Re: mechanical cochlear model ("reinifrosch@xxxxxxxx" )


Subject: Re: mechanical cochlear model
From:    "reinifrosch@xxxxxxxx"  <reinifrosch@xxxxxxxx>
Date:    Sat, 6 Mar 2010 15:15:43 +0000
List-Archive:<http://lists.mcgill.ca/scripts/wa.exe?LIST=AUDITORY>

Dear Alain, In my opinion it would be better to start a beginners' course on cochlear mechanics with a treatment of surface waves on the ocean. Their propagation velocity is much smaller than the sound velocity in water. The water particles move on closed (elliptical) trajectories in vertical planes. The liquid motion is limited to a surface layer of thickness ~1/k, where k = 2pi/wavelength is the wave number. The wavelength of tsunamis on a 3 km deep ocean is about 100 km, so in their case there is liquid motion down to the ocean floor. In the case of the passive cochlear wave, the surface wave is mass-loaded (mass = BM and cells attached to it), and the wave is not gravity-driven, but spring-driven (springs = fibres of the BM), so that a surface wave with liquid on both sides of the "surface" (i.e., of the cochlear partition) is possible. Etc. -- see the already mentioned Chapters 4 and 5 of the book "The Cochlea", or also Chapter 2, "Cochlear Structure and Function", by the late Graeme Yates, of the book "Hearing", B.C.J. Moore (Ed., 1995), Academic Press, San Diego. Reinhart Frosch, Dr. phil. nat., r. PSI and ETH Zurich, Sommerhaldenstr. 5B, CH-5200 Brugg. Phone: 0041 56 441 77 72. Mobile: 0041 79 754 30 32. E-mail: reinifrosch@xxxxxxxx . ----Ursprüngliche Nachricht---- Von: Alain.de.Cheveigne@xxxxxxxx Datum: 06.03.2010 02:54 An: <AUDITORY@xxxxxxxx> Betreff: Re: mechanical cochlear model Strangely I didn't see Jan's message on the list. I agree with his comments. The controversy seems partly due to people using the term "travelling wave" with different meanings. The way I explain things to students is the following. If this account is fundamentally flawed (as opposed to inaccurate) I'd be glad to know. It goes like this: " This is a simple explanation of cochlear selectivity. We know that the cochlea is divided into two ducts separated by the basilar membrane, surrounded by hard bone, and filled with fluid. Both ducts are closed at their base by flexible windows, the oval window for the upper duct (scala vestibuli), and the round window for the lower duct (scala tympani). The ducts communicate at their apex via a hole, the helicotrema. Our explanation assumes four ingredients. First, the fluid within the cochlea is incompressible. Second, the bony walls that surround it are rigid. Third, the fluid has mass. Fourth, the basilar membrane is stiff at the base (where it is narrow) and compliant at the apex (where it is wide). If the pressure in the outside air is slowly increased (relative to middle ear) the displacement of the ear drum, transmitted via the ossicles to the oval window, causes that window to flex inwards. Fluid flows up scala vestibuli, through the helicotrema, and down scala tympani, causing the round window to flex outwards. Acceleration being small, the effect of fluid inertia is negligible, and the pressure at all points in the ducts on both sides of the membrane is equal. The membrane does not move. For faster vibrations, force is required to accelerate the column of fluid in the ducts, implying a pressure gradient along the duct and a pressure difference across the membrane. Deformation of the membrane may then be an easier option than displacement of the fluid. Whether deformation or displacement occurs depends on the balance between the inertia of the fluid to move on one hand, and the stiffness of the membrane to bend on the other. The terms of this balance depends on frequency of vibration and place along the cochlea. At the base of the cochlea stiffness is maximal, and so deformation requires a large trans-membrane pressure. This occurs only with the relatively large accelerations involved in vibration at high frequencies. Hence the base of the cochlea responds only to high frequencies. For lower frequencies the inertia/compliance balance occurs where the membrane is less stiff, ie apically. Thus the apex responds to lower frequencies. This explanation is a rough first-order explanation. A more complete explanation would involve detailed modeling of the mechanics of the basilar and tectorial membranes, of the fluid mechanics of the fluid in both ducts, and of the electrical and chemical phenomena that interact with cochlear mechanics at the outer hair cells. In particular, we know that the moving parts of the inner ear are immersed in fluid and therefore resonances should be very much damped (the "underwater piano" metaphor). To compensate for damping and produce the exquisitely fine tuning and high sensitivity of the cochlea requires a form of "negative damping" by which energy is injected into the system to compensate for dissipation. This is ensured by active processes involving the outer hair cells, that interact with cochlear mechanics under control from the central nervous system. " Is this account reasonably correct? For example if the balance of membrane compliance and fluid inertia did not play a role I'd consider it to be flawed. Alain


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