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THE EFFECT OF THE "THIRD WINDOW" ON THE RESPONSE OF A 2D MODEL OF THE COCHLEA Alice Halpin, Stephen Elliott, Guangjian Ni ISVR, University of Southampton, Southampton, SO17 3BJ email: [email protected] There are multiple mechanisms in which the healthy cochlea can be excited, including air con-duction, bone conduction and direct non-volumetric excitation of either of the windows. A 2D box model of the passive cochlea, solved using a finite difference method, has been used to investigate the basilar membrane response and fluid pressure in the cochlea due to several of these mechanisms. It is shown that all the investigated mechanisms of stimulating the cochlea produce a similar pattern of travelling wave along the basilar membrane, however, the magnitude of this response varied with excitation mechanism. It is also shown that the cochlea is still responsive to some of these mechanisms when it is diseased in such a way that the oval window becomes stiff. The effect of including a cochlear aqueduct and vestibular aqueduct, or ’third window’, in the model was investigated and shown to have a significant effect to the magnitude of the basi-lar membrane response when excited by some of these mechanisms when the oval window is assumed to be stiff. 1. Introduction It has been shown that the mammalian hearing organ, the cochlea, can be excited by a number of mechanisms in order to produced a hearing sensation, including the conventional route of air conduction hearing, bone conduction, BC, and excitation at the round window [1], [2]. The cochlea has a coiled complicated 3D geometry, with many delicate structures inside. It has been suggested that some of these structure which are fluid outlets from the cochlea, collectively know as the "third window", could have a significant effect on the basilar membrane, BM, response, and therefore hearing response, when the cochlea is excited by certain mechanisms [3]. A 2D finite difference box model of the cochlea has been used here to invesitgate a number of mechanisms of exciting the cochlea and the effect of two of these cochlea structures, the vestibular aqueduct, VA, and cochlear aqueduct, CA, on the BM response. The simple design of the model allows it to be easily adapted to simulate the difference cochlear excitation mechanisms as well as to compared the BM response within one framework. Air conduction hearing involves a pressure waves travelling down the ear canal, vibrating the middle ear ossicles and causing the stapes to vibrated the oval window. It is often assumed that the stapes has a piston-like motion, even though this motion has been shown to be complex at high frequencies [4]. Therefore, two motions of exiting the cochlea at the oval window have been examined here, a piston-like motion and a rocking motion of the stapes, which has previously been investigated ICSV22, Florence, Italy, 12-16 July 2015 1 The 22nd International Congress of Sound and Vibration using a finite element model [5]. Bone conduction, BC, hearing is where vibrations of the skull are transferred by various pathways to the cochlea. BC hearing have been separated into five pathways [1], the two inertial pathways are simulated here: the fluid inertia and the middle ear component. The middle ear inertia component is when the vibration of the skull, and hence the temporal bone, cause a relative motion between the cochlea and the stapes. While the fluid inertia component is due to the vibrations of the temporal bone imparting inertial forces on the the cochlear fluid. Conductive hearing loss can be due to diseases such as otoscelerosis, which cause bone to grow around the oval window area, immobilising the stapes. It is clear that any pathway that would excite the cochlear via the oval window would not be possible in patients with these conditions, however, other mechanisms of exciting the cochlea could still be possible. It is a well documented phenomena that the BC hearing thresholds of patients with otoscelerosis are typical of a healthy ear, apart from around 2 kHz where the threshold increases, called Carharts notch. It also has been shown that driving the round window with a transducer that only partially occludes the window, can excite the cochlea, and is one possible mechanism of exciting a cochlea with an immobile oval window [2]. Here, two mechanisms of exciting the cochlea, when a stiff oval window is defined, are investigated; the fluid inertia component of BC and local excitation of the round window, where only the middle section of the round window is driven. The 2D model was solved and the BM responses due to each of the excitation mechanisms calculated. The VA and CA are modelled as complaint elements in the cochlear walls. The effect of these aqueducts on the BM frequency response, due to the various cochlear excitation mechanisms, was investigated. 2. The Model The complex 3D coiled geometry of the cochlea is simplified to a 2D box model, where a passive BM separates a fluid filled box into two chambers, the scala vestibuli and scala tympani, terminating at the helicotrema which is located at the apex of the cochlea, such that the BM does not extend to the end cochlea wall. The oval window and vestibuli aqueduct is located on the upper cochlea wall and the round window and CA on the lower. The CA is located in very close proximity, 0.2 mm, to the round window, to reflex the real geometry [6] while the vestibuli aqueduct is located close, but slightly further away, 0.5 mm, from the oval window, as shown in Fig. 1. ,ǀ nj KǀĂůtŝŶĚŽǁ sĞƐƚŝďƵůĂƌ ƋƵĞĚƵĐƚ pd x,Ν mo vBM x,Ν Ϭ Ͳ,ƚ Ϭ > ZŽƵŶĚtŝŶĚŽǁ ŽĐŚůĞĂƌ ƋƵĞĚƵĐƚ dž cBM x kBM x vw x,Ν Figure 1: Left: diagram of the 2D box model of the cochlea, where the thick black line is the basilar membrane, the thick black dotted lines are the cochlear windows and the brown dotted line the aqueducts. Right: diagram of one single degree of freedom micromechanical model for this passive basilar membrane. It is assumed that the cochlear fluid is inviscid and compressible in both directions, and so the pressure at every location satisfies, (1) 2 ∂2 ∂2 p(x, z) + p(x, z) − ∂x2 ∂z 2 ω co 2 p(x, z) = 0, ICSV22, Florence, Italy, 12-16 July 2015 The 22nd International Congress of Sound and Vibration Table 1: List of parameters used in the cochlear model. Variable L Hv Ht N M kow cow mow Aw co ωB mo madd l Qo kBM (x) cBM (x) mm km cT Lr Parameter Length of cochlea Height of scala vestibuli Height of scala tympani Number of nodes in x-direction Number of nodes in z-direction Stiffness of the oval window Resistance of the oval window Mass of the oval window Area of cochlear windows Speed of sound in cochlear fluid Natural frequency at the base of the cochlea Physical mass of basilar membrane Inferred fluid mass on the basilar membrane Frequency distribution characteristic length Q factor of passive cochlea Stiffness variation along basilar membrane Damping variation along basilar membrane Effective inertial mass of middle ear Effective stiffness of the middle ear Effective resistance of the middle ear Lever ratio Value 35 mm 1.5 mm 1 mm 295 22 830 N m−1 0.1 N m−1 s 4.5 × 10−6 kg 3.2 × 10−6 m2 1500 ms−1 20 kHz 0.02 kgm−2 0.27 kgm−2 7 × 10−3 m 2.5 −2x 2 (mo + madd )e l ωBp Q−1 kBM (mo + madd ) o 5.5 × 10−5 kg 6300N m−1 0.12N m−1 s 1.4 where ω is the angular frequency and co the speed of sound in the cochlear fluid. The model was separated into 294 elements in the x-direction and 21 in the z-direction, in order to solve this equation, Eq. 1, where ∆x and ∆z are the sizes of the elements in the x and z directions respectively. The pressure at location, (n∆x, m∆z), which is written here as [n, m] for convenience, is solved using the finite difference approximation of Eq. 1, (2) p[n+1, m]−2p[n, m]+p[n−1, m] p[n, m+1]−2p[n, m]+p[n, m−1] + ∆x2 ∆z 2 2 ω − p[n, m] ≈ 0. co The boundary conditions in the x and z directions are imposed by relating the velocity at the boundaries, vbc , to the pressure gradient in the x and z directions, (3, 4) ∂p(x, z) = −iωρvbcz (z), ∂z ∂p(x, z) = −iωρvbcx (x). ∂x These equations are used to describe the velocity of the BM, cochlear windows, cochlear walls and aqueducts. All cochlear walls are assumed to be rigid, and for most cochlear excitation mechanisms the cochlea is stationary, hence the velocity perpendicular to cochlear wall is equal to zero. The velocity of the complainant structures in the cochlea are related to the admittance of the structure. The BM is modelled as a series of locally reacting single degree of freedom systems, as ICSV22, Florence, Italy, 12-16 July 2015 3 The 22nd International Congress of Sound and Vibration shown in Fig. 1, where the stiffness, kBM , and resistance, cBM , varies along its length, but the physical mass, mo , remains constant. The values and variation of the parameters are shown in Table 1. The admittance of the BM, at location x, which is equal to, YBM (x) = (5) iω , iωcBM (x) + kBM (x) − ω 2 mo which is related to the BM velocity, vBM , relative to the horizontal cochlear wall velocity, vw , by, vBM (x) = −YBM (x)pd (x) + TBM (x)vw (x). (6) pd (x) is the pressure difference across the BM at location x and, TBM (x), is the transmissibility of the wall velocity to the relative BM velocity, which is given by, TBM = (7) ω 2 mo . iωcBM (x) + kBM (x) − ω 2 mo The admittances of the cochlear windows and aqueducts are calculated as if looking out from the cochlea, and represented here a series of single degree of freedom systems, similar to the BM. The admittance of the oval window is dependant of the effect mass, stiffness and resistance of the middle ear looking out from the cochlea while the round window admittance is modelled to only have a stiffness, with a value of a tenth of the stiffness of the oval window, [7], (8, 9) Yow = iAw ω , (kow + iωcow − ω 2 mow ) Yrw = 10Aw iω . kow The parameters used to determine the admittance of the oval window were calculated from those measured by Puria [8], and can be found in Table 1. The velocities of the oval window, vow , and round window, vrw are related to the their respective admittances by, (10, 11) vow = vw + Yow p(x, Hv ), vrw = vw − Yrw p(x, Ht ). The CA and VA are modelled as fluid filled cylindrical tubes, where their admittances, YAi , are dependant on the dimensions of the tube. The values can are the inverse of the impedances defined by Stenfelt [3]. The velocities at the VA, vAV and CA vAC are defined as, (12, 13) 2.1 vAV = vw + YAv p(x, Hv ), vAC = vw − YAc p(x, Ht ). Methods of Exciting the Model Described above is the complete cochlear model with all cochlear structures outlined. The model is excited by applying a velocity at specific locations, either cochlear windows or walls, in order to simulate five different mechanisms of cochlear excitation. When a velocity is applied to a cochlear window, in order to excited the cochlea, the velocity is no longer proportional to the admittance of the window but is now a defined input condition of the model. A piston-like motion of the stapes is simulated here by applying uniform velocity at the oval window locations in the model. This is a volumetric excitation mechanism because it produces a net volume velocity through the oval window. A rocking-types motion of the stapes, pivoting about its 4 ICSV22, Florence, Italy, 12-16 July 2015 The 22nd International Congress of Sound and Vibration short axes has been examined here, which has also been considered by Edom et al. [5]. The rocking motion of the stapes is modelled here by applying a graded velocity to the oval window, such that there is a zero net volume velocity through the oval window, creating a non-volumetric excitation mechanism. The middle ear inertia BC component is modelled by applying a piston-like velocity at the oval window. The magnitude of this velocity is frequency dependant and was calculated from a lumped parameter model of the middle ear. The middle ear is represented by a damped lever connected by springs and dampers to a surrounding box, which represents the temporal bone. The velocity of the stapes, vs , due to the displacement of the surrounding box, vE , is calculated by, vs = (14) ω 2 mm vE , Lr km − ω 2 mm + iωcT where the values of these parameters can be found in Table 1. The fluid inertia component of BC is simulated by assuming that the cochlear model is harmonically vibrated in the z-direction. This is modelled by defining an equal upper and lower cochlear wall velocity. This mechanisms is examined with both a flexible and stiff oval window. When a stiff oval window is assumed, the stiffness, kow , is 1030 times greater than that of the flexible oval window. Local excitation at the round window was modelled by keeping the 0.6 mm at either edge of the window complaint but forcing the middle 1.1 mm with a proscribed velocity. This excitation method is modelled with an immobile oval window. The effect of the CA and VA on all of the mechanisms described above is investigated here by comparing the BM response with and without the aqueducts present. Equation 2, along with the constraints of the boundary conditions and input conditions for a specific mechanisms, was solved using a matrix equation that relates the vector containing the pressure at each node, p, to the vector defining the external excitation, q, Ap = q, (15) so that p = A−1 q, for the pressure distribution in the cochlea. The BM velocity, due to the excitation mechanism, was then determined using Eq. 6. In order to be able to compare the BM responses due to the different cochlea excitation mechanisms, the BM velocity was divided by the excitation velocity. This was either the maximum window input velocity or, for the BC components, the assumed excitation velocity of the surrounding bone. The frequency response of the BM due to the different excitation mechanisms was calculated by determining the peak BM velocity for a range of frequencies. 3. Results It was found that every mechanism of exciting the cochlea investigated produced a travelling wave along the BM, even when an immobile oval window was assumed. An instantaneous BM velocity was found for each mechanism, which for ease of comparison, was normalised to its peak value and its phase altered to so that the peaks all aligned, which in shown in Fig. 2. The shape of the travelling wave is very similar at locations towards the apex of the cochlea, for all mechanisms of excitation. However, when excited by the rocking motion of the stapes or the local excitation of the round window there is a small deviation. This is due to the relatively high near field pressure produced at the excitation window for these mechanisms. The BM response is shown at 1 kHz here, however, a similar shapes of response is produced at other frequencies but the location of the peak velocity is dependant on the excitation frequency. ICSV22, Florence, Italy, 12-16 July 2015 5 The 22nd International Congress of Sound and Vibration 1 Fluid Inertia Piston−like Stapes Motion Rocking Motion of the Stapes Local Excitation at the Round Window Fluid Inertia With a Stiff Oval Window Normalised Basilar Membrane Velocity 0.8 0.6 0.4 0.2 0 −0.2 −0.4 −0.6 −0.8 −1 0 5 10 15 20 25 Distance along Cochlea, mm 30 35 Figure 2: A comparison of the instantaneous basilar membrane velocity, normalised to the peak velocity, for multiple mechanisms of exciting the cochlea, all at 1 kHz 3.1 Basilar Membrane Response when Excited at the Oval Window or via the Inertia Components of Bone Conduction Fig. 3 shows the frequency response of the peak relative BM response due to excitation via the oval window by a piston-like motion and by a rocking motion, as well as the two inertia components of BC, with and without the inclusion of the aqueducts in the model. It was found that the presence of the aqueducts had little effect on the overall BM response for all the mechanisms at frequencies above 500 Hz. However, at frequencies less than 500 Hz, the aqueducts increased the BM response by a maximum of 15 dB when the cochlea was excited by the rocking of the stapes and 20 dB when excited by the fluid inertia component. 20 Middle Ear Aqueducts Middle Ear No Aqueducts Piston−like Stapes, No Aqueducts Piston−like Stapes, with Aqueducts Rocking Stapes, Aqueducts Rocking Stapes, No Aqueducts Fluid Inertia, No Aqueducts Fluid Inertia, Aqueducts Peak Relative Basilar Membrane Velocity, dB 10 0 −10 −20 −30 −40 −50 −60 −70 0 500 1000 1500 2000 2500 3000 Frequency, Hz 3500 4000 4500 5000 Figure 3: Comparison of the peak value along the cochlea of the relative BM velocity for multiple excitation mechanisms, with and without the inclusion of the aqueducts. 3.2 Basilar Membrane Response with an Immobile Oval Window Fig. 4a shows the BM frequency response, when an immobile oval window is assumed, due to the excitation by the fluid inertia component of BC and the local excitation of the round window, when both aqueducts are included, only the CA and when no aqueducts are modelled. The BM response was found to be very low, due to excitation via both of these mechanisms, when no aqueducts were modelled. For this condition, the only outlet in the cochlea is the round window and therefore no volumetric flow can be generated, only non-volumetric. It has already been shown that a non-volumetric excitation mechanism, the rocking motion of the stapes, produces a low BM response which is of similar magnitude to the BM response in Fig. 3, without the aqueducts, since 6 ICSV22, Florence, Italy, 12-16 July 2015 The 22nd International Congress of Sound and Vibration Total Excitation Volumetric Excitation Non−Volumetric Excitation 10 10 0 0 Peak Relative Basilar Membrane Velocity Peak Relative Basilar Membrane Velocity, dB Local Excitation at Round Window, Both Aqueducts Local Excitation at Round Window, No Aqueducts Local Excitation at Round Winodw, Only Cochlear Aqueduct Fluid Inertia, Both Aqueducts Fluid Inertia, Only Cochlear Aqueduct Fluid Inertia, No Aqueducts −10 −20 −30 −40 −50 −60 −70 −80 −10 −20 −30 −40 −50 −60 −70 0 500 1000 1500 2000 2500 3000 Frequency, Hz 3500 4000 4500 (a) 5000 −80 0 500 1000 1500 2000 2500 3000 Frequency, Hz 3500 4000 4500 5000 (b) Figure 4: (a) Comparison of the relative BM velocity due to the fluid inertia and local round window excitation mechanisms, with no aqueducts, only the CA and both aqueducts. (b) Comparison of the total excitation due to local excitation at the round window and the separate non-volumetric and volumetric components. these excitation are due to the near field pressure distribution [5], [2], [9]. The CA is on the same side of the cochlea as the round window and including it in the model does not significantly alter the magnitude of the BM response in Fig. 4a. However, when both aqueducts were included, the BM response is significantly changed, for the both excitation mechanisms modelled here. It was found that the BM response also has a very similar shape and magnitude to those in Fig. 4a when only the VA was modelled, however, for clarity this has not been shown. The VA is on the opposite side of the cochlea, creating a complaint structure either side of the BM, and hence allowing for a volumetric excitation of the cochlea to be generated. The BM response due to local excitation at the round window is increased by more than 50 dB at low frequencies by the inclusion of the VA, however, at around 2 kHz, the BM response is reduced by up to 10 dB and then at higher frequencies the response becomes comparable to a model with no aqueducts present. This suggests that the excitation mechanism is volumetric at low frequencies and non-volumetric at high frequencies. The total BM response, when both aqueducts are present, was separated into its volumetric and non-volumetric components, which is shown in Fig. 4b. At low frequencies the volumetric component dominates, while at high frequencies, this switches and the non-volumetric dominates. Between about 1 kHz and 3 kHz the magnitude of the components are similar, however, there is a phase different between the BM responses due to each component, which causes the combined response to be lower. The BM response, due to the fluid inertia component, could not be separated into a volumetric and non-volumetric component, in the way the local round window excitation can be. However, it was concluded that the introduction of a volumetric component of excitation, via the addition of the VA, significantly increased the BM response due to this excitation mechanism. 4. Conclusion A simple 2D finite difference model of the cochlea was used to investigate multiple mechanisms of cochlear excitation and, in particular, the effect of the aqueducts on the BM response was considered. The simplicity of the model allowed the model to be easily adapted to simulate the various excitation mechanisms; the piston-like and rocking motion of the stapes, the two inertia components of BC ICSV22, Florence, Italy, 12-16 July 2015 7 The 22nd International Congress of Sound and Vibration hearing and local excitation at the round window, as well as simulating a diseased ear, which has an immobile oval window. Because all the mechanisms were simulated on the same model, the BM responses could be easily calculated and justifiably compared. It was found that all mechanism of exciting the cochlea produced a similar travelling wave along the BM. When the cochlea was excited at the oval window or by the fluid inertia component of BC with a flexible oval window, the presence of the aqueducts made little difference to the peak BM frequency response, apart from a low frequencies. Below 500 Hz, the BM response was increases by the addition of the aqueducts when excited by the rocking motion of the stapes or by fluid inertia component. However, when the oval window was assumed to be immobile, and local excitation at the round window or the fluid inertia component of BC were simulated, the addition of the VA had a considerable effect on the BM response. When no aqueducts or just the CA was modelled the model could only be excited by non-volumetric flow, however, the inclusion of the VA in the model allowed for volumetric excitation of the cochlea. The increase in BM response when excited by the fluid inertia component was due to the ability to generate a volumetric excitation. The BM response, due to the local excitation at the round window, was separated into its volumetric and nonvolumetric components. It was shown that the response at low frequencies was mainly volumetric, while at high frequencies the non-volumetric component dominated, however, at mid frequencies the two components added destructively producing a low BM response. Acknowledgement This research was supported by the EU project SIFEM Grant No. FP7-600933. REFERENCES 1. S. Stenfelt and R. L. Goode, “Bone-conducted sound: physiological and clinical aspects", Otol Neurotol 26, 1245-61, (2005). 2. T. D. Weddell, Y. M. Yarin, M. Drexl, I. J.Russell, S. J. Elliott and A. N. Lukashkin, “A novel mechanism of cochlear excitation during simultaneous stimulation and pressure relief through the round window", J R Soc Interface 11, 20131120, (2014). 3. S. Stenfelt, "Inner ear contribution to bone conduction hearing in the human ", Hearing Res, (2014). 4. N. Hato, S. Stenfelt S and R. L. Goode, “Three-Dimensional Stapes Footplate Motion in Human Temporal Bones", Audiol Neurotol 8, 140-152, (2003). 5. E. Edom, D. Obrist and R. 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