J. Physiol.

(I949) IIO, I-I7 3

THE MECHANICS OF THE SEMICIRCULAR CANAL

BY A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
From the University Clinic for Diseases of the Ear, Nose and Throat, Utrecht
(Received 26 August 1948)
When Steinhausen (1931) discovered that the cupula terminalis in the semi-
circular canal reaches the top of the ampulla, fitting hermetically in it, and
when he moreover observed that a deviation of the cupula, caused by an
endolymph flow, gradually decreases because of the directional force of the
cupula, the theory of the semicircular canal underwent a significant change.
Up till then there was held to be a narrow canal in which an endolymph-flow
would soon be stopped by frictional damping. After a clinical turning test the
endolymph would flow for about 0 5 sec. only, and then every part of the canal
would be at rest. The prolonged after-sensations of 30 sec. or more could not be
explained by the mechanical properties of the canal. They then had to be
ascribed to a 'central origin', the only possible means of surmounting this
difficulty (Mach, 1875; Gaede, 1922).
Steinhausen gave the cupula its proper prominence. The endolymph after
a sudden arrest of the turning chair will indeed flow on for about 05 sec., but
the cupula is forced to bend and after having attained its maximal deviation,
without further external forces acting, it will try to recover its former position
of zero deviation. This process may take from 1 to 60 sec., depending on the
rate of the angular velocity just before the sudden arrest. The prolonged after-
sensations appear to have a very simple mechanical origin. As Steinhausen
pointed out, the cupula-endolymph system has to be considered as a heavily
damped torsion pendulum. It thus has a differential equation with solutions
available in every handbook on mechanics. This statement, important as it may
be, brings us no further if the constants of the equation for this case cannot be
determined, but if they are known this equation ought to predict any conduct
of the cupula found in practice.
Steinhausen did not establish the values of the constants for his test animal,
the pike. So he was unable to prove the hypothesis of the torsion pendulum
with its consequences. We have determined these values for the human subject
by means of experiments based on mechanical principles.
PH. CX. I
 2 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
METHOD
A much clearer idea of the function of the semicircular canals can be obtained, using small stimuli,
than is possible with the usual Barany test.
The subjectis placed in a turning chair, with the head inclined forward about 300. The movement,
either clockwiseor anticlockwise,starts smoothlywitha subliminal acceleration, whichis maintained
until the angular velocity needed is attained. It is halted quite suddenly (braking time 1-3 sec.,
depending on the angular velocity). The test subject has a sensation of turning round. This sensation
expires after a certain period. The duration of the sensation is expressed as a function of the
anguJar velocity. This relation is plotted in a diagram, which is called the cupulogram.
The test is performed with the test subject in darkness, lest he obtain his sensations from the
surroundings. The determination of the duration of the nystagmus is performed with Frenzel's
spectacles.
It is important that the test subject has no sensations of turning up to the time of braking.
The time, necessary to obtain the required velocity, depends on this velocity, but as a rule does
not exceed 3 min. The angular velocities used lie between 1 and 60'/sec. The non-physiological
velocities of BArSny of 180°/sec. and more were not used.
Comparison of the graphs of sensation and nystagmus duration gives a far better indication of
any abnormality of function than the earlier methods.
For the vertical canals the same procedure can be followed, with the head in the appropriate
position.
RESULTS
The differential equation
The differential equation of a torsion pendulum where there are no external
forces is fI A
E)e+TIIe+/ =O or 1+jejs+_=O. (i)
The constants to be determined are II/@ and A/(:
E0 = moment of inertia of the endolymph.
II = moment of friction at unit angular velocity.
A = directional momentum at unit angle caused by the cupula.
=the angular deviation of the endolymph in relation to the skull.
=the angular velocity of the endolymph in relation to the skull.
=the angular acceleration of the endolymph in relation to the skull.
(All these at the centre of the canal.)
The approximate solution of equation (i) is
s ='y1 [e-AtIl - e ] (ii)

with the limiting conditions: t= 0, =00 and =y.

The angular velocity just before the sudden arrest of the turning chair is y.
It will be called 'the impulse', though it has not all the properties of a mechanical
impulse.
Equation (ii) concerns the clinical turning test. After a period of turning
with a constant velocity y the subject is stopped: the endolymph will thus have
the initial velocity y, whereas the cupula had no deviation and was in perfect
 THE SEMICIRCULAR CANAL 3
equilibrium. The cupula is pushed forward and attains its maximal deviation
at tmax.. 0 fl2
tmax. = -i log <A (approx.). (iii)
It then has a deviation emax., where
fmax. =y 1I (approx.). (iv)
At this moment the second term in equation (ii) e-1t110 is almost zero and
may be neglected. The cupula then tends to regain equilibrium (zero deviation).
This prolonged period is determined by the first part of equation (ii) e-At/l.

075

L. 05

0
c-0 0-25.
2
in

0-1 0 10 t,, 20
Time (sec.)
Fig. 1. Deviation t of the cupula produced by sudden arrest of an angular velocity y0/sec. Maximum
deviation at t =0*5 sec.; minimum deviation passed at t = 14 sec., giving rise to sensation.
Ordinate logarithmic. Slope of curve gives 11/A.
The subject has a sensation of declining angular velocity with a direction
opposite to the original rotation of the chair. This is quite plausible from
a mechanical point of view. The sensation expires after t. sec.

/\log ( ) (approx.), (v)

where emin. denotes the smallest deviation of the cupula giving rise to a just
perceptible sensation of rotation. In Fig. 1 tu is shown where the curve inter-
sects with the emin. line. This equation gives us a means of determining a ratio
between two unknown constants of the differential equation. The value of tu
has to be determined as a function of y and the slope of the curve gives IH/A,
if the impulse is plotted logarithmically. This is done in Fig. 2. It appears that
l/A = 10 sec.
The expectation that the curve is a straight line is confirmed. The smallest
perceptible impulse Ymin. is about 1-50/sec. This figure was obtained from
a normal sensitive subject. All the sensitive subjects give values very close to
1-2
4 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
those of Fig. 2. The average values for all the normal subjects tested, the
insensitive ones included, are
II/A =8 sec. and Ymin. = 25°/sec.
From equation (v) it follows that
Ymin. = emln.J1/0 (approx.). (vi)
This agrees with {max. as expected.
Unfortunately, equation (iii) has no practical value for human subjects, as
tma,. has to be of the order of 0x5 sec. and this is too short a time to be estimated
with satisfactory accuracy. Otherwise, with Il/A known, the differential
equation would be determined.
100

10

ED

1 A1 f I I
0 10 20 30 40
Tlme (sec.)
Fig. 2. Duration of sensation as a fimetion of the impulse. Normal sensitive subject. Slope gives
Il// = 10 see. Minimal impulse (Ymin.) = 1-5°/sec.
Resonance vibrations on the torsion swing
A second experiment, independent of the former one, is needed to establish
the value of Il/0. Resonance vibrations on a torsion swing enable us to impose
upon the subject harmonic vibrations about a vertical axis. In this manner the
horizontal canals are studied. The sensation of the subject is also a harmonic
vibration, but there is a phase difference between sensation and swing. Only
when the frequency of the swing is equal to the natural frequency of the canal
is the phase difference zero. The natural frequency is that frequency which the
cupula-endolymph system would have without damping, and is a theoretical
value which cannot be demonstrated directly. The system does not vibrate
when undisturbed. When a pendulum, damped or not, is forced to vibrate by
means of a harmonic external momentum, it has a phase difference of 90°
(lagging) when it is in resonance. When the external frequency is higher the
phase difference becomes more than 90°, and in case of a lower frequency it
 THE SEMICIRCULAR CANAL 6
becomes less than 90°. When the swing reaches the turning point, the cupula,
in case of resonance, is at zero deviation and the subject has the sensation of
rest. The swing, coming from its turning point, will pass its zero deviation at its
maximal velocity. The cupula at that moment has reached its turning point
(and so its maximal deviation) and the subject has a sensation of maximal
velocity. Thus sensation and vibration always coincide in case of resonance.
But when the swing frequency is too high, the sensation of rest at the turning
point will lag behind; when the swing frequency is too low the sensation will
go ahead.
The theory in this case demands
A
+ + aC sin cot, (vii)
where oc denotes the maximal angular acceleration and w the frequency of the
swing. oc is connected to the amplitude A, and w to the period T of the swing in
the following way: 2A (viii)
2HI
211
T (ix)
The approximate solution of equation (vii) is
{= - sin (cot +. .. terms (neglgible). (x)
The angle ¢ is determined by
tan + = eOs _s, '(xi)
where c0 is the natural frequency of the canal, and
2 A

If it is possible to determine c0, A/E is calculable. When co >,wo, tan + <0;

thus f> > 900 and the subject lags behind the swing. When co <coo, tan > 0,
k <90°: the subject goes ahead of the vibration of the swing. The resonance
frequency ought to lie between these two cases. Coincidence of sensation and
swing itself does not suffice as an experiment, for it could be ascribed to a faulty
observation. The phase differences to be expected are measurable as very short
time differences of the order of 0 3 sec. at most. This is only possible with
sensitive subjects. As an average value we found: coo=10 sec.-1. Thus
A/( = 1-0 sec.-2. The differential equation (i) becomes
e+10e+5e=O. (xii)
The probable error in I1/0 is about 25% and in A/0 about 20% (Egmond,
Groen & Jongkees, 1943, 1948).
The measurements mentioned above may be checked as follows.
6 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
Test 1
In case of a circular movement with constant acceleration cx rad./sec.2, the
behaviour of the cupula-endolymph system ought to be as follows:
t+ne {+e{=o. (xiii)

.j 01_* ,..--

610
o.

o r 5 10 1
Time (sec.)
Fig. 3. Deviation 6 of the cupula under influence of a constant angular acceleration (l°/sec.)2.
The latent period Tr seconds lies between start and the moment when the cupUla passesStmn
After infinite time (about 30 sec. in practice) the cupula attains equilibrium.
It has an approximate solution
5t=oc pv (1-e-At/lI), ~(xiv)
with the limiting conditions: t = O, 0 , 0 , =oc. The movement starts from
rest; gradually the velocity y will increase. After infinite time
tO
0a_0 (xv)
When in the case of the human subject an angular acceleration of 1°/sec.2 is
used, the deviation of the cupula with time may be estimated (Fig. 3). The
formula is et = 1 - e-10 degrees.
At first the subject feels at rest until the cupula passes the emin. value; then
the first sensation of rotation will be perceived. Thus there is a latent period of
,r sec. From equation (xiv) follows

emin.= cx II or 0t7r= emin. <).(xrvi)

As the right side of equation (xrvi) has a constant value for a certain subject,
the product ocz also has that property, as was first mentioned by Mulder (1908).
We propose to call wr the 'Mulder product' (Buys & Rylant, 1939; Buys,
1940).
 THE SEMICIRCULAR CANAL 7
With a sensitive subject:
ac 7 OCT
(0/sec.2) (sec.) (0/sec.)
10 2-0 2*0
3*6 0*5 1-8
5*0 0*3 1.5
The error in the -r column is rather large for the shorter times, but, never-
theless, OCT is evidently of the order of 2°/sec.
More may be said of the ocT product, for from equation (vi) it appears that
OCT =Ymin. (xvii)
As these results were obtained from approximated equations, it is to be expected,
in this case, that ocx will be larger than 6min., but it remains true that ocr must
have a fixed proportion to Ymin..
After infinite time (in practice after about 30 sec.) equilibrium between the
directional momentum of the cupula and the inertia of the endolymph will be
attained. The cupula is then at rest with a deviation e., and
to 0 or, as ---=
=a E0
1, =o. (=xviii)
This means that the numerical value of the cupula deviation should be equal to
that of the acceleration, but only after infinite time. We were able to confirm
that the maximal sensation is indeed attained at about 30 sec. As physio-
logical movements last for 3 sec. at most this numerical equality is never
reached. This is hardly surprising, because the function of the semicircular
canals is to control bodily movements by indicating the attained angular
velocity and the angle of rotation executed.
As a certain subjective angular velocity corresponds to a certain deviation of
the cupula, the problem is twofold: (a) to establish the correspondence between
cupula deviation and sensation of rotational velocity; (b) to prove that during
a physiological movement the sensation of angular velocity is equal to the
velocity attained at that moment and that the angle covered by the movement
corresponds to the subjective estimation.
(a) Some subjects are able to estimate the angle covered by their sensation
of angular velocity after the turning test. An angle commonly stated is 3600,
or one turn. When the time to cover this angle is measured, the mean subjective
angular velocity during this time can be obtained. The subject mentions the
moments when he has completed his subjective turns and the observer times
them with a stop-watch (Groen & Jongkees, 1948). An example of this kind
of experiment is given in Fig. 4. The impulse administered was 40°/sec. and
the subjective angular velocity extrapolated to t = 0 gives 40'/sec., a very
good agreement.
In Fig. 4 the slope is again HI/A. Here II/A = 10 sec.
8 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
It is evident that the subjective velocity is equal to the objective one y,
connected to the cupula deviation { by

6 I=I or = y, for human subjects, (xix)

and that the determined 11/A corresponds to that established in Fig. 2.
The numerical value ofthe cupula deviation is one-tenth ofthat ofthe impulse.
There is not always such a thorough agreement between objective and
subjective y, but sensitive subjects give many results with such agreement, so
that the statement is probably true. When there are deviations, they mostly
tend to too high a subjective velocity.
100

~~50
20

, 10

bo S

tI 2 '

1 l l l l l l l l
0 10 20 30 40
Time (sec.)
Fig. 4. Subjective angular velocity against time. The impulse administered was 40°/sec., as is the
extrapolated subjective velocity. The slope gives H/A = 10 sec.

(b) The physiological movements consist mostly of an accelerative and

a following decelerative phase. The maximal velocity is reached between these
two phases. The statement is now that the cupula deviation ought to give rise
at any moment to a sensation of rotation equal to the acquired objective
velocity. From equation (xvi) it is already known that during the first part of
a movement with constant angular acceleration the cupula has a deviation
proportional to i-r, otherwise the threshold values of the sensation would never
give rise to the constancy of the oTr product. As y =oar'/sec. the cupula should
have a deviation proportional to y. This proportionality factor was determined
to be I . Within the limits of the validity of the Mulder product oCr, the theory
mentioned above may be generalized for non-constant accelerative movements:
{-+Y={cadt, (xx)
,where e = I1 y and T is the moment considered.
 THE SEMICIRCULAR CANAL 9
In the decelerative phase of the movement the cupula deviation is abolished,
but only within the period of validity of Mulder's law, for the attained maximal
velocity is reduced to zero, and when the cupula deviation remains proportional
to the velocity they decrease together.
If there is a space of time with constant angular velocity between the
accelerative and the decelerative phase, the cupula tends to recover zero
deviation during this period, with the result that at the end of the decelerative
phase the cupula is forced past its zero position and that an after-sensation is
caused (clinical turning test, prolonged waltz in the same direction). The condi-
tions imposed by nature on the semicircular canal are therefore confirmed by
the theory.
Too short a movement has an opposite effect, i.e. an after-sensation in the
same direction as the original movement (in contrast to the results from the
clinical turning test). An example of this is the turning of the head.
Turning of the hed. This movement lasts so short a time that the ar law has
not begun to apply when the head has already covered a considerable angle.
The Mulder law then comes into action and for the remainder of the movement
applies rather strictly. When the head stops the cupula lags behind (a 'fault'
originating from the slow start of the cupula) and there is a slight after-sensa-
tion for a very short time owing to the peculiarities of this movement. An
example of a turning of the head may illustrate this statement. Suppose the
head turns from right to left over an angle of 900 in 0-6 sec. The acceleration
generally follows a sine function, as may be shown by measurements. The head
obeys the equations:
x=oc sin cot =2A sin wt,j
xt = coA (I -cos cot), (Xi)
x = A(t-sin cot-7r. J
The angle of 900 gives: A = 290 = 140 approx., t = 0-6 sec., thus w =10 sec.-1.
The differential equation of the cupula endolymph is
e+ (i e + A =oa sin ct, (xxii)
with the limiting conditions at t = 0, = = =0
=
co V/(022+n2/92esin(ct-f)+
4(
+ 112/02) C0
2
+
(0/l)
12/E02~w(e/0)
- C04)
'~ + (11O/0)2~

and tan = Co l
(approx.). (xxiii)
With co=10sec. 1, oc=1400'/sec.2=co2A, r1/0=-I/A =10sec.-A, and A/E =
1 0 sec.-2 the result may be simplified to
=-7 [V2 sin (lOt+450)+e-'Ot2] degrees.
10 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
The behaviour of the cupula deviation 6 with time is shown in Fig. 5. There is
a slow start with the result that the maximal cupula deviation is reached after
the middle of the movement. At the end there still remains a deviation of 70,
which is neutralized in about 0 3 sec., for the cupula not only has a deviation,
but also a velocity which drives it to zero deviation in accordance with the
equation
= 7e-10I degrees.
To test the law of Mulder it is sufficient to consider the behaviour of the cupula
in relation to the velocity of the head. At a maximal velocity of the head of
280'/sec., the maximal deviation of th-e cupula is 240, corresponding to a sub-
jective angular velocity of 10 x 24 = 240'/sec. There is thus a slight discrepancy
between objective (2800) and subjective (2400) angular velocity, but in general
it may be said that the law of Mulder is valid.
300 Velocity of the head
tSubjective velocity
200
End of the movement
100

0 03 0-6 09
Time (sec.)

Fig. 5. Turning of the head 900 in 0-6 sec. The angular velocity is maximal at t =03 sec. and
attains the value 280'/sec. The subjective velocity lags behind, but reaches almost the same
value, 2400/sec.
The ballet dancer. When pirouetting a ballet dancer will far surpass the period
of Mulder's law. When coming to a standstill the dancer would be very dizzy
and control of the situation would be impossible if his head had followed the
continuous turning of his body. However, he divides the total turning into
parts, keeping his head at rest by looking at a fixed point, and when his body,
turning on beneath his head, goes too far he jerks his head to the next halting
point. Every jerk with the short rest after it lies within the range of the law of
Mulder, so that he minimizes the tendency to dizziness during the turning. This
practice is now plausible from the theoretical point of view.
Test 2
When the torsion swing vibrates with a gradually decreasing amplitude, the
test subject will have a declining sensation. Ultimately he will only feel the
 THE SEMICIRCULAR CANAL 11
maxima of the swinging. Then sin (cot+o) in equation (x) will be unity
(Fig. 6). Thus amin. 0 09
{min. = n = Amin. n (approx.). (xxiv)
This result, compared to equation (xvi), gives
n
Ymin. = at = woAmin. = gmin. (approx.). (xxv)
With equation (xxv) there is a check for the three results:
(a) ymim. determined by experiment and graphically.
(b) a-r determined by experiment.
(c) co Amin. determined by experiment.

Time-.

Fig. 6. Deviation of the cupula on the torsion swing in the neighbourhood of the minimum. Only
the tops of the declining sine wave are associated with sensation.
As the torsion swing gives an easy controllable'sensation, the C Amin. measure-
ments may be trusted far more than the ocr values, which are difficult to obtain:
Period of swing Frequency of Sensation with regard Amin. coA
(sec.) swing (sec.-') to swing (degrees) ('/sec.)
5-8 1-08 Goes ahead 1-3 1-4
50 1-25 Equal 0-8 1.0
40 1-57 Lags behind 07 1.1
average 1-2
These results were obtained in a verysensitive subject. For him, ymmn. =1-1°/sec.,
ocr= 1-50/sec. and wA = 1.20/ sec.
His natural frequency oo= 1-25 is higher than usual (1.0) but it is, neverthe-
less, a common value. On the other hand, there are sensitive persons with
coo==08 sec.-'. Thus far the theory contains no contradiction.
Test 3
It ought to be possible to administer, directly after the end of the clinical
turning test, an angular acceleration which keeps the cupula in its deviated
position, from which it would have retreated to zero in the case of a single
impulse. The impulse throws the cupula to
fmax. = y
12 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
and an adequate acceleration keeps it there. Then

E)= Ej V11
and 0 0=Yn or -=- sec. (xxvi)
In practice an acceleration is chosen and the impulse is adapted to it. The
subject has the sensation of increasing, constant or decreasing angular velocity
when the impulse is too small, well chosen (equation (xxvi)) or too large
respectively.
The best way to determine equality is to find two adjacent y values, one of
which is too small, the other too large. A sensation of constant angular velocity
is not sufficient, for it might be due to a faulty observation. An example of
this experiment may be given:
Angular acceleration Impulse Impulse =y/OC
(o0/sec.2) (y /sec.) Sensation Acceleration
3*7 40 Grows 35
36
30
Constant
Decreases
-7 = 10 sec. approx.
This test confirms the value of [I/A as of the order of 10 sec.
Test 4
The nerve responses of the semicircular canal of the ray. An unexpected proof
of the theory has arisen from the measurements of Lowenstein & Sand (1940 a, b).
They cut the nervus ampularis of the horizontal canal in the ray and by
applying a micro-electrode to one of the fibres of the crista end of the nerve,
they measured the frequency of the nerve responses as a function of the applied
stimulus. Every fibre has its own resting frequency. When the cupula is at
rest, there is apparently a permanent nerve activity which is possibly the
origin of the permanent tone originating from the labyrinth. The frequency
increases when the canal is stimulated adequately (Ewald) and decreases in
the opposite case. Less than zero frequency is not possible and the sensitivity
of the canal should therefore be different for opposite directions. In the neigh-
bourhood of the equilibrium position (zero deviation) of the cupula, the sensi-
tivity will be equal for both directions, but not for larger deviations. In Fig. 7
the impulse frequencies given by L6wenstein & Sand are plotted logarithmically
with the resting frequency subtracted from the measured frequency. The result
is a straight line with a slope I/A of 10 sec., as was the case with the human
subject. A resting activity of 30 discharges/sec. was assumed here in contrast
to the value which L6wenstein & Sand measured (i.e. 25/sec.).
It follows that something must have happened to the cupula-nerve mechanism
as a result of the impact of the endolymph. It is most probable that this rise
(or fall) has existed from just after the impact.
 THE SEMICIRCULAR CANAL 13
As Liwenstein & Sand did not make use of a torsion swing, the determination
of the unknown constants has to be derived from equation (iii). From their
measurements it may be deduced that the moment when {max. is reached
should be about 0-6 sec. Thus tm.. = 0-6 sec. From equation (iii) it follows that
IH/0 has to be about 10 sec.-1 so that A/E0 = 1 sec.-2. The differential equation
becomes e+ Oe+6=0,
o
20

0~~~~~~1

1 I

Time (sec.)
Fig. 7. Response of a single receptor unit of the left horizontal canal to ipsilateral rotation at
a constant speed of 36°/sec. Ordinate logarithmic. Replotted from Lowenstein &G Sand
1940b), after subtraction of resting frequency of 30/sec. from observred frequencies.

IV'
VW

° 0-5
0
3
0.
dW
10 15,,I
10 is
Time (sec.)
Fig. 8. As Fig. 3, using the figures of Lowenstein & Sand (1940b) for the ray. x, ipsilateral;
A, contralateral stimulation. The former agrees well with the theoretical curve, but the latter
deviates considerably.

which is the same as in a human subject. This is not surprising since the
dimensions of the canals of man and ray are almost the same. The probable
error is about 30% for both constants. The uncertainty of 11/@ is the larger
as (i/A could be determined with a reasonable accuracy. The uncertainty is
therefore imposed by the 1I/E factor.
14 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
L6wenstein & Sand gave their test animals (the ray) constant acceleration
for a short period and recorded the nerve impulses. In Fig. 8 a mean value of
twelve measurements is plotted. It is reduced to an acceleration of 1I/sec.2, a
procedure which is permissible, because it alters nothing of the essentials of the
measured quantities. As far as the measurements allow us to compare, there is
no difference between theory and experiment. The contralateral stimulation is
also given. It deviates from the ipsilateral curve except for a small region at
the beginning. We may therefore conclude that the ray follows the theory of
the torsion pendulum for ipsilateral stimulation and for contralateral to
a limited extent.
Theoretical value of 1/0
When the anatomical data concerning the semicircular canal are known, the
value of 11/0 may be calculated. The moment of inertia 0 is governed by the

Fig. 9. Schematic diagram of the semicircular canal. When the endolymph moves over an angle of
5 the cupula is forced over an angle of 0; e and 1 are of the same order of magnitude.

endolymph ring, including the part in the utriculus (Fig. 9) and the diameter of
the cross-section of the ring is determined by the canal itself. Then 0 is given by
0 = 2arg2r2R3,
where R is the radius of the canal and a the density of the endolymph. 11, the
moment of friction at unit velocity, is derived from Poiseuille's law. As the
friction is dictated by the canal and not by the utriculus, the length of the
damping canal will be about half the circumference of the circle, 7rR. The
quantity of fluid flowing through a canal with a circular cross-section, radius r,
length 7iR, under influence of the constant pressure P, the viscosity of the
endolymph being -1 is V_ r4P
during the period -r. When the angular velocity is 1 rad./sec. the quantity of
fluid is V,-=rr2R.
 THE SEMICIRCULAR CANAL 15
The pressure needed is 8nR2
pi P=r22
The momentum (pressure x surface x radius) is
r2ir2 R=-8vjr2R3.
87-s
This is the moment of friction at unit angular velocity H.
Thus r = 8ipr2R3.
From these values follows I _
ar2
As -9 = 0.006 (c.g.s.) at 370, r = 0-03 cm. and a = 1 0 approx., 11/= 27 sec.-1
approx.
Such a high value is never obtained with normal persons. As was to be
expected there is a slight difference between the original mathematical basis
and the experimental results. The most obvious assumption is that there is
a leak between cupula and ampulla, an assumption which is confirmed by
results obtained from insensitive subjects, who suffer from a large leak or from
such symptoms as suggest a leak. Where such a defect exists, the theory has
to be modified. The theory thus revised hints at deviations from the ideal case
with a close-fitting cupula, the very deviations which are found in practice in
insensitive subjects, where there is no close-fitting cupula, but a degree of leak
in every ampulla. The cupula-endolymph system is neither a pure torsion
pendulum nor a Mach-canal. Sensitive subjects have almost perfectly fitting
cupulae and the insensitive less well-fitting ones. A large leak means a gap of
about 01 mm. at most, whereas a close-fitting cupula has a clearance of not
more than 0*01 mm.
Artificial leak. The cupula is only a weak valve. It may be deformed by
a small force, i.e. an impact of the endolymph during the clinical turning test.
It will then (a) permit an increased leak, and (b) tend to regain its shape.
(a) The artificial leak may be demonstrated by comparing the slope of the
graph of subjective angular velocity against time before and after a large
impulse. In Fig. 10 impulses of 45°/sec., 180'/sec. and 45°/sec. have been
administered to the subject; thus the first and last are of the same magnitude.
The second impulse has deformed the cupula, causing it to leak. This is expressed
in the smaller - H/A value (2-8), whereas H/A is 6-2 before deformation. Where
the deformation is not too severe, the process of restoration will last only 3 hr.
In severe cases it will take weeks, or even months before the original value is
regained. This has been observed by us in several cases. This is why impulses of
not more than 60'/sec. average have usually been used in this work. In
sensitive subjects they do not exceed 450/sec. and in insensitive cases 900/sec.
16 A. A. J. VAN EGMOND, J. J. GROEN AND L. B. W. JONGKEES
As initial test impulse, 30°/sec. is used. A sensation lasting about 20 sec. may
then be expected for sensitive persons, whereas 6 sec. is a common value for
insensitive ones. The result then obtained in the unknown subjects decides on
the procedure to be followed.
(b) The deformation of the cupula is in most cases only temporary. There is
some reason to believe that the process of restoration begins immediately after
the deformation.
The impact of the endolymph deforms the cupula so as to compress one side
of it whilst the other is overstretched. In con-
sequence, the sensory hairs terminating in fine 100
canals in the cupula on one side retreat from i
the canal wall and on the other are pressed r4
-
/
further into the canals. The recovery of the /
cupula restores the hairs to their original . I I
attitudes by a series of successive stretchings, 10 -
and so brings about secondary after-sensations. ,WV I/
These have nothing to do with a normal after- Iv ,
sensation which follows an exponential law. E /
The curling of the cupula does not follow a
fixed law; nor do the secondary after-sensations.
Usually they are alternate, but sometimes two
consecutive sensations have the same direction. 0 10 20 30
This is a strong argument for the assumption Time (sec.)
of the peripheral origin of these secondary Fig. 10. Time-course of sensation be-
after-sensations. Furthermore, the secondary fore and after Ba'rany test, showing
after-sensations occur only when the cupula slow restoration of cupula. There is
considerable diminution of sensi-
is deformed (after impulses > 60°/sec., not after tivity even after 4 days. before;
smaller ones). In addition, secondary after- 1 hr. after; -- 4 days
sensations only occur with sensitive subjects. after.
All secondary after-sensations are not peripheral, though most are.
The foregoing arguments have led to the abolition of the Barany test in our
clinic, a test which may be said to yield at best values concerning a deformed
system and never any information on the original physiological properties of
the observed canal.
SUMMARY
The semicircular canal with its cupula-endolymph system is best considered as
a torsion pendulum with a high degree of damping. There is always a slight leak
between cupula and ampulla, which causes deviations from the theoretical
behaviour described by the solution of the differential equation of the torsion
pendulum. It is possible to calculate the movements of the cupula, taking the
leak into account, and the results then correspond to those obtained from
insensitive subjects. It is not asserted that the degree of leakage decides
 THE SEMICIRCULAR CANAL 17
whether a subject is sensitive or not, but it is a likely explanation; though
central causes cannot be excluded.
The calculations show that the semicircular canal acts as an 'impulsometer'.
For physiological movements it is a most appropriate instrument, comparable
to a flux-meter (ballistic galvanometer) used for the measurement of electric
charges. It normally causes no after-sensation. Excitation of the canal gives
rise to a very accurate subjective estimate of the angle of turning, provided that
the movement lasts not more than 3 sec. (Mulder's law). If a turn lasts too long
or has a period of constant velocity, the cupula no longer acts as an impulso-
meter, but is forced back to equilibrium by its own directional property.
A decelerative phase following one of constant velocity not only annihilates the
deviation, but drives it past equilibrium in the opposite direction, from which
the cupula will slowly return, giving rise to an after-sensation. In the clinical
turning test this is done intentionally.

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Buys, E. (1940). Ann. otolaryng. 3/4, 109.
Buys, E & Rylant, P. (1939). Arch. int. Phy8iol. 49, 101.
Egmond, A. A. J. van, Groen, J. J. & Jongkees, L. B. W. (1948). J. Laryng. 62, 63.
Egmond, A. A. J., Jongkees, L. B. W. & Groen, J. J. (1943). Ned. Tijdschr. Geneetk. 87, 1793.
Gaede, W. (1922). Arch. Ohr.-, Na8.-, u. KehlikHeilk. 110, 6.
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Mach, E. (1875). G9rundlinien der Lehre von den Bewegungsempfindungen. Engelman, Leipzig.
Mulder, J. (1908). Thesis, Utrecht.
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PH. CX. 2