Sunteți pe pagina 1din 16
Analysis of the Swimming of Microscopic Organisms Author(s): Geoffrey Taylor Reviewed work(s): Source: Proceedings of

Analysis of the Swimming of Microscopic Organisms Author(s): Geoffrey Taylor Reviewed work(s):

Source: Proceedings of the Royal Society of London. Series A, Mathematical and Physical

Sciences, Vol. 209, No. 1099 (Nov. 22, 1951), pp. 447-461 Published by: The Royal Society

Accessed: 20/03/2012 16:06

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.

information about JSTOR, please contact support@jstor.org. The Royal Society is collaborating with JSTOR to digitize,

The Royal Society is collaborating with JSTOR to digitize, preserve and extend access to Proceedings of the Royal Society of London. Series A, Mathematical and Physical Sciences.

http://www.jstor.org

Analysis of the swimming of microscopic organisms

BY

SIR GEOFFREY TAYLOR, F.R.S.

(Received 25 June 1951)

in air or water make use of inertia in the surrounding

the fluid backwards, while the resistance of the body gives

the

propulsive organ and the resistance can be thought about as acting separately. This conception

cannot be transferred to problems of propulsion in microscopic bodies for which the stresses due

No case of self-

propulsion in a viscous fluid due to purely viscous

to viscosity may be many thousands of times as great

Large objects which propel themselves

fluid. The

the fluid a forward momentum.

propulsive organ pushes

The forward and backward momenta exactly

as those due to inertia.

and A the wave-length. a viscous

through

and

stress

in the

tail can move a body The energy dissipation

balance, but

forces seems to have been discussed.

displacement

The motion of a fluid near a sheet down which waves of lateral

is described.

of propagation of the waves.

seems to explain on reaction

calculated.

work

relying

The

are propagated

tail

are also

It is found that the sheet moves forwards at a rate 27'2b2/A2

Here b is the amplitude

how a propulsive

inertia.

due to

is extended

times the velocity

This analysis

fluid

without

to explore the reaction between the tails of two neighbouring

tails

in the fluid between them than when the waves

of the wave in one tail lags behind

reaction, due to the viscous stress in the fluid between them,

It is found that if the waves

phase

down neighbouring

It

is in fact

observed that the tails of

small organisms with propulsive tails.

are in phase very much less energy is dissipated

are in

opposite phase. that in the other there is a strong

which tends to force the two

spermatozoa wave in unison when they are close

It is also found that when the

wave trains into phase.

to one another and pointing the same way.

INTRODUCTION

The manner in which a fish swims by causing a wave of lateral displacement to travel down its body from head to tail seems to be understood through the work of James Gray and his colleagues. This movement gives rise to circulations round the

body which, in a fluid of small viscosity like water, are necessary to produce a forward

force by dynamical reaction.

itself entirely to the inertia forces set up in the surrounding fluid by its muscular

movements. Viscosity is important only in so far as it plays a part in the mechanics

of the boundary layer, which in turn plays a part in determining the magnitude of the circulations with which the inertia reaction of the water is associated.

In other words, the creature owes its ability to propel

The propelling organs of some very small living bodies (spermatozoa for instance)

by

sending waves of lateral displacement down a thin tail or flagellum. The direction of movement of the organism is, like that of the fish, opposite to that of propagation of the waves of lateral displacement. The dynamics of a body as small as a sperma-

tozoon-say 5x 10-3cm. long with a tail 10-5 cm. diameter-swimming in water

must, however, be completely

istic length defining the size of a body moving in water with velocity

densityp, and viscosity It, the Reynolds number, R = L Vp/ju,expresses in numerical

form the order of magnitude

bear a superficial resemblance

to those of fish, in that propulsion is achieved

different from that of a fish. If L is some character-

of the ratio

V, in a fluid of

stress in fluid due to inertia

stress due to viscosity

Vol. 209.

A.

[

447

]

29

448

Sir Geoffrey Taylor

In most fishes R is of order many thousands, in a tadpole it is perhaps of order 102,

and in bodies of the size of spermatozoa it is of order 10-3 or less.

therefore, that the forces due to viscosity,

comparison with inertia forces, in studying the motions of fish, may be thousands of times as great as the inertia forceg in the case of the smallest swimming bodies.

Reynolds number is usually defined in relation to a body moving steadily through

It will be seen, be neglected in

which may legitimately

a fluid with velocity V. In cases where the body is vibrating the inertia stresses arise from the reaction between the vibrating surface and the surrounding fluid. The number which corresponds with Reynolds number in describing the order of

magnitude of the ratio stress due to

stress due to inertia .

inetia

stress due to viscositys

nL2p

where n is the frequency of vibration. In the case of a spermatozoon n is of order

100 c./sec., and Iu for water is 10-2. The length which is of importance in consider- ing the stress in the fluid is the diameter of the tail rather than its length, so that L is of order 10-5 cm. and nL2p/,u is of order 10-6. In considering the motions of spermatozoa therefore it is necessary only to take account of viscous forces. Inertia

forces may legitimately be neglected. These considerations naturally give rise to the following question. How can a body propel itself when the inertia forces, which are the essential element in self- propulsion of all large living or mechanical bodies, are small compared with the forces due to viscosity? An attempt will be made to answer this question by showing that self-propulsion

which

is possible in a viscous fluid when bodies immersed in it execute movements

bear a strong resemblanceto those which spermatozoa are known to make.

Self-propulsion in a viscous fluid

The

only problems

concerning the motion

of solids in viscous fluids which have

so far been solved relate to bodies which aremoved by the application of an external

forcelike gravity. The motion of spheres and ellipsoids in an infinitely extended fluid under external forces or couples has been analyzed. It has been found that such

bodies tend to move along with them a very large volume of the surrounding fluid. Long cylindrical bodies move so much fluid that the whole volume, extending to infinity, moves with the body. The fact that a cylinder in steady motion gives rise to finite fluid velocity at an infinite distance was discussed by Stokes (i85 ) who also obtained the solution to the problem presented by an oscillating cylinder in which

inertia stresses are comparable with those due to viscosity. He pointed out that as the frequency of oscillation decreases the volume of fluid which moves with the

cylinder increases till, as the frequency approaches zero, the disturbance tends to extend to infinity.

When

large

bodies

like

ships

or aeroplanes

are propelled by some internal

mechanism through a fluid, the mechanics of their motion is always analyzed by

consideringseparately (a) a propelling mechanism like a paddle wheel or airscrew

which develops a forward force by pushing fluid backwards, and (b) resistance which

arises because the body entrains some of the surrounding fluid and thus gives it

Analysis

of the swimming

of microscopic organisms

449

a forwardmomentum.When the self-propelledbody is moving-at a steady pace it is clear that the backward momentum considered under (a) exactly balances the forwardmomentum of (b).

When a body propels itself in a viscous fluid it is still true that the total rate of production of momentumis zero. In other wordsthe resultant force which the fluid exerts on the body must be zero. On the other hand, it is clear that when inertia

stresses are negligible

to use the conception of propulsion as being due to the separable effects of a pro- pulsive unit and fluid resistance. The truth of this statement is at once obvious if

the body consideredis two-dimensional, in the form of an infinitely long cylinder, for the effect on the fluid of moving the cylinder-considered independently of the propulsive unit-would, as Stokes showed, be to move the whole fluid in which it was immersed.Thereis no reasonto suppose that a self-propellingbody would move

a great volume of the

solution of which fills most of this paper, the influence of a self-propellingbody

extends only a very short distance from it.

compared with those due to viscosity it is no longer possible

fluid surroundingit, in fact in the particular problem the

Provided that no attempt

is made to separatepropulsion from resistance, but the

motions of the whole fluid and the body are considered as inseparable, Stokes's difficulty disappears. Though microscopicswimming creaturesare certainly three- dimensional, yet the great simplicity of two-dimensional, compared with three- dimensional analysis, makes it worth while to discussthe problem of self-propulsion

in a viscous fluid in two dimensions.

The propellingorgan of a spermatozoon is a thin tail down sendswaves of lateral displacement. Whetherthis tail moves in

sionsis not clear (Rothschild195I). The analogous two-dimensional problem is that of

a sheet down which waves of lateral displacement are propagated. This problem will be investigated with a view to finding out whethersuch waves can give riseto viscous stresses which drive the sheet forwards.

Waves of small amplitude in sheetimmersedin a viscous fluid Takeaxes whicharefixedrelativeto the mean position of the particles of the sheet. The waving surfacewill be taken representedby

which the organism two or three dimen-

(1)

The velocity of the wave is or/k and it moves in the directionx positive. The wave-

b, the amplitude, will be assumed small

length is 27r/k= At represents time.

compared with A. If the sheet is inextensible and the amplitude of the wave small,

material particles will oscillate in a path which is nearly parallel to the axis of y,

are narrow figures of 8. The com-

ponents of velocity of a particle of the sheet are u0,v0where

though, as will be seen later, their actual paths

yO = bsin (kx - ct).

Uo=

,

v V =

ay0

= -bo

cos (kx -

t).

(2)

The problem is therefore to find a motion in a viscous fluid which satisfies (2) as

a boundary condition on Y0 = bsin (kx - ct).

in two dimensionssatisfies when inertia is neglected is

V4-1 = o,

(3)

The field equation which viscous flow

29-2

450

where

Sir Geoffrey Taylor

fris a stream function and the components of velocity are

aVr

-

ay

aVr

9ax

u=

V-=

 

(4)

Vy.

(5)

As a first approximation when bk is small assume for / -

This function satisfies (3). The velocity of the fluid at infinity is V, so that if V has a finite value the particles of the waving surface will move relative to the main body of viscous fluid with velocity - V. The conditions to be satisfied at the surface Yo = b sin (kx - at) are

f

= (Aly + B1) e-ky sin (kx - at)

-x = v

ax

= -ba

cos (kx-

t),

-

=0

ay

=

0.

(6)

To the first order (when bk is small) the values of u and v at y = 0 will be the same

as those at y = bsin (kx -

t), so that the boundary conditions

(6) are satisfied if

-V

+ (A1-Blk)sin

t) =

and

(7) and (8) are satisfied if

B, k cos (kx -

(kx -

vo =

-b

t) =

-u

cos (kx -

V=0,

A1 =B1k=-ba.

=

0

t).

(7)

(8)

(9)

Inserting values of A1 and B1 in (5) it is found that

1 =

-

-k (

+ ky)e-k

sin (kx-

t),

(10)

and rf represents the flow near a sheet down which waves of small amplitude are

It will be noticed that since V = 0 the waves in the sheet do not propel it

through the fluid. This conclusion, however, will be modified when the equations are treated, using a higher order of accuracy than that which led to (7) and (8).

The dissipation of energy can be found by calculating the work done per unit area of the sheet against viscous stress. Its mean value is

travelling.

W =

dyo

dt y

(11)

where Yy is the stress normal to the sheet and (Lamb 1932)

,

=-p

+2u

+

dv

(12)

Here -p is the mean value of the principal stress components. p is described as pressure. The pressure associated with the stream function (10) is

(13)

p = 2abkc e-ky cos (kx - at),

and since at the surface u = 0, au/lx

=

0 so that

av/ay = 0. Hence

W = 2b2ro2ktzcos2 (kx - at) = b2o2kCt.

(14)

Analysis

of the swimming

of microscopic organisms

451

Since the motion vanishes at infinity, it is clear from what has been said that the

total force on the sheet must be zero. In fact the forward component

which the pressure exerts on the plate is

dyo

(15)

of the force

1 = Px

dX -?Urb2k2. -

This is negative so that the pressure tends to drive the sheet in the direction - x. The

To the first

stress on the sheet is

force, due to the tangential component of stress at any point, is 1(au/ay).

order of small quantities the mean value F2 of the tangential

zero, but it must be remembered that the tangential

wavy surface yo = bsin (kx- ot). Thus taking the variation in tangential stress due

to this fact into account, the mean stress is the mean value of .(Su/ay)

surface is

(16)

putting y = b sin (kx - crt) in (16), and remembering that sin2 (kx-

(17)

stress actually

acts over the

over this

=

F2 = ,uobk(ky-

1)e-ky sin (kx-ot);

ot)

F2 =- /Cb22.

The total mean force per unit area exerted by the fluid on the surface is F1+ F2.

From (15) and (17) it is seen

general principles.

Propulsive

that F1+

F2 = 0, a result which was anticipated

on

effect of waves which are not small

It has been shown that waves of small amplitude travelling down a sheet do not give rise to propulsive stresses in the surrounding viscous fluid. It is now proposed to discuss the effect of waves whose amplitude is not so small that terms containing

b2k2can be neglected.

waves whose amplitude is unlimited, but it is possible to consider the effect of waves

of finite amplitude by expanding the various terms in the mathematical expressions representing the disturbance produced by the waving sheet in powers of bk. This

expansion will be carried to include terms containing powers of bk as high as (bk)4.

To simplify the analysis the equations will be written in non-dimensional

taking k = 1. If z is written for x - ct the appropriate form to assume for ??

It does not seem to be possible to discuss by analytical methods

form by

.

r is

(18)

1 -

t

oo

E

n even

Vy

f'

=

E

Xnodd

(A,y+

B)e-

sinnz+

(Cy+

D,)e-ncosnz-

This satisfies V4f1 = 0, and the disturbance rapidly decreases with distance from the sheet. The term Vy/lo is again inserted to allow for the possibility that the waving sheet may move relatively to the fluid far distant from it with velocity - V.

Boundary conditions

It will be assumed that the form of the sheet is

Yo = bsin z,

(19)

even when b is not small. The boundary condition which must be satisfied by the fluid in contact with the sheet is that there is no slip at its surface. The fact that the

452

Sir Geoffrey Taylor

sheet is in the form Yo = b sin z controls the component of velocity normal to its surface, but some further physical assumption must be made about the sheet before

the component parallel to its surface can be expressed in mathematical assumption will be that the sheet is inextensible.

form. This

Velocity of particles in an inextensible sheet disturbed by transverse waves

shape can be reduced to rest by

imparting to the whole fluid a velocity -cr. The velocity of particles of an in-

extensible

The velocity

of the waves is o. Their external

sheet moving along the fixed curve y = bsin z is

Q (length of a curve in one wave-length(20) one wave-length

This ratio is

Expanding

1

r2"

(1

(21) in powers of b up to b4

1_= l

b2-cos2z) dz.

b2--d3b4.

(21)

(22)

The velocity the waves are

components of particles in the sheet relative to axes which travel with

 

ul=-Qcos

,

vl =-QsinO,

where

tan 6 = dy dz

=b cos z.

(23)

(24)

After some reduction it is found from (22), (23) and (24), retaining all terms up to those containing b4,that

u+l

1

-=

b4 + (ib2-

_

b4) cos 2z- Ab4 cos 4z,

(25)

!-

-

(b -

b3) cos z - (gb3) cos 3z.

(26)

Since u1+ o-,v, are the components

to the original axes the boundary conditions for bfare

of velocity

of the particles of the sheet relative

_1[

]-

=_

-3-

Lay y-=b slnz

b4+(ib2-_b4)cos2z-b4cos4z,

(27)

_[W~]=]

JO'

y=b

sin z

= -

(b -

X

b3) cosz- -b3 cos 3z.

(28)

It will be noticed that if only terms containing b are retained the particles oscillate in the lines parallel to the axis y. If terms containing b2and b are retained particles of the sheet traverse paths in the form of figures of 8.

on the boundary. For this purpose

it is convenient to expand ?f near y = 0 in powers of y. Thus

It remains to find the values of 8f/a8z and a/lay

(Any+Bn)e-

B=

+ (A,

- nBn)y+

(Ay+Ben2\

.

-nAy

)

+\

y

++

2

2

n

An-By+

yX

(

+

3 +

Analysis of the swimming of microscopicorganisms

and

(Any+ne- = An-nB+2(-nAn+

At y = y0= bsinz

Bn)Y+3

An-

! B)y2+

a B + y(2A

+ {C2 -

22

+ yo( -

+ B)+

y2(A-iB

4C2 + 4D ) + y2(62

1)y3

2(-AA + B1)}sin z

-

4D2)} cos 2z

453

+ {A3 - 3B3 + y(-

6A3 + 9B3)}sin 3z

+ {C4- 4D4} cos 4z-

V/,

(29)

ar

=[B1 + yo(Al

B1)+ yo(-A1 + ?B1) + y3(JAI -

B1)] cos z

+

+ [Bg + yo(A3 - 3B3)] (3 cos 3z)

+ D4(-4sin 4z).

[D+ yo(C2- 22)

+ y( - 2C2+ 2D2)](-

2 sin 2z)

(30)

In order that the boundary conditions may be satisfied for all values of z it is

)mz in (29) and (30) in the form EA,( . )lz, I

necessary to express terms like y sin .

being an integer. The coefficientsof

in the expressions for the boundary

conditions (27) and (28) may then be equated. The expressions necessary for

developing the expansionsup to terms containing b4are given in table 1. table, the coefficients given in table 2 may be equated to zero.

\sin (i?lz

Using this

TABLE

1.

RELATIONS

NECESSARY

FOR EXPANDING

BOTNDARY

CONDITIONSIN POWERSOF b UP TO b4

Yo = b sin z

yo sin z =

Yocos

yosin2z =

jb(l -cos

2z)

z = lb sin 2z

= b(sin

b(cos z -cos

3z-sin

b(cos 2z - cos 4z)

3z)

z)

yo cos 2

Y0 sin 3z =

Yocos 3z = lb(sin 4z-sin

2z)

y

yo sin z = b2(3 sin z -sin

yc

ysin

3z)

b2(cos

- coscos 3z)

4z)

2z = Jb2(2 sin 2z-sin

cos 2z =

b2(_

+ 2 cos 2z-

cos 4z)

y

y

sin z = =

cos

b3(3 - 4 cos 2z + cos 4z)

-

b3(2 sin 2

sin 4z)

TABLE 2.

COEFFICIENTSTO BE EQUATEDTO ZEROIN THE DEVELOPMENTOF (27)

AND (28)

1

sinz

cos 2z

sin 3z

cos 4z

cos z

sin 2z

cos 3z

sin 4z

( -

(A

-

2A

+ Bl)? b +

1

( -

- B)+()+ (A

(-

2A

+ B)

-B)

lb -

A1 Ax2 + -B

~(6) -

-2)

b2 -ib(-

4C2 + 4D2)

1b3(- wA, + -B1)

+i

-

2

+

-

3(6-4D)

+(

6A.+ 9B3) + b

- b2(iAI

- 1BB) + (-

4C2+ 4D2) Ib + A3- 3B3

 

+ ib3(-

A1 + iBI) - i2(C2

- 4D2) -

b( -

6A3 + 9B3)+ C - 4D4 -

3-b4

-

b4

(a)

(a)

(b)

(c)

(d)

(e)

B1 +,

Ib(Ax-

b2(-A1 + IB )-

B1) +

(IA1 -

2(C - 2D2) b

B1) {b3 -

2D2 -

+ b-

b3

(f)

b2(

2C2 + 2D2) -

-b(A3 -

3B3)

(g)

-

-

b2(

A1 + B1) + b(C2- 2)2) + 3B3 +

b3

8b3(-A1 - WB1) + b2( -C2

+ D2) + 1b(A3 -

33)

-

4D

(h)

(i)

454

It will be noticed D2, A3 and B3 may

equation

noticed that in order that (b) and (f) may be satisfied it is necessary that B1 shall be of the form

A1 = - b+ higher powers of b

and

In fact, in order that the six equations may be satisfied for all values of bit is necessary that A1, B1, C2,D2, As and B3 shall be of the form

Sir Geoffrey Taylor

that C4 and D4 occur only in (e) and (i). The constants A1, B1, C2,

be obtained

by equating

(b), (c), (d), (f),

(g), (h) to zero. The

V has a particular value.

It

will be

A1 and

(a) can then only be satisfied when

B1 = -b + higher powers of b.

A1=-&b(l+ab2),

D2-=

B=-

b2+5'2b4,

b(l +,b2),

A,3=eb3,

C2= yb2 +72b4,

B 3-b3.

It remains to determine o, fi,Y,,y7,

and (h). It can be verified that the appropriate values are

81,

22, e, y from the equations

(b), (c), (d), (f), (g)

X-2

fi-=,

t

72--6

1-

2-12,

=O=?

Inserting these in (e) and (i) it is found that

04=

292b4 D4=

b4.

Propulsive effectof propagating transverse waves in a sheet (a) may be written in the form

-=

b2+----(

o-fl+33-1

) b4.

=-12.

(31)

(32)

(33)

Hence from (31)

-

= lb2(1-9b2)

In the non-dimensional

units the velocity

of the wave relative to the particles of

the sheet is o-.When dimensional units are used (33) is written

= 1b2k2(l1-tIb2),

or if the velocity of the waves of lateral displacement relative to the material of the

sheet is V

V

27T2b2

19 72

(34

U A2

4

A2(34)

V is the velocity

to the material of the sheet.

V is positive the sheet moves with velocity - V relative to the fluid at infinity

of the fluid at infinity

relative

Since

when waves of lateral displacement

travel with velocity

+ U down the sheet.

Viscous fluid on bothsides of the sheet

In the foregoing discussion the reaction of the viscous fluid on one side only of the

waving sheet has been considered. In applying the results to the swimming of microscopic organisms it is necessary to suppose that the sheet is in contact with the fluid on both surfaces. In that case for a wave of given amplitude the sheet will move

Analysis of the swimming of microscopic organisms 455

relative to the fluid at infinity at the same speed V that has been calculated when fluid on one side only was contemplated. On the other hand, the rate of dissipation of energy is 2W instead of W where W has the same meaning as in (14). It has been proved therefore that when small but not infinitesimal waves travel down a sheet immersed in a viscous fluid they propel the sheet at a rate which is 27F2b2/A2 times the wave velocity and in the opposite direction to that of propagation of the waves. It would have been less laborious to calculate only the first term of the expression for V/U. The second term containing the factor b4/A4 was calculated in order to form some idea of how large the amplitude might be before a serious error might be expected in the analysis. The outside limit at which the formula might be expected to give reasonably accurate results would be when the second term was, say, one-quarter, as big as the first. That is when

In that case

head (symbolic)

= --

V (/-U =

= 0.073.

) =

0 079.

FIGURE 1. Symbolic representation

tail when kh= 025,

of microscopic organism swimming.

Shape of waving

b/A = 0-073; ->, direction of propagation of waves in tail.

The shape of the tail in this case is shown in figure 1. A tail of the shape shown in

figure 1 would have to oscillate 1/0.079 = 12*7 times in

length. It will be noticed that the wave shown in figure 1 is not very large. It may

well be that waves of larger amplitude would propel the sheet more than 1/12.7 of a wave-length per oscillation, but the method of analysis here adopted could hardly be used in discussing such a case without great labour. The Southwell's relaxation

order to progress 1 wave-

technique might perhaps be employed.

Stress in the tail The internal mechanism necessary to produce lateral motion can only be due to tensions and compressions acting across each normal section of the tail so as to produce a couple M. This couple varies along the tail. Its magnitude can be calculated when the distribution of pressure along the tail is known. In the case of a waving

sheet which has fluid on both sides the equilibrium equation is

dM=

- dF=

-

dx

dx

(35)

where P is the difference of pressure on the two sides of the sheet. The pressure variations are equal in magnitude but of opposite signs on the two sides of the sheet. Equation (13) therefore gives P = 4o'bkl/ cos (kx - at)

456

so that (35) becomes

Hence

Sir Geoffrey Taylor

d2M

2

X2

=-

= -

4obk

b

k

cos (k

cos (kx -

-

t).

The maximum value of M is

4orb#

or

4nb,uA,

t).

where n is the frequency of vibration'of the tail. The magnitude of the maximum stress can only be calculated if the thickness d of the tail is known. The minimum possible value of the maximum stress is then

4Ml

d2

or

Taking the case when A = 10-3cm., this stress is 2 x 104or 20g. weight/sq. cm.

I

=

16#ubnA/d2.

10-2,

b -=

A, d

10-5cm.,

n=

50c./sec.,

Mechanical reaction between neighbouring waving tails It has been observed that when two or more spermatozoa are close to one another there is a strong tendency for their tails to vibrate in unison. James Gray (1928) writes: 'Numerous authors have observed that when the heads of individual sperma- tozoa are in intimate contact their tails beat synchronously and a very striking

example of the phenomenon can be observed in Spirochaeta balbianii.' Figure 2, which is reproduced from figure 78, p. 119 of James Gray's book Ciliary movement (1928), shows his idea of the way in which aggregates of these organisms which vibrate in unison are formed. Rothschild (I949) attributes certain comparatively

large-scale motions in dense suspensions

of bull or ram spermatozoa

to 'periodic

aggregation of spermatozoa the tails of which probably beat synchronously

aggregations'.

in the

FIGTURE 2.

Spirochaeta balbianii formingaggregates, the individualsin which soon

(Reproduced from Gray'sCiliarymovement.)

establish synchronous movements.

Among the various possible explanations of this phenomenon it might be supposed that the stresses set up in the viscous fluid between neighbouring tails may have a component which would tend to force their waves into phase. It is of interest

Analysis of the swimming of microscopicorganisms

457

therefore to analyze the field of flow between two waving sheets when their waves are not in phase in orderto find out whetherthe viscous stressesare of such a nature as to tend to force them into phase. Taking axes of co-ordinates midway between the two sheets which are at y = + h

it will be assumedthat waves of the same amplitude, b, travel down each sheet with

the same velocity crjk.It will be assumedalso that the phase of the sheet at y = + h

lags behind that of the sheet at y = - h by an angle 25. All cases will be covered if

q is taken

to lie in the range 0 < 0 < ffr.

The equations to the two sheets are then

y = h+yt

and

where Figure 3c shows the sheets when

y= -k+y2

2h I

= h+bsin(z+),

z

=-h+bsin(z-0~),

kx - ct.

= =4545? so that y, lags 90? behind y2

a

b

WYz~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~1

-

(36)

FIGURE 3. a, waves in phase,

0

=

b/A = 0*073, 2h/A =

3-. c, wave

of wave propagation.

0, b/A = 0.073, 2h/A = |.

Y1 lags behind

b, waves in opposite phase,

= Sr,

y2, A= qTr. The arrows indicate

the direction

458

Sir Geoffrey Taylor

The stream function is assumedto be ~r = (Aly sinh ky + B1cosh ky) cosq sin z + (A2y cosh ky + B2 sinh ky) sinq cosz. (37)

The condition

r/lay = 0 at y = + h is satisfied provided

B

A1

=

-(kh coth kh+ 1),

A

A2

-

(khtanh kh+ 1);

(38)

the second condition to be satisfiedat y = + h is

and at y = -his

a_ay, atez

ax

az 8t

ar=-'

ax

Both these are satisfiedif

ayl

az'

a=-

Alh sinh kh+ B1cosh kh = -bo/k,

From (38) and (39)

A2hcosh kh +B2sinh

kh = - bo/k.

1 =

ba sinh ckh

csinhk

h

h

'A

bcoshh

2 sinh khcoshkh-kh'

bcr/kh coshkh+ sinh kh\

2

khcoshbbkhkh hsinhsh

cosh kh + kh

+ cosh

k

coshkh-khJ5sinhh

(39)

(4

(40)

It is now possible to calculatethe stress which the viscous fluidexerts on the sheets. The componentperpendicular to the sheet is as in (12)

Y=

-p+2#

=-p.

(41)

The pressurep corresponding with the stream function (37) is given by

p _ sinh ky sinh kh cos 0 cos z

2/1kb -

sinh kh cosh kh + kh

At y = h the pressure is Pi, where

cosh