Characterization of rigid open-cell foams using direct ultrasonic simulation.

An ultrasonic simulation technique based on the direct fluid model is proposed as an alternative to the analogous experimental technique to determine the tortuosity and characteristic lengths for high pore-density foams. It is beneficial as it reduces cost and almost eliminates the signal-to-noise issues encountered in the experiment. The proposed method is demonstrated for periodic microlattices with three different unit-cell configurations, 75%-90% porosity, and a pore size of about 200 microns. The technique is also applicable to high-resolution computed tomography (CT) scans of open-cell foams with a priori unknown microporous structure. An acoustic simulation software, ACTRAN® (Hexagon AB, Stockholm, Sweden), is used to model and perform analysis of the ultrasonic pulse propagation through the foam. Based on through-transmission by foam saturated with two different mediums, the tortuosity, and characteristic lengths are estimated from the high-frequency asymptotic behavior of the square of the propagation index (Nr2) versus the inverse square root of frequency (1/f). The predicted parameters are validated by comparing them with those determined by solving the electric conduction boundary value problem for the same configuration. Further, detailed parametric sensitivity analysis reveals the sensitivity of the Johnson-Champoux-Allard parameters to errors in Nr2 and so the effect of these errors on the acoustic absorption behavior of the rigid porous sample.

Microstructure-based modeling to characterize low pore density open-cell foams and its experimental validation.

Microlattices with large pore sizes are involved in many multifunctional applications, so it is essential to understand their acoustic properties. However, for these low pore density microlattice foams, the classical homogenization or "equivalent fluid" methods fail abruptly. This paper proposes and discusses a microstructure-based direct fluid model (DFM) that would help to predict the acoustic performance of low pore density periodic open-cell foams with spherical pores. The DFM is simulated directly, including the microscale geometric features inherent in the unit cell. A comparative study is performed for designed three-dimensional (3D) body-centered-cubic (BCC) porous foams having pores per inch (PPI) ranging from 1 to 12 over the frequency range of 500-4100 Hz with equivalent fluid models and experiments. The study shows the extent of deviation in homogenization-based methods from the experiment for PPI < 5. On the other hand, the acoustic performance parameters predicted with the DFM agree well with experiments on 3D-printed samples fabricated by additive manufacturing of varying PPI starting from 1. This study shows that the DFM is a valid method to predict the acoustics of low PPI microlattices. Furthermore, the gradual transition from dissipative to the reactive regime with a decrease in PPI is also brought out.

Enhanced absorption in structural acoustic silencers using piezo-shunting.

Structural acoustic silencers (sas) consist of a thin flexible plate that is introduced into a wall of an otherwise rigid-walled duct in which acoustic noise is propagating. Transmission loss (TL) in sas arises from reflection due to impedance mismatch, as well as partial absorption due to material damping in the plate. Reflection-based TL is less preferable than absorption-based TL, especially where back-pressure is undesirable. In this study, the TL arising out of absorption is increased by increasing the effective plate damping using the piezo-shunting method. Experiments are conducted on a custom-made sas with copper sheets of two different thicknesses. Piezo-shunting to increase the damping of the copper plate is done by connecting an electrical resistor to the external circuit of a thin piezoelectric bimorph attached to the copper plate. The effect of piezo-shunting on the sas TL is analyzed using a model. Piezo-shunt increases the fraction of power absorbed and decreases the reflected power fraction. The dependency of these fractions on the value of the electrical resistor and the impedance of the plate are discussed. This study demonstrates that piezo-shunting can be used to improve absorption-based TL in sas.

Minimal basilar membrane motion in low-frequency hearing.

Low-frequency hearing is critically important for speech and music perception, but no mechanical measurements have previously been available from inner ears with intact low-frequency parts. These regions of the cochlea may function in ways different from the extensively studied high-frequency regions, where the sensory outer hair cells produce force that greatly increases the sound-evoked vibrations of the basilar membrane. We used laser interferometry in vitro and optical coherence tomography in vivo to study the low-frequency part of the guinea pig cochlea, and found that sound stimulation caused motion of a minimal portion of the basilar membrane. Outside the region of peak movement, an exponential decline in motion amplitude occurred across the basilar membrane. The moving region had different dependence on stimulus frequency than the vibrations measured near the mechanosensitive stereocilia. This behavior differs substantially from the behavior found in the extensively studied high-frequency regions of the cochlea.

Optogenetic Control of Mouse Outer Hair Cells.

Normal hearing in mammals depends on sound amplification by outer hair cells (OHCs) presumably by their somatic motility and force production. However, the role of OHC force production in cochlear amplification and frequency tuning are not yet fully understood. Currently, available OHC manipulation techniques for physiological or clinical studies are limited by their invasive nature, lack of precision, and poor temporal-spatial resolution. To overcome these limitations, we explored an optogenetic approach based on channelrhodopsin 2 (ChR-2), a direct light-activated nonselective cation channel originally discovered in Chlamydomonas reinhardtii. Three approaches were compared: 1) adeno-associated virus-mediated in utero transfer of the ChR-2 gene into the developing murine otocyst, 2) expression of ChR-2(H134R) in an auditory cell line (HEI-OC1), and 3) expression of ChR-2 in the OHCs of a mouse line carrying a ChR-2 conditional allele. Whole cell recording showed that blue light (470 nm) elicited the typical nonselective cation current of ChR-2 with reversal potential around zero in both mouse OHCs and HEI-OC1 cells and generated depolarization in both cell types. In addition, pulsed light stimulation (10 Hz) elicited a 1:1 repetitive depolarization and ChR-2 currents in mouse OHCs and HEI-OC1 cells, respectively. The time constant of depolarization in OHCs, 1.45 ms, is 10 times faster than HEI-OC1 cells, which allowed light stimulation up to rates of 10/s to elicit corresponding membrane potential changes. Our study demonstrates that ChR-2 can successfully be expressed in mouse OHCs and HEI-OC1 cells and that these present a typical light-sensitive current and depolarization. However, the amount of ChR-2 current induced in our in vivo experiments was insufficient to result in measurable cochlear effects.

Filtering of acoustic signals within the hearing organ.

The detection of sound by the mammalian hearing organ involves a complex mechanical interplay among different cell types. The inner hair cells, which are the primary sensory receptors, are stimulated by the structural vibrations of the entire organ of Corti. The outer hair cells are thought to modulate these sound-evoked vibrations to enhance hearing sensitivity and frequency resolution, but it remains unclear whether other structures also contribute to frequency tuning. In the current study, sound-evoked vibrations were measured at the stereociliary side of inner and outer hair cells and their surrounding supporting cells, using optical coherence tomography interferometry in living anesthetized guinea pigs. Our measurements demonstrate the presence of multiple vibration modes as well as significant differences in frequency tuning and response phase among different cell types. In particular, the frequency tuning at the inner hair cells differs from other cell types, causing the locus of maximum inner hair cell activation to be shifted toward the apex of the cochlea compared with the outer hair cells. These observations show that additional processing and filtering of acoustic signals occur within the organ of Corti before inner hair cell excitation, representing a departure from established theories.

Non-uniform distribution of outer hair cell transmembrane potential induced by extracellular electric field.

Intracochlear electric fields arising out of sound-induced receptor currents, silent currents, or electrical current injected into the cochlea induce transmembrane potential along the outer hair cell (OHC) but its distribution along the cells is unknown. In this study, we investigated the distribution of OHC transmembrane potential induced along the cell perimeter and its sensitivity to the direction of the extracellular electric field (EEF) on isolated OHCs at a low frequency using the fast voltage-sensitive dye ANNINE-6plus. We calibrated the potentiometric sensitivity of the dye by applying known voltage steps to cells by simultaneous whole-cell voltage clamp. The OHC transmembrane potential induced by the EEF is shown to be highly nonuniform along the cell perimeter and strongly dependent on the direction of the electrical field. Unlike in many other cells, the EEF induces a field-direction-dependent intracellular potential in the cylindrical OHC. We predict that without this induced intracellular potential, EEF would not generate somatic electromotility in OHCs. In conjunction with the known heterogeneity of OHC membrane microdomains, voltage-gated ion channels, charge, and capacitance, the EEF-induced nonuniform transmembrane potential measured in this study suggests that the EEF would impact the cochlear amplification and electropermeability of molecules across the cell.

Outer hair cell somatic electromotility in vivo and power transfer to the organ of Corti.

The active amplification of sound-induced vibrations in the cochlea, known to be crucial for auditory sensitivity and frequency selectivity, is not well understood. The outer hair cell (OHC) somatic electromotility is a potential mechanism for such amplification. Its effectiveness in vivo is putatively limited by the electrical low-pass filtering of the cell's transmembrane potential. However, the transmembrane potential is an incomplete metric. We propose and estimate two metrics to evaluate the effectiveness of OHC electromotility in vivo. One metric is the OHC electromechanical ratio defined as the amplitude of the ratio of OHC displacement to the change in its transmembrane potential. The in vivo electromechanical ratio is derived from the recently measured in vivo displacements of the reticular lamina and the basilar membrane at the 19 kHz characteristic place in guinea pigs and using a model. The ratio, after accounting for the differences in OHC vibration in situ due to the impedances from the adjacent structures, is in agreement with the literature values of the in vitro electromechanical ratio measured by others. The second and more insightful metric is the OHC somatic power. Our analysis demonstrates that the organ of Corti is nearly optimized to receive maximum somatic power in vivo and that the estimated somatic power could account for the active amplification.

Persistence of past stimulations: storing sounds within the inner ear.

Tones cause vibrations within the hearing organ. Conventionally, these vibrations are thought to reflect the input and therefore end with the stimulus. However, previous recordings of otoacoustic emissions and cochlear microphonic potentials suggest that the organ of Corti does continue to move after the end of a tone. These after-vibrations are characterized here through recordings of basilar membrane motion and hair cell extracellular receptor potentials in living anesthetized guinea pigs. We show that after-vibrations depend on the level and frequency of the stimulus, as well as on the sensitivity of the ear. Even a minor loss of hearing sensitivity caused a sharp reduction in after-vibration amplitude and duration. Mathematical models suggest that after-vibrations are driven by energy added into organ of Corti motion after the end of an acoustic stimulus. The possible importance of after-vibrations for psychophysical phenomena such as forward masking and gap detection are discussed.

The biophysical origin of traveling-wave dispersion in the cochlea.

Sound processing begins at the peripheral auditory system, where it undergoes a highly complex transformation and spatial separation of the frequency components inside the cochlea. This sensory signal processing constitutes a neurophysiological basis for psychoacoustics. Wave propagation in the cochlea, as shown by measurements of basilar membrane velocity and auditory nerve responses to sound, has demonstrated significant frequency modulation (dispersion), in addition to tonotopic gain and active amplification. The physiological and physical basis for this dispersion remains elusive. In this article, a simple analytical model is presented, along with experimental validation using physiological measurements from guinea pigs, to identify the origin of traveling-wave dispersion in the cochlea. We show that dispersion throughout the cochlea is fundamentally due to the coupled fluid-structure interaction between the basilar membrane and the scala fluids. It is further influenced by the variation in physical and geometrical properties of the basilar membrane, the sensitivity or gain of the hearing organ, and the relative dominance of the compression mode at about one-third octave beyond the best frequency.

A mechanoelectrical mechanism for detection of sound envelopes in the hearing organ.

To understand speech, the slowly varying outline, or envelope, of the acoustic stimulus is used to distinguish words. A small amount of information about the envelope is sufficient for speech recognition, but the mechanism used by the auditory system to extract the envelope is not known. Several different theories have been proposed, including envelope detection by auditory nerve dendrites as well as various mechanisms involving the sensory hair cells. We used recordings from human and animal inner ears to show that the dominant mechanism for envelope detection is distortion introduced by mechanoelectrical transduction channels. This electrical distortion, which is not apparent in the sound-evoked vibrations of the basilar membrane, tracks the envelope, excites the auditory nerve, and transmits information about the shape of the envelope to the brain.

Minimally invasive surgical method to detect sound processing in the cochlear apex by optical coherence tomography.

Sound processing in the inner ear involves separation of the constituent frequencies along the length of the cochlea. Frequencies relevant to human speech (100 to 500 Hz) are processed in the apex region. Among mammals, the guinea pig cochlear apex processes similar frequencies and is thus relevant for the study of speech processing in the cochlea. However, the requirement for extensive surgery has challenged the optical accessibility of this area to investigate cochlear processing of signals without significant intrusion. A simple method is developed to provide optical access to the guinea pig cochlear apex in two directions with minimal surgery. Furthermore, all prior vibration measurements in the guinea pig apex involved opening an observation hole in the otic capsule, which has been questioned on the basis of the resulting changes to cochlear hydrodynamics. Here, this limitation is overcome by measuring the vibrations through the unopened otic capsule using phase-sensitive Fourier domain optical coherence tomography. The optically and surgically advanced method described here lays the foundation to perform minimally invasive investigation of speech-related signal processing in the cochlea.

Thin and open vessel windows for intra-vital fluorescence imaging of murine cochlear blood flow.

Normal microvessel structure and function in the cochlea is essential for maintaining the ionic and metabolic homeostasis required for hearing function. Abnormal cochlear microcirculation has long been considered an etiologic factor in hearing disorders. A better understanding of cochlear blood flow (CoBF) will enable more effective amelioration of hearing disorders that result from aberrant blood flow. However, establishing the direct relationship between CoBF and other cellular events in the lateral wall and response to physio-pathological stress remains a challenge due to the lack of feasible interrogation methods and difficulty in accessing the inner ear. Here we report on new methods for studying the CoBF in a mouse model using a thin or open vessel-window in combination with fluorescence intra-vital microscopy (IVM). An open vessel-window enables investigation of vascular cell biology and blood flow permeability, including pericyte (PC) contractility, bone marrow cell migration, and endothelial barrier leakage, in wild type and fluorescent protein-labeled transgenic mouse models with high spatial and temporal resolution. Alternatively, the thin vessel-window method minimizes disruption of the homeostatic balance in the lateral wall and enables study CoBF under relatively intact physiological conditions. A thin vessel-window method can also be used for time-based studies of physiological and pathological processes. Although the small size of the mouse cochlea makes surgery difficult, the methods are sufficiently developed for studying the structural and functional changes in CoBF under normal and pathological conditions.

Half-octave shift in mammalian hearing is an epiphenomenon of the cochlear amplifier.

The cochlear amplifier is a hypothesized positive feedback process responsible for our exquisite hearing sensitivity. Experimental evidence for or against the positive feedback hypothesis is still lacking. Here we apply linear control theory to determine the open-loop gain and the closed-loop sensitivity of the cochlear amplifier from available measurements of basilar membrane vibration in sensitive mammalian cochleae. We show that the frequency of peak closed-loop sensitivity is independent of the stimulus level and close to the characteristic frequency. This implies that the half-octave shift in mammalian hearing is an epiphenomenon of the cochlear amplifier. The open-loop gain is consistent with positive feedback and suggests that the high-frequency cut-off of the outer hair cell transmembrane potential in vivo may be necessary for cochlear amplification.

In vivo outer hair cell length changes expose the active process in the cochlea.

BACKGROUND: Mammalian hearing is refined by amplification of the sound-evoked vibration of the cochlear partition. This amplification is at least partly due to forces produced by protein motors residing in the cylindrical body of the outer hair cell. To transmit power to the cochlear partition, it is required that the outer hair cells dynamically change their length, in addition to generating force. These length changes, which have not previously been measured in vivo, must be correctly timed with the acoustic stimulus to produce amplification. METHODOLOGY/PRINCIPAL FINDINGS: Using in vivo optical coherence tomography, we demonstrate that outer hair cells in living guinea pigs have length changes with unexpected timing and magnitudes that depend on the stimulus level in the sensitive cochlea. CONCLUSIONS/SIGNIFICANCE: The level-dependent length change is a necessary condition for directly validating that power is expended by the active process presumed to underlie normal hearing.

A mechano-electro-acoustical model for the cochlea: response to acoustic stimuli.

A linear, physiologically based, three-dimensional finite element model of the cochlea is developed. The model integrates the electrical, acoustic, and mechanical elements of the cochlea. In particular, the model includes interactions between structures in the organ of Corti (OoC), piezoelectric relations for outer hair cell (OHC) motility, hair bundle (HB) conductance that changes with HB deflection, current flow in the cross section and along the different scalae, and the feed-forward effect. The parameters in the model are based on guinea-pig data as far as possible. The model is vetted using a variety of experimental data on basilar membrane motion and data on voltages and currents in the OoC. Model predictions compare well, qualitatively and quantitatively, with experimental data on basilar membrane frequency response, impulse response, frequency glides, and scala tympani voltage. The close match of the model predictions with experimental data demonstrates the validity of the model for simulating cochlear response to acoustic input and for testing hypotheses of cochlear function. Analysis of the model and its results indicates that OHC somatic motility is capable of powering active amplification in the cochlea. At the same time, the model supports a possible synergistic role for HB motility in cochlear amplification.

A theoretical study of structural acoustic silencers for hydraulic systems.

Theoretical studies show that the introduction of an in-line structural acoustic silencer into a hydraulic system can achieve broadband quieting (i.e., high transmission loss). Strategies for using structural acoustic filters for simultaneously reducing reflection and transmission by tailoring the material properties are studied. A structural acoustic silencer consists of a flexible layer inserted into nominally rigid hydraulic piping. Transmission loss is achieved by two mechanisms--reflection of energy due to an impedance mismatch, and coupling of the incoming acoustic fluctuations to structural vibrations thereby allowing for the extraction of energy through losses in the structure. Structural acoustic finite element simulations are used to determine the transmission loss and evaluate designs. Results based on the interaction of orthotropic and isotropic plates with variable geometry, operating in heavy fluids like water and oil, are presented.

Structural acoustic silencers--design and experiment.

The effectiveness of introducing flexible structural layers into air conveying ducts for controlling noise is investigated through theoretical and experimental means, focusing at low frequencies where conventional passive silencing technology is least effective. Previous theoretical work has shown that using flexible rather than rigid walls has the potential to achieve high transmission losses. The physical mechanisms responsible for structural acoustic silencing, including the relation between transmission loss peaks and structural resonance corresponding to different transverse structural modes, are presented. Sensitivity of the performance to acoustic and structural boundary conditions is discussed. To eliminate radiated noise from these walls (breakout noise), a rigid walled cavity is introduced under the flexible plate. The challenge is to find means to reject plane waves in the two-duct system. Designs that overcome these issues and achieve appreciable transmission loss are investigated. Results based on three-dimensional finite element simulations are compared with experimental results.