108 J. Acoust.
Soc.Am.72(1),July1982 0001-4966/82/070108-23500.80 ¸ 1982Acoustical
Society
ofAmerica 108
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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
. "•
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
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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
110 J. Acoust.Soc.Am.,Vol. 72, No. 1, July1982 Lynchet al.: Inputimpedanceof the cat cochlea 110
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detectedphotonrate (counts/s), STAPES VELOCITY (Ps•R (PvTRANS[X.•E:R
= asymptotic
valueof r whenIv - V•I•F, OUTPUT VOLTAC-.-.•
) OUTPUT VOLTAGE)
111 J. Acoust.Soc.Am.,Vol. 72, No. 1, July1982 Lyncheta/.: Inputimpedanceof the cat cochlea 111
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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
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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
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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
z
IO0 1000 IOO00
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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
115 J. Acoust. Soc. Am., Vol. 72, No. 1, July 1982 Lyncheta/.: Input impedanceof the cat cochlea 115
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_ /•_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.
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.
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+90
O
. •'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
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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
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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 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).
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03 +90 tu +90
tu 0 tu 0
i x_•_ •- •
0- I/ '•.•_ •.t/ x• .
I
.
• -I0 - I .
o
ß
..
-20 - .
LYNCH
et
ol.
N=zn
j
-40 - TONNDORF
8(
m---m KHANNA(1967)
N= N
(1966)
.
.
• 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
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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
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
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
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Zsc=Ps/Us more de ta//ed network
122 J. Acoust. Soc. Am., Vol. 72, No. 1, July 1982 Lyncheta/.' Input impedance of the cat cochlea 122
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MECHANICAL NETWORKS Zc:Z•,, ZRW ELECTRICAL NETWORKS
US ZS
PsC • STAPES • ANNULAR'
LIGAMENT
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
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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 C^i• 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}
sinceC^i• 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 C^i• 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 C^i• 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
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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
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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
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/
-- LYNCH et al.,
• 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
-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
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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
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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.
independentof frequencyfor constantsoundpressureinput to the ear. Soc. Am. 24, 519-523.
Measurementsof cochlearpotentialsin cat indicatethat basilar mem- Fletcher,H. (1981)."On the dynamicsof the cochlea,"J. Acoust.Soc.Am.
branedisplacementshouldbe approximatelyproportionalto frequency 23, 637-645.
for frequencies
between0.2 and 1.0kHz (Dallos,1970;Weisset al., 1971). Franke, R., and Dancer, A. (1980)."Cochlearmicrophonicpotentialand
løDallos
(1970)actuallyproposed
hismodeltorepresent
thetotalinputim- intracochlearsoundpressuremeasurements at low frequenciesin guinea
pedanceof the cochleaZ c . In the absence of any intracochlearpressure pig," Proceedingsof theConference onLow FrequencyNoiseand Hearing,
measurements at that timeheassumed that pressure in scalatympaniwas editedby H. M•ller and P. Rubak (Aalborg,Denmark).
negligibleand thus did not includethe effectof the round-windowmem- Fung, Y. C. (1965).Foundationsof Solid Mechanics(Prentice-Hall, Engle-
brane in his theoretical considerations.However, since his CM measure- woodCliffs,NJ).
mentsare presumedproportionalto pressuredifference,thesemeasure- Fung, Y. C. (1967)."Elasticityof softtissuesin simpleelongation,"Am. J.
mentsshouldrelatedirectly to Z •. Physiol.213, 1532-1544.
llM. J. Mulroy (1977),usingscanning
electronmicroscopic
techniques, Geisler,C. D. (1977}."Mathematical modelsof the mechanicsof the inner
measuredhelicotremadiametersof 250 q- 10/am in two unsectioned cat ear," in Handbook of SensoryPhysiology,Vol. V/3 Auditory System,
cochleas; we assumethat thisdiameteris roughlyequalto that of the api- Clinical and Special Topics,edited by W. D. Keidel and W. D. Neff
cal portionof scalatympani.[Thishelicotremadiameteris abouthalf the (Springer-Verlag,New York), pp. 391-415.
valuereportedby Dallos (1970).] Geisler,C. D., and Hubbard, A. E. (1972}."New boundaryconditionsand
12 The "large tube" expressions, R = L (2/apco)l/2/(•ra3)
and resultsfor the Peterson-Bogertmodel of the cochlea,"J. Acoust. Soc.
M = pL [1 + (2/aw/pa2)l/2]/(rra2),
arevalidfora•or.o//a)l/2•
1.BothR and Am. 82, 1629-1634.
M dependon frequencybut, sincetheseelementsinfluenceZc predomin- Geisler,C. D., and Hubbard, A. E. (1978)."The compatibilityof various
antly in a smallfrequencyrange(40 to 100 Hz), we assumew = 50/2•r. measurements on the ear asrelatedby a simplemodel,"Acustica33,220-
Witha -- 125/am, p -- lg/cm3,/a-- 10-• g/cm-s(Steer,1967;Moneyet 222.
al., 1971),
w/(2•r)-- 50Hz, a(pw//a)•/• = 2.2,whichisnot• 1andthere- Gilad, P., Shtrikman, S., Hillman, P., Rubinstein, M., and Eviatar, A.
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