Input Impedance of The Cochlea in Cat

Input impedance of the cochlea in cat
ThomasJ. Lynch,Ill,b) Victor Nedzelnitsky,

c) and WilliamT. Peake
Departmentof ElectricalEngineeringand ComputerScienceand ResearchLaboratoryof Electronics,
Massachusetts
Institute of Technology,Cambridge,Massachusetts 02139 and Eaton-PeabodyLaboratoryof
AuditoryPhysiology,MassachusettsEye and Ear Infirmary, Boston,Massachusetts02114
(Received24 June 1981;acceptedfor publication17 March 1982}

Toneswere delivereddirectly to the stapesin anesthetizedcatsafter removalof the tympanic
membrane,malleus,and incus.Measurements weremadeof the complexamplitudesof the sound
pressureon the stapesœs,stapesvelocityVs, and soundpressurein the vestibulePt. From these
data,acousticimpedanceof the stapesand cochleaZsc--•Ps/Us, andof the cochleaalone
Zc --•Pv/Us werecomputed(Us • volumevelocityof the stapes= Vs X areaof the stapes
footplate).Somemeasurements weremadeon modifiedpreparationsin which (1) holeswere
drilled into the vestibuleand scalatympani, (2) the basalend of the basilarmembranewas
destroyed,(3) cochlearfluid wasremoved,or (4) staticpressurewasappliedto the stapes.For
frequenciesbetween0.5 and 5 kHz, Zsc •Zc; this impedanceis primarily resistive
(IZcl • 1.2x 106dyn-s/cm
5)andisdetermined
bythebasilarmembrane
andcochlear
fluids.For
frequenciesbelow0.3 kHz, IZsc] > IZcl andZsc is primarilydetermined by thestiffness
of the
annularligament;drying of the ligamentor changesin the staticpressuredifferenceacrossthe
footplatecanproducelargechanges in IZscl. For frequencies
below30 Hz, Zc is apparently
controlledby the stiffnessof the round-windowmembrane.All of the resultscan be represented
by an networkof eight lumpedelementsin which someof the elementscan be associated with
specificanatomicalstructures.Computationsindicatethat for the cat the soundpressureat the
input to the cochleaat behavioralthresholdis constantbetween1 and 8 kHz, but increasesas
frequencyis decreasedbelow 1 kHz. Apparently,mechanismswithinjthe cochlea(or more
centrally)havean importantinfluenceon the frequencydependence of behavioralthresholdat
low frequencies.
PACS numbers: 43.63.Kz, 43.66.Gf, 43.80,Lb, 43.63.Hx
INTRODUCTION soundpressurehavebeencombinedwith measurements of

Airborne acousticsignalsare normally transmitted ossicularmotionto estimatecochlearinput impedancefor
from the externalear canal to the cochleathrough the me- thecat(Nedzelnitsky,1974a,b, 1980)andguineapig(Franke
chanicalsystemof the middleear. To understandthe oper- and Dancer, 1980).The primary goal of the work reported
ation of the middle ear, it is necessaryto know the mechani- hereisto contributeto a morecompleteunderstanding of the
cal constraintsimposedby the cochleaon the relation middleearby measuringthe impedances of the cochleaand
betweenforce and motion at the oval window. If the stapes stapesin the cat and by determiningexperimentallyhow
motion is one dimensionaland the systemis linear, these variousanatomicalstructuresinfluencetheseimpedances.
constraintsare characterizedby the "input impedance"of
I. MECHANICAL MODEL AND DEFINITION OF
the cochlea,whichrepresents the mechanical"load" driven VARIABLES
by themiddleear.To understand thequantitativevariations
of middle-earresponseamongindividualsand specieswe Acousticstimulationat the tympanicmembranepro-
alsoneedto knowhowthe mechanicalpropertiesof various ducespistonlikemotionof the stapes(Guinanand Peake,
anatomicalstructurescontributeto the cochlearinput im- 1967; Dankbaar, 1970; H$gmoen and Gundersen,1977;
pedance. Rhode, 1978).The forcesthat act on the stapesresultfrom
Somemeasurements of cochlearinput impedancehave the structuresadjoiningit and from the soundpressures in
beenreportedfor cats(Tonndorfet al., 1966;Khannaand themiddle-earcavityandthevestibule(Fig. 1).If weassume
Tonndorf, 1971) and for human cadavers(B6k6sy,1942, that (1)thestapesisa rigidbodyand(2)eachof thepressures
1960,pp.435-436;Onchi,1961).All of theseresultsareseri- Ps andœv is uniform over the relevantsurface,then summa-
ouslylimitedin their accuracyby technicalproblemsin the tion of the forcecomponents
that are alongthe directionof
measurements.Recently measurementsof intracochlear motion yields
alA portionof thisworkwassubmitted

by T. J. Lynch,III asa mastefts ArppS
+fs =f^L + Arppv+ M•bs, (1)
thesisto the Departmentof ElectricalEngineeringandComputerScience, where
Massachusetts Institute of Technology,September1974 and a prelimi- Aep= effective
areaofthestapes
footplate,
]
nary presentationwasgivenat the 91st meetingof the AcousticalSociety
of America (Lynchet al., 1976). M •' = (mechanical)
massof the stapes,and
hipresent
address:
M.I.T. LincolnLaboratory,RoomL-110,Lexington, bs = dvs/dt = accelerationof the stapes.
MA 02173.
tiPresentaddress:National Bureau of Standards,SoundBuilding (233), If we further assumethat this systemis linear for the
Room A149, Washington, DC 20234. small displacementsassociatedwith acousticstimulation,
108 J. Acoust.
Soc.Am.72(1),July1982 0001-4966/82/070108-23500.80 ¸ 1982Acoustical
Society
ofAmerica 108
Redistribution subject to ASA license or copyright; see http://acousticalsociety.org/content/terms. Download to IP: 134.225.1.226 On: Mon, 22 Dec 2014 07:15:28
ß
fAL I----•-Velocity
I Detector:' Signal
Absorber--i Processo•r
ß.•i
'•J Plastic
Connector - ....... ' ................... ß
••
•" ,•
I•
Jg] ii •........:..•...•:....•
........
•
'
[!'g"•:i
....................................................
P•obeAX '•!F•ad•otx•!
•Cavity
Tube
• i .X :M•,ba•rl
I• Stapes•X.... •ce•\
ement'
FIG.1.Diagram
indicating
pressures
•d forces
acting
onthe ß
stapes
a •d []•
•alant
("•ltrat•')
?.:."•::•
Drilled
Hole
the
resulting
stapes
motion.
Ps=soundpress•e
onthe
middle-ear
surface
of the sta•s, Pv = soundpress•eon the vestibular
from
the
incud•stap•ialjoint,fn•
= force
from
the
•nular
ligament,
and
surface,
• = force '•/3••
Vs
=velocity
ofthe
stapes.
Forces
pe•endicular
tothe
assumed
direction
ofmotion,
including
that
from
the
stapedi•
tendon,
have
been
omitted.
--
thenforthe sinusoidal
steady
state
thevariabl• canberepresented
(of
angularfrequency•)
bytheircomplexamplitudes Ps
Ps,Pv,F•, FAC,l/s;theforcesof theannularligament and Zsc
=A---•pVs
=IMPEDANCE
OF
STAPES
I•COCHLEA
cochleacan be expressed in termsof impedances with
FAC=•Z AL
m VsandAfpPv•Z m[/r m andZ •n
S,whereZ AL c are = Pv=IMPEDANCE
OFCOCHLEA
c
Zc AfpVs
themechanical impedances [force/(lineal
velocity)]
dueto
theannularligamentandcochlea, respectively.
With these FIG. 2. Configuration of themeasurement systems.Thesoundpressure on
thelateralsideof thestapes Ps wasmeasured withan air-filledprobe-tube
substitutions
œq.(1)canberearranged
to give microphone system (leftside).ThelinealvelocityofthestapesVs wasmea-
(AfpPs
+ Fj)/Vs= Z• +jwM• + Z• suredusingtheM/Sssbauer technique,in whicha gamma-ray sourceisat-
=z
rn
+z (2) tachedto thestapes andtherateat whichphotons aredetected througha

resonantabsorberis usedto determinethe velocity.Soundpressureon the
wherewehavedefinedZ Srn='r'"
A 7 ALrn .•_jwMS'rnDivisionof Eq. medialsideof the stapes Pv wasmeasured with a transducer (rightside)
havinga fluid-filledprobetubethat wasintroduced into the vestibule
(2)byA •pyields througha holedrilledintothetemporal bone.The lower'portionof the
+ = g " + Z "--•Z" (3) figurewastracedfroma projection ofa histological
section ofthetemporal
boneof oneof theexperimental cats.(Thesection wasapproximately in a
where horizontalplane,butwasslightlyinclined soastointersect boththestapes
footplateandthedrilledhole.)Theprobetubesandacoustic cavityaround
Us= AtpVs= complex
amplitude
ofstapes
volume thestapes aredrawnapproximately to scale,buttheirlocations havebeen
velocity, distortedto presenta clearerpicturein twodimensions.
. "•
Zs Z •/A •p= acoustic
impedance
ofthestapes
(and madewith a probe-tubemicrophone.From measurements
annularligament}, of the threecomplexamplitudes
Ps, Pv, and Vs and of the
Zc"--•Z •/A •p= acoustic of thecochlea, stapesfootplateareaAlp,the acoustic
impedance impedances
canbe
and calculated
asZsc = Ps/(AfpVs)andZc = Pv/(AfpVs).
Z •c = acoustic
impedance
ofthestapes
andcochlea.
B. Animal preparation
In therestofthispaperwewilldealexclusively
withacoustic
impedances
andthesuperscripts
inZ •, etc.,willbeomitted. Adult catsweighingbetween1.6and3.9kg wereinitial-
ly anesthetized withanintrapcritoneal injectionof Dial (0.75
II. METHODS ml/kg}. Additionaldosesof 0.2 ml were administeredas
A. General plan of the measurements needed.Each cat received250 000 units of penicillinintra-
Equation(3)indicates
thatthestapes
canbedriven
both muscularly and 50 ml of physiological salinesubcutaneous-
by forceappliedfromtheincusFj andby soundpressure ly. A cannula was inserted
into the trachea.Rectaltempera-
applied
tothestapes
surface
Ps.In theexperiments
reported ture was monitored and maintained at 37*_+ 2øC with a
here most of the middle-ear structureswere removed and heatingpad.
acousticstimuliweredeliveredto a cavityaroundthe stapes The pinna,earcanal,tympanicmembrane, bullawall,
(Fig.2) sothatF• --0 andPs is thesoledrivingvariable. bony septurn,and ventralsegment of thetympanicringwere
This method,whichhasbeenusedpreviously(Weverand removedto exposethemiddleear.The incuswasthensepa-
Lawrence,1950;Tonndorfet al., 1966;KhannaandTonn- rated from the stapesat the incudo-stapedial
joint with a
dorf,1971),hastheadvantagesthatnorigidmechanical at- miniaturescalpel.The tensor-tympani tendonwascut, and
tachmentneedbe madeto the stapesand that the driving the remainingsegmentof the tympanicring, the malleus,
forcecan be inferredfrom measurementsof soundpressure and the incus were removed.
109 J. Acoust.
Soc.Am.,Vol.72, No.1, July1982 Lynchet al.' Inputimpedance
ofthecatcochlea 109
In orderto providean unobstructed
viewof thestapes diaphragm.A theoreticalanalysisindicatesthat the sound
footplatesomenearbystructureswereremoved.The tensor pressures at theprobetip andat themicrophone
diaphragm
tympaniwascompletely excised.
The stapediustendonwas differby lessthan 1 dB for frequenices
below17 kHz. The
severednearitspointof attachmentto thestapes.
(Measure- calibrationcurvewasstoredin thecomputersystemsothat
mentsof [Zsc[ in onecatbeforeandafterstapedius detach- outputvoltagesfrom the probe-tubemicrophonecouldbe
mentshowedno significant changes.)Usuallythebonecov- convertedto soundpressures.
etingthefacial-nervecanalposterodorsalto thestapeswas In one experimentstaticpressurewas alsointroduced
removedalongwith a shortsectionof the facialnerve.Both into the cavityaroundthe stapes.An otoadmittancemeter
endsof the canalwerethenpluggedwith cottonto restrict (Grason-Stadler1720)wasusedasthestaticpressuresource
fluidseepage.
The stapediusmusclewasalsoremoved.Dur- andconnectedby a Tjunction to our sounddeliverytube.A
ingtheseproceduresit wasimportantto avoidbreaking into specialhousingfor thePs condenser microphoneventedthe
the lateral semicircular canal. staticpressure to the backsideof the diaphragmsothat the
To providea goodsurfaceforsecure bondingofcement, microphonesensitivitywas nearly independentof static
the periosteum wasremovedfrom the petrousbonesur- pressure.
roundingtheovalwindowandtheexposed bonewasallowed
to dry. A few dropsof physiological salinesolutionwere D. Velocity measurements
placedaroundthestapes footplateto keeptheannularliga- I./ntroduction
mentmoist(seeSec.III).
A cavity was then constructedaroundthe stapes(Fig. Our useof the M6ssbauereffectfor velocitymeasure-
2). The baseof the cavity was made with dental cement mentsdifferssomewhat fromthemethods of othergroups
("Grip," S.S. White Co.) whichwasappliedto the dry bone (Giladetal., 1967;Johnstone
etal., 1970;Rhode,1971;Hel-
surroundingthe oval windowsoasto fill the spacesformed fenstein,1974;Gundersen et al., 1978).Figure3 showsthe
by the removalof the facialnerveandstapedius andtensor- relationof thephotonrateat thedetector to thevelocityof
tympanimuscles.Thin layersof the cementwerebuilt up to theM/Sssbauer source.
Thisrelationcanbeexpressed as
form a flat surfacearound the oval window. During this r=Roo(1--a/[1 + [(v--Vi)/F]2}), (4)
processthe endof thePs probetubewaspositionedlessthan where
2 mm from the dorsaledgeof the stapesfootplateand the
cementcavitywall wasbuilt aroundthe tube.A cylindrical,
rigid, plastictube(3 mm i.d., 7 mm long)wasthencemented
PHOTON
to thebaseto form a cavitycontainingthestapesandoneend
RATE, r
of the probetube. The M/Sssbauer sourcewasaffixedto the
stapesheadusingeither petrolatumor zinc-oxidecement. -R m I
Usually after this portion of the procedurehad been • OUTPUT:

completed,we connectedthe soundsourceto the cavity and ,-_-/ \
madeinitial measurements of pressurePs and velocity Vs. [ / r(t)
-I- --I-A-- AVERAGE
These measurements included determination of the stimulus
levelIPslrequiredto producea constant
velocityamplitude
overa rangeof stimulusfrequencyfsothat IZsc(f)l could (]R•o
be estimated.
C. Stimulus generation and measurement

Rapid measurementof stapesvelocitywith the M/Sss-
bauermethodrequireshigh sound-pressure levelsat the sta- VELOCITY, v
pes.The stimulussystemconsistedof an oscillator,attenua- INPUT:
tor, poweramplifier,and eithera JensenDD- 100or an Atlas VELOCITY

)
WAVEFORMS
PD-60 driver coupledto the stapescavitywith rigid plastic v(t)

tubing.Thissystemcouldproduceup to 150dB SPL from20
to 5000 Hz. For some of the measurements the oscillator and
attenuatorwerecontrolledby a computersystemwhich al-
lowed sweepingof the frequency while keeping either a
stimulusor responsevariableconstant(Weisset al., 1969).
FIG. 3. Essential features of the Mfssbauer method for measurement of
Themicrophone (12mm diamBrfiel& Kjaer4134)and
velocity.The curver(v),represents
thedependenceof photondetectionrate
probetubeusedforPs measurements werecalibrated(Weiss r on velocityv. Two sinusoidalvelocitywaveformsandthe resultingrate
and Peake, 1972)in a speciallyconstructedcavity. A con- waveformsare showm The smaller velocity waveform(peak ampli-
densermicrophone(25 mm B&K 4132)wasusedasa sound tude-- 0.5 I V•l:dottedcurve)produces
a ratewaveform
thatisapproxima-
sourceto drive a cylindricalcavity 3 mm in diameterand 7 telysinusoidal
with an averagevalueaboutequalto Ro,theratewith v -- 0.
The largervelocitywaveform(peakamplitude- 3 I V•I: solidcurve)pro-
mm long.A calibratedcondenser microphone (3 mm B&K duces
a nonsinusoidal
ratewaveform withanaverage
value(indicated
by
4138)terminatedthiscavityandtheprobetubewasinserted thedashed
line)thatislargerthantheresting
rateRo.(Positive
velocity
sothat its tip was 1.5mm from the centerof the microphone indicates
motion
oftheM6ssbauer
source
toward
thestationary
absorber.)
110 J. Acoust.Soc.Am.,Vol. 72, No. 1, July1982 Lynchet al.: Inputimpedanceof the cat cochlea 110
detectedphotonrate (counts/s), STAPES VELOCITY (Ps•R (PvTRANS[X.•E:R
= asymptotic
valueof r whenIv - V•I•F, OUTPUT VOLTAC-.-.•
) OUTPUT VOLTAGE)
fractionaleffect= (Roo-- Rr }/R oo, where

= minimum (resonant}rate,
velocityof the M6ssbauersource, /
= isomershift (velocityfor maximumabsorption},
and TJL 25 Pt. 8
F = linewidth(or half-widthat half-height).

Iv, rms d/cm
2rms IPvI=8i d/crn
•' rms
The relationbetweenr and v isdeterminedby four para- •.Vs=200ø &Ps
=-197ø 6.Pv=-i23ø
metersR oo,I/i, a, andF, whichmustbeknownto determine Afp
=1.26
xI(•:'
cmz
v from measurements of r. R oodependson the sizeand spe-
cificactivityof the sourceand on the configurationof the Zsc=
•-•pVs
=2.0
xlOS/),
,/-:37
ø Zc=a-•,:
•pVs
I.Ix106Q,/'+37
ø
absorberand detector(e.g.,source-to-detector distance,de-
tectorsize,and efficiency).Vi is determinedby the source FIG. 4. Examplesof averagedwaveformsof response variablesand imped-
and absorbermaterials.F anda are primarily determinedby ancecomputations.In the velocitywaveform,negativeVsis upwards.The
othertwo waveformsaretheamplifiedvoltagesfrom thetwo pressuretrans-
the materialsand configurationof the sourceand absorber, ducers;pressuresare computedby taking into accountthe transfer func-
but they can also be influencedby materialsbetweenthe tionsof the transducers and amplifiers.The anglesshownfor Vs, Ps, and
radioactivesubstance and the absorber(e.g.,sourcematrix Pr are relativeto the electricinput to the stimulussystem.Impedancesare
material,acousticcavitywalls,bone,fluid) and by the pho- computedfrom the fundamentalcomponents of the waveforms.This veloc-
ity waveform(tenpoints/period)wascomputedover63 846 periods;about
tondetectionscheme(e.g.,thewidthof thepulse-height win- 5 min wererequiredto obtainthiswaveform.The pressurewaveforms(100
dow). points/period)were averagedfor 1000 periods;about 5 s were requiredto
Use of nonzeroisomershift hastwo advantages.(1) As obtain eachof thesewaveforms.Stimulusfrequency-- 0.4 kHz.
the velocitydecreases, the sensitivityAr/Av doesnot ap-
proachzero(asit doesif Vi = 0). (2)Positiveandnegative angleof the fundamental,second,and third harmoniccom-
velocitycanbedistinguished, therebyavoidinga "180øambi- ponentsof the waveformswere computed.
guityin phase"(Johnstone andSellick,1972,p. 7; Robleset The time requiredto make a velocity measurementby
al., 1976,p. 929)., this methodis primarily determinedby the random process
We usedtwo proceduresto measurevelocity.(1) The associated with the emissionof photons.The measuredrate,
velocitywaveformv(t) wasobtainedfrom the instantaneous which is an estimate of the instantaneous rate of the random
rater(t ).(2)For sinusoidal
waveforms thevelocitymagnitude process, is a randomvariable;to increasethe accuracyof the
I vI wasobtainedfromtheaveragerate r(t ). estimate,the photoncountsare averagedover many stimu-
lus repetitions.The variability in the velocityestimateis a
functionof the velocitymagnitude,the photonrate, and the
2. Veloci• waveforms
averagingtime.
Equation(4) canbe solvedfor v to give The limitationsof the "velocitywaveform"methodcan
i)/Vi = 1+__
[(R• -- Ro}(r
-- Rr}/(Ro
-- Rr)(Roo
-- r)]1/2. be illustratedby consideringthe time required to make a
(5) particular measurement.With our M6ssbauersourceof
57Coin palladiumanda stainless
steelabsorber
enrichedin
If the four parametersRr, Ro, R oo,and Vi are known,mea-
57Fewe obtainedparametervaluesof V• = --0.2 ram/s,
surements ofr(t )canbeconvertedto v(t) by Eq. (5)exceptfor
F = 0.3 ram/s, and a = 0.3. The detectorwasusuallyplaced
the ambiguityassociatedwith the signof the squareroot.
soasto giveRo• 103/s.For sinusoids
with peakamplitude
This ambiguitywas avoidedby contrainingthe stimulusto
near0.5 Via few minutesof averagingwasrequiredto obtain
levelssuchthat Ivl < IVi I, whichensures
thatthesignof the
relativelyclearvelocitywaveforms(e.g.,Fig. 4} with a reso-
squareroot in Eq. (5)is negative.
lution of ten pointsper stimuluscycle.Sincepeak ampli-
Our methodof determiningvelocitywaveforms wasba-
tudesgreaterthan producedthe ambiguitymentioned
sicallythat usedby Gilad et al. (1967).We useda laboratory
above(andlargeerrorsbecause of therelativeinsensitivity
of
computersystemto (a)accumulatephotoncountssynchro-
rateto velocityfor Iv - Vi ), it wasundesirable
to work
nizedto the stimuluscycles,(b)performthe computationof
Eq. (5), and (c) display a velocity waveform (Lewis and at largervelocities.Smallervelocitiesrequiredlongeraver-
Peake, 1971). aging times so we usually made velocity measurements
To calibratethissytemRo,Rr, andR ooweremeasured (magnitude
and angle}at only onelevel(ca.0.5 Iil peak
amplitude}.Obtainingvelocitywaveformsat 20 frequencies
in eachexperimentwith the source,absorber,anddetectorin
place.Ro was measuredwith no stimulus.A stimuluslevel
with the associatedprandps waveformstookapproximate-
ly 10h.
highenoughto driveIvl> I vii wasthenusedto estimate Rr
fromminimaof r(t ) (seesolidwaveformin Fig. 3). An even 3. Velocitymagnitudefrom average rate
higherstimuluslevelwasusedto estimate R oofrommaxima
ofr(t ).Aftercompletion ofthiscalibration
procedure, wave- The meanrate, r(t), canbe estimated morerapidly
formsofv(t)lVi werecomputedaccording to Eq. (5)asillus- thanthewaveform
r(t}.If v(t} = X/ZVsin(cot
},integration
of
trated in Fig. 4. For sinusoidalstimulithe magnitudeand Eq. (4}yields
111 J. Acoust.Soc.Am.,Vol. 72, No. 1, July1982 Lyncheta/.: Inputimpedanceof the cat cochlea 111
A EXPERIMENTAL(f=708 Hz) M6ssbauermeasurementswere comparedwith those ob-
-- THEORETICAL
tainedfromtheaccelerometer. Thesemeasurements agreed
800- (within ___
2.5 dB and q- 15ø)indicatingthat a thin layerof
petrolatum was a suitable adhesive for the M6ssbauer
aY70
o source. A similar test of the constant F method from 10 to
z
20 000 Hz demonstrated the sameaccuracyin velocitymag-
TJL-24 nitude.To further testthe adhesion(in three cats}a M6ss-
tu 600
•oo • •2o •o •4o •5o • o bauersourcewasattachedto the headof the stapesfirstwith
STIMULUSLEVEL,IPsl(dBSPL) petrolatum and then with zinc-oxidedental cement;mea-
I I I I I I '•
surements
of Igscl werenotsignificantly
different.Sincethe
-50 -20 -I0 0 IO 20 50 cementbondsthesourceto thestapesveryrigidly,thisresult
PEAK
VELOCITY,
•/•V (dSre Ivil) indicatesthat the petrolatumis alsoan adequatebonding
agentand hasthe advantageof easein placementand remo-
FIG. 5.Meanphotonrate(counts/s)
versusstimulus
andvelocity
level.The val.
theoretical
curve(7vspeakvelocity,Vr2'V)wasdetermined
fromEq. (6)
usingparametervalues(Rr = 595,Re= 658,R•o= 820counts/s) that The M6ssbauersourcewas made (New England Nu-
weremeasuredduringan experiment.The experimental
pointsare mea- clear}fromS7Coplatedon palladium
foil (12/•m thick}.A
suredmeanrateversusstimulussound-pressure
level.The twoplotswere rectangularpiece{300X 380/•m) wascutto fit on theheadof
positioned
horizontally
soasto minimizethemeansquareerrorbetween the stapes.The estimatedmassof the source{20/•g}is about
experimental
andtheoretical
points.I V•l - 0.2mm/s.
4% of thestapesmass{seeSec.III D1}. Sincethe stapesmass
itselfseemsto make only a small contributionto the normal
?= { ] -a[(y + (6) Zsc, the additionof the sourcemassshouldhavea negligible
effect.
where x:y2+ b, y: [(2F2- F/2)//"2]q-l, and
The M6ssbauer system detects the velocity of the
b = (2Y'i/I' }2.FromEq. {6)wefind{Fig.5)thatF increases source relative to the absorber. For determination of the im-
fromReto closeto R oowhen11/'2V/Vii increasesfrom0 to pedancesZsc and Zc we needto know the velocityof the
30dB.For I1/'JV/V• I intherange10to20dB,Fisa sensitive stapesrelativeto theskull(petrousbone).To testwhetherthe
indicatorof thevelocityamplitude.To determineimpedance velocityof eitherthe petrousboneor theabsorberwassigni-
magnitude overa widefrequency range,it is convenient
to ficantly differentfrom zero, in one preparationthe M6ss-
adjustthestimuluslevelat eachfrequency sothattheveloc- bauersourcewasplacedon the petrousboneadjacentto the
ityremains
intherangewhere
Fissensitive
tochanges
in I Fl. oval window. With the highestsoundpressuresthat our
Thecomputersystemwasprogrammed
to adjustthestimu- acousticsystemcouldgenerate,we lookedfor increases in F
lus levelsothat the measuredvalueof F waswithin a given at frequenciesspanningthe range that we used.We were
toleranceof a specifiedvalueand to displaythe resulting unable to detect any motion with this method. Since the
stimuluslevelversusfrequency.The curverepresentingEq. samemethod was able to produceincreasesin F at lower
{6}wasusedto convert thespecified
valueoff intoa velocity soundpressures whenthe sourcewason the stapes,the mo-
magnitude (Fig. 5) sothat impedancemagnitude couldbe tion of the other structures, such as skull or absorber, was
computed. negligible.
Thephotonratewasaveraged witha "ratemeter"hav-
inga timeconstantof 0.5 s. With thiscomputerizedsweep 6. Summary of velocitymeasurement methods
systemwe couldmakemeasurements at 20 frequencies
The rangesof velocityand frequencyover which these
(between30 and 10000 Hz) in 5 min.
two methodswereappliedare indicatedin Fig. 6 alongwith
4. Determination of isomer shift V• a curveindicatingan upperlimit for predominantlylinear
behaviorof the intact ossicularchain.The figureshowsthat
Both the velocity-waveformand mean-ratemeasure- the velocitylevelsusedin the work reportedherewerewith-
mentsproducedvelocityvaluesexpressed in termsof the in the rangein which the fundamentalcomponentof the
isomershift V,..To determinethe valueof F'i we measured motion showslinear growth with stimuluslevel, with the
the magnitude of the motionof a vibratorat onefrequency possibleexceptionof the F measurements at the lowestfre-
(56Hz}(1} withtheM6ssbauer waveformmethod,(2}witha quencies.
calibratedaccelerometer,and {3} with a microscope,eye-
piecemicrometer,
andstrobescopic illumination.
Thelatter
E. Measurement of sound pressure in the vestibule
two measures
agreedwithin20%. To makethe M6ssbauer
waveformmagnitudeequalto the averageof the othertwo The transducersusedfor measuring
intracochlear
pres-
measuresrequiredthat F• = -0.2 mm/s. sureand their calibrationhavebeendescribedby Nedzel-
nitzky (1974a,1980}.
5. Validation of the method
1. Calibration
The systemwas testedby measuringvelocitywave-
formson a vibrator(B&K 4290 or 4810)for sinusoidal
mo- A fluid-filledvial wasmountedon a vibratorto gener-
tion at frequencies
from 30 to 30000 Hz. The M6ssbauer ate an approximatelyuniform pressurefield for calibration
sourcewas attachedto an accelerometer
with petrolatum. purposes.
Thetip of thetransducer's
probetubeanda refer-
112 J. Acoust.Soc. Am., Vol. 72, No. 1, July 1982 Lynchet &l.' Inputimpedanceof the cat cochlea 112
apparent damage to the annular ligament or the basilar
membrane.)With the P•, pressuretransducermountedon a
micromanipulatorthe probetubewasinsertedinto the hole.
To providea sealaroundthe probetubea mixtureof algin-
ate-basedentalimpressionmaterial("Jeltrate,"L. D. Caulk
Co.) was placedon the bone around the probe tube. This
liquidsetwithina fewminutesintoa firmgelwhichprovided
a sealthat preventedperilymphleakagearoundthe probe
tube.
INSTANTANEOUS
RATE Measurements of IZsclwereusedto testthequalityof
theseal(e.g.,Fig.9).If IZsc[wasnotthesame(withina few
dB)with the holesealedasit hadbeenwith the labyrinth
j I I I I IIIIj I I I i illill I i I I IIIIj I I I intact, the Jeltratewas removed,the bonearoundthe hole
I0 I00 I000 I0000 wasdried,andanotherapplicationwastried.In mostcasesit
FREQUENCY (Hz) waspossible to achievea goodsealthatwasstablefor at least
a few hours.
FIG. 6. Ranges
of thetwovelocitymeasurement
methods
compared
with
3. Yalidityof measurements
an estimateof the linearrangeof operationfor the intactossicularchain.
The velocityandfrequency rangesusedfor eachmeasurement methodare In orderto ensurethat the transducerwasmeasuring
indicatedbytheshaded rectangles.
Thecurve(basedonanaverage transfer
the pressureat the tip of the probetube,measurements
were
functionofthecat'smiddleear,GuinanandPeake,1967)givesthestapes
velocityfor a soundpressure of 130dB SPL at the tympanicmembrane madein thevestibule(intwo cats)beforeandafterthe probe
(withthebullaopenandthebonyseptum removed).Theresults of Guinan tip wasmechanicallyplugged.Also, the outputof the mea-
andPeake(1967)indicated thatthefundamental component ofstapes dis- suring system(the noise floor) was determinedwith the
placement isa linearfunctionofstimulus
sound-pressure levelatleastupto
thispressurelevel.In thisandallthefollowing
figures theordinate isloga-
stimulusoff. For all resultsreportedherethe measurements
rithmicwithtickmarksat equallogarithmic intervals. areat least 10dB aboveboththenoisefloorandtheplugged-
tip output.
ence transducer were immersed in the fluid to the same
F. Impedance computations and accuracy
depthandtheoutputsofboth weremeasuredto determinea
calibrationcurvefor theprobe-tube transducer.
[Therefer- To calculateacousticimpedancesfrom the pressure
encetransducer, whichhadanexposed diaphragm, wascali- andvelocitymeasurements,
the stapes
footplate
area,4rp
must be known. The area of the oval window was measured
bratedin airusinganelectrostaticactuator
andpistonphone
(Briiel,1964,1965).]At the highfrequencies
thismethodis in temporalbonesof eightof the catsusedin experiments.
A
moreaccurate thanthemethodreported previously(Nedzel- meanof 1.20mm2wasobtained
witha rangeof 1.00to 1.38
nitsky,1980),because it doesnotrequireabsolute
rigidityof mm 2 and a standard deviation of 0.13 mm •. Similar results
the attachment of the vial to the vibrator. havebeenreportedby Wever et al. (1948)and Guinan and
The probewascalibrated at thebeginningandendof Peake(1967).In the resultspresentedherethe footplatearea
eachexperiment. In general,measurements are reported wastaken(somewhat
arbitrarily)as 1.26mm• in all imped-
onlyfor frequencies wherethesetwo calibrations agreed ancecalculations.
We estimatethaterrorsin acousticimped-
within -[-5 dB. (Thelargestdifferences
usuallyoccurredat ancemagnitudeof approximately-[- 1.5 dB may be intro-
high frequencies and the low-frequencydifferenceswere ducedbyignoring
intercat
variations
of,4rp.
considerably
smaller.)In order to confirmthat the trans- Zc and Zsc were calculatedfrom data obtainedwith
ducerwasresponding
tothepressure
attheprobetip,thetip both of the velocitymeasurementmethodsdescribedabove.
waspluggedat theendof theexperiment andmeasurements Basedonworst-case
esimates
of errorsin thequantities
used
in thevialwererepeated.
Theoutputmeasurement wascon- (,4fp,q- 1.5dB;IPsl, _+ 1.5dB;I s l, _+3 dB)we calculate
sideredvalidonlywhenit wasat least10dBlargerthanthe thelimitson errorin IZscl as q-6 dB withthewaveform
"plugged-tip"output.ThiscriterionusuallylimitedthePv method.Because of largerinaccuracyin absolute
calibration
measurements
to frequenciesbelow 10 kHz. and instabilityin the P•, transducer,we estimatethe error
limitsin IZcl as _+10dB. For theconstant • methodlimits
2. Transducerplacement onerrorin Vsareaboutq- 5dBsothatfor IZsclerrorlimits
are -t- 8 dB. Note that the actual measurement errors are
Priorto the insertionof thePv pressure probe,a hole likely to be substantiallysmallerthan theseworst-case
esti-
(usuallyabout0.3 mm diam)wasdrilledinto the vestibule mates of error limits.
anteroventrallyto the oval window. perilymphalways
flowedfromthe openhole.If therewasbleedingfrom the
III. RESULTS
vestibule,
physiological salinewasusedto washawaythe
bloodbeforeit clotted.(Todetermine whetherthedrilling A. Stability of preparation
produced grossdisruptionsin thelabyrinth,temporalbones In mostof ourexperiments
wemadeaninitialmeasure-
fromfour experiments werepreparedfor histological sec- mentof the sound-pressure
magnitude IPsl required
to
tions. In eachcasethere was a clean hole in the bone without maintain
a constant
meanrate• overa rangeoffrequencies.
113 J. Acoust.Soc.Am.,Vol.72, No.1, July1982 Lyncheta/.:Inputimpedance
ofthecatcochlea 113
TIME (HR:MIN)
I00 - o 0.'00 (INITIAL MEASUREMENT)
ß 5'.20 (LIGAMENT APPEARS DRY)
A 5•40 (AFTER SALINE PUT ON
LIGAMENT) m+90]
•
,-, --90
0
I I I IIII1! I I I IIII1! I I I IIII1! I

.J
z
IO0 1000 IOO00
10- iP,i (ds SPL)

+ •o
o 120
--
o •:•o
TJL- 9
-10-
i' I I I I I III I I I I I I IIII I I I I I IIII I I
I0 I00 I000 I0000
-20-
FREQUENCY (Hz)
-30-
i
FIG. 7. Changesin the magnitudeof the impedanceof the stapesand coch- I00 I000 I0000
leaZsc associatedwith dryingof the annularligament.The initial measure-
ment was obtainedwith salineon the ligament.Approximately5 h later FREQUENCY (Hz)
IZscl had increasedat mostfrequencies and the ligamentappeareddry.
After additionof a few dropsof saline,[Zscl returnednearlyto its initial FIG. 8. Ratio of the pressure
in the vestibulePV to stimuluspressure
values. P$ versusfrequencywith IP$1asa parameter.Neitherthe magnitude
or angle showssignificant dependenceon stimuluslevel. Four curves
are plottedin eachsectionof the figure: one for eachof the IP$1
Fromthesemeasurements themagnitude of theimpedance levels. Data points axeevenly spacedon a logarithmic frequency scale
of thestapes
andcochleaIZsc(f)l wascomputed. The "ini- with 40 points/decade. The symbolsare used only to identify the
tial measurement"
in Fig. 7 is typicalof theseresultsin that curves. The sharpdips at 1800 Hz are an artifact which resulted from
an instability in the acousticsystem. No measurementswere obtained
IZsclhasa negative
slope(about-- 6 dB/oct}forfrequen-
in the region near 8 kHz becausethe stimulussourcecould not gener-
ciesbelow300 Hz and is approximately constantat a value
ate the specifiedIP$1 . For the dataplottedabove8 kHz the upper
near 1 M/2 for frequenciesabove500 Hz. magnitudepointswereobtainedwith IP$1=110 dB SPL;thedatafor
In preliminary experiments themagnitude of IZsclof- 120 and 130 dB SPL are indistinguishableand they paxtially overlap
tenincreased substantiallyovera fewhours(Fig.7}.When with the points for 140 dB.
thisoccurred, thestapes footplateandannularligament ap-
linear growth of velocitywas observed.For instance,the
pearedto be dry asseenthrougha dissecting microscope.
The additionof a few dropsof physiological salineto the impedance magnitudes derivedfrombothvelocity-measure-
exposed surfaceof the annularligamentusuallyresultedin ment methodswere essentiallyequal (e.g.,Fig. 16), even
though the velocitymagnitudesdifferedby 14 to 21 dB.
thereturnof IZscltoapproximately itsoriginalvalues (Fig.
7}. Thus it seemeddesirableto keepthe tissuearoundthe Also,thewaveforms of all thevariables
usuallyappeared
to
be sinusoidal;
secondand third harmonicsfor the velocity
footplate moistat all times.Salineevaporatedtooquicklyso
waveformswere at least 18 dB belowthe fundamental,and
mineraloil wasusedroutinelyin laterexperiments. If theoil
disappeared or IZscl increased, moreoil wasadded.This thePs microphoneandp r transduceroutputsappearedto
procedureresultedin measurementswhich were stablewith- belessdistortedthanthevelocitywaveforms.
Thuswefound
in a fewdB overaslongastwoor threedaysin theliving predominantly
linearbehavior
ofresponses
at thestapes
for
animal. stimuliIPsl up to 140dB SPL.2
In onecase,measurementsof IZscl afterdeathshowed
a gradualincreasewhichreached20 dB at low frequencies C. Effects of modifications of the system
after5 h eventhoughthefootplatewaskeptmoist.No mea- In one cat the stapeswas manually rockedback and
surements fromcadaverearsareincludedin thisreport. forthsoasto rupturemostof theannularligamentwith the
stapes remaining in theovalwindow.The measured IZscl
B. Tests of linearity wassmallerthan normalfor frequencies below600 Hz and
In characterizing
thesystemin termsof impedances [as wasessentiallynormalfor frequencies above600 Hz. This
in Eqs.(2}and(3}]wehaveassumed that thesystemislinear. resultand thoseobtainedwhenthe annularligamentwas
Since it is difficult with the M6ssbauer method to make accu- allowedto dry (Fig.7)suggest thatat lowfrequencies
Zsc is
rate velocitymeasurements
over a large dynamicrange, controlledby the annularligament.
measurements of Pv?Ps providethe mostconvenientmeth- In someearly experimentsthe lateral semicircularca-
od for measuringthe dependenceof a response
on stimulus nal wasinadvertently openedduringthe surgicalprepara-
level. None of thesemeasurements {e.g.,Fig. 8} showeda tion. IZscl in thesepreparations wasgenerallylowerthan
significantdeviationfrom lineargrowthof the fundamental normal for frequenciesbetween1 and 10 kHz. This result
componentof Pv with increasing levelof Ps. The velocity suggests that in this frequencyregionZsc is determinedby
measurements were consistent with this conclusion in that the cochlea;the holeprovideda pathfor the stapesvolume
114 J. Acoust.Soc.Am.,Vol.72, No. 1, July1982 Lyncheta/.' Inputimpedance

of thecatcochlea 114
10--
..
• •--,o;o.z
=• L'OdB
-30
/-20 -I0
i 0 I0
i20 :30
STATIC PRESSURE IN STAPES CAVITY
PDc
(cm H:,O)
• o NORMAL
(%c=0)
0--0}COCHLEA
INTACT
"•:--x' x•.__."'• A poc:7+lcmH20
x--x} VESTIBULE
HOLE
OPEN '• • x RW
REMOVED
(PDc=O)
Z•:-.-..A+}
VESTIBULE
HOLE
PLUGGED %_
IO IOO IOOO
FREQUENCY (Hz) TJL-3I
I I I IIIII I I I I IIIII] I I I IIII1]

I00 I000 I0000
FIG. 9. Theeffectof a holeintothevestibuleonthemagnitude of theimped- FREQUENCY (Hz)
anceof thestapesand cochleaIZscl. Fivecurvesare shown.The firstwas
obtainedwiththecochleaintact(¸), thesecond afterdrillinga hole(approx-
imately1/3 mmin diameter)intothevestibule (r•),thethirdwitha mechan- FIG. 10.Changesin themagnitudeof the impedance of thestapesandcoch-
icalpluginserted intothehole(A), thefourthwith theplugremoved(X), lea ZscI associated with changesin staticpressures
on thestapes.The up-
andthefifthwithit reinserted ( 4- ).Theplugwasheldbya micromanipula- per plot is cochlear-potential
magnitude(CPI) for IP•I - 130dB SPL at
tor andcouldbequicklymovedin or out of the hole.Sevenhourselapsed 100Hz versusthestaticpressure in thestapescavityPoc. (A curveof similar
betweenthe first and secondcurve.For the secondthroughfifth curves, shapewasobtainedfor ]Ps = 120dB.)CP wasmeasured betweena wireon
approximately20 min elapsedbetweensuccessive curves.Similar results
the roundwindowandtheheadholder.For thesestimulusconditionsCP
were obtainedin anotherpreparationin which the hole was repeatedly
wasapproximately 2/zVrmswhenPoc = 0. The lowerplotis Zscl versus
closedand openedusing"Jeltrate"to form a gelatinoussealoverthe hole.
The elevatedvalueat 4 kHz is nottypical(seeFigs.12and 13)anddid not frequencyfor threestaticpressure conditions:Poc = 0, the NORMAL ex-
appear in all the "intact" measurementsfor thisanimal. perimentalcondition;Poc = 7 -t- 1 cm H20, the valueof Poc that yielded
themaximumCP[ in theupperplot;Poc = 0 and theround-window mem-
brane removed.
velocitywhichby-passed
the cochleaandreducedIZscl by
partially "short circuiting" the cochlear impedance.(At-
temptsto plug theseholeswere generallynot effective;no per).Subsequently,
measurements (notshown)of ICPI ver-
further data from thesepreparationsare includedin our re- susstaticpressurewererepeatedafter (a) openinga hole in
sults.) the dura mater over the cerebellum so as to reduce cerebro-
More systematicmeasurements of the effectsof open- spinal-fluid(csf)pressureandperilymphaticpressureand (b)
ingsin the bony wall of the labyrinthwere obtainedwhen removingthe round-windowmembranesoasto reducethe
holesweredrilledfor insertionof a pressure probe.Results perilymphaticpressuredirectly.Theseprocedures produced
obtainedwith a holedrilledinto the vestibule(Fig. 9) show a horizontalshiftin curvessuchasFig. 10 (upper)sothat the
that pluggingthe holeduplicatesthe "intact" conditionand location of the maximum moved toward zero pressure.In
that the differences
betweenthe "open"and "plugged"con- contrast to these resultsat 0.1 kHz, with f= 1 kHz only
ditionswererepeatable. The reductionof IZscl for frequen- smallchanges(
< 3 dB)in IePI occurredwithstatic-pressure
ciesbetween0.3 and 3 kHz with the hole openpresumably variations. These results are consistent with the idea that
resultsfrom the ½ochlearimpedancebeing"shortcircuited" pressurevariationsaffectprimarilythe annularligament,
bythehole.Thereductionin IZscl forfrequencies below0.3 whichcontrolsZsc for the lowerfrequencies.
kHz is moredifficultto explain,sincethe impedanceis sup- Measurements for three conditions(Fig. 10, lower)
posedlydeterminedby the annularligamentin thisfrequen- showedthat IZc(f)l withtheround-window membrane re-
cy range.Similarreductions in IZscl at low frequenciesoc- movedis approximatelythe sameasthat obtainedwhen the
curred when a hole was made in either the vestibule, scala roundwindowisintactanda positivestaticpressure of 7 cm
tympani of the basalturn, or the round-windowmembrane. H20 is appliedon the lateralsideof the stapes;at low fre-
To account for the low-frequency (f<0.3 kHz) quencies
iZcl underthese
twoconditions
islessthanIZcl
changes,it was hypothesizedthat the static perilymphatic with the cochleaintact. Thesechangesat low frequencies
pressure
in theunopened
labyrinthstretches
theannularlig- canbe interpretedasresultingfrom staticdisplacements of
ament so as to increase its incremental stiffness. To test this the stapeswhich producechangesin its incrementalstiff-
hypothesis,measurementswere made with different static ness.When the staticpressuredifferenceacrossthe annular
pressures
in the cavityaroundthe stapes.Stapesvelocitywas ligamentis zero (eitherfrom balancingthe perilymphatic
monitoredby the M6ssbaueraverage-ratemethodand also pressurewithanexternalpressure or fromreductionof peri-
changesin velocityweredetectedby measuringchangesin lymphaticpressure to zero),thestiffness
of theannularliga-
cochlearpotentialsrecordedfrom the round-windowmem- ment is a minimum.The magnitudeof the pressure(7 cm
brane.With a fixed stimulusPs at 0.1 kHz, a maximumin H20 ) determinedin thisonecatis outsidethe range(11.4to
cochlear-potential
magnitude(andin M6ssbauermeanrate) 18.0cm H20) and only about 1/2 the meanvalueof csfand
occurredat a positivepressureof 7-8 cm H20 (Fig. 10, up- perilymphaticpressures reportedfor catby Beentjes(1972).
115 J. Acoust. Soc. Am., Vol. 72, No. 1, July 1982 Lyncheta/.: Input impedanceof the cat cochlea 115
_ /•_NORMAL
'Zsc'
•,,•-• HOLE
OPEN
--'X-•i••/•/- • REMOVED
z <[
<[
0.1-
I
LINE• I
• i i'l'l"fiq I ! !l!•ll
•! • I I•l ' TJ L- 30
! !
I0 I00 I000 I0000
FREQUENCY (Hz)
FIG. 11.Magnitude
oftheimpedance
ofthestapes
andcochlea
[Zsclfora series
ofcochlear
manipulations.
Theinitialmeasurement
islabeled
"NOR-
MAL." A holewasthendrilledintoscalatympanineartheroundwindowandIZsclwasdetermined
withthis"HOLE OPEN."A pressure
transducer
was
inserted
intotheholeandsealed(measurement
of IZsclundertheseconditions
isnotshown butissimilarto theNORMALmeasurement) andfrom
pressure
measurementsinscala
tYmpani
(PRw) theimpedance magnitude
oftheround-windowmembranewascomputed I/•wl - IP•w/Usl(assuming
URw= Us).Nexttheround-window membrane andapproximately2 mmofthebasalendofthebasilar
membrane wereremoved(RW+ BASALBM
REMOVED) and[Zsclwasmeasured.
Finally,
aspirationoftheperilymph
inthebasal
regionofbothscalae
yielded the"FLUIDREMOVED"condition.
Thestapes
wasexcised
andweighed
(wetweight
= M •' = 617/•g);
theimpedance
ofthismass
isgiven
bythecurve
Ijo)Msl,
where
Ms = M •'/A•p.
In two preparations(with similar results)more drastic frequencydependence isessentially

thesamein all cats.At
modificationsof the cochlea were carded out (Fig. 11). frequenciesbelow300Hz, IZsclhasa slopeof - 6 dB/oct;
Openinga hole in scalatympani produceda decreasein forhigherfrequencies
it isalmostconstant
witha magnitude
IZcl forf< 0.3 kHz thatissimilarto thedecrease produced near 1.5 MI2. In six of these 14 cats,we alsoobtainedboth
by a hole in scala vestibuli(Fig. 9) or by removal of the themagnitudeandtheangleofZsc frommeasurements of
round-windowmembrane.IZscl for f> 0.6 kHz was un- Ps andVswaveforms (Fig.13).For frequencies
below0.3
changedin this casebecausea scalatympanihole doesnot kHz, whereIZsclhasa slopeof - 6 dB/oct,theangleap-
"short circuit" the cochlea as does a hole in the vestibule. proaches- 90ø.Near1kHz, IZsclisrelatively
constant
and
Notethat IZRwI<lZscl. SinceZRW wasshortcircuited
by theangleisnear0ø.Above6 kHz theangleandmagnitude
the hole, subsequent
removalof the round-windowmem- tend to increasewith frequency.
braneprobablydid not appreciablyalter IZscI. However, To determinehow the stapesmassaffectsZsc, six
removalof the most basalregionof the basilarmembrane stapes
wereremoved
afterexperiments,
stored
insaline,
and
reducedIZscl for0.3 <f< 7 kHz, indicating
thatthebasilar
membraneis normally an importantfactorin determining
14 CATS
cochlearinput impedance.When fluid was removedfrom
thecochlea,IZscl decreased furtherfor frequenciesabove1
kHz indicating that the fluid alone providesa significant
load on the stapes.With the fluid removed,IZscl at high
frequencies(f> 5 kHz} isapproximately threetimesthe im-
pedanceof the measuredstapesmass.The difference canbe
accounted
forbythemass
(equivalent
tothatofabout1mm3
-6 dB/OCT.,'x
of H20} associated
with theaddedmineraloil andper!lymph SLOPE
remainingon the footplate.
'I i i illill I i i illill I i ! illill I i i
D. Impedance measurements I0 I00 I000 I0000
FREQUENCY (Hz)
1. Impedance of the stapes and cochlea Zsc
Zsc magnitudescalculatedfrom constant? data are FIG. 12.Summary ofmeasurementsof themagnitudeoftheimpedance of
the stapesandcochlea[ZscI madewith the constant
? method.In each
shownin Fig. 12 for all experimentsin whichthe measure- curvedatapoints
arespacedevenly
ona logarithmic
frequencyscale
andare
ment was madebeforeopeningthe labyrinth.At eachfre- connected
bystraight-line
segments.
Pointdensity
varies
among
curves
and
quencytheinteranimalrangeof IZcl isabout10dBandthe is either7, 10,or 20 points/decade.

+90
O
I0 I00 I000 I0000

c• IO0-- A
A 22 I
. •'oo•
+o+O
+ IO- SYMBOLCAT i
• 2õ
x 7
• x,•OAo•+
* 27
I0-
+A.
0 o+t 25
o +o•- . ,,"%
* oA+ • A + •
x 0+• A A+• _•o•
-•B/OC•.• • • oo • •+ • •0 •
I' x x 13_
>
-20
a, a,O
I OO IOOO I OO• -30- o
I I I I lllll I I I I I I I III1!
FREQUENCY (Hz)
IO IOO IOOO IOOOO
FREQUENCY (Hz)
FIG. 13. Impedanceof the stapesand cochleaZsc from six cats.Cat 25 is
theonlyanimalin this groupin whicha holehadbeendrilledinto the vesti- FIG.14.Ratio
ofpressure
inthevestibule
tostimulus
pressure
P•,/Ps:
mea-
bulebeforethemeasurements;
IZsclmeasurements bytheconstant • meth-
surements
fromfourcats.Thecurves(markedbyX, +, O,and&) are
odbeforetheholewasmadewerecloseto thoseobtainedafterthepressure made upofstraight-line
segmentsconnecting
points
witha density
of40
transducer was inserted and sealed into the vestibule.
points/decade.
Theuntilledsymbols
atlowfrequencies
represent
dataob-
tained
fromaveraged
waveforms forconditions
where
P•,wasbelow the
noise
flooroftheusualmeasurement system.
Absolutecalibrations
oftwo
weighed.The weightrangedfrom 422 to 618/•g with an transducers
areinvolved
inthedetermination
ofthispressure
ratio.Because
averageof 521/•g. This averagevaluecorresponds
to an ofpossible
errors inthese
calibrations,
themagnitude
oftheratioatmiddle
acousticimpedancemagnitudeat 10kHz of 0.2 M/2, which andhighfrequencies
maynotbesignificantly
different
from1(0dB).
isaboutonetenthof IZscl. Thusthemassof thestapesdoes
weobserved
that,whendryingof theannularligament
notappearto contributemuchto Zsc evenat highfrequen- caused
anincrease
in IZscI, IPv/Psl decreased.
cies.
Measurements
ofZc areshown
inFig.15.At frequen-
2. CochlearinputimpedanceZc
From measurements ofPv/Ps, Zc canbecomputedby
multiplying Zsc by Pv/Ps. The Pv/Ps measurements
shownin Fig. 14 are representative
of thoseobtainedfrom .A_:.;•' " -:,'.;•--o' "o..-o
ten cats. The four cases shown are all those in which the
.•..•.:•...
d' 7x
........
effectiveness
of the sealaroundthePv probewasverifiedby -9o' I I I I IIIII I I I i iiiii I i i i i ii11 I
z
measuring IZsclbeforeandaftertheprobewasplacedin the IOO IOOO IOOOO
vestibule.
For frequenciesabove1 kHz themagnitudeis ap-
SYMBOL CAT
proximatelyconstantnear0 dB andthe anglevariesaround ß 7
zerosothatPv •Ps. For 50 <f< 300Hz the magnitudede- o 18 _
o 25
creaseswith frequencyandtheangleapproachesd- 90*.For A 27
frequenciesbelow50Hz, IPv/Psl isapproximatelyconstant 0 ' -0- - -0

at - 25 dB and the angleapproaches0'.
In one preparation we obtained measurementsthat 0..... / ' "'.. ..0
wereatypicalin that IPvI wasapproximately
equalto IPs[
evenat low frequencies.It wasfoundthat the cementbaseof
... .o.-
the acousticcavityaroundthe stapeswasnot bondedto the
bonein onespotanteroventralto the ovalwindow. Sincethis
regionof the cementwasin contactwith sealantaround the I I I I IIIII I I i I I IIII I I I I IIIII I
vestibulehole and pressureprobe,apparentlydirect cou- I0 I00 I000 IOOOO
plingofPs tothepressure probeoccurredthroughthispath- FREQUENCY (hz)
way. Thusthe rigidityof the enclosurearoundthe stapes
FIG. 15.Impedance
ofthecochlea
Zc fromfourcats.Magnitude
andangle
seemsto beof greatimportancefor theaccuracyof Pv mea-
were determinedfrom the productof Zsc and P•,/Ps for three cats
surements. Thiskind of couplingwasapparentlynot an ap- (18,25,27).For the fourth cat magnitudewasdeterminedfrom measure-
preciablefactorin our reportedPv measurements,because mentsof IPvl with I•'sl heldconstant.

ciesbelow50 Hz thedatasuggest
that asfrequencydecreases IV. DISCUSSION
the angleapproaches-- 90øand the magnitudeincreases. A. Comparisonwith previouslyreported measurements
Forfrequencies
between 50and500Hz IZcl increaseswith 1. Impedanceof the stapesand cochleaZsc
a slopeofapproximately
4 dBoctandtheangleincreases
toa
maximum value near q-45 ø and then decreases.For fre- The measurements
of Zsc in the cat reportedby Tonn-
quencies
above500Hz, the angleaverages
aboutzeroand doffet al. (1966}andKhannaandTonndorf(1971} alsouti-
themagnitude
is approximately
constantnear2M•2. lized acousticstimulidelivereddirectlyto the stapes.Tonn-
doff et al. (1966} used a capacitiveprobe to measure
displacement of a regionof the round-windowmembrane,
3. Comparisonof measurements
whereasKhanna and Tonndorf(1971}usedtime-averaged
Figure 16 is a summaryof Zsc andZc measurements holographicreconstructions to determineround-window
obtainedin oneanimalutilizingboth velocity-measurement volumedisplacement. The averaged resultsfor IZscl from
methods.Theseresultsillustratethe agreementof the two eachof thesestudiesare shownin Fig. 17 alongwith our
methodsof measurementand they allow comparisonof Zsc averagedresults.The mostprominentdifferences occurat
with Zc. lowfrequencies wheretheirvaluesfor IZscl area factorof
The impedancemagnitudes obtainedfrom the two ve- threeto fivelargerthanours.Sinceweobserved thiskindof
locity-measurement methodsagreewithin 4 dB. This is differencewhen the annular ligamentwas not kept moist
within the limits of error of the methodsand is typicalof the (Fig.7},it maybethatsomedryingof theligamentoccurred
agreementobtainedin all preparations. In thispreparation in their experiments.
resultsfrom both methodsindicatethat duringthe measure- Theaccuracy of the IZsclvaluesobtainedbyTonndorf
mentstherewasa changein IZcl at verylow frequencies. et al. (1966}is limitedby theassumptions thatwereusedto
Comparisonsof Zsc to Zc in otheranimalsweresimi- converttheir lineal displacement measurements to volume
lar in all cases.For frequencies
below0.3 kHz IZscl is displacements. In theirdiscussionof thisproblemKhanna
greaterthan IZc I. For frequencies
between0.5 and 5 kHz andTonndorf(1971,p. 1475}statethat"all oftheseassump-
Zsc •Zc. At the highestfrequencies the angleof Zsc tends tionswereratherpoor."In addition,Tonndorfet al. (1966,
to be more positivethan the angleof Zc. (An interpretation p. 759}reported"someuncertainty" in theabsolutecalibra-
of the high-frequencydifferencesbetweenZsc and Zc is tion of the displacement measurements. Theseproblems
presentedin Sec.IV B3.} withthe 1966results(alongwiththepresumed dryingofthe
annularligament} couldeasilyaccountfor the discrepancy
betweenour averageIZscl andtheirs.
The holographicmeasurements {Khanna and Tonn-
doff, 1971} provideda descriptionof the spatialdistribution
of round-windowmotionsothat volumedisplacement could
be obtained directly. However, in this study "variations
betweenanimals...wererather large" (Khanna and Tonn-
doff, 1971,p. 1480}asisshownby the IZscl rangeplottedin
Fig. 17;KhannaandTonndorf(1971,p. 1482}alsoreported
that IZsclva•d withstimulus levelandtime.Exceptforthe
ß ß Zsc changesthat we found to be associated with drying of the
z I0 ::: Zc
annularligament(Fig.7}or with ineffective pluggingof holes
, into the labyrinth, we havenot encounteredsuchvariations.
It seemslikely to us that the reportedvariationswith level
• '... ."• .. • ' . t and time, and interanimal variations resulted from inade-
• I ,'-,:
.......
•:•,,•;'• - A. quatecontrolof experimental
variables.
3 Their suggestion
(Khannaand Tonndorf,1971,p. 1482}that IZscl wasal-
I i I i i i iii I i i i i i iii I i i i i i iii I i i i i teredby uncontrolledchangesin csfpressureis not support-
I0 I00 I000 I0000 ed by our observations,
sincethey report largevariationsin
FREQUENCY (Hz)
IZscl at 1.9 kHz, whichis abovethe frequencyrangefor
FIG. 16. Comparisonof impedanceresultsfrom onecat. Zsc is the imped- whichmoderatestatic-pressure
changesacrossthe footplate
anceof the stapesand cochlea;Z c is the impedanceof the cochlea.Filled alterIZscI (Fig. 10}.
symbolsrepresentresultsobtainedusingvelocitywaveforms.The solid, M½ller's(1965}measurements of input impedanceat
dashed,and dotted curvesare impedancemagnitudesobtainedwith the
constant• method. No symbolsare plotted for thesecurves;points are
the tympanicmembranein cat canalsobe usedto estimate
equally spacedon a logarithmic frequencyscaleand are connectedby Zsc, sincehemeasured impedance beforeandafterinterrup-
straight-linesegments.
The dashedIZcl curvewasobtainedbeforethe tion of the incudø'stapedialjoint. The circuitmodelof the
waveformmeasurements. The dottedIZcl curvewastakenafter 15wave- middleear shownin Fig. 18 indicatesthe assumptions in-
formdatapointsweredetermined; the 15thdatapoint(markedbyanarrow} volvedin the computation.If it is assumedthat (a} at all
lieson thedottedcurve.Thusa changein IZcl at verylowfrequencies
oc-
curredduringthe timerequiredfor the waveformmeasurements. The other
frequencies the ossicles
moveasa rigidbody,(b)tympanic-
data pointsobtainedin the latter part of this time interval were at higher membranevolumevelocitythat isnotcoupledto themalleus
frequencies,
wheresignificant
changes
in IZcl did notoccur. canberepresented by a paralleladmittance,and(c}possible
118 J. Acoust.Soc.Am.,Vol.72, No.1, July1982 Lynchet al.:Inputimpedance

I'T
• 180-
b.I -
• 90-
e'• _
0-
-.I -
z -90 -
I i ' ''''"1 I i i lllll I i w illill I i I FIG. 18. A simplecircuit modelfor the middleear which wasusedto esti-
I0 I00 I000 I0000 mate the impedanceof the stapesand cochleaZsc from M•ller's measure-
LYNCH et ol. N=14 mentsof admittanceat thetympanicmembraneYD. In thismodelvoltageis
o....o TONNDORFet el analogousto soundpressureand currentto volumevelocity.
I00 - (1966) ß N=.3
•___• KHANNA
TONNDORF(1971)
N=5 a short circuit. Thus
• x M•LLER(1965) N= I
Zsc= T2[(Y:v- YMB)

--1-- (YxJ
-- YMB)--I], (7)
'. x
c- 10- where the admittanceat the tympanicmembraneYz• was

measuredfor the followingconditions:
o \, .oq
C,1 -
Yz• = Ys = normal admittance,
Yz• = Y•s = admittancewith the malleusblocked,
Yz•= Y•j = admittancewith incudo-stapedial
joint
interrupted,and
T = the effective transformer ratio.
The resultsof thiscomputationwith M•ller's measurements

O.I--
I I i illill I I I illill I I I illill I I I
of YN, YMS,and YEJshow(Fig. 17)that for T = 60 the esti-
I0 I00 I000 I0000 matedZsc agreesquitecloselywith our resultsfor frequen-
:30- . cies from 0.25 to 4 kHz. (For f< 2 kHz, I
.,
< 1/41Y•l < I Y.l sothat YMs haslittle effecton thecom-
20-- putedZsc. However,if the effectof YMS is not included,the
-
' estimatedangleof Zsc deviatesappreciablyfrom our mea-
io surements for frequenciesfrom 2.5 to 4 kHz.} Above5 kHz
' ß
..,
..
Zsc estimatedfrom M$ller's resultschangesrapidly with
0 ' frequency,unlike our measurements. It seemslikely that
I I I I I IIIII I I I I IIII I I I I I IIII I I I
M$ller's measurements are relativelyinaccuratein this fre-
I0 I00 I000 I0000
quencyrange{Lynch,1981}.In addition,if relativemotionof
FREQUENCY (Hz) the malleusand incusoccurs,as hasbeenreportedin this
frequencyrangeby GuinanandPeake(1967},differences are
FIG. 17.Comparisonof averagedmeasurements of impedanceof thestapes
expectedat thesefrequencies, sincethe modelof Fig. 18 is
andcochleaZsc in the cat. Our curveswereobtainedfrom the data shown
in Figs. 12and 13.Magnitudewascomputedusingthe pointsin Fig. 13and
inadequate.
data from Fig. 12 for thosecatsnot in Fig. 13. Both magnitudeand angle
werecomputedby averagingall pointsin a 1/2 octavewindowwhosecenter
2. Ratioofpressurein the vestibuletopressureat the stapes
frequencywas incrementedin 40 stepsper decade.The Tonndorfet al. Pv/Ps = Zc/Zsc
(1966)curvesare basedon data for three eatsshownin their Fig. 2. The TheratioPr/Ps wasmeasured directlyin ourprepara-
KhannaandTonndorf(1971)resultsarebasedondatain theirFigs.7 and8.
All their data from two live and three dead cats were included, becausethe tions(Fig. 14}.TonndorfandKhanna(1967}determinedthe
measurements from the live catstend to be the extremes,and the oneprep- pressure requiredoutsidethe ovalwindowto producea giv-
arationthat is reportedbothin live anddeadstatesshowslittle change.For en levelof ½ochlearpotentialboth with and without the sta-
bothTonndorfetal. (1966)andKhannaandTonndorf( 1971),averages were
pes in place. The condition of equal ½ochlearpotential
computedfor eachfrequencyof measurement. Intercat variabilityin the
resultsfrom the threestudiesis indicatedin the lowestplot. In Figs. 17, 19, shouldcorrespond at eachfrequency to equalvolumeveloc-
and20"RANGE" is20 log[Zma
x/Zmin1:noattempthasbeenmadeto take ity through the oval window, if the cochleaitself is un-
accountof the different number of cats involved in each study. RANGE changedby removalof the stapes.In that case,the ratio of
was computedat all frequencies for which data existedfor two or more pressures obtainedin the two situationswould be Zc ?Zsc
animals;interpolationwasusedwhennecessary. In computingthe imped-
macefrom M½ller's(1965)results,data weretakenfrom his Fig. 6 (Y•v, Yu ) and shouldbe the sameas our measurements of Pr/Ps.
and Fig. 8 (Y•a) eachcontainingresultsfrom onecat. Comparisonof results(Fig. 19}indicatesquitecloseagree-
mentin that IP/Psl is near0 dB for frequencies above300
Hz, and hasa positiveslope(about30 dB/decade}for fre-
changes in themodeof motionof theossicles andtympanic quenciesbetween50 and 300 Hz. The differencebetweenthe
membrane thatresultfrominterruptionof theincudo-stape- two setsof resultsincreases
with decreasing
frequencyfor
dialjointorblockingofthemalleusdonotsignificantly alter frequenciesbelow 0.4 kHz. The sourceof this difference
Z•, Z2, andZMB, thenthemodelof Fig. 18is appropriate. couldagainbe dryingof the annularligamentas suggested
Interruption of theincudo-stapedial
jointreplacesZsc with for thelow-frequency
differences
in IZscl(Fig. 17).
119 J. Acoust.Soc.Am.,Vol.72, No.1, July1982 Lyncheta/.:Inputimpedance

ofthecatcochlea 119
03 +90 tu +90
tu 0 tu 0
-• -90 'J -90

z
io ioo iooo ioooo z
i i i iiiii I i i i ii!111 i i i iiiii I
I0- I00 I000 I0000

I0--
.
i x_•_ •- •
0- I/ '•.•_ •.t/ x• .
I
.
• -I0 - I .
o
ß
..
-20 - .
-- LYNCH et ol. N=4

a->-$0 - ...... NEDZELNITSKY N=25 --
O.I--
(1980)
--- TONNDORF et el. N=3
LYNCH
et
ol.
N=zn
j
-40 - TONNDORF
8(
m---m KHANNA(1967)
N= N
(1966)
.
.
I I I I I IIII I I I I I IIIll I I I I IIIII

-50 - I00 I000 I0000
I I I I IIIII I I I I IIIII I I I I
I00 I000 I0000 • 30-
• 20 20-
z IO
o
I I I IIIII1 I I I IIIII I I I I IIIII I I I I I I IIIII I I I Illill I I I IIIIII
100 1000 10000 io ioo iooo ioooo
FREQUENCY (Hz)
FREQUENCY
FIG. 19.Comparisonof averagedmeasurements of the ratio of pressurein

the vestibuleto stimuluspressurePv/Ps. Our curveswerecomputedfrom FIG. 20. Averagesand rangesfor measurements of cochlearinput imped-
the data of Fig. 14 by the followingprocedure:(1) all the pointsin Fig. 14 anceZc from threestudiesin cat. Our curvesare basedon Fig. 15 of this
connectedby straight-linesegmentswereaveraged(with a 1/2 octavewin- paper.Thecurves
forTonndorf
etal. (1966)wereobtained
fromthedatain
dow);(2) the "waveformpoints"(opensymbols)whichwereat frequencies Fig. 3 of that paper.In botheasesdatain a 1 octavewindowwereaveraged.
not includedin the first averagewereaveragedwith a 1 octavewindow;(3) Nedzelnitsky(1980) curvesare mediansof the magnitudeand angle of
the two averageswere then combined.The Tonndorf and Khanna (1967) Zv = Psi/Us from Fig. 16 of his paper.Note that for the Nedzelnitsky
curveis an averageof data from threecatsin their Fig. 10. (1980)measurements the indicatedRANGE is the rangein measuredpres-
suresothat someof the variabilitymay resultfrom inter-animalvariations
in middle-eartransferfunctionratherthan variationsin IZcl.
3. Coch/ear /nput /ropedanceZc
Three setsof measurements are plottedin Fig. 20. The ments.If the Nedzelnitskycurveis translatedupwardby 6
Nedzelnitsky(1980}curve (whichwas termedZ• in that dB the two curves are within 5 dB of each other over the
paperto distinguishit fromZc }isbasedonmeasurements of frequencyrangefrom 15 to 2000 Hz. The frequency-inde-
soundpressurein scalavestibuliof the basalturn in 25 cats pendentdifferencecould resultfrom severalfactorsinclud-
with intact ossicularchains.Measurementsof stapesveloc- ing errorsin the absolutecalibrationsof the pressuretrans-
ity were not made in theseanimals;Z• was obtainedby ducersand of the velocitymeasurement.Both setsof results
dividingthe medianpressuremeasurements by the average indicatethat at verylow frequencies,
IZcl increaseswith
stapesvolumevelocityof GuinanandPeake{1967}.Because decreasingfrequencyandtheangleapproaches- 90ø.This
Nedzelnitsky'spressureprobeswere placed4-6 mm from eornpliancelike
behaviorofZc occursin thesamefrequency
the basal end of the basilar membrane,the measuredpres- rangewhereIPg/Psllevelsoff(Fig.19}.Nedzelnitsky (1980}
suremightbe expectedto differfrom that at the stapesfor hasproposeda modelin whichZc is controlledby the im-
high frequencies;Nedzelnitsky(1974a, p. 348} infers on pedanceof the round-window membranein this frequency
theoreticalgroundsthat for frequencies below 1 kHz, Z• range.,
shouldnot differfrom the cochlearinput impedance{Zc } by The differences
betweenour averageIZcl and the
more than about2 dB in magnitudeand 10øin angle.For Tonndorfet al. (1966}averageare probablynot merelythe
frequenciesbelow 1 kHz the principaldifferencebetween resultofinteranimalvariations,becauseat somefrequencies
IZl of Nedzelnitsky
{1980}and IZcl measured
by usis a the rangeof variationfoundin eachseriesof measurements
frequency-independent
differenceof about6 dB, whichis is lessthan the differencein the averages.SincetheseTonn-
within the estimated error limits of the two sets of measure- dorf et al. (1966}motion measurements were made with a
120 J. Acoust.Soc.Am.,Vol.72, No. 1, July1982 Lynchet al.:Inputimpedance

capacitive
transducer
ontheroundwindow,thelimitations u) +90
on accuracypreviouslymentionedabovecouldaccountfor
the differences.
The magnitudeof cochlearinput impedancein guinea
pig has beeninferred from measurements of intracochlear '-9 '•
pressureand malleusmotionby Dancer and Franke (1980, <[ I0
I I I !llll I
I00
I I I Illll
I000
I I I I Illll
I0000
I
Fig. 2}.Theirresultsaresimilarto oursin that IZcl isrelati-

vely independentof frequency{+ 5 dB} over a broad fre-
.....
NEDZELNITSKY
(1980)
N=6
quencyrange.Also IZcl-0.45 x 10• dyn-s/cm
5 whichis
H DALLOS (1970) N=3
about one-half of our mid-frequencyvalue. The positive
slopeof IZcl vsf, whichwe haveseenfor 50 <f< 500 Hz
(Fig. 20), doesnot appearin the guineapig results,which is •_
consistent with the cochlearpotentialmeasurements of Dal-
o
los (1970). Further resultsfrom guinea pig (Franke and ½D -
Dancer, 1980,Fig. 5) indicatethat at frequenciesbelow 20

Hz, Zc behavesasa compliance,whichis alsosimilarto our o 0.1
results in the cat.
4. Inputimpedanceacrossthe cochlearpart/t/onZ•
If one considersthe stapesvolumevelocity Us as the FIG. 21. Data relevantto theinputimpedanceacrossthecochlearpartition
inputto theinnerearandthesoundpressure in thevestibule, Z •. The Nedzelnitsky{1980}curves{leftordinate}arebasedon the median
Pr as the response,thenZc = Pv/Us is the transferfunc- pressure difference
fromsixcats{hisFig. 15}.The Dallos{1970}curves{fight
ordinate}arebasedon mediansof differentiallyrecordedCM from thebasal
tion.On theotherhand,theinputto themechanical system turn of threecats{hisFig. 3}. In both of thesestudiesacousticstimuliwere
oftheinnerear(i.e.,.
thecochlearpartition}
ispresumably the deliveredto thetympanicmembrane. TheGuinanandPeake{1967,Fig. 14}
differencein pressurebetween
scalavestibuli
andscalatym- averagemiddle-eartransferfunctionwasusedto computethe ratiosof the
pressuredifferenceand CM to stapesvolume velocity.The vertical axes
paniat thebasalendofthecochlea,Pv - Paw• Pc, andthe
werepositionedto approximatelysuperimpose the plotteddata.
relevant transfer function is
z = - = P/Us, suredifference(Pc) at low frequencies.

The measurements of
whichwe will call the inputimpedance
across
the cochlear Dallos(1970}havebeenusedto plot the ratio of basalturn
partition. CM to stapes volumevelocityin Fig.21.4As pointedoutby
Measurements from thebasalturn of cat {Nedzelnitsky, Dallos {1974},the frequencydependence of theseCM mea-
1980)indicatethat for frequenciesabove100 Hz the sound surementsparallelsthe Nedzelnitsky{1974b}pressurere-
pressurein scalavestibuliis considerablygreater than the sultsquiteclosely.Theseresultsclearlysuggest themasslike
behavior.
soundpressurein scalatympaniand thereforethe pressure
differenceacrossthe cochlearpartition is approximately In guineapig,low-frequency
masslikebehaviorof Z •:
equalto the pressurein scalavestibuli.For this frequency is not indicatedby CM measurements{Dallos,1970};even
rangethe input impedanceZc •Z •. However,at frequen- forfrequenciesaslowas3 Hz (FrankeandDancer,1980}the
ciesbelow40 Hz the pressuresin scalavestibuliand scala angleof Z •: {thoughit is positive}doesnot approach90ø.
tympaniarenearlyequalandZc isprimarily determinedby
the round-windowmembraneimpedance{Nedzelnitsky, B. Network models for impedances
1. Goal
1980}.To obtainZ • the round-windowimpedancemustbe
subtracted,
i.e.,Z • = Zc -- Zi•w. {Thisapproach
hasbeen We have chosen mechanical network models to fit the
described
byGeislerandHubbard,1975.) impedancemeasurements of Sec.IV A. Thesenetworksde-
The measurements reportedheredo not permitus to scribethe mechanical(or acoustic}systemin terms of con-
determineZ •:, sincepressurewasnot usuallymeasuredin nected masses,resistances,and compliances.Since many
scalatympani.However,Nedzelnitsky{1980}hasreported middle-ear models (Onchi, 1949, 1961; Zwislocki, 1957,
measurements of the ratio of pressuredifferenceacrossthe 1962, 1963; Mq•ller, 1961}have beenpresentedas electric
cochlearpartitionto soundpressure at the tympanicmem- circuit analogsof the mechanicalsystem,we will also in-
brane,Pc ?PD. If this ratio is dividedby the cat middle-ear cludeelectriccircuitsin whichvoltageis analogousto sound
transferratio, Us?PD,an estimateof Z •: is obtained{Fig. pressureand current is analogousto acousticvolume veloc-
21}. As mentionedabove,the principaldifferencebetween ity. The two formsfor the networksarecompletelyequiva-
Z •: and Zc (Fig. 20} occursat frequenciesbelow 40 Hz lent: both are included as a convenience for readers with
whereZc is compliancelike
[i.e.,Zc •-l/(jcoCRw}],
while differentbackgrounds.
Z $ is morenearlymasslike{i.e.,Z • =•jcoMo}.
Masslikebehaviorfor Z •: at low frequencieswas im- 2. A simplenetworkfor the impedanceof the stapesand
pliedby studiesofcochlearpotentialsin thebasalturn of cats cochlea Zsc
{Dallos,1970;Weissetal., 1971},in whichit wasarguedthat The frequencydependence of the Zsc measurements
cochlearmicrophonic potential{CM}isproportionalto pres- canbeapproximated bya three-elementnetwork{Fig.22}in
121
J. Acoust. Sec. Am., Vol. 72, No. 1, July 1982 Lynch et aL' Input impedance of the cat cochlea 121
o
Zsc=Ps/Us more de ta//ed network
u, u, Althoughthe three-element networkof Fig. 22 is ade-

quatefor representing
thepropertiesof Zsc, it doesnotfully
Ps Ms
ø Rec representour knowledgeof its components. Zsc is the sum
oftheimpedance ofthestapes {andannularligament} Zs, the
Rs0
=1,4x IOs DYNE-SEC/CM
5 Msc
=2-5GRAM/CM
4 inputimpedance acrossthe ½ochlear partitionZ •, and the
Cs•=0.$6x I• e CMS/DYNE
•n +90 -'"'
round-window membrane impedance Zaw [Fig.23{a}or (b}].
Our goal in this sectionis to representeachof thesethree
• 0 ß impedances by a simplenetworkof lumpedelementssuch
that the overall network has characteristics that mimic all
._m_ 90 themeasurements.
In a followingsection{Sec.IV C}wewill
relate the network elements to anatomical entities.
< I0 I00 I000 I0000
First we must chooserepresentative setsof measure-
eee AVERAGED MEASUREMENTS
mentsto be matchedby the networkmodel.The averaged
• I00- •
---
SIMPLE MODEL (R$c,M$C, C$C)
MORE COMPLETE MODEL Zsc results{Fig. 22} are taken as one set. For Zc we have
combined ourmeasurements andthoseof Nedzelnitsky {Fig.
20}to givea grandaverage Zc thatisconsistent withtheZsc
o 10- set{Fig.24}.5
The networkusedto representthe averagedsetsof Zc
and Zsc is shownin Fig. 23 [{c}or {d}].After the circuit
topologyandelementtypeswerechosento providethe main
featuresin the Zc and Zsc curves,the elementvalueswere
'
IO '"'"'" IOO IOOOO " chosen as follows.
FREQUENCY (Hz)
Consider first the impedanceof the round-window
FIG. 22. Comparisonof our averagedmeasurements of acousticimpedance membraneZaw. Nedzelnitsky(1980)has concludedthat
of the stapesand cochleaZsc to the input impedanceof a three-element Z Rw behavesasa compliancefor frequencies
below0.3 kHz.
networkmodel.The averagedmeasurements are from Fig. 17. In Figs.22, Assuming that Zaw = 1/{/coCaw) we chose
23, and24 networksareshownin two equivalentformsin theupperpartof Caw= 10X 10-9 cmS/dyn
tofit IZc I forfrequencies
below
the figurewith a mechanicaldiagramon the left and an electriccircuiton
40 Hz whereZ c
theright.In bothnetworksthelabelsrepresent acousticvariablesandpara-
meters(Ps,Us; Rsc,Msc,Csc), which are analogousto either mechanical Considernext the impedanceof the stapesZs. The
quantities(force,velocity;resistance,
mass,compliance) or to electricquan- measurements
with perilymph removedfrom the cochlea
tities(voltage,current;resistance,
inductance,capacitance}. The arrowson {Fig. 11}indicatethat Zs is compliancelike
up to 1 kHz,
the complianceelementsindicatethe dependence of this elementvalueon resistivefor frequencies
between1 and 4 kHz, and masslike
staticpressuredifferenceacrossthestapesfootplate.The maximumvalueof
Csc is presumablyaboutthreetimeslargerthan the valueusedhere(Fig. at higherfrequencies.
Thisindicates
thatZ s canberepre-
10).The dashedlines(whichare visiblydifferentfromthe solidfor frequen- sentedbya three-element
networkof compliance,
resistance,
icesbetween50 and 1000 Hz) indicateZsc for the more completemodel andmass.The compliancefor our "normal"conditionmust
shownin Figs. 23 (c}and (d). be chosensothat whencombinedwith Caw the net com-
pliancewill equalCsc (i.e., I/CAL = 1/Csc-- 1/Caw},
whichthe compliance
valueCsc is chosento fit IZscl for whichyieldsC^L= 0.37X 10-9 cmS/dyn. Thestapes mass
frequenciesbelow0.1 kHz, the resistancevalueR sc is cho- M s is basedon theaverage
weightfor excised
stapes (0.52
sento fit IZscl for frequencies
between0.5 and7 kHz, and mg}yielding
anacoustic
mass
Ms = M •/,4 •p= 3.3g/cm4.
the massMsc is chosento fit for frequenciesabove10 kHz. Theresistance
R ^L mustbe0.2X 106dyn-s/cm 5toyieldthe
The root-mean-square {rms}differences {i.e.,the meanover minimumvalueof IZscl (e.g.,Fig. 11}.
all the frequenciesof the "AVERAGED MEASURE- Considernowtheelements necessary
to representZ
MENTS"} betweenmodeland averagedmeasurements are Thedominantelementat mid-frequencies
isRe, whichmust
1.2 dB in magnitudeand 12øin angle;the maximumdiffer- be chosen such that Rc = Rsc -- RAL = 1.2X 106
encesat any frequencyare 2.8 dB and 29ø. Insofar as this dyn-s/cm
5.To account
forthemasslike
behavior
ofZ • at
modelfits the Zsc measurements, it providesa satisfactory very low frequencies
(Fig. 21) a parallelmassMo mustbe
descriptionof Zsc which shouldbe adequatefor modeling included;its valuemustbechosento fit Zc for frequencies
the normal middle ear. between50and100Hz {Fig.24)yieldingMo = 2250g/cm4.
The elementsof thismodelcanbe associated primarily ThevalueforRoischosen to matchIgcl nearthefrequency
with eitherthe stapesZs or the cochleaZc. Sincewe know of itsminimum(40Hz). Sinceat highfrequencies {f> 4 kHz)
that the annularligamentcontrolsthe low-frequencybehav- Zs-•/coMs,Zsc•/coMsc, andZc•jcoMw,Mw•Msc-Ms
ior of Zsc (Figs.7 and 10},Csc mustrepresentprimarilythe -- 25 -- 3 = 22 g/cm4.
complianceof the annularligament.Becausedestructionof The behaviorof the resultingeight-element
network
the basalendof the basilarmembraneandremovalof peri- [Fig.23(c}or (d}]fitstheaveraged
ZscandZc (andP•,/Ps}
lymphreducesIZsclforfrequencies
above0.3kHz {Fig.11}, measurementsquitewell{Figs.22 and24, dashedlines}.The
R sc and Msc mustboth be primarily determinedby struc- rmsdifferences
are 1.2dB in magnitude
and 13øin anglefor
tures within the cochlea. Zsc and 1.2dB and 16øforZc (2.0dB and24øforP•,/Ps}.
122 J. Acoust. Soc. Am., Vol. 72, No. 1, July 1982 Lyncheta/.' Input impedance of the cat cochlea 122
MECHANICAL NETWORKS Zc:Z•,, ZRW ELECTRICAL NETWORKS
(a) Zsc:Zs,Zc (•)
US ZS
PsC • STAPES • ANNULAR'
LIGAMENT
FIG. 23. Networkrepresentationsof theimped-

anceof the stapesand cochlea.The uppernet-
works[(a)and (b)]indicatethe relationshipof
the three main componentimpedances.The
lower figures are the detailed networks that
(c) (d) have been used to fit the measurements. Ele-
Us CALMS RAL ment valuesare CA•.----0.37X10-9 cmS/dyn,
Ms = 3.3 g/cm4, RAL= 0.2X 106 dyn-s/cm5,
MV My = 22 g/cm4, Rc -- 1.2X 106 dyn-s/cm5,
Ro-- 0.28X 106dyn-s/cm 5,Mo -- 2250g/cm4,
CRW--' 10X 10-9 cmS/dyn.
Ro
Re
Mo
g I,
Zc=Pv/U
s The maximum differencesat any frequencyare 2.8 dB and
29øfor both Zsc and Zc (5.9 dB and 66øfor P•,/Pc).
Thereis an apparentdisagreement betweenbehaviorof
Us
• ^ Rc ..___.Ro the network model and the measurementsat high frequen-
ciesin that the angleof Zc for the network(Fig. 24, P•,/Us,
dashedline) is significantlymore positivethan the averaged
- CR
w CRW
measurements (Fig. 24, dots).This suggests that Me (which
was neededin addition to Ms to accountfor the masslike
Rc=l.2x IOe DYNE-S/CMs M0=2250 GRAM/CM4 behaviorof Zsc at high frequencies) shouldbe a component
Ro=O.28
x IOe DYNE-S/CM
5 My=22 GRAM/CM
4 of Zs ratherthan of Zc (whichwouldchangethe Zc model
• +90 CRw=
I0x10
.9 CMS/DYNE to the solidline in Fig. 24 with a reductionin rms differences
to 1.1 dB and 11 ø and maximum differences to 2.8 dB and
27ø).However,thissuggestionconflictswith the results(Fig.
11)whichshowthat IZscl is diminished at highfrequencies
' '90
by removalof perilymph.This conflictcanberesolvedif it is
assumed
thatthepressure
measured
in thevestibule
•,
< I0 I00 I000 I0000 differsfrom the actual pressureon the medial face of the
ß .. AVERAZED MEASUREMENTS footplateP•, (Ref. 6). Even thoughthesetwo locationsare
'- I0 - -- MODELPv/Us -- separated by onlya smallfractionof a wavelength(forsound
in water), there could be significantpressuredifferences.
z
Sincethe stapesfootplatecoversonly a fractionof the sur-
face of the vestibule,somenonuniformityin the soundfield
will exist within the vestibule. We assume that this nonuni-
ml'O-
•
• -
z <(
formitybecomessignificantat highfrequencies and contrib-
- utesa masscomponent(seeSec.IV C3). Thus M•, is inter-
pretedas belongingto the cochlearinput impedance,even
-• - o.i- _ thoughthe measuredZc doesnot appearto be masslikeat
I0 I00 I000 I0000
high frequencies.In effectwe assumethat for high frequen-
FREQUENCY (Hz) cies,Ps is a better estimateof the averagepressureon the
vestibularsideof the stapesP•,, than the measuredpressure
FIG. 24.Comparison
ofacoustic
impedance
ofthecochlea
Zc frommea-
surements
andnetworkmodel.The averagedmeasurements
arc thoseder-
ivedfromtheresultspresented
inFig.20(29catstotal).Thenetworks
inthe C. Implications for structures that contribute to stapes
upperpartof thefigurearethecochlearportionsof thoseshownin Figs.
impedance Z$ and cochlear impedance Zc
23(c)
and(d).Thesolid
curves
arethemodel
impedance
•,/Us, which
ex-
cludes
themass
My' thenetwork
impedance
Pv/Us (dashed
curves)
in- This discussionis organizedin termsof the elementsof
eludesMy. the network model of Fig. 23.
123 J. Acoust. Soc. Am., Vol. 72, No. 1, July 1982 Lynch eta/.: Input impedance of the cat cochlea 123
1. Stapesand annularligamentimpedanceZs Viidik, 1973,for review).Samplesfrom bovineligamentum
nuchae
showa compliance
increase
bya factorofalmost103
a. •Innularligamentcompliance
CAL.Our resultsindi- with an increasein water contentfrom 0% to 60% (Gotteet
cate that between 10 and 1000 Hz, Zs behavesas a com- al., 1968}.Our findingof changesby a factor of 30 with
pliance{Fig.11}withtheannularligamentbeingthestruc- partial drying(Fig. 7} is well within this range.
ture that controls this behavior. Quantitativecomparison of annularligamentelasticity
i. Effectsof staticperilymphaticpressure on middle-ear to that of other ligamentscan be attemptedusingresults
transmission. The demonstration{Fig. 10} that the incre- obtainedwith displacements that are smallenoughto yield
mentalvalueof Cî• is sensitiveto changesin the staticpres- linearbehavior.To comparethe differentmaterialsan elas-
sure differenceacrossthe footplatesuggests a mechanism tic modulus(i.e.,a characteristicof the ligamentmaterial
throughwhichpressure in the csf-perilymphsystemmight whichis independent of the ligamentshape}is used.Mea-
affect middle-ear transmission. Measurements in the cat surements of ligamentelasticpropertiesmadewith a tensile
havedemonstrated that changesin perilymphaticpressure stressin one direction yield values of Young's modulus
tendto reducethe amplitudeof cochlearpotentialswith the {E = stress/strain}
that varyovera considerable
range.For
largestchangesoccurringfor low-frequency stimuli {i.e., the anterior cruciateligament from the knee (of dogs}
f< 1 kHz} (Allenet al., 1971}.Thisfeatureof theseresultsis EA½= 109dyn/cm2(HautandLittle,1969},whereas forthe
qualitativelyin agreementwith our proposedmechanism, ligamenturn nuchae from the neck (of cattle}
sinceCî• controlsZsc onlyat low frequencies. It isdifficult ELN----10 7dyn/cm•- (Krafka, 1939;Cartonet al., 1962;
to makea morequantitativecomparison because the small- ApterandMarquez,1968;Yamada,1970}.ThevaluesforE
est pressureusedby Allen et al. is largerthan the largest are apparentlycorrelatedwith the relativecontentof the
pressurethat we used.However,it seemspossiblethat, at fibrousproteinscollagenandelastin.Ligamentsthat arere-
leastfor moderatestaticpressures in the cat, a decrease of lativelycompliantcontainmainlyelastinandthosethat are
incrementalannular-ligament complianceis the causeof a stifferarelargelycollagen
(Hardy,1951}.Forindividualcol-
middle-eartransmissionlosswhich secondarilycausesa re- lagenfibersE = 109to 1010dyn/cm
2 (Harkness,
1961,p.
ductionof cochlearpotential,ratherthanothermechanisms 428;Stromberg andWiederhielm,1969},whereas forelastin
that havebeenproposed{Allenet al., 1971,p. 393}. fibersE = 7X 106dyn/cm•-(Cartonetal., 1962}.
The proposedmechanismmay alsohavesignificance To estimate an elastic modulus for the material of the
for clinicalstudiesthat havedemonstratedheatinglossasso- annularligamentwe mustmakesomeassumptions about
ciatedwith increasedintracranialpressure{Saxenaet al., ligamentconfigurationandstapesmotion.If weassume{a}
1969}.Followingsurgeryto reducethe pressure, significant thatthedimensionsof theligamentarethesameastheannu-
improvementin heatingoccurredonly for low frequencies lar space
betweentheedgeofthefootplate andtheovalwin-
{Johnetal., 1979}.Predictionsof theinfluenceof thismecha- dow,and{b}thatstapes
motion
ispurely
translational
ina
nismin the intact ear are difficultbecauseknowledgeof the directionperpendicularto the planeof the footplate,then
static strain of the annular ligamentis lacking.The incus theannularligamentissubjected to a pureshearstress.
If we
probablytransmitssomestaticforceto thestapeswhichin- furtherassume thatthismaterialishomogeneous, 7isotrop-
teractswith the perilymphaticpressureto determinethe ic, and relativelyincompressible,
the compliancecanbe ex-
staticstrainof the annularligament. pressed
sin termsofthedimensions
oftheannularspace
and
ii. Comparison of dynamicand staticcompliance. Kiri- Young'smodulusEAi•'
kae {1959,p. 90} hasreportedmeasurements of staticdis-
placementof the stapesin excisedtemporalbonesof cats.
where we have assumed that the thickness t and width to of
Theseresults
suggest
anacoustic
compliance
of0.16X 10-9
cmS/dyn.Thisvalueis aboutone-halfof ouraveragevalue the annularspaceare uniformoverthe wholeperimeterpof
for Cî• andaboutone-sixthof our valuewith no staticpres- the ovalwindow.From Eq. (8),with dimensions
to = 20/•m,
sure differenceacrossthe stapesfootplate.However, the t = 0.2 mm, p = 4 mm (seeGuinan and Peake, 1967, Fig.
combinationof the way in which Kirikae's mechanicalload 11),Afp= 1.26mm•, andCAi•= 10-9 cmS/dyn
(where
we
wasapplied,thepost-mortemstateof theligaments,andthe assumethat the unstressedCAi• is three timesthe valuecho-
ratherlargedisplacement magnitudeusedcouldeasilymake senfor thenetwork},
we obtainEAi• = 105dyn/cm2.This
the complianceestimatesfrom his measurements smaller resultindicatesthat if the materialwhich controlsthis com-
than our values.It seemspossiblethat the differences
arenot pliancehasthe configurationof the annularspace,the mate-
significantand that the dynamicand staticcompliances of rial is more compliantthan anterior cruciateligamentby a
the annular ligament are equal for stapesdisplacements factorof 104andmorecompliant thanligamenturnnuchae
within the rangeassociated with normalacousticstimuli. by a factorof 102.
iii. Comparison
tomechanical
properties•
ofotherliga- It seemslikelythat erroneousassumptions
maycontri-
ments.Two propertiesof Cî• are similarto thoseof other butesignificantly to thelargediscrepancybetweenapparent
ligaments,at leastqualitatively:{1}Many kindsof connec- EAi• andE for otherligaments.Certainlythe fibrousstruc-
tive tissue have nonlinear elastic behavior in which the incre- tureoftheligamentwouldtendto makeitselasticproperties
mental compliancedecreases with increasingstrain {e.g., anisotropic. Probablymoreimportantisthelikelihoodthat
Fung, 1967}.(2}Stiffeningof ligamentsand tendonsasa re- the mechanicallysignificantcomponentsof the ligament
sultof dryingis well knownfrom in vitromeasurements {see maynotbeuniformlydistributed;theymaynot evenoccupy
124 J. Acoust.Soc. Am., Vol. 72, No. 1, July 1982 Lyncheta/.' Input impedanceof the cat cochlea 124
the annularspace.Studieswith specific
stainsindicatethat 3. Inputimpedanceacrossthe cochlearpartitionZ•
the elastictissuein humanannularligamentis primarilyon Some of the theoretical treatments of cochlear dyna-
the vestibularand middle-earsurfacesof the footplate and
mics{Zwislocki,1948;Steele,1974;Geislerand Hubbard,
extends on the surfaces of the bone for some distance from
1972}predictthat Z • is resistiveand constant{for some
theannulus(Davies,1948;Harty, 1953;WolffandBellucci,
rangeoffrequency between 0.1and10kHz}sothattheyare,
1956).Theseinhomoõeneities in theconfiguration of thelig- in this respect,essentiallyconsistentwith our measure-
amenthavenot beendescribed completelyenoughto be use- ments. Other theoretical results have rather different behav-
ful in makingquantitative estimates of elasticmoduli.How-
ior {Bogert,1951;Fletcher,1951;Dallos,1970;Steele,1974;
ever, it seemsclear that if this configurationof the elastic
Sondhi,1978}.{Mostof theseanalyses weremadeto inter-
tissueoccursin cat, the effectivedimensionsof the control-
pretresultsobtainedonspecies otherthanthecat.}It should
ling structurewouldbe muchdifferentfrom thoseof the
bepossibleto relatetheelementsof ourdescriptivenetwork
annularspace[i.e.,in Eq. {8)w wouldbemuchlargerandt
modelto propertiesof cochlearstructuresthroughthese
smaller]andtheactualmodulus,EAL couldbemuchlarger models.In mostof the studiesof cochleardynamicsthe rela-
than that calculatedabove.Thus more precisedetermina-
tionsarenotevidentuponinspection because computational
tion of the configuration
and composition of the annular
methodswereusedto obtainsolutionsfor a particularsetof
ligamentis requiredto makea meaningful comparison of
valuesfor cochlearparameters. However,Zwislocki's(1948,
annularligamentpropertiesto thoseof otherligaments.
1965}analysisdoesyielda simple{approximate} expression
b. Annular ligamentresistance
RAL. For frequencies
between1 and 4 kHz, Zs is apparentlyresistive{Fig. 11}.
for the dependence of Z • on cochlearproperties[seealso
Sondhi(1981}].
Althoughwe haveno directevidenceas to the structures
involvedin this effect,it seemslikely that the annular liga- a. Mid-frequencies,
Z • •tlc. The resultthatZ • is ap-
mentis theprimarycontributor.Apparently,properties
of proximated by a constantresistance
overa ratherbroadfre-
otherligamentshavenot beenmeasured at frequenciesin quency range{0.1< f< 3 kHz}isconsistentwithsomerather
different views of cochlear mechanics.
thisrange(Apterand Marquez, 1968}.
c.StapesmassMs. We havemeasuredan averagestapes It is sometimessuggested that theinnerearhasa high
mass
of0.52mg,whichistheequivalent
ofanacoustic
mass inputimpedance because it is filledwith a fluidhavingthe
of 3.3g/cm4.Becausethisisconsiderably
lessthantheequi- acoustic properties
of water;theacoustic impedance isthen
assumedto be that which would resultfrom a uniform plane-
valentmassofthestapesandcochlea (Msc= 25g/cm4,Fig.
22},weconcludethat responses
of thenormalmiddleearin wavepropagating withoutreflectionin an infinitetube(see
catsshouldnot be sensitiveto small changesin stapesmass. Schubert,1978,pp.45-49, for somediscussion of thisissue}.
In thiscase
Rc = pc/,4fp,
wherep
andcare,respectively,
the
massdensityandspeedof soundfor thecochlearfluid.Com-
pellingtheoretical
reasonshavebeenpointedoutfor reject-
2. Round windowimpedance ZRW
ing thispointof view(e.g.,WeverandLawrence,1954,p.
381; Von Gierke, 1958;Killion and Dallos, 1979}.First,
Nedzelnitsky's(1980)resultsindicatethat the imped- sincethe cochleais enclosedby bonywallsgivingit dimen-
anceof the round-windowmembranecan be representedby
sionsthat {forthe frequencies
of interest}
are onlya small
a compliancefor frequenciesbetween20 and 300 Hz and the
fractionof a wavelengthof soundin water,it shouldnot
resultsreportedhere(Fig. 11} supportthisconclusion.How- behaveas an infinite tube. Second,in theoretical treatments
ever,this representation
may not be accuratefor higherfre-
of cochleardynamics(e.g.,Zwislocki,1965;seeGeisler,
quencies{Nedzelnitsky,1974a}.For lower frequencieswe
1977,for review}the cochlearpartitionplaysa key rolein
canuseKirikae's(1959,p. 89}measurements of staticdeflec-
determiningpressures and velocitiesin the cochlea,and
tion of the round-windowmembranein cat temporalbones thereforethe mechanicalpropertiesof the partitionshould
to compute
a compliance
of9 X 10-9 cmS/dyn,
whichisap- havean importantinfluenceon the input impedance.Our
proximatelythe valuewe haveobtained.Perhapsthe round- resultsadd experimentalevidencethat conflictswith the
window membranecan be representedby a frequency-inde-
viewthatRc = pc/,4fpin tworespects:
(a)thevaluewefind
pendentcompliancefor all frequencies
below 300 Hz.
for Rc is approximately
one-tenth
of pC/Alp----1.2X
107
The visualappearance of theround-windowmembrane dyn-s/cm5 and {b} obliterationof the basilarmembrane
frequentlychangedduringour experiments. The changesin whileleavingthefluidin thecochlea
makesa largechangein
IZcl at lowfrequencies
observed
in oneexperiment
{Fig.16} IZscl{Fig.11}.Thus,althoughtheconceptionthatthecoch-
can be interpreted as a decreasein Caw. Nedzelnitsky learinputimpedance results
fromthewaveimpedance ofthe
{1974a, p. 151} also observedchangesduring some of his cochlearfluid alonedoespredicta constantresistance
for
experiments
that couldbecausedby a decrease in Caw. Re- Z •, the conceptionis demonstrably incorrect.
movalof periosteumfrom the surfaceof the cochleacould Thelong-wave treatmentofcochlear dynamicsof Zwis-
affect the vascular circulation to the outer round-window
locki{1965}yieldstheresultthatfor a broadfrequencyrange
membranewith resultantchangesin its mechanicalproper- theacoustic
impedance
at a distance
x fromthebasalendof
ties.Suchchanges
couldeasilyhaveoccurredwithoutnoti- the cochleais closelyapproximatedby
ceableeffect on our measurements,since the round-window
impedancenormally contributeslittle to Zsc and only in- Z(x) = [p/[S( x)C( x)]]'/•', (9)
fluencesZc at very low frequencies. whereC { x) istheacousticcompliance
perunit lengthof the
125 J. Acoust. Soc. Am., Vol. 72, No. 1, July 1982 Lynch et al.' Input impedance of the cat cochlea 125
basilarmembraneat the locationx, and $ (x) is an effective parametersvaryappreciablyeitheracrossspecies or through
area resultingfrom the areasof scalatympaniSt and scala experimentalmanipulation.
vestibuliSo i.e., b. High frequencies,Z •--•jcoM•.. At high frequencies
thebehaviorofZ • isapproximatelythat of an acousticmass
S= •/(• =4-•), (10) My=22 g/cm4.Zwislocki{
1962,p. 1520}hassuggested
that
a masscomponentin the cochlearimpedancecould result
whereboth areasare evaluatedat the positionx. Zwislocki
from "the column of perilymphbetweenthe oval window
(1975,p. 46) arõuedthat the availabledatafrom human,cat,
and the cochleaproper," and our interpretationof the Zc
and õuineapiõ were consistentwith his theoreticalimped-
andZsc measurements
ance.Recentlyfurther resultshavebeenreportedwhich al- (Sec.IV B3}suggests thatMy results
low computationof the variablesin this expression. from the fluid near the footplate.If it is assumedthat this
In the cat, measurements
of intracochlear
pressure massresultsfrom a cylindricalplug of perilymphwith the
(Nedzelnitsky,
1980)andbasilarmembranedisplacement cross-sectional area of scala vestibuli at the basal end
(EvansandWilson,1975) at theap- (•1• = 2 mm2,Dallos,1970,Fig.10),thecalculated
9 havebeenreported lengthof
proximatelocationx = 6 mm.Ratiosof thesemeasurements, thisplug{•1•.M•./p = 4 mm}ismuchlargerthanthedistance
(at 1 kHz)yield a "compliance" of 4X 10-9 cm•/dyn.To from the footplateto the basalendof the basilarmembrane
convertthis to an acousticcomplianceper unit lenõthone {• 1 mm}. It is conceivablethat becauseof nonuniformve-
must multiply by the effectivewidth of the basilar mem- locity distributionin the vestibulean additionalmasscom-
brane:Theanatomical widthat thislocationis 200/•m (Ca- ponentexistsat high frequenices.Sincethe stapesdrivesa
bezudo, 1978). If we arbitrarily assumethat the effective volume{thevestibule}which is largerin crosssectionthan
widthis half thisvalue,thenC(6 ram)= 40X 10-•= cm•/ the footplate,a massloadingoccurs{Burkhardand Sachs,
dyn. The area of scala vestibuli at this location is about 1977;Ingard, 1948}.Becauseof the complicatedshapeof the
7X 10- • cm=(Dallos,1970)andscalatympaniissomewhat vestibule it is difficultto estimatethis effectquantitatively,
larõer,sowe(somewhat arbitrarily)let$ (6 ram)= 5X 10-• but rough approximationsindicate that this mechanism
cm=.Withp = 1g/cm• thesevalues õiveanacoustic imped- could introduce a masseffectof the right magnitude.
anceof 1.6X 10• dyn-s/cm•. Thus,with theseestimates of c. Low frequencies,Z • --•jcoMo+ Ro. In our network
cochlearparameters,Eq. (9) õivesan impedanceat x = 6 model(Fig. 24) the pathwaythroughRo and Mo providesa
mm thatisabout1.3timesourRe. Onemiõhttry to evaluate low impedancefor volumevelocityfrom the ovalwindowto
Eq. (9)at x = 0 to findZ at the stapesfootplate.Becausethe the round windowat low frequencies (f< 100Hz). In discus-
anatomyof the cochleain the mostbasalreõionis quitedif- sions of cochlear mechanics it is usually assumedthat the
ferentfrom the confiõurationusuallyassumedin the theo- helicotremaprovidessucha pathway.A simpleviewof low-
reticaldevelopments, it maynot be usefulto applythe for- frequencycochleardynamicsmight be that the volumeve-
mulain thisway.Because(a)the volumevelocityof thebasal locity of the stapesis approximatelyequal to the volume
6 mm of the basilar membrane is small relative to the volume velocitythroughthehelicotremawith thevolumevelocityof
velocityof the footplate(exceptat hiõhfrequencies)
and (b) the cochlearpartitionbeingrelativelysmall.In thiscasethe
the perilymphaticscalaein thisreõionaremuchlarõerthan perilymphwouldflowprimarilythrougha tubeconsisting of
in moreapicalreõions,the acousticimpedanceat the foot- scalavestibuli,the helicotrema,and scalatympani.(A simi-
platefor frequenciesbelow1kHz miõhtbeaboutthe sameas lar picture for low frequenciesis indicatedby Geislerand
that at the 6 mm position[astheoreticalestimatesof Nedzel- Hubbard, 1972).The considerations and calculationsthat
nitsky(1974b,p. 348)suõõested]. Thuswe concludethat the follow are intendedto test whetherthis view is compatible
variousmechanicalmeasurements in thebasalreõionof the with the network elementvaluesrequiredfor Ro and Mo.
cat cochleaare not inconsistentwith Eq. (9) evaluatedat Dallos (1970)considered variousaspectsofcochlearan-
x= 6mm. - atomy which might affectinput impedanceat low frequen-
Resultsfrom mechanicalmeasurements on guineapigs ciesandheproposeda networkmodelfor Zc that isidentical
can alsobe comparedwith evaluationsof Eq. (9). From the in topologyto the onethat we haveusedfor Z • (Ref. 10).We
basilar-membrane-displacement measurementsof Wilson have,however,chosenquitedifferentelementvaluesandour
and Johnstone(1975, Fig.7), the pressuremeasurements of interpretationof the relative roles of the helicotremaand
Dancer and Franke (1980, Fig. 2), and the basilar mem- perilymphaticscalaeis basicallydifferent.Dallos (1970)as-
brane-widthmeasurements of Fern•mdez(1952),onecanes- sumedthat for low frequenciesthe cochlearinput imped-
timate the compliance of the basilar membrane, C(2 ance is dominated by the impedanceof the helicotrema.
mm)= 40X 10-• cmn/dyn.With S(2 mm)= 0.65 mm• However, from our observationsof unsectionedcat cochleas
(Fern•mdez,
1952)Eq. (9) yieldsZ = 1.0X 10• dyn-s/cm
•' (embeddedin eponand with the bonycapsuleremoved)un-
about two times the value reportedby Dancer and Franke der a dissectingmicroscope, the helicotremadoesnot appear
(1980,Fig. 2). Againthepredictionsof Eq. (9)arelargerthan to bea dominantconstrictionin thispath;thecross-sectional
the experimentally determinedIZscI. However,the impre- area of scala tympani in the most apical half-turn is not
cisionsinvolvedin the measurementsandcomputations are greatlydifferentfrom that of the helicotrema.Also, the mea-
largeenoughthat wecannotconcludethat Eq. (9)isinconsis- surements reportedby Dallos(1970,Fig. 10)showtheareaof
tent with the measurements. scalavestibulioverits upperhalf to be only abouttwo times
A bettertestof the validityof Eq. (9)wouldbeto obtain the helicotremaarea.Thus it seemslikely that the imped-
accurate measurements in situations in which the cochlear anceof this pathway can receivesignificantcontributions

ofthecatcochlea 126
/
I I I I IllIll I I I IllIll I I I IIII1| I I I. lllll! I I

from the perilymphaticscalae.
From the dimensions of the helicotrema and scalae it
shouldbepossible to estimatethenetimpedanceof thispath-
way. Unfortunately, the anatomicaldata are incomplete,
120 - • • BEHAVIORAL
{ßß
CAT
(MILLER
et
al.,
EVANS,
1974)
1965) THRESHOLDS • HUMAN (YEOWART 8[
-- LYNCH et al.,
and the lack of publishedmeasurementsof scala tympani I10- ESTIMATES
area is particularly bothersome,sincethe apical region of OF PD/Pc xx BALLOS

(19TO)
•oo-
• leol
for
ICMI
=50nV
scalatympaniis apparentlythe mostconstrictedpart of the
path. If for a roughapproximationwe assumea circular tube 90-
witha radiusof 125/•m,TM
thevalues
ofRoandMothatfitour
measurements
both requirea tubelengthof approximately7 80-
mm.•2In thecata distance
of 7 mmfromtheapicalendof
the basilar membraneencompasses the upper 1 and 1/2 70-
turns of scalatympani (Schuknecht,1960).Although per- 60-
hapsonly the upper half-turn of scalatympani has a cross-
sectionalarea this small, the remainder of the pathway 50-
x
shouldalsoprovidesignificantadditionsto the impedance.

40-
Thus the impedanceof the perilymphatictubecouldbe ap-
proximatelyequal to R o +icomo. These calculationsindi-
30-
catethat the valuesof M oandR ousedin our networkmodel
couldbe consistentwith a pictureof flow at low frequencies 20-
whichis primarily controlledby the perilymphatictube.
I0-
O-
-IO-
D. Relationship of peripheral mechanical system to the I ' ' ' ' ''"1 ' ' ' '""1 ' ' • •''"1 ' ' • ' ''"1 ' '
threshold of hearing I IO IOO IOOO IOOOO
FREQUENCY (Hz)
The frequency-dependent transferfunctionsof the out-
er and middle ear and of the mechanicalsystemof the inner
ear must influencethe frequencydependenceof the beha- FIG. 25. Comparisonof the frequencydependence of behavioralthreshold
and sound pressureat the input to the cochlea.Two sets of behavioral
vioralthresholdfor tones.Rathercontradictoryconclusions threshold measurements are summarized. Cat behavioral threshold data
havebeenreachedon the relativeimportanceof thesecom- (monaural,Miller et al., 1963,Table 3, p. 22}weremodifiedby the transfor-
ponents(Guinanand Peake,1967,p. 1258;Dallos, 1973,p. mation from free-fieldsoundpressureto soundpressureoutsidethe tym-
126;Zwislocki,1975,p. 54; Khannaand Tonndorf, 1977; panicmembrane(Weineret al., 1965,Fig. 7}, sothat the plottedpointsre-
presentthe soundpressureat the tympanicmembrane(Po} at behavioral
Franke and Dancer, 1980).In Fig. 25 physiologicaland be-
threshold.The dashedline segments extendingfrom 2 to 100Hz summarize
havioraldataare plottedtogethersothat the frequencyde- resultsfor humansubjectsreportedby Yeowart andEvans(1974};their bin-
pendenceof behavioralthresholdcan be comparedto rel- aural curvehasbeenraised3 dB to approximatethe monauralthreshold.
evantphysiologicalvariables. Two estimatesof Po/Pc (Pc = pressureacrossthebasalendof thecochlear
The behavioral threshold measurements for the cat are partition}from physiologicaldata are plotted. Our curve was obtainedby
combiningthe averagedZ c = Pv/Us of Fig. 24 (usingZsc for Zc at the
plottedin termsof the sound-pressure levelat the tympanic highest frequencies}with the ½1osed-bulla middle-ear transfer function
membranePo. Sincewe wouldlike to makecomparisons at Us/Po from Fig. 21 of GuinanandPeake(1967}to givePo/Pv. The Dallos
frequencies below thoseat which the cat behavioialthre- (1970}data weretakenfrom measurements of cochlearmicrophonicpoten-
sholdshavebeendetermined,a summarydescriptionof hu- tial corrected(with Fig. 20 of Guinan and Peake, 1967}for the effectof
openingthebullaandbonysepturn.The verticalpositionsof the two physio-
man low-frequency behavioral threshold measurements logicalmeasurements were chosento make the pointsat 1 kHz coincide
(Yeowart and Evans, 1974)is included. with the behavioralthreshold.As a result,the IPol for IeMI curveis for
From physiologicaldata we havecomputed(for the in- IeMI -- 50 nV, andthe IP•l for IP•l curveisfor IP•l - 28 dB SPL.
tact middle ear) two estimatesof the ratio Po/Pc, where
Pc --Pv- Pr is the sound-pressure differenceacrossthe
basal end of the cochlearpartition. Our curve is basedon
mechanical measurements. The computation, Below1 kHz it seemsclearthattheslopes of thebeha-
Po/Pc -- 1/[ Z •(Us/Po)], wascarriedoutusingmeasure- vioraland physiological resultsdiffer.•3 For frequencies
ments of the middle-ear transferfunction Us/Po and the between20and100Hz thephysiological resultsbehaveap-
averagedZc curve (Fig. 24), sinceZc •Z •. The Dallos proximatelyasf-2, whereasthebehavioralthresholddata
(1970) curve is basedon cochlearmicrophonicpotential (twopointsfor thecatandthelinefor human}arecloserto
(CM) measurements with the assumptionthat CM is propor- f-3. Thus,asfrequency
is lowered
below1 kHz, Pc at
tionalto Pc (DancerandFranke, 1980).Thesetwo estimates threshold
increases;if weassume anextrapolation
of thecat
of Po/Pc havethe samefrequencydependence for frequen- thresholdparallelto the linesuggested
for human,Pc at
ciesbetween30 and 2000 Hz, thus supportingthe assump- behavioralthresholdwill be38 dB largerat 20 Hz thanat 1
tion of proportionalityof CM andPc for low frequencies. kHz.
127 J. Acoust.Soc. Am., Vol. 72, No. 1, July 1982 Lyncheta/.: Input impedanceof the cat cochlea 127
For frequencies
between1 and 8 kHz the behavioral port from the MIT Whitaker Health SciencesFund and
and physiological
data haveapproximately the samefre- iNIH Training Grants.
quencydependence.[Thesharppeakat 4 kHz iscaused
bya
resonancein the middle-earcavities{Guinanand Peake,
1967}.]Thusbehavioralthresholdapparentlycorresponds lit iseasilydemonstrated
thatthenetforceexerted
bythepressure
Psact-
to constant(i.e.,within 3 dB} sound-pressure differenceat ingoverthemiddle-ear
surface
ofthestapes
is•4fpps.
Consider
theequilib-
riumcondition
obtained
(neglecting
gravitational
forces)
withthestapes
the input to the cochleain thisfrequencyrange.
subjected
toauniform pressurepsoveritsentire
surface.
(Assume thatthe
At the high frequencies (i.e., 10 kHz and above}it is netforceactingonthesmallfraction of thesurface
thatabutstheoval
difficult to arrive at firm conclusions on the basisof available window isnegligible
in thissituation.)
Thesurfaceintegral
of theforce
data.Oneproblemistherelativelackof validdata.A second acting
onthestapes canthenberesolved intotwocomponents resulting
frompressure(1)onthevestibularsurfaceofthefootplate
and(2)onthe
problemresultsfromthe uncertainties associated with the middle-earsurface
ofthestapes.Sincethestapesisin equilibrium,thenet
measurements of diffractionby the cat'sheadat thesefre- forceexertedonthemiddle-earsurfaceisequalandopposite to thatexert-
quencies. Stillanotherproblemresultsfromourinterpreta- edon thevestibular
surface,
i.e.,•4rpPs.
tionof theapparentincrease in IZscl at highfrequencies. If 2Althoughmeasurementsreportedhere(Fig.8)andthose
ofNedzelnitsky
thisincreaseresultsfrom an impedancecomponentcontrib- (1980)haveindicated
thatintracochlearpressuresgrowlinearlywith
stimulus
sound-pressure
level,neither
studyhasinvestigated
thepromin-
utedby the perilymphnearthe ovalwindow,someof the enceof distortioncomponents
of the kindsreportedby Kemp(1978,
associated pressure dropwill not contributeto thepressure 1979a,b}or Kim et al. {1980}.
differenceacrossthe cochlearpartition,anda component of 3Wedonothavedatathatdirectlydisagree
withthereported
dependence
of
Z • shouldbe removedin calculations of Po/Pc at high IZsclonsound-pressure
level(Khanna
andTonndorf,
1971,Fig.10)be-
cause
wemadenomeasurements
at thehighSPL'sthattheyused(140-160
frequencies. Because of theuncertainties involved, wehave dBSPLat thestapes).
However,
theresults
ofNedzelnitsky
(1980,Fig.9)
not attemptedto resolvethis issue. demonstrate lineargrowthof intracochlear
soundpressure
upto compar-
In summary,the ideathat "behavioralsensitivity...(is} ablelevels(ina catwithintactossicular
chain}.
4TheDallos(1970)measurements madewithdifferential
electrodes
and
determined largelybytheproperties ofthemiddleear"(Dal- those
ofWeissetal. (1971
}frommicropipets
inscalamediaareinessential
los,1973,p. 126}apparently holdsfor a restricted frequency agreement.The cochlearpotential
measurements of Tonndorf and
range(perhaps 1to 10kHz forthecat}.Theresults in Fig.25 Khanna (1967)
froma round-windowelectrode exhibit
a largerpositive
clearlyindicatethatthebehavioral threshold at lowfrequen- slope
atlowfrequencies
thantheotherdata;thisdiscrepancy
couldresult
froma significant
neuralcomponentin theirround-windowrecords.
ciesis significantly influencedby morecentralmechanisms. 5Therationale
andprocedureforobtainingthegrandaverageZc isasfol-
Becausethe minimumcharacteristic frequencyfor auditory lows.We thinkthatZc isnearlyequalto Zc forf> 0.3kHz. Thereisa
nervefibersin the catmaybeapproximately 100Hz (Liber- problemindeciding
precisely
howclosetheratioIZc/Zscl= [Pv/Pslis
man, 1978},onewouMexpectthresholdto increase below tounitybecause
themeasured
pressure
ratiodependsontheabsolute
cali-
100Hz. Zwislocki(1975}hasconsidered othermechanisms bration
accuracy
oftwotransducers.
However,
wecanobtainanestimate
fromthemeasurements
of Zsc beforeandafterremovalof basilarmem-
thatmightbeinvolved.Sinceavailable evidence is quitein- braneandcochlear fluid(Fig. 11).For the regionwherethe angleof
complete, it is difficultto quantifytheimportance of differ- ZscnO (i.e.,1< f< 4 kHz),theaverage IZscl- 1.4M/2;measurements
ent mechanisms. withperilymph
removed
(averaged
overtwocats},
indicate
thatIZsclhas
To determine
definitively
whyheating
sensitivity
de- a valueofabout0.2M/2 inthisfrequency
region.
Thesefigures
leadtothe
creases
at low andhighfrequencies
it isnecessary
to obtain conclusionthatIPv/Psl- IZc/Zsclis(1.4- 0.2)/1.4= 0.86or - 1.3
dBinthisfrequencyrange.However,
inouraveraged measurements (Fig.
accuratemeasurements outsidethe frequencyrangethat is 19)IP•/Pslisabout4-2dBinthisrange.
Thissmall
(i.e.,3.3dB}inconsis-
usuallythoughtof as "important"[asYeowartand Evans tencybetween
theZscandZc measurements
probably
results
fromthe
{1974}havedone].Throughunderstanding
of themechan- combinedinaccuracies
of the measurements.
Becausewe havethe least
ismsthat contributeto the lossof heatingsensitivityat the confidence
inthePv absolutecalibration,
wechose, somewhat arbitrarily,
tomakeoneadjustment in theabsolutelevelof thesemeasurements.
We
extremefrequencies,we may be able to understand how assumed thattheaverage ]Pv]is toohighby 3.3dB andaccordingly
morphologicalvariationsarerelatedto thefrequency
range loweredtheaverage
IZcl (solid
curve,seeFig.20)bythisamount.Wealso
ofhearingfordifferent
species
in bothnormalandpathologi- raisedtheNedzelnitsky
curveby 2.5 dB to put it in agreement
(onthe
cal states. average}
withourshiftedresults
forf< 1kHz.Thegrand average(magni-
tudeandangle}
ofthetwosetsofmeasurementsisthencomputed weight-
ingeachaccordingto thenumber ofcatsinvolved.
Becausetheoretical
ACKNOWLEDGMENTS estimates
suggestthattheNedzelnitskyresults
differsignificantly
from
thecorrect
inputimpedance
forhighfrequencies
(seeNedzelnitsky,
1974a,
ProfessorL. Grodzinsof the MIT PhysicsDepartment p. 323),theywereusedonlyforf<0.7 kHz.
6Section
I defined
Pv asthepressure
acting
overthevestibular
surface
of
providedhelpfuladviceon M/Sssbauer techniques.S. M. Li-
thefootplate.
However, succeeding
sections
haveusedPv forthemea-
bermandevelopedsurgicaltechniquesand madenumerous suredpressure
inthevestibule.
Pvisintroduced
heretodenotemeasured
other contributions. Others who have contributed are D. G. pressure
andtoclarifytheissues
concerning
My. Thisdistinction
isun-
Beil, J. E. Bell, J. Cross, D. H. Johnson,M. C. Liberman, E. necessary
forotherdiscussions
inthispaper,
andPv isusedformeasured
pressurein followingsections.
M. Marr, L. P. Miller, C. R. Northrup, W. M. Rabinowitz, 7Inthecattheannularspacecontains nofluid-filledarticularcavities(Bolz
M.D. Silverstein,and J. S. Wiley, III. Helpful commentson and Lim, 1972)and it is apparentlyfilledwith fibers(WolffandBellucci,
the manuscriptwerereceivedfrom J. J. Guinan,Jr., N.Y. S. 19S6).
8Equation
(8)canbe obtainedsimply.A pressure
P actingon the stapes
Kiang, W. M. Rabinowitz,J. J. Rosowski,and T. F. Weiss.
produces
a shearstress
P•4fp/(tp).
If thelineardisplacement
isx, theshear
This work wassupportedin part by the National Insti- strainisx/w (assuming
x<w). Thustheshearmodulus(shearstress/shear
tutes of Health and in part by the ResearchFund of the strain)isn = wP•4fp/(tpx).
Theacoustic
compliance,
C^• = x•4fp/P,can
American OtologiealSociety.T. J. Lynch, III receivedsup- beexpressed
interms
ofn asC^•= w•4f2p/(tpn).
Forincompressible,
iso-
128 J. Acoust.Soc. Am., Vol. 72, No. 1, July1982 Lynchet al.: Inputimpedanceof the cat cochlea 128
tropic,homogeneous
materialsE is 3n (Fung,1965,p. 131). rent basilar membraneand singlenerve fiber measurements,"Science
9There
maybeaproblem
withthese
EvansandWilsonmeasurements
since 190, 1218-1221.
theyappearto showthat the basilar-membrane displacementis relatively FernJndez,C. (1982)."Dimensionsof the cochlea(guineapig)," J. Acoust.
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thetotalinputim- intracochlearsoundpressuremeasurements at low frequenciesin guinea
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130 J. Acoust. Soc. Am., Vol. 72, No. 1, July 1982 Lyncheta/.' Inputimpedanceof the cat cochlea 130

Input Impedance of The Cochlea in Cat

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Input impedance of the cochlea in cat

ThomasJ. Lynch,Ill,b) Victor Nedzelnitsky,

(Received24 June 1981;acceptedfor publication17 March 1982}

INTRODUCTION soundpressurehavebeencombinedwith measurements of

alA portionof thisworkwassubmitted

+z (2) tachedto thestapes andtherateat whichphotons aredetected througha

Usually after this portion of the procedurehad been • OUTPUT:

C. Stimulus generation and measurement

tor, poweramplifier,and eithera JensenDD- 100or an Atlas VELOCITY

PD-60 driver coupledto the stapescavitywith rigid plastic v(t)

fractionaleffect= (Roo-- Rr }/R oo, where

F = linewidth(or half-widthat half-height).

I I I IIII1! I I I IIII1! I I I IIII1! I

10- iP,i (ds SPL)

114 J. Acoust.Soc.Am.,Vol.72, No. 1, July1982 Lyncheta/.' Inputimpedance

I I I IIIII I I I I IIIII] I I I IIII1]

In two preparations(with similar results)more drastic frequencydependence isessentially

116 J. Acoust.Soc.Am.,Vol.72, No. 1, July1982 Lyncheta/.' Inputimpedance

I0 I00 I000 I0000

frequenciesbelow50Hz, IPv/Psl isapproximatelyconstant 0 ' -0- - -0

117 J. Acoust.Soc.Am.,Vol.72, No. 1, July1982 Lyncheta/.' Inputimpedance

118 J. Acoust.Soc.Am.,Vol.72, No.1, July1982 Lynchet al.:Inputimpedance

Zsc= T2[(Y:v- YMB)

c- 10- where the admittanceat the tympanicmembraneYz• was

The resultsof thiscomputationwith M•ller's measurements

119 J. Acoust.Soc.Am.,Vol.72, No.1, July1982 Lyncheta/.:Inputimpedance

-• -90 'J -90

I0- I00 I000 I0000

-- LYNCH et ol. N=4

I I I I I IIII I I I I I IIIll I I I I IIIII

FIG. 19.Comparisonof averagedmeasurements of the ratio of pressurein

120 J. Acoust.Soc.Am.,Vol.72, No. 1, July1982 Lynchet al.:Inputimpedance

Fig. 2}.Theirresultsaresimilarto oursin that IZcl isrelati-

Dancer, 1980,Fig. 5) indicatethat at frequenciesbelow 20

z = - = P/Us, suredifference(Pc) at low frequencies.

u, u, Althoughthe three-element networkof Fig. 22 is ade-

(a) Zsc:Zs,Zc (•)

FIG. 23. Networkrepresentationsof theimped-

126 J. Acoust.Soc.Am.,Vol.72, No. 1, July1982 Lyncheta/.' Inputimpedance

I I I I IllIll I I I IllIll I I I IIII1| I I I. lllll! I I

and the lack of publishedmeasurementsof scala tympani I10- ESTIMATES

area is particularly bothersome,sincethe apical region of OF PD/Pc xx BALLOS

shouldalsoprovidesignificantadditionsto the impedance.

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