted to hyperpolarized potentials. The sensory processing role of hair cells in general and outer hair cells in particular could be disrupted by the presence of a regenerative voltage-dependent current. The functional role of the OHC sodium channels is puzzling, particularly as they may be silent in vivo.
Annual International Conference of the IEEE Engineering in Medicine and Biology - Proceedings, 2007
The electromotility of cochlear outer hair cells (OHCs) is a required process for normal hearing,... more The electromotility of cochlear outer hair cells (OHCs) is a required process for normal hearing, and involves a membrane-based mechanism in which the transmembrane protein, prestin, plays a central role. We have investigated the contribution of prestin to the mechanics and electromechanics of the cell membrane using membrane tethers formed from human embryonic kidney cells. Our results suggest that prestin appears to change membrane tension and amplify electrically-evoked force generation, while a single point mutation of alanine to tryptophan in prestin reduces electrically-evoked force generation without affecting the membrane tension. We propose that prestin and membrane work in synergy to produce the electrical and mechanical changes that are required during OHC electromotility. I. INTRODUCTION N the mammalian auditory system, the cochlear outer hair cells (OHCs), located within the organ of Corti, exhibit rapid electrically-induced movements known as electromotility [1, 2], a process required for the sensitivity and frequency-resolving capabilities during normal hearing. Recently, a transmembrane protein, prestin, was discovered in the OHCs of the mammalian cochlea [3]. Prestin (SLC26A5) is a member of the solute carrier family 26 (SLC26) of proteins, which are structurally related and have been shown to transport one or more anions including sulfate, chloride, bicarbonate, and iodide [4]. When prestin is expressed in non-auditory mammalian cells it endows the transfected cells with electromotility [3], and voltagedependent non-linear capacitance (NLC) [3], which serves as a reliable "signature" of electromotility [5]. Atomic force microscopy experiments have demonstrated that upon electrical stimulation of rat prestin-transfected human embryonic kidney (HEK) cells, mechanical forces up to a stimulus frequency of 20 kHz are generated [6]. Since electromotility involves the coupling between electrical and Manuscript received April 16, 2007.
Pflügers Archiv - European Journal of Physiology, 2011
Changing the concentration of cholesterol in the plasma membrane of isolated outer hair cells mod... more Changing the concentration of cholesterol in the plasma membrane of isolated outer hair cells modulates electromotility and prestin-associated charge movement, suggesting that a similar manipulation would alter cochlear mechanics. We examined cochlear function before and after depletion of membrane cholesterol with methyl-βcyclodextrin (MβCD) in an excised guinea pig temporal bone preparation. The mechanical response of the cochlear partition to acoustic and/or electrical stimulation was monitored using laser interferometry and time-resolved confocal microscopy. The electromechanical response in untreated preparations was asymmetric with greater displacements in response to positive currents. Exposure to MβCD increased the magnitude and asymmetry of the response, without changing the frequency tuning of sound-evoked mechanical responses or cochlear microphonic potentials. Sodium salicylate reversibly blocked the enhanced electromechanical response in cholesterol depleted preparations. The increase of sound-evoked vibrations during positive current injection was enhanced following MβCD in some preparations. Imaging was used to assess cellular integrity which remained unchanged after several hours of exposure to MβCD in several preparations. The enhanced electromechanical response reflects an increase in outer hair cell electromotility and may reveal features of cholesterol distribution and trafficking in outer hair cells.
The apical surface of vertebrate inner ear sensory cells is characterized by a bundle of giant mi... more The apical surface of vertebrate inner ear sensory cells is characterized by a bundle of giant microvilli commonly known as stereocilia. Stereocilia bend about a neck-like thinning near their base and more than three decades of research has established that the direction and magnitude of sideways bundle deflection is the basis of the mechanoelectrical signalling that initiates sound perception. Aside
The effect of decreasing membrane cholesterol on the mechanical response of the cochlea to acoust... more The effect of decreasing membrane cholesterol on the mechanical response of the cochlea to acoustic and∕or electrical stimulation was monitored using laser interferometry. In contrast to pharmacological interventions that typically decrease cochlear electromechanics, reducing membrane cholesterol increased the response. The electromechanical response in untreated preparations was asymmetric with greater displacements in response to positive currents and cholesterol depletion increased the
The mechanism responsible for electromotility of outer hair cells in the ear is unknown but is th... more The mechanism responsible for electromotility of outer hair cells in the ear is unknown but is thought to reside within the plasma membrane. Lipid lateral diffusion in the outer hair cell plasma membrane is a sigmoidal function of transmembrane potential and bathing media osmolality. Cell depolarization or hyposmotic challenge shorten the cell and reduce membrane fluidity by half. Changing the membrane tension with amphipathic drugs results in similar reductions. These dynamic changes in membrane fluidity represent the modulation of membrane tension by lipid-protein interactions. The voltage dependence may be associated with the force-generating motors that contribute to the exquisite sensitivity of mammalian hearing.
Tethers are thin membrane tubes that can be formed when relatively small and localized forces are... more Tethers are thin membrane tubes that can be formed when relatively small and localized forces are applied to cellular membranes and lipid bilayers. Tether pulling experiments have been used to better understand the fine membrane properties. These include the interaction between the plasma membrane and the underlying cytoskeleton, which is an important factor affecting membrane mechanics. We use a computational method aimed at the interpretation and design of tether pulling experiments in cells with a strong membranecytoskeleton attachment. In our model, we take into account the detailed information in the topology of bonds connecting the plasma membrane and the cytoskeleton. We compute the force-dependent piecewise membrane deflection and bending as well as modes of stored energy in three major regions of the system: body of the tether, membrane-cytoskeleton attachment zone, and the transition zone between the two. We apply our method to three cells: cochlear outer hair cells ͑OHCs͒, human embryonic kidney ͑HEK͒ cells, and Chinese hamster ovary ͑CHO͒ cells. OHCs have a special system of pillars connecting the membrane and the cytoskeleton, and HEK and CHO cells have the membrane-cytoskeleton adhesion arrangement via bonds ͑e.g., PIP2͒, which is common to many other cells. We also present a validation of our model by using experimental data on CHO and HEK cells. The proposed method can be an effective tool in the analyses of experiments to probe the properties of cellular membranes.
The outer hair cell can be divided into three domains: the apex, the base, and the lateral wall. ... more The outer hair cell can be divided into three domains: the apex, the base, and the lateral wall. With the use of filipin, a polyene fluorescent antibiotic that binds to cholesterol, we found under fluorescence microscopy that the lateral wall membranes were less intensely stained than the apical and basal membranes. This difference in filipin fluorescence between the lateral walls and the ends diminished when cells were incubated in water-soluble cholesterol before staining, suggesting that exogenous cholesterol enters the lateral wall. Under confocal microscopy, we studied the incorporation pattern of a fluorescent cholesterol analogue, NBD-cholesterol. NBD-cholesterol did not stain the apical membranes whereas it intensely labeled the lateral wall. The micropipette aspiration technique was used to assess the effect of cholesterol on lateral wall stiffness. The lateral wall stiffness parameter of cells treated with water-soluble cholesterol ( n= 23) was significantly higher than th...
Hearing relies on mechanical stimulation of stereocilia bundles on the sensory cells of the inner... more Hearing relies on mechanical stimulation of stereocilia bundles on the sensory cells of the inner ear. When sound hits the ear, these stereocilia pivot about a neck-like taper near their base. More than three decades of research have established that sideways deflection of stereocilia is essential for converting mechanical stimuli into electrical signals. Here we show that mammalian outer hair cell stereocilia not only move sideways but also change length during sound stimulation. Currents that enter stereocilia through mechanically sensitive ion channels control the magnitude of both length changes and bundle deflections in a reciprocal manner: the smaller the length change, the larger is the bundle deflection. Thus, the transduction current is important for maintaining the resting mechanical properties of stereocilia. Hair cell stimulation is most effective when bundles are in a state that ensures minimal length change.
The Journal of the Acoustical Society of America, 1983
We have developed a method for measuring current density within the fluid spaces of the cochlea a... more We have developed a method for measuring current density within the fluid spaces of the cochlea and report the existence of stimulus evoked radial currents in scala tympani of the guinea pig cochlea. The spatial distribution of electrical potentials in scala tympani was measured along a radial path parallel to the basilar membrane. Click evoked potentials were recorded at successive points separated by a fixed increment as the electrode was either advanced from the spiral ligament or withdrawn from a position near the modiolus. Potential differences were found to exist between recording points and gradients were calculated from the evoked potential measurements. Evoked potential gradients are observed at the same position along the path of the electrode both on advancing and on withdrawing the electrode. The largest potential gradients are located beneath the organ of Corti. Condensation and rarefaction clicks produce radial currents in opposite directions at a given location along the electrode's path. The magnitude and spatial distribution of radial currents is a function of stimulus intensity. Potential gradients of small magnitude are observed at locations other than below the organ of Corti in some penetrations. Control experiments suggest the smaller gradients are artifactual and may result from displacement of the spiral ligament by the recording electrode. The locations, magnitude, and direction of intracochlear ionic flow relate directly to the mechano-electrical transduction process in the organ of Corti.
The Journal of the Acoustical Society of America, 1995
Brownell et al. [Science 227, 194-196 (1985)] observed that an isolated, cylindrically shaped coc... more Brownell et al. [Science 227, 194-196 (1985)] observed that an isolated, cylindrically shaped cochlear outer hair cell can change its length when an electric field is applied. In their experiments, the cell was fixed at one end, and located between two electrodes which lie on the cell axis but were positioned far from the cell. Kachar et al. [Nature 322, 365-368 (1986)] had suggested that the cell's electrically evoked elongation could be caused by pressure gradients resulting from electro-osmosis of the intracellular fluid. A mathematical model is developed which predicts the length change that would result from electro-osmotically generated pressure gradients inside the cell. Estimated parameter values are included to demonstrate that the pressures generated by electro-osmosis inside the cell would result in elongations that are at least two orders of magnitude below the experimentally measured values.
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Papers by W. Brownell