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Peripheral Mechanisms of Hearing

Peter Dallos

10.1002/cphy.cp010314

Source: Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes

Originally published: 1984

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Overview and Gross Anatomy
2 Evolution of Ears and Their Variety
2.1 Evolution of the Inner Ear
2.2 Evolution of the middle ear
3 Sensory Receptors
3.1 Cochlear Hair Cells in the Mammal
3.2 Hair Cells as Transducers
4 Function of Cochlear Hair Cells
4.1 Events at Receptive Region of Cells
4.2 Anatomy of Mammalian Cochlear Duct
4.3 Generation of Receptor Current
4.4 Micromechanics of Organ of Corti
5 Input to Organ of Corti: Basilar Membrane Mechanics
5.1 Structure and Properties of Cochlear Canal and Basilar Membrane
5.2 The Traveling Wave
5.3 Nonlinearities
5.4 The Second Filter
6 The Middle and Outer Ear
6.1 Evolutionary Necessity
6.2 The Middle Ear
6.3 The Outer Ear
7 Electrical Signs of Transduction: Cochlear Potentials
7.1 Recording Methods
7.2 Intracellular Recording
7.3 Gross Recording
8 Output of Receptor Cells
8.1 Transmitter Substances
8.2 Site of Impulse Initiation
8.3 Action of Efferent Nerve Fibers
9 Summary
10 Note Added in Proof
Figure 1. Figure 1.

Block diagram of auditory system. Center, anatomical subdivisions, arrows indicating flow of information. Top, functions of the various segments. Bottom, the dominant mode of operation.

From Dallos 18
Figure 2. Figure 2.

Reproduction of Brödel's classic drawing of cross section of human ear.

From Brödel 13
Figure 3. Figure 3.

Schematic diagram showing basic morphological components of common hair cell. At apex of cell body several stereocilia emerge from a cuticular plate, and a single kinocilium is anchored to the intracellular basal body. Presynaptic specializations such as the synaptic bar (dark circle) and vesicles (small open circles) are seen adjacent to afferent nerve terminal, while efferent endings themselves are full of vesicles (small circles).

From Flock 50
Figure 4. Figure 4.

Cross section of mammalian cochlear duct with insets showing detailed structure of outer (left) and inner (right) hair cells. All important structures within scala media are identified; scala media space itself contains endolymph that is probably free to flow into subtectorial space and inner sulcus through noncontinuous marginal net. Intercellular spaces within organ of Corti are filled with cortilymph, which is probably not markedly different from perilymph. The morphological specializations depicted within the 2 hair cell types are described in detail in text.
Figure 5. Figure 5.

Scanning electron microscope picture of fracture in organ of Corti. Comparison with Fig. 4 identifies the various structures, although shape of organ is different because this figure depicts a more apical location where volume of organ of Corti is greater and outer hair cells and Hensen's cells are longer than at the more basal location depicted in Fig. 4.

From Bredberg et al. 187
Figure 6. Figure 6.

Scanning electron microscope picture from vantage point of tectorial membrane. Cilia from the single row of inner hair cells are seen on top, the top of pillar cells below them, and W‐shaped groups of cilia on top of 3 rows of outer hair cells below pillar cells. On bottom foreground, phalangeal processes of Deiters' cells are seen, and behind them the cell bodies of outer hair cells are discernible.

[From Bredberg et al. 187
Figure 7. Figure 7.

Schematic diagram showing hair cell and associated nerve endings as well as structural (center) and functional (right) block diagrams of system. Insets (left) demonstrate wave forms of the various quantities that are indicated in functional block diagram. It is assumed here that a brief sinusoidal burst stimulates the hair cell with a sinusoidal mechanical deformation (top). This results in 2 receptor potential components, an AC (cochlear microphonic) and a DC (summating potential) response. A wave form of postsynaptic potential, presumably arising in dendritic region of auditory nerve, and a spike train traveling in myelinated axon are also shown.

Adapted from Dallos 18
Figure 8. Figure 8.

Various conceivable modes of hair bundle deflection (stipples). A: bundle at rest; B: highly compliant bundle would bend; C: very stiff bundle could be displaced laterally in its entirety; D: stiff bundle with strongly adhering individual hairs could rotate about its insertion into cuticular plate; E: bundle of stiff individual hairs only weakly coupled to one another could fan out, with hairs rotating around their point of insertion. In all cases it is assumed that force is exerted at tip of tallest cilium.

From Flock et al. 52
Figure 9. Figure 9.

Cross section of cochlea with a plausible, but simplified, electric circuit superimposed upon it. Batteries represent voltage sources located in stria vascularis and in hair cells. Resistances are those of membranes separating various cochlear compartments; variable resistance, located within schematized receptor cell, is assumed to be controlled by mechanical deformation of cell. Change in this resistance produces a modulation of resting current through it and thus a modulation of potentials that can be measured at the various nodes of circuit.

From Davis and Silverman 190. In: Hearing and Deafness (3rd ed.), edited by Hallowell Davis and S. Richard Silverman. Copyright 1947 © 1960, 1970 by Holt, Rinehart and Winston, Inc. Reprinted by permission of Holt, Rinehart and Winston
Figure 10. Figure 10.

Pattern of excitation and inhibition of output of hair cell as function of direction of ciliary displacement. If θ is angle between direction of excitation and axis of morphological polarization through kinocilium, then the approximate dependence of output upon this angle is cos θ. This is figure‐eight pattern shown on top.

From Flock 49
Figure 11. Figure 11.

Schematic diagrams showing aspects of mechanics of basilar membrane organ of Corti complex. A: framework of organ of Corti is provided by basilar membrane (at bottom) and rigid pillar cells and reticular lamina (at top). As basilar membrane is displaced upward, reticular lamina shifts right. B: Tectorial membrane (heavy black line) is rigidly anchored only at its periphery; therefore, as basilar membrane moves up, less rigid structures above it are pulled by tectorial membrane toward periphery. C: in longitudinal dimension, tectorial membrane is anchored throughout its length; thus a displacement of basilar membrane‐tectorial membrane complex causes a rotation of content between the two but no shearing force.
Figure 12. Figure 12.

Volume compliance of cochlear partition (measured as volume of basilar membrane displacement over length of 1 mm when distention is caused by pressure exerted on one side of membrane by water column 1 cm high) as function of distance from oval window, ; , width of basilar membrane as function of distance from oval window, [, Data from von Békésy 6; , data from Fletcher 48.]
Figure 13. Figure 13.

Spatial patterns of traveling‐wave envelope at various driving frequencies. Dashed portions of curves are inexact because the more apical segments of cochlea were destroyed so that vibrations in base could be observed.

From von Békésy 6
Figure 14. Figure 14.

Simulated instantaneous wave forms of cochlear partition at driving frequency of 2,000 Hz. t, Time delay expressed as phase angle; interval between successive plots is one‐eighth cycle.

From Khanna et al. 91
Figure 15. Figure 15.

Frequency‐response patterns of 2 points, 1.5 mm apart, on squirrel monkey's basilar membrane (T2 and θd2 represent responses at point closer to stapes). Amplitude (top) and phase (bottom) of ratio of basilar membrane displacement to malleus displacement are plotted. Note that for amplitude plot the frequency scale is logarithmic, while for phase plot it is linear.

From Rhode 121
Figure 16. Figure 16.

Computational results showing the fit by Zweig's model of Rhode's basilar‐membrane frequency‐response patterns.

From Zweig et al. 178
Figure 17. Figure 17.

A: configuration for computational model of acoustic transmission loss, assuming that oval window, having surface area Ss, (denoted in C and D) and equivalent radius a, is flush with skull (represented by an infinite baffle). B: equivalent circuit showing transformer (with turns ratio N:1) needed to match input impedance of cochlea (ZC) to radiation impedance (ZR) seen by small piston; PC, pressure across cochlear input impedance. C: ossicular chain seen from interior of ear, showing different surface areas of drum (Sd) and stapes footplate (Ss), and arms of ossicular lever (lengths lm and li). Entire chain rotates around axis (). D: schematic of ossicular lever, hydraulic amplifier system; Ps, pressure at oval window; Vd, velocity of drum; Vs, velocity of stapes.
Figure 18. Figure 18.

Sound pressure level (SPL) as function of stimulus frequency required to produce constant DC intracellular recording (summating potential) from single inner hair cell of guinea pig. Parameter is recorded summating potential magnitude, in mV.

From Russell and Sellick 127
Figure 19. Figure 19.

Cochlear microphonic (CM) responses from turn 1 (T1) and turn 3 (T3) of guinea pig's cochlea, recorded with the differential electrode technique. Stimulus is triangular displacement pattern of stapes. Note that T3 response follows T1 potential by a time lag of approximately 0.9 ms, reflecting travel time in cochlea. Both T1 and T3 responses roughly reflect time derivative of stimulus.

From Dallos and Durrant 189
Figure 20. Figure 20.

Magnitude and phase of cochlear microphonic potential (CM) as function of frequency at constant stapes velocity (2,500 nm.s−1) from three locations: turn 1, 2, and 3 (T1, T2, and T3) in guinea pig's cochlea. Recording is made by using differential electrode technique.

Adapted from Dallos 19
Figure 21. Figure 21.

Comparison of basilar membrane displacement (○) and cochlear microphonic (CM) potential (•), both measured in guinea pig's cochlea at approximately the same location.

From Dallos et al. 25
Figure 22. Figure 22.

Cochlear microphonic (CM) potentials measured with the differential electrode technique from 3rd turn of guinea pig's cochlea. Left: magnitude of CM as a function of frequency at various sound pressure levels (SPL), shown in dB reference level 20 μN.m−2. Center: same data as at left, but normalized by shifting each plot by an appropriate multiple of 10 dB to compensate for input intensity changes. If CM were a linear function of sound level, all these plots would superimpose. Right: 3 input‐output functions at frequencies of 100, 900, and 1,400 Hz derived from data in panel at left.

Adapted from Dallos 19
Figure 23. Figure 23.

Various wave forms of summating potential recorded from guinea pig's cochlea. Recordings are made between indifferent tissue and an intracochlear electrode (location indicated by ST, scala tympani, and SV, scala vestibuli; subscript shows cochlear turn). Stimulus conditions for the 4 pairs of recordings are indicated above traces. Black bar at bottom shows duration of 40‐ms stimulus tone burst. A: polarity of responses from the 2 scalae are opposite, constituting at these stimulus conditions the positive summating potential. C: the contrasting negative summating potential, is shown, which is produced at same cochlear location as positive summating potential, but by different stimulus parameters. B and D emphasize that polarity of summating potential can be the same in both scalae if stimuli are chosen appropriately. Note further that the 2 responses in B are virtually identical, suggesting a zero potential difference across cochlear partition and a response conducted from a romote site of generation. This response pattern is identified as a postsynaptic potential.

From Dallos et al. 26. Copyright 1970 by the American Association for the Advancement of Science
Figure 24. Figure 24.

Recordings of difference (DIF) and average (AVE) summating potential (SP) components from guinea pig's cochlea at 3 locations (T1, turn 1; T2, turn 2; T3, turn 3) as function of frequency. All plots are obtained at constant (10 Å) stapes displacement. Note transition from positive to negative DIF SP and from negative to positive to negative AVE SP as frequency increases.

From Dallos 188, by permission of S. Karger AG, Basel
Figure 25. Figure 25.

25. Comparison of frequency dependence of cochlear micro‐phonic (CM) and the difference (DIP) summating potential (SP) component. Note how much more sharply the dominant SP component (negative DIP) is tuned. Both plots are obtained from turn 1 of guinea pig's cochlea at constant (50 dB) sound pressure level (SPL).

Adapted from Dallos 19
Figure 26. Figure 26.

Schematic diagram showing changes in diameters of afferent nerve fibers innervating inner (IHC) and outer (OHC) hair cells. Fibers acquire their myelin sheath (shaded portion) central to habenula perforata. Dendrites connecting to outer hair cells average 0.7 mm in length. Probable locations of spike initiation are denoted by large open arrows. For inner hair‐cell fibers, all‐or‐none discharges are most likely excited at level of habenula, whereas corresponding location for outer hair‐cell fibers is more uncertain. It may be at habenula but could be at some more peripheral location (broken arrows).

Adapted from Spoendlin 145


Figure 1.

Block diagram of auditory system. Center, anatomical subdivisions, arrows indicating flow of information. Top, functions of the various segments. Bottom, the dominant mode of operation.

From Dallos 18


Figure 2.

Reproduction of Brödel's classic drawing of cross section of human ear.

From Brödel 13


Figure 3.

Schematic diagram showing basic morphological components of common hair cell. At apex of cell body several stereocilia emerge from a cuticular plate, and a single kinocilium is anchored to the intracellular basal body. Presynaptic specializations such as the synaptic bar (dark circle) and vesicles (small open circles) are seen adjacent to afferent nerve terminal, while efferent endings themselves are full of vesicles (small circles).

From Flock 50


Figure 4.

Cross section of mammalian cochlear duct with insets showing detailed structure of outer (left) and inner (right) hair cells. All important structures within scala media are identified; scala media space itself contains endolymph that is probably free to flow into subtectorial space and inner sulcus through noncontinuous marginal net. Intercellular spaces within organ of Corti are filled with cortilymph, which is probably not markedly different from perilymph. The morphological specializations depicted within the 2 hair cell types are described in detail in text.



Figure 5.

Scanning electron microscope picture of fracture in organ of Corti. Comparison with Fig. 4 identifies the various structures, although shape of organ is different because this figure depicts a more apical location where volume of organ of Corti is greater and outer hair cells and Hensen's cells are longer than at the more basal location depicted in Fig. 4.

From Bredberg et al. 187


Figure 6.

Scanning electron microscope picture from vantage point of tectorial membrane. Cilia from the single row of inner hair cells are seen on top, the top of pillar cells below them, and W‐shaped groups of cilia on top of 3 rows of outer hair cells below pillar cells. On bottom foreground, phalangeal processes of Deiters' cells are seen, and behind them the cell bodies of outer hair cells are discernible.

[From Bredberg et al. 187


Figure 7.

Schematic diagram showing hair cell and associated nerve endings as well as structural (center) and functional (right) block diagrams of system. Insets (left) demonstrate wave forms of the various quantities that are indicated in functional block diagram. It is assumed here that a brief sinusoidal burst stimulates the hair cell with a sinusoidal mechanical deformation (top). This results in 2 receptor potential components, an AC (cochlear microphonic) and a DC (summating potential) response. A wave form of postsynaptic potential, presumably arising in dendritic region of auditory nerve, and a spike train traveling in myelinated axon are also shown.

Adapted from Dallos 18


Figure 8.

Various conceivable modes of hair bundle deflection (stipples). A: bundle at rest; B: highly compliant bundle would bend; C: very stiff bundle could be displaced laterally in its entirety; D: stiff bundle with strongly adhering individual hairs could rotate about its insertion into cuticular plate; E: bundle of stiff individual hairs only weakly coupled to one another could fan out, with hairs rotating around their point of insertion. In all cases it is assumed that force is exerted at tip of tallest cilium.

From Flock et al. 52


Figure 9.

Cross section of cochlea with a plausible, but simplified, electric circuit superimposed upon it. Batteries represent voltage sources located in stria vascularis and in hair cells. Resistances are those of membranes separating various cochlear compartments; variable resistance, located within schematized receptor cell, is assumed to be controlled by mechanical deformation of cell. Change in this resistance produces a modulation of resting current through it and thus a modulation of potentials that can be measured at the various nodes of circuit.

From Davis and Silverman 190. In: Hearing and Deafness (3rd ed.), edited by Hallowell Davis and S. Richard Silverman. Copyright 1947 © 1960, 1970 by Holt, Rinehart and Winston, Inc. Reprinted by permission of Holt, Rinehart and Winston


Figure 10.

Pattern of excitation and inhibition of output of hair cell as function of direction of ciliary displacement. If θ is angle between direction of excitation and axis of morphological polarization through kinocilium, then the approximate dependence of output upon this angle is cos θ. This is figure‐eight pattern shown on top.

From Flock 49


Figure 11.

Schematic diagrams showing aspects of mechanics of basilar membrane organ of Corti complex. A: framework of organ of Corti is provided by basilar membrane (at bottom) and rigid pillar cells and reticular lamina (at top). As basilar membrane is displaced upward, reticular lamina shifts right. B: Tectorial membrane (heavy black line) is rigidly anchored only at its periphery; therefore, as basilar membrane moves up, less rigid structures above it are pulled by tectorial membrane toward periphery. C: in longitudinal dimension, tectorial membrane is anchored throughout its length; thus a displacement of basilar membrane‐tectorial membrane complex causes a rotation of content between the two but no shearing force.



Figure 12.

Volume compliance of cochlear partition (measured as volume of basilar membrane displacement over length of 1 mm when distention is caused by pressure exerted on one side of membrane by water column 1 cm high) as function of distance from oval window, ; , width of basilar membrane as function of distance from oval window, [, Data from von Békésy 6; , data from Fletcher 48.]



Figure 13.

Spatial patterns of traveling‐wave envelope at various driving frequencies. Dashed portions of curves are inexact because the more apical segments of cochlea were destroyed so that vibrations in base could be observed.

From von Békésy 6


Figure 14.

Simulated instantaneous wave forms of cochlear partition at driving frequency of 2,000 Hz. t, Time delay expressed as phase angle; interval between successive plots is one‐eighth cycle.

From Khanna et al. 91


Figure 15.

Frequency‐response patterns of 2 points, 1.5 mm apart, on squirrel monkey's basilar membrane (T2 and θd2 represent responses at point closer to stapes). Amplitude (top) and phase (bottom) of ratio of basilar membrane displacement to malleus displacement are plotted. Note that for amplitude plot the frequency scale is logarithmic, while for phase plot it is linear.

From Rhode 121


Figure 16.

Computational results showing the fit by Zweig's model of Rhode's basilar‐membrane frequency‐response patterns.

From Zweig et al. 178


Figure 17.

A: configuration for computational model of acoustic transmission loss, assuming that oval window, having surface area Ss, (denoted in C and D) and equivalent radius a, is flush with skull (represented by an infinite baffle). B: equivalent circuit showing transformer (with turns ratio N:1) needed to match input impedance of cochlea (ZC) to radiation impedance (ZR) seen by small piston; PC, pressure across cochlear input impedance. C: ossicular chain seen from interior of ear, showing different surface areas of drum (Sd) and stapes footplate (Ss), and arms of ossicular lever (lengths lm and li). Entire chain rotates around axis (). D: schematic of ossicular lever, hydraulic amplifier system; Ps, pressure at oval window; Vd, velocity of drum; Vs, velocity of stapes.



Figure 18.

Sound pressure level (SPL) as function of stimulus frequency required to produce constant DC intracellular recording (summating potential) from single inner hair cell of guinea pig. Parameter is recorded summating potential magnitude, in mV.

From Russell and Sellick 127


Figure 19.

Cochlear microphonic (CM) responses from turn 1 (T1) and turn 3 (T3) of guinea pig's cochlea, recorded with the differential electrode technique. Stimulus is triangular displacement pattern of stapes. Note that T3 response follows T1 potential by a time lag of approximately 0.9 ms, reflecting travel time in cochlea. Both T1 and T3 responses roughly reflect time derivative of stimulus.

From Dallos and Durrant 189


Figure 20.

Magnitude and phase of cochlear microphonic potential (CM) as function of frequency at constant stapes velocity (2,500 nm.s−1) from three locations: turn 1, 2, and 3 (T1, T2, and T3) in guinea pig's cochlea. Recording is made by using differential electrode technique.

Adapted from Dallos 19


Figure 21.

Comparison of basilar membrane displacement (○) and cochlear microphonic (CM) potential (•), both measured in guinea pig's cochlea at approximately the same location.

From Dallos et al. 25


Figure 22.

Cochlear microphonic (CM) potentials measured with the differential electrode technique from 3rd turn of guinea pig's cochlea. Left: magnitude of CM as a function of frequency at various sound pressure levels (SPL), shown in dB reference level 20 μN.m−2. Center: same data as at left, but normalized by shifting each plot by an appropriate multiple of 10 dB to compensate for input intensity changes. If CM were a linear function of sound level, all these plots would superimpose. Right: 3 input‐output functions at frequencies of 100, 900, and 1,400 Hz derived from data in panel at left.

Adapted from Dallos 19


Figure 23.

Various wave forms of summating potential recorded from guinea pig's cochlea. Recordings are made between indifferent tissue and an intracochlear electrode (location indicated by ST, scala tympani, and SV, scala vestibuli; subscript shows cochlear turn). Stimulus conditions for the 4 pairs of recordings are indicated above traces. Black bar at bottom shows duration of 40‐ms stimulus tone burst. A: polarity of responses from the 2 scalae are opposite, constituting at these stimulus conditions the positive summating potential. C: the contrasting negative summating potential, is shown, which is produced at same cochlear location as positive summating potential, but by different stimulus parameters. B and D emphasize that polarity of summating potential can be the same in both scalae if stimuli are chosen appropriately. Note further that the 2 responses in B are virtually identical, suggesting a zero potential difference across cochlear partition and a response conducted from a romote site of generation. This response pattern is identified as a postsynaptic potential.

From Dallos et al. 26. Copyright 1970 by the American Association for the Advancement of Science


Figure 24.

Recordings of difference (DIF) and average (AVE) summating potential (SP) components from guinea pig's cochlea at 3 locations (T1, turn 1; T2, turn 2; T3, turn 3) as function of frequency. All plots are obtained at constant (10 Å) stapes displacement. Note transition from positive to negative DIF SP and from negative to positive to negative AVE SP as frequency increases.

From Dallos 188, by permission of S. Karger AG, Basel


Figure 25.

25. Comparison of frequency dependence of cochlear micro‐phonic (CM) and the difference (DIP) summating potential (SP) component. Note how much more sharply the dominant SP component (negative DIP) is tuned. Both plots are obtained from turn 1 of guinea pig's cochlea at constant (50 dB) sound pressure level (SPL).

Adapted from Dallos 19


Figure 26.

Schematic diagram showing changes in diameters of afferent nerve fibers innervating inner (IHC) and outer (OHC) hair cells. Fibers acquire their myelin sheath (shaded portion) central to habenula perforata. Dendrites connecting to outer hair cells average 0.7 mm in length. Probable locations of spike initiation are denoted by large open arrows. For inner hair‐cell fibers, all‐or‐none discharges are most likely excited at level of habenula, whereas corresponding location for outer hair‐cell fibers is more uncertain. It may be at habenula but could be at some more peripheral location (broken arrows).

Adapted from Spoendlin 145
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Article Title: Peripheral Mechanisms of Hearing
 

How to Cite

Peter Dallos. Peripheral Mechanisms of Hearing. Compr Physiol 2011, Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes: 595-637. First published in print 1984. doi: 10.1002/cphy.cp010314