Power Dissipation in the Subtectorial Space of the Mammalian Cochlea Is Modulated by Inner Hair Cell Stereocilia  Srdjan Prodanovic, Sheryl Gracewski,

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Power Dissipation in the Subtectorial Space of the Mammalian Cochlea Is Modulated by Inner Hair Cell Stereocilia  Srdjan Prodanovic, Sheryl Gracewski, Jong-Hoon Nam  Biophysical Journal  Volume 108, Issue 3, Pages 479-488 (February 2015) DOI: 10.1016/j.bpj.2014.12.027 Copyright © 2015 Biophysical Society Terms and Conditions

Figure 1 Fluid dynamics in the STS. (A) The organ of Corti. The red rectangle indicates the modeled part of STS. (B) Two different modes of STS motion resulting in squeezing and shear flow. (C) Model of the STS. The model consists of a rectangular fluid domain with boundaries representing the second-row outer hair cell stereocilia bundle, tectorial membrane, and reticular lamina, and a rigid rod hinged at the root representing the IHC stereocilia bundle. The torsional spring at the hinge represents the flexural rigidity of the IHC stereocilia. Hensen’s stripe is not modeled (see Discussion). The displacements are exaggerated for illustrative purposes. To see this figure in color, go online. Biophysical Journal 2015 108, 479-488DOI: (10.1016/j.bpj.2014.12.027) Copyright © 2015 Biophysical Society Terms and Conditions

Figure 2 Power dissipation in the STS and CFSTS. (A) Tall (apical) model. (B) Short (basal) model. Top panels: Normalized power dissipation. Bottom panels: Phase of the hair bundle with respect to the tectorial membrane. Labels on the curves indicate the size of the gap between the IHC stereocilia bundle tip and the tectorial membrane (in μm). Stereocilia bundle stiffness is 10 and 100 mN/m for the tall and short models, respectively. The broken horizontal lines represent power obtained from the same model without the IHC stereocilia bundle (P0). The broken vertical lines indicate CFSTS, which correspond to half power dissipation or 45 degrees of phase difference. Biophysical Journal 2015 108, 479-488DOI: (10.1016/j.bpj.2014.12.027) Copyright © 2015 Biophysical Society Terms and Conditions

Figure 3 Effect of stereocilia bundle stiffness and gap size on STS power dissipation (Tall model). (A) Normalized power dissipation as function of frequency for three different values of the IHC stereocilia bundle stiffness; the gap size is 0.25 μm. Solid square symbols indicate CFSTS. Contour plots of normalized power dissipation. (B) Effect of the gap size and the stimulating frequency; stiffness is 10 mN/m. (C) Effect of the stereocilia bundle stiffness and the stimulating frequency; the gap size is 0.25 μm. (D) Effect of the stereocilia bundle stiffness and the gap size; the stimulating frequency is 1 kHz. The circles (B–D) indicate the standard parameter values of the apical model (see text). The base model result is shown in Fig. S4. Biophysical Journal 2015 108, 479-488DOI: (10.1016/j.bpj.2014.12.027) Copyright © 2015 Biophysical Society Terms and Conditions

Figure 4 Effect of different stimulation modes (short model). Snapshots of pressure (hotter color indicates higher pressure) and velocity (arrows) fields from (A) the first squeezing mode, (B) the second squeezing mode, and (C) the shear mode. (D) Normalized power dissipation from all three modes. The squeezing modes’ power dissipation is normalized by the dissipation from the same model, but without the IHC stereocilia bundle. (E) Comparison of the absolute dissipated power from the three modes. Boundary velocities were chosen so the volumetric flux through the IHC stereocilia bundle plane is constant. To see this figure in color, go online. Biophysical Journal 2015 108, 479-488DOI: (10.1016/j.bpj.2014.12.027) Copyright © 2015 Biophysical Society Terms and Conditions

Figure 5 Active IHC stereocilia modulate STS power dissipation. (A) Normalized IHC transduction current (Δp0) of the tall model. (B) Normalized STS power dissipation of the tall model. (C) Normalized IHC transduction current of the short model. (D) Normalized STS power dissipation of the short model. Gap size and bundle stiffness are 0.5 and 0.1 μm and 10 and 100 mN/m, for the tall and the short models, respectively. Results in (A) and (B) are without fluid-interaction. Four cases were simulated: three different tuning levels of the active IHC and the passive IHC. Biophysical Journal 2015 108, 479-488DOI: (10.1016/j.bpj.2014.12.027) Copyright © 2015 Biophysical Society Terms and Conditions

Figure 6 Power dissipation is determined by phase relation. (A) Highly tuned IHC stereocilia bundle (Q3dB = 16); stimulating frequency is 0.55 kHz. (B) Untuned transduction IHC stereocilia bundle; stimulating frequency is 0.55 kHz. (C) Highly tuned IHC stereocilia bundle (Q3dB = 16); stimulating frequency is 0.1 kHz. (D) Moderately tuned (Q3dB = 1.6) IHC stereocilia bundle, stimulating frequency is 0.9 kHz. Top row in all panels: force exerted by the stereocilia bundle on the surrounding fluid. Middle row in all panels: velocity of the stereocilia tip. Blue color indicates power transmitted to the fluid from the stereocilia bundle; red color indicates dissipated power (shaded area above and below the horizontal line, respectively). To see this figure in color, go online. Biophysical Journal 2015 108, 479-488DOI: (10.1016/j.bpj.2014.12.027) Copyright © 2015 Biophysical Society Terms and Conditions