PATTERNS

OF

COCHLEAR

MICROPHONICS
APPENDIX

519

(3) The third "form of motion" is an dastic wave that dependson the mutual forcesbetweenadjacent elements in the cochlear partition. The pressure

Table of Mathematical Symbols and Notations B: width of the basilarmembrane(a functionof x). F: coefficient for the frictional force acting upon an

differences acrossthe partition, whether delivered directly(firstform) or asa surface wave(second form), set the cochlear partition in motionaccording to Eq. 3. Suppose, as an approximation, that over a certain rangeof frequencies the driving force, (pt--p-o)Bin Eq. (3), is restrictedto a regionof the cochlea near the stapes. Then the moredistantpartsof the partition
will carry a transverse wave described by the equation

M (&y/ O)+ R( Oy/Ot)qKy = r(a-oy/Ox)-U(O'y/Ox). (7)

This transverse wave depends solelyupon the solid structureof the partition and not upon the difference in the hydrostatic pressure across the partition. In this respect it differsfrom the second form of wavemotion

described above.Our failureto modifythe patternof the actual wave in the cochlea (observedby the electricalmeans)by producing an "acoustical short circuit" supportsour view that this third form of motioncannotbe neglected in considering the actual motion in the cochlea.When the driving frequency

elementarybox of fluid in scalavestibuli in its longitudinal motion. F-o:similarvalue of the fluid in scalatympani. F: average of F and K: coefficientfor the elastic force acting upon the cochlear partition. M: mass of the cochlear partitionper unit length. pt andp.o: pressure at point x in the scalavestibuli and tympani,respectively. R: coefficientfor the frictional force acting on the cochlear partition. Sx,S2: cross-section area of the scalavestibuli and scalatympani,respectively.

S: a{erage of S andS. T: longitudinal tension of thecochlear partition.

It isourpresent opinion that theamazing complexity t: time. of thephase andamplitude patterns of cochlear motions p: densityof the fluid. of the sound. can be understood, at least qualitatively, in termsof c0: 2r timesthe frequency co,: (K/M) , namely 2r times theresonance frequency Ihe three differenttypes of motion: direct driving, surfacewaves,and transverse elastic waves. at point x.

U: coefficientfor the shear force acting upon the cochlear partition. m, u-o: averagelongitudinal velocity of the fluid in scala vestibuli and tympani, respectively. is high and the wavelengthshort, the elastic waves x: distance along the cochlear partition. should becomeprominent becausethe effects of the displacement of the cochlear partitionat "mutualforces" in the partitiondepend on the curva- y: average point x. ture of the structures alongthe partition.

THE JOURNAL

OF THE

ACOUSTICAL

soCIETY'

OF AMERICA

VOLUME

24. NUMBER

SEPTEMBER.

1052

Dimensionsof the Cochlea (Guinea Pig)*

CentralInstitute for theDeaf, St. Louis,Missouri (Received April 27, 1952)

Measurements havebeen made underthe microscope of manystructures in thecochlea of theguinea pig and the results plottedgraphically as a function of position alongthe cochlea. The structures measured are thespiral lamina,Rosenthal's canal,the basilar membrane (length, widthand thickness), the channels
and windows of the cochlea (cross-section area) and the hair cells(angles of orientation).

thecochlea2 More recently theyhavebecome important also for calculations of the probable distribution of HE dimensions of thestructures of thecochea electrical current flow and potential fields within the and the way in whichthey vary as a function of Wever a has published many usefulmeasurepositionalongthe cochlea have long beena matter of cochlea.: ments on the human cochlea. The presentstudy was interestbecause of their bearingon the dynamics of undertaken to supplysimilardata for the guineapig,
* This work was earfled out under Contract N6onr-272 between the Office of Naval Research and the Central Institute for the Deaf. The author is indebted to Dr. Walter P. Coyell of the

INTRODUCTION

Department of Otolaryngology of Washington University for the

H. Fletcher,J. Acoust.Soc.Am. 23, 637 (1951). a G. v. B6kisy, J. Acoust.Soc.Am. 23, 18 (1951). aE.G. Wever, Theoryof Hearing (John Wiley and Sons,Inc.,

useof laboratoryradiities and for helpful criticismand guidance. NewYork, 1949).

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$20

CISAR

FERNANDEZ

Drawingsat magnification of about 250 diameters weremadewith a camera lucidaof the cochlear passages

(scalae vestibuli, tympani, andmedia)andof the organ of Corti, including the innerand outer tunnelof Corti and Nuel's space. The cross-sectional areas of these structures were measured with a plankmeter. The organof Corti was also drawn with the cameralucida at high maguification, and lines parallel to the axes of the innerand the outerhair cells weresuperimposed as in the sketchin Fig. 6. The angles formedby these
two lines with one another and with the basilar mem-

FIG. 1. Projectionof the basilar membraneof guineapig on a plane perpendicular to the axis of the modiolus.Line AB is a
mid-modiolarsectionwhich cuts the cochleaat 8 points. Line CD is also a mid-modiolarsectionbut at right anglesto AB; it cuts the cochlea at 8 pointswhichalternatewith the pointson Line A B.
The numerals indicate the distances in millimeters of thes intermembrane under the tunnel of Corti.

braneweremeasured with a protractor. The total length of the basilarmembrane and also the distancealong the basilar membranefrom the round windowto each point of intersection with the axesindicatedin Fig. 1 were measured by the reconstruction method of Guild.4 The length of the basilar
PERCENTAGE OF LENGTH

20

40

60

80

100

sectionsfrom the round window, measuredalong the basilar

the animal

which serves so well for studies of the

electrical phenomena of the ear.

METHOD

0.4: o.3-

O-5 .
.o...-'

i-.-"'?
.y .... ' \

Six adult guinea pigs were perfusedwith fixing

solution;their temporalboneswere then removedand
mounted in celloidin. Four cochleas were sectioned

'o.-""
I. SA /

!
i JI

:.
'm' [ APICAL

seriallywith the axis of the modiolus orientedin the horizontalplane and turned so that the sections were MILLIMETERSON BASlEAR MEMBRANE (CIINEA parallelto line AB in Fig. 1. Two other cochleas were sectioned parallelto the plane indicatedby CD. The F/o. 3. Width of the basilarmembrane in the guineapig. The X and represent our own measurements madeaccording sections were stainedwith hematoxylin-eosin and the curves to the criteria shoa n in the sketch. The criterion Y' was used in most representative mid-modiolarsectionof each ear the upper tums where the bony spiral lamina doesnot extend
was selected for measurement.

DISTANCE CANAL
IN

ACROSS AND SPIRAL

ROSENTHALJ LAMINA
PIG

GUINEA

beyond the limbus. The lowestcurve, X--X, represents Guild's data for the guinea pig measuredfrom spiral ligament to the spiral lamina. The upper curve (HUMANI) showsWever's measurements. For these data use the scale for percentageof length (at the top). To convert these percentages into millimetem multiply by the factor 0.315.

membranewas also determineddirectly as described elsewhereS: The basilar membranewas carefully dissectedout undera dissection microscope and measured with a calibrated eyepiece.

,.ojc L

The width and thickness of the basilarmembrane, the width of Rosenthal's canaland of the spirallamina weredetermined by measurements, madeby the useof a calibrated eyepiece,on the mid-modiolarsections
described above.
RESULTS

Spiral Lamina
.

The osseous spirallaminaof the guinea pig does not have a uniformwidth alongthe cochlea. It tapersfrom
IN MILLIMETERS

DISTANCE

ON

SILAR

MEMBRANE

a broad lamina 0.5 mm wide at the middle of the first

. 2. he wid of e oens spiralla murcd between the ts indit by A in'the inner fire. In the t three
quem of e t turn e is a secondreal (C) betwe e

turn until it disappears at the beginning of the apical

S. R. Guild, Science 65, 67 (1927). s C. Femtindez,Laryngoscope 61, 1152 (1951).

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DIMENSIONS

OF

THE

COCHLEA

521

turn (Fig. 2). In the apicalturn the nervefibersare containedin channels formed by fibroustissueand
continuations of the fibers of the basilar membrane.
CROSS SECTION AREA OF'

The part of the lamina lying betweenthe limbusand habenulaperforatais so thin that it seems probable that it canvibrateas part of the basilarmembrane.
RosenthaPs Canal

THE

THREE

CHANNELS OF

OF THE

COCHLEA

I
!

THE GUINEA PiG

Rosenthal's canal has an averagewidth of 0.30 mm

in the first turn and in the basal half of the second turn.

From the middleof the second turn it tapersoff until the apical turn is reached. In the apical turn the canal is no longer clearly defined.Here the spiral ganglion
consists of a clusterof bundlesof nerve cellsand fibers, sometimes,but not consistently,separatedby bony
walls. In the first half of the first turn there is an

additionalchannel(C in Fig. 2) about 0.35 mm wide

that lies between Rosenthal's canal and the modiolus.

This channel plus Rosenthal'scanal proper have a

total widthof 0.7 mm in thispart of the cochlea.

0
7

I% .... ' I
DISTANCE

' ' I0 ....

IN

fi , l]I , 1,5 ' ' 20

MILLIMETERS

Fro. 5. The abscissa is distance in millimeters alongthe basilar

membrane. Thepoints represent averages of measurements in six

must be substracted.

ears. The broken lineswereobtained by extrapolation. To obtain the areaof scalavestibuli at any point,the areaof scalamedia

3.5

10.5

12.5

14.25

16.5

I,,

laminadid not extendbeyondthe limbusthe latter was usedas the innerboundary (Y in Fig. 3). The
width of the basilar membranemeasured to the habenula

LENGTH OF THE BASILAR MEMBRANE (MM)

Fro. 4.

perforata (X) increases rapidly overthe'first 2 mmof

the basilar membrane from 0.07 to 0.15 ram. Thereafter

Basilar

Membrane

it increases moreslowlyuntil it reaches its maximum

RATIO OF CROSS SECTION
I

(.4)
Direct measurements and the method of reconstrucI

tion agreein showing that the average lengthof the basilarmembrane in the guineapig is 18.8 mm with extremeindividualdifferences of +0.5 ram. (In all of
our references to distances from the round window the

AREASOF SCALAVESTIRUI..I i
TO SCALA TYMPANI

(GUINEA PiG)

distances are measured along the basilar membrane underthe tunnelof Corti.) The average lengths of each of the four turns, as indicated in Fig. 3, are 8.5, 4.8, 3.4, and 2.1 mm, respectively.
(.) Wit,
It is difficult to measure the width of the basilar

"!
Jh . .... : .......... Sv

membrane because the boundaries are not clearly

defined. We undertook to measure the width of the

sectionthat would probably participate in vibratory

motion. We therefore chose as our outer reference

I 0/

DISTANCE IN MLIMETERS ALONG BASILAR point the junctionbetween the basilarmembrane and Fro. 6. The brokencue represents the ratio of the crossthe spiral ligamentas indicatedin Fig. 3. The two to sla tyminner reference pointswere (1) the outer end of the ctional areasof scalavestibuliplusscalamedea pani,andthesolid linetheratioof ala vestibu alone to la habenulaperforata,and (2) the edgeof the osseous tympani. The broken portion of_e solidle w obted by spirallamina (Y in Fig. 3). When the osseus spiral teolation.

....

. . . . ., . .* . , . . .
u

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522

CiSAR

FERNANDEZ mm 2. The cross-sectional area of the scala vestibuli

I00-

first increases slightly and then diminishesrapidly

until it reaches an area of 0.36 mm 2 at the middle of

thefirstturn (Fig. 5). Fromhereon it tapers, with some irregularity and slight variations from one animal to
another, until it reachesan area of 0.10 mm2 closeto
the helicotrema. These measurements the area of the scala media. do not include

The cross-sectional area of scalamedia is practically uniform throughoutthe entire length of the cochlea. It tapersvery graduallytoward the helicotrema over
0 . I0 lift ,
DISTANCE IN MILLIMETERS ALONG BASILAR MEMBRANE

as measured in two ears.The pointsand triangles represent aver- media is 0.10 mm 2. age measurements. No measurements were made between0 and The area of the roundwindow,calculated from major 3.5 ram, but examinationof other sections (not cut exactly at right angles to the tunnelof Corfi) shows that A does not change and minoraxes,is 1.02mm and the cross-sectional area significantly while B seems to increase and then to decrease of scala tympai at this level is 1.20 mm . Scala tympani slightlyand graduallytoward the roundwindow.

the upper8 mm of the cochlea. The final taper, both at the roundwindowand at the helicotrema, is very Fro. 7. The anglesof the hair cellswith the basilat membrane abrupt. The typical cross-sectional area of the scala

first increases and then diminishes rapidly until it

reaches an area of 0.48 mm at a distance of 5 mm from

of 0.25 mm at the middleof the apieal turn whereit then rapidly tapersoff. The measurements madeto the osseous spiral lamina or to the limbus(Y or Y') showa similarcurve,althoughof course the width is always greater than the width measuredto the habenula perforata.The maximumwidth of Y is 0.34 mm and is reachedat about the same position as for X in the apical turn. The width of the basilarmembrane in the guineapig was measured by Guild. s His data are alsoplotted in Fig. 3 as the lowestcurve. He statesthat he measured from the edgeof the spiral lamina to the attachment to the spiral ligament, and also that "some were measured from the foramina nervosa" (habenula pefforata).The agreement with our curvelabeledX is very satisfactory if we assume that by the phrase "edge of thespiral lamina" hemeans thefirstthickening of the membrane underthe foot of the internalpillar.
Wever's a data for the width of the basilar membrane

the round window. From this point on it tapers on gradually to an area of 0.05 mm closeto the helicotrema (Fig. 5).

In Fig. 6 is plotted the ratio of the areasof scala

vestibuli and also of scala vestibuli plus scala media to that of scala tympani. At approximately 1.5 and
again about 6.0 mm from the round window the

cross sections of scalavestibuli and scalatympani axe equal.From 0.15 mm scalavestibulipredominates with a maximum ratio of 2.2 at the opening of scala vestibuli.
From 1.5 to 6.0 mm from'the round window scala

tympani predominates. The maximum ratio of 3.5 is

reached at 3.5 mm from the round window.Beyond 6 mm the scala vestibuli againpredominates, reaching
a maximum ratio of 2.8 near the helicotrema. If the area of scala media is included with that of

scalavestibulithe effecton the ratio is very slightfor the first turn, but, as shown in Fig. 6, the ratio is altered more and more in favor of vestibuli-plus-media in man is alsoplotted in Fig. 3. The basilarmembrane is approached. of humansand of the guineapig is the samewidth for as the helicotrema The most recent comparablemeasurements of the the first 30 percent of their lengths. After that the cross-sectional areas of the human cochlea were made width in man increases more rapidly until it reaches The humanscalas tympani and vestibuli its maximum of about0.50 mm at the same percentage by Zwislocki. of its lengthfrom the roundwindowas the position of do not showthe great and abrupt reductionin area in the first half of the basal turn that is so evident in the maximumwidth in the guineapig. guineapig. The openings of the scalasare of the same (C) Thickness orderof magnitude (2 to 3 mm ) as in the guinea pig, The thickness of the basilar membrane is 7.4 at but the taper is lessabrupt and lessregular. the beginning of the first turn. It tapersgraduallyuntil Organ of Corti it reaches 1.34 at the apicalturn (Fig. 4). This measThe total cross-sectional area of the organof Corti, urement does not include the organ of Corti or the mesothelial cells that lie on the oppositeside of the includingthe inner and outer tunnel of Corti and Nuel's
membrane.

Channels

and Windows

of the Cochlea

The area of the oval window, calculated from the

space,was measured.The measurementincluded the fiupporting cellsof the inner spiralsulcus and alsothe supporting cells of the outerspiralsulcus, asfar astheir junctionwith the spiral ligament.The total surface
area amounts to at least 0.012 mm x. The measure-

major and minor axes,by the formulafor an ellipse, is 1.41 mm2. The opening of the scalavestibuliis 3.27

i j. Zwlslockl, Acta Oto-Laryngol. Suppl.72, 1-76 (1948).

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DIMENSIONS

OF

THE

COCHLEA

523

ment increases graduallyover the second and third

turns to a max'mum of 0.017 mm 2 at a distance of 17.5 mm from the round window and diminishes to 0.015 mm 2 near the helicotrema. The cross-sectional area of

Orientation

of the Hair

Cells

Figure7 shows the angles formedby the axesof the

outer and inner hair cells with one another and with

the basilarmembrane. The anglebetweenthe inner the tunnels in the organof Corti alsoincreases slightly ha/r cellsand the basilarmembrane (alphain Fig. 7) from 0.002 mm 2 in the first turn and in the first half of is constant at about 65 throughout the cochlea. The the secondturn to 0.005 mm2 at the beginningof the angle(beta) formedby the outerhair ceilsdecreases apicalturn. The total cross-sectional areaof tissue (total progressively from baseto apex and consequently the area minus the area of the tunnels) thus varies from angle betweenthe outer and the inner hair ceils in0.010 mm throughout the first turn to a maximumof creases. This apica] angle (gamma) is about 60 at 0.013 mm 2 at the beginning of the third turn. It then 3.5 mm from the round window and increases to 92, 0.5 mm from the he]icotrema. graduallyfalls to 0.011 mm near the helicotrema.

THE

JOURNAL

OF THE

ACOUSTICAL

SOCIETY

OF AMERICA

VOLUME

24. NUMBER

SEPTEMBER,

1952

The Masking of Tones by White Noise as a Function of the Interaural Phases of Both Components.*I. 500 Cycles
L.o1) A. JEFlSS, HUaR C. BLODOErr,Am BRUCEH. DAl'tmRV.

Defense Research Laboratory andDepartment of Psychology, TheUniversity of Texas, Austin,Texas (Received March 3, 1952)
Work by Hawkins,Hirsh, Licklider,Stevens, Webster, and others hasshown the considerable reduction in masking whichoccurs whentheinteraural phase of eitherthesignal or thenoise is reversed, or whenthe interaural phase of thesignal isshifted by various amounts. Thepresent paper extends thisworkby shifting theinteraural phases of the masking component of thenoise andof the toneby various amounts between q-180 and- 180 ,andalso by shifting thenoise timewise by amounts upto4.0milliseconds. Theinteraural phases of both the noise and the 500 tonalsignal are therefore parameters of thisstudy. The resultsare in agreement with thosealreadypublished, but showin additionlarge reductions in threshold evenwhenthe noise and the tonedifferin their interaural phase positions by amounts lessthan
180 .They alsoshowa periodicfall and riseof the maskedthreshold as the Enteraural time difference for the noise is varied.Finally, they show theimportance of interauralcorrelation in determining theextentto which
the binaural threshold will be lower than the mortaural.

R-C network (Experiment 1), or by displacing the whole noise bandin timeby means of a delayintroduced HErecent literature contains a number ofpapers of a binauraltape reproducer (Exuponthe relationbetween the masking effectof into one channel 2). The R-C network could produce shifts up a noiseand the interauralphases of the noiseand the periment . Larger shiftswere obtainedby reversing the masked signal.In mostcases - the phasechange is a to 90 to one channel(180 ) and subtracting as reversalof phaseof the noise,the tone, or both. In one connections necessary by means of the R-C network. study the tonal signalhas alsobeenshiftedthrough In both experiments the stimuli were presented angles otherthan 180 . The present work extends the investigation to a greater variety of noiseand tone automatically,two hundredat a sitting--twenty at eachof ten intensitylevels1.5 db apart. The signals phase-combinations.
METHOD

INTRODUCTION

The present experiments employed a tonal signalof

500 and a wide-band white noise. The interaural

were 150 milliseconds in duration,and were presented at intervals of three seconds against a continuous noisebackground. Each was preceded by a 150 ms
The phase-shift networkwasadapted from that described by H. J. Reich,Theory andApplications of Electron Tubes (McGrawHill BookCompany, Inc., New York, 1944),p. 460.It consists of the low impedance secondary of a transformer, the centertap of which constitutes one terminal, and the commonconnection between a condenserfrom one leg and a variable resistor from the

phasepositionof the signalwas adjustedby an R-C network. The noise position was altered, either by shiftingthe 500 masking component by means of an
* Resultingfrom research doneunder Bureau of ShipsResearch

and Development ContractNObsr-52267. xI. J. Hirsh, J. Acoust.Soc.Am. 20, 536 (1948). J. C. R. Licklider,J. Acoust. Soc.Am. 20, 150 (1948). aI. J. Hirsh and F. A. Webster,J. Acoust.Soc.Am. 21, 496
(1949).

otherleg constitutes the other terminal.The phase shift for the 50(P,masking component of the noisewasdetermined by substitutinga 500 tone for the noise,and then adjustingthe network to give the appropriate Lissajoufigureon an oscilloscope.
Calculation shows that this will shift the extremes of the critical

J. E. Hawkins, Jr. and S.S. Stevens, J. Acoust. Soc.Am. 22,

6 (1950).

band throughvery nearly the samephaseanglesas the middle

F. A. Webster, J..Acoust. Soc. Am.23452(1951).

frequency whenthe maximal phase shiftis no greater than90. A similar device wasused in the circuitfor the signal tone.

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