Einstein and 1905

(2005 was Einstein Year – celebrating 5 key papers published

that year.)
• In 1905 the atomic hypothesis was
not fully accepted.
• Despite Brownian motion having
been known about for 75 years, its
significance was not appreciated.
• The kinetic theory of gases was
thought of as a 'mechanical
analogue', but implied reversibility.
• The 2nd law of thermodynamics
required irreversibility.
• Einstein understood that taking a
statistical approach, and assuming
atoms existed, reconciled the paradox.
• His paper, the second of the 5, was
written in April 1905.
• A macroscopic particle – such as the
pollen particle – would be buffeted by the
atoms/molecules in the surrounding water.
• The particle would undergo diffusion and
measuring the diffusion constant (or
equivalently the displacement) should
show an increase with t (not linearly).
• Perrin's subsequent experiments on
sedimentation showed how all this hung
together.

1
 Brownian Motion
Nelson; Dill and Bromberg
• Brownian motion of a particle arises due
to its constant bombardment by
molecules(e.g as in the first observation
by Brown in 1827 of pollen grains by
the water molecules).
• Net force, averaged over time, on the
particle is zero.
• But at any moment there is a constantly
fluctuating net force and this gives rise
to the observed Brownian motion.
• The motion of the particle follows a
random walk
• For a sphere of radius r in a liquid of
viscosity , we have seen from Stokes
Law the force is 6rv
• This can also be written in terms of the
drag coefficient, defined so that
• Force = drag coefficient × velocity
= 6r
• Equation of motion for particle of mass m
d 2R dR
m 2   Frandom
dt dt
where R(t) is the position coordinate, and
Frandom is the random force due to
collisions.
2
 Brownian Motion
Jones
• Forces are random so each • We can also use the identity
direction behaves in the same way 2
d  dx   dx  d 2x
and x     x 2
dt  dt   dt  dt
 x 2  y 2  z 2 
2 2 • Thus
 R  3  x  2
2
•
d  dx   dx   d ( x )
Work in 1D for simplicity m  x   m    xFrandom
dt  dt   dt  2 dt
d 2x dx
m 2   Frandom
dt dt • Rearranging and taking an average
yields
• Use the substitution
 d  (x2 ) 
d (x2 ) dx 
 2x 2 dt
dt dt d dx
2

 dx 
xF  m x    
random  dt dt  dt  
 
• Then d 2 x  d (x2 )
m 2   Frandom
dt 2 x dt
3
 Brownian Motion cont
 d  (x2 ) 
 • And hence the total squared
2 dt
displacement in 3D is
d dx  dx 
2

xF  m x    
random  dt dt  dt   6 k BT
 
R 2
3 x 2
 t
• The direction of the random force on 
the particle is not correlated with the
• The motion is diffusive with a
particle's position, so the first term is
diffusion coefficient given by the
zero. Similarly, there is no
Einstein relation
correlation between the particle's
position and its velocity, so the D=kBT/
second term is also zero.
• The third term can be rewritten via
• We will explore the implications of
the theorem for equipartition of
this later, but note the inverse
energy, since
relation between drag () and the
• mvx2/2=kBT/2 or vx2=kT/m diffusion constant D.
d x 2  2 k BT
 
dt  4
 Aside on Ensembles
(for the theoretically inclined)
Waldram:The theory of thermodynamics

• On the previous slide we saw averages
<….>.
• What are we averaging over?
• is the same as the average over time.
Those of you who took TP1 will
probably assume this is a time average
– and it may be.
• But the ensemble average can also be
over a set of replicas at a given instant.
• ensemble average, which could mean
In your Thermal Physics course you
have been taught about the different
types of ensembles.
• You have also been taught about the
Principle of Equal Equilibrium
Probability,namely that for an isolated
system, all microstates compatible with
the given constraints are equally likely
to occur.
• If the system obeys this Principle, then an
average over the probabilities of the
system being in a particular configuration
• But, formally, one should always specify
what sort of average is being taken by the
<…..>.
• Frequently is is simply referred to as an
either.
• Underlying all this is the ergodic
assumption – that the system can go
anywhere within the allowed energy range.
• In which case in equilibrium, the
fluctuation distribution is identical with the
ensemble average distribution.
5
 The Diffusion Coefficient
• When the diffusing particle is a sphere,
• For a molecule such as a protein, the
Stokes Law gives
radius R may not equate to the actual,
 =6r, so physical size.
k BT • A hydrodynamic radius RH can be
D
6r defined, which is the effective radius
presented to the fluid by the molecule.
• This is known as the Stokes- Einstein Additionally, the molecule may or may
equation. not permit the fluid to drain through it,
• For other shapes the drag coefficient, and depending on the density of chain
hence diffusion constant, take different packing.
forms. • Thus the detailed hydrodynamics of e.g.
• E.g. disc moving randomly globular proteins diffusing through a fluid
 = 12r can be complex.
• Ellipsoid (major and minor axes a and b) • We will return to these concepts when
=6a/ ln(2a/b) for random rotation discussing proteins and polymers in more
detail.
(other expressions for motion sideways
or lengthwise)

6
 Typical values for D in water

Molecule T(oC) MW D (cm2s-1)

(g mol-1)
Oxygen 25 32 2.1 x 10-5

Sucrose 25 342 5.23 x 10-6

Myosin 20 493,000 1.16 x 10-7

DNA 20 6,000,000 1.3 x 10-8

Tobacco mosaic 20 50,000,000 3.0x 10-8

virus

Quantitatively one would expect D ~ m-1/3, but because a hard sphere

model for the molecules is not accurate, this precise dependence is not
found.

7
 Diffusion Equation
Dill and Bromberg

• Fick's first law states that the flux J of • Specific solutions depend on
particles is proportional to the boundary conditions (recall your
concentration gradient. 2nd year maths!).
• e.g for point source diffusing in 1D
J   Dc along x, starting at x=0
• And conservation of particles requires
c
 .J
t
• Hence in 1D and assuming D is a
constant, then

c   2c 
 D 2 
t  x 
• which is Fick's second law in its
Numbers
simplest form.
correspond
to values of
8
Dt
 Diffusion Equation cont
• Compare with data from Wall Street
• Note that D need not be a constant,
although in many simple situations
it is.
• If it is not, then the equation must
be modified to allow for the fact
that D can be a function of
position.

Distribution of monthly returns for c     c  

   D( x)  
a 100-security portfolio 1945-1970 t  dx   x  

• Over time stock prices also exhibit a random

walk with drift!
• Individual whims can lead to the statistical
movement.
• Baseline drift comes from the fact that
overall shares do make a profit….
9
 Diffusion Control

• Diffusion may limit: 1. Diffusion of molecules which react at

– Growth of 2nd phase particles
– Supply of nutrients to organisms
– Colloidal aggregation
the surface.
If these are transformed/lost during the
reaction, then c(r = a) = 0. If the
concentration well away from particle is
c solution is

a a
c(r )  c (1  )
r
• We will model as spherically
symmetric, so work in spherical
coordinates.
• Assume steady state so dc/dt=0. • For such a concentration profile, the flux
• Diffusion equation becomes is given by
2 1 d 2 (rc ) dc Dc a
 c 0
r dr 2 J (r )   D  2
dr r
• Solution depends on boundary 10
conditions.
 Diffusion Controlled Collisions and Reactions
• Hence the number of collisions at the 2 Growth of particles post-nucleation
surface per unit time I(a) is
• In this case the molecules diffusing to
the surface are causing growth of the
I (a)  J (a)4a 2  4Dc a particle – a process known as Ostwald
• Minus sign implies flux is towards ripening or coarsening.
particle (-r direction). • Flux J(r) as before but now we have
• This is the fastest possible rate at which
any process can occur, i.e. when it is 1 dr
J (r )    
limited by diffusion. r2 dt
• If the reaction at the surface itself is rate
• Integration yields
limiting, a process will be slower.
• If the diffusing molecule is a nutrient to r3  t or
an organism, then we also have to think
about its consumption and this will be r  t 1/ 3
proportional to the volume. • This is known as the Lifshitz-Slyozov
• Thus if the bacterium etc is too large, law, and is empirically found to hold as
cannot get sufficient supply of the long as diffusion is the limiting factor.
nutrient by diffusion, and so this will set
an upper limit on size.
11
Diffusion Controlled Aggregation
• During colloidal aggregation, can have either
diffusion or 'reaction' being the limiting factor.
No salt • Which is dominant depends on the conditions
of aggregation
• E.g. in the examples shown here of 250nm
polymethyl methylacrylate particles, it is the
salt concentration which is changing.
• This changes the interparticle potential.
With salt
• The shape of the aggregates can provide
insight into the processes occurring.
• Very often fractal structures form.
• In these the structure is the same at all
lengthscales
1/ d f
R~M
Watching paint
dry; a new paint
formulation; • Where R is the size, M is the mass and the
fractal structures fractal dimension 1<df3 (equality for compact
again visible, object)
but clearly with 12
rearrangement
 Aside on Fractals
Jones
• Computer simulations show such fractals should
have a fractal dimension of 1.71 when diffusion
limited aggregation occurs.
• If clusters are allowed themselves to diffuse, the
dimension is larger at 1.78 implying a slightly more
dense structure.
• In practice, the rate at which rearrangements and
sticking occur will all affect the fractal dimension.
• But it always lies between 1 and 3.
• When aggregation occurs – of
individual molecules or particles –
the structures that result depend on
the probability of sticking
• i.e. whether diffusion or sticking is
the limiting factor.
• If rearrangements can then occur
the structure will be able to
compact.
• If not, a fractal structure results,
over a range of lengthscales from
the size of the particle up to the
size of the aggregate.
13
 Implications of the Einstein Relation
Dill and Bromsgrove

• We have derived the Einstein equation • The ensemble average of v is given by

k BT
D
 dt
• In this, the drag coefficient  (dissipation)
and the diffusion coefficient D (fluctuations
about equilibrium) are inversely related.
• We will see shortly that this is a very
general result.
• In general, if there are random fluctuating
forces f(t) and a drag force v acting
together on a particle, we can write
dv
m  f (t )   v(t )
• <f(t)> =0 since this is a fluctuating force
arising from the many collisions.
• So the solution for <v(t)> is
v(t )  v(0) exp  t / m
• So we can define a characteristic time
 = ma viscous relaxation time.

• This is known
dv as the Langevin equation • Example: For a sphere of radius 10nm
(strictlymthe A-Langevin v
 f (t ) equation, as dealing (typical size for a large polymer) in water,
dt
with acceleration). this leads to ~10-10 s.

14
 Velocity Correlation Function
(Doi, Balescu – see the latter for a rigorous and general discussion)
• More generally, look at the velocity
correlation function defined as
<v(0)v(t)>.
• Recall the <…> imply a suitable ensemble
average.
• Correlation functions in general measure
the extent to which there is any correlation
between a function at two separate times.
• It can be applied to position, force,
velocity etc, and can be defined in the
same way as for the velocity correlation
function.
• The larger the value of the function is, the
greater the extent of correlation.
• For random motion, the velocity
correlation function equals <v2(0)> for t~0,
and = 0 at large t, when all correlations
lost.
• For so-called stationary stochastic processes,
any autocorrelation function only depends on
the time interval involved
i.e <v(t1)v(t2)> = Cv(t1- t2) with Cv a function
• Thus more specifically we can write
v(t )v(0)  2 D (t )
• Where D is the diffusion coefficient.
• This can be seen by considering, the mean
square displacement (MSD) in 1 dimension
about an average position x = 0 for diffusive
motion is
t t
• x(t )   dt1  dt2 v(t1 )v(t 2 )
2
0 0
t t
x(t )   dt1  dt2 2 D (t1  t 2 )  2 Dt
2
0 0
15
 Velocity Correlation Function cont
• Returning to the Langevin equation • By equipartition of energy
1 1
m v 2 (0)  k BT
dv 2 2
m  f (t )  v kT
dt  v(0)v(t )  B exp ( t / m)
m
multiply by v(0) and take the ensemble average
Velocity correlation function
dv(t )
m v ( 0)  v(0) f (t )   v(0)v(t ) <v2(0)>

<v(0)v(t)>
dt
<v2(0)>e-t/m
• Now v(0) and f(t) are uncorrelated so that the
first term on RHS = 0, and
t
d  • The figure confirms that the longer the
v(0)v(t )  v(0)v(t )  0
dt m time interval, the less the correlation
 v(0)v(t )  v 2 (0) exp ( t / m) between v(0) and v(t) .
• For timescales shorter than the
characteristic time) the velocity retains
• So we have obtained an equation for the correlation, but not for time scales much
velocity autocorrelation function. longer than this.
16
 Fluctuation-Dissipation Theorem
Dill and Bromsgrove

• Conceptually, the fluctuation-dissipation • We have seen that the velocity

theorem states that the fluctuations in a
system are correlated (inversely) with
the energy dissipation, so generalises
our conclusion from the Einstein
equation.
• In the Einstein relation
k BT
D
 •
we see that the drag coefficient 
(dissipation) and the diffusion
coefficient D (directly related to the
fluctuations in mean position as we have
just seen) are inversely related.
• Large dissipation leads to small
fluctuations about equilibrium.
autocorrelation function indicates how fast
a particle 'forgets' its initial velocity due to
the effect of Brownian motion and
collisions leading to randomisation.
• If this timescale is long, then clearly there
is little dissipation, there are few
collisions, and equilibrium is slow to be
achieved.
Thus low dissipation means it takes a long
time to establish equilibrium.
•  is m/  and is typically in the
picosecond range for a small protein.
17
 Fluctuation-Dissipation Theorem cont
• Integrate the time correlation function over
all possible lag times t <v2(0)>

k BT  k BT

<v(0)v(t)>

0
v(0)v(t ) dt 
m 0  exp ( t / m ) dt 

D

• This integral equals the area under the

curve.
• If the area is small it implies that kT/ is
small and D is also small: the dissipation is
large, and the diffusion coefficient/
transport coefficient is small.
• Equilibrium is rapidly reached as there are
many collisions.
• And the fluctuations about equilibrium are
small, as we saw from the MSD.
t
• Conversely, if the area is large, the
dissipation is small, and the fluctuations
are large.
• So we have a statement about an
equilibrium property – the fluctuations –
related to a non-equilibrum property , in
the form of dissipation.
• The magnitude of equilibrium
fluctuations is related to how fast the
system reaches equilibrium.

18
 Further thoughts on the Fluctuation
Dissipation Theorem cont
• Further Examples of how the FDT applies
• Consider a pendulum moving about an
equilibrium position, due to fluctuations in air • Nyquist's formula in electrical circuits
compared with its motion in a viscous fluid. says that the larger the resistance (giving
• At a fixed temperature, the mean square rise to dissipation) the larger the
displacement will be the same in both cases, (thermal) noise emf present.
corresponding to the magnitude of the
fluctuation in the displacement. • Thus this theorem translates into many
• However since the damping in the second case different situations where fluctuations
is much greater, then the random forces f(t)
giving rise to the fluctuations must be greater
and dissipation are present.
too.
• An alternative form of the equation expressing • BUT It only applies to equilibrium
this relationship is given by systems.
• Recent work has been directed at trying
to understand how far it can be pushed
e.g. for glasses which have quenched in
force correlation function disorder

1

2 k BT 

f (0) f (t ) dt

19
Using Brownian Motion to obtain Local
Viscosities
• Beads moving around in viscous
fluids provide a means of probing
local viscosities.
• In some materials (e.g. liquid
crystals, see later) these may be
anisotropic.
• Other systems (e.g gels) may be
heterogeneous.
• A generalised version of the Stokes
Einstein equation allows the
complex moduli to be determined
by tracking the motion of the
particles.

20
 Using Particle Tracking in Practice
• The mean square displacement as a function
of time can be followed eg. during some
process.
m
2 dk BT
r ( )  
3a
• (d is dimensionality, a is particle radius,  is
lag time)
m • Data for hectorite clay 1 hour and 24 hours
after the start of the experiment, showing how
Brownian motion becomes restricted as the
systems 'sets' or 'gels'.
m
• From the displacements the viscoelastic
moduli can be determined (via Laplace
transforms).

dk BT
G ( s) 
m 3as r ( s ) 2 21
 Chemical Potential and Free Energy
• • Model situation:
Recall the definition of the Helmholtz free
energy F for a simple system
F=U-TS A B
• If one has a reservoir containing N
atoms/molecules, the chemical potential 
is defined as • Particle (species i) and energy
exchange is possible between two
 F  boxes A and B.
  
 N T ,V • Total change in entropy
• The chemical potential is the change in
free energy when one particle moves from  S   S 
one position, phase etc to another. dS tot   A dE A   B dE B 
• Equivalently  E A   E B 
 S A   S B 
 G 
   
 U 
 
 H   S 
  T   i  N dN Ai  i  N dN Bi
 
 N T , P  N  S ,V  N  S ,P  N  E ,V  Ai   Bi 

• In this field of soft condensed matter, F where summation is over species i.

and G are often used interchangeably since
22
the PV term is usually negligible.
 Chemical Potential and Equilibrium
• Now both energy and particle number
must be conserved, and the net change
in entropy must be zero or greater.
• Using the relationship between  and S
and recall 1/T = dS/dE and dEA+dEB=0
this equation reduces to
 1 1     
  dE A     Ai  Bi dN Ai  0
 TA TB  i  TA TB 
and this must be true for all the species.
• This implies (obviously) that at
equilibrium TA=TB and
 A =B
• When systems can exchange particles,
not only the temperature but also the
chemical potential for each species
must be equal at equilibrium.
• Can now extend all the thermodynamic
functions to include the possibility of
Ni varying.
dF  SdT  pdV   i dN i
i
dH  TdS  Vdp   i dN i
i
dG   SdT  VdP   i dN i
i
23
 Aside on the Clausius Clapeyron Equation

• Since we must have gas=liquid on

Liquid the phase boundary, then since dG
P = 0 also across the boundary
Solid
 S gas dT  Vgas dp   S liq dT  Vliq dp
Vapour

T
• But across the boundary T and p
• Recall this tells us about how must be equal so
properties change along the phase
boundary, eg between gas and dp S L
liquid.  
dT V TV

24
 Chemical Potential of a Perfect Gas
• Recall F= -kBTlnZ • Now z/N can be written in terms of the
pressure as
where Z is the partition function.
z zk BT
• For N indistinguishable particles, as in a 
N pV
perfect gas,
Z = zN/N! • Thus  = o + kBTlnP
where z is the individual particle partition where o represents the chemical
function. potential of a standard state.

• • If you have a solution in equilibrium

Then
with its vapour soln= vap
  z 
F  k BT  N ln   N   so ln   o  k BT ln P
 N  • Hence
Hence • Equivalently  so ln   o  k BT ln c
 F   z
    k BT ln  • Where c is the concentration in the
 N T ,V N
solution.
• This result is generally true.

25
 Membranes
• Membranes are the envelopes
around all our cells in our bodies.
• They are complex, dynamic
structures comprising so-called
lipid bilayers (see later).
• They have to be able to respond to Vesicles – simple
significant shape changes as our approximation to
bodies move or an organism biological cells - easily
crawls.
distort to allow for shape
• Initially let us treat the membrane changes, but do not
as a simple fluid interface with
associated surface tension, and exhibit short wavelength
look at its fluctuations. fluctuations as these cost a
• This is far too simple, because in lot of energy.
fact there is a curvature elasticity to
be taken into account (to be
discussed later).

26
 Monge Representation of a Membrane Surface
Boal
• The Monge representation is a common • We can construct tangent vectors
way of representing an undulating along x and y, represented by rx
surface, such as a membrane. and ry by taking a unit step in the x
• A point r on a surface is represented by (or y) direction, and a step
• r = (x,y, h(x,y)) h  h 
 hx  or  hy 
x  y 

in the z direction, to give

rx  (1, 0, hx ) ry  (0, 1, hy )
h(x,y)
z y • These are not unit vectors, and are
not generally orthogonal.
x
• They define the plane tangent to
• h represents the 'height' away from the
the surface at (x,y,h(x,y)) by
x,y surface.
• For complex surfaces, h may not be n
r r 
x y

 h ,h ,1x y

single valued (e.g if overhangs), but in

the Monge representation it is
rx  ry 1  h 2
x  hy2 
1/ 2

27
constrained to be single-valued.
 Monge Representation cont
• The surface can also be defined by
n rydy its curvature – these are simply
z dA geometrical relationships,
hx rxdx discussed in maths texts.
• The curvature of a surface is
1 defined as
x
  2r 
• The element of area dA is defined
C  n. 2 
 s 
by
where s is the arc length.
dA  rx  ry dxdy  1  h  h
2
x y
2 1/ 2
dxdy • In general there will be two
principal curvatures (the two
• And the metric g of the surface is
extremal values) C1 and C2.
defined by
• The combination (C1+ C2) is the
mean curvature .
g  (1  hx2  hy2 )
• And C1. C2 is the Gaussian
curvature .
So that dA=g dx dy

28
 Fluctuations in Membranes treated as a
Simple Fluid
Safran
• Hence, writing the undulations as a
Fourier sum
h(r )   h(q)e iq.r
h(x,y) q

h(q)   h(r )e iq.r dr

fluctuating surface
• For a simple fluid surface there will • The free energy F is given by
be an associated surface energy 

per unit area. F  
2 q
q 2 | h(q) |2
• The change in free energy
associated with the undulations of
the surface h(x,y) is • And therefore by equipartition of
 energy (for each mode q)
• where
 (h
2 2
F  x  hy )dxdy
2
k BT
h h 2
| h(q) |  2
hx  ; hy  q
x y 29
 Interpreting these Fluctuations
Safran
• Thus the mean square fluctuation
• Consider diverges as the logarithm of the
system size.
1
h 2 (r ) 
(2 ) 2  | h(q) |2 dq • Also, since

2 k BT
• In two dimensions the logarithm of
| h(q) |  2
q
the integral diverges at small q,
and so we must think carefully small wavelength fluctuations in
about the limits. real space (ie large q) have far
smaller fluctuations associated with
• Set the upper limit as/a, where a them than large wavelength (small
is the atomic size. q) fluctuations.
• Set the lower limit as  /L, where L • Or equivalently higher energy is
is the size of the interface. associated with small wavelength
• Then fluctuations.
k T L • Thus largescale distortions of
2
h (r )  B log membranes are more favourable
2 a than local rumpling.

30
 Real Cells
• Example: Bovine pulmonary arterial endothelial
cells imaged in the confocal laser scanning
microscope
• Dual labeled with a green stain for actin
microfilaments (FITC-Phalloidoin) and a red stain
for mitochondria.
• The stain for actin is actually derived from a toxic
compound found in mushrooms.
• The stain for mitochondria only becomes
fluorescent when the stain is activated by enzymes
which reside in the mitochondria.
• For this reason, only the mitochondria appear red,
Bone cells not the other organelles.
• Example: Bone cells imaged in the environmental
scanning electron microscope (ESEM).
• Normal scanning electron microscopes work in high
vacuum, and so require cells to be dehydrated.
• The ESEM does not, and so potentially provides a
new route to high resolution examination of cells.
31
 Real Cells cont
• Real cells are a lot more complex than simply an interface undergoing
fluctuations, but the basic ideas are correct.
• Have to take into account
– The physical structure of the membrane
– The (bio)chemical structure of the membrane
– The contents of the cell, particularly the cytoskeleton.
– What is surrounding the cell
– Osmotic effects
Example: what happens
when the cell is placed in
different media which
cause different degrees of
stress? The example shown is for
changing the osmolarity of the
surrounding medium, when the cell
shrinks or stretches according to
whether molecules have flowed in or
out.

32
 Aside on Osmotic Pressure
• The comment on osmolarity
introduces the idea of osmotic
pressure.
• You will be familiar with this
through the idea of what happens
with a semi-permeable membrane.
A B
• If there is solute in A, then water will
flow across the semi-permeable
membrane until the chemical
potentials on the two sides are equal.
 mixture (T , p   , ci )   pure (T , p,0)
• The osmotic pressure, , is the pressure which
will just stop the flow occurring.
• In general
V  RT  ci
• Which for low concentrations approximates to
N
 RT
V
• This has a form equivalent to the case of a perfect
gas.
• This effect is crucial in maintaining cells healthy.
• For instance, when a plant 'wilts' it has lost water
from its cells and the external pressure causes the
cells to collapse.
• By watering the plant, water is sucked back into
the cells to return the plant to full 'turgor' (in this
context one talks about 'turgor pressure', and the
cells are then said to be turgid).
33
 Brownian Motion within Cells- current (and
preliminary!) Research

Two dimensional mean square

displacements of beads in this cell. A
selection of particles have been
highlighted in red, for clarity. The
Fluorescence image of an individual human slopes exhibit significant scatter, which
dermal fibroblast cell showing clustering of the we attribute to local variations in the
beads in the perinuclear region, with a few rheology according to cytoskeletal 34
beads in the periphery of the cell. arrangements.
(Preliminary) Microrheology

It is clear very different responses can be

measured.
From the mean square displacements one
can then extract the local moduli, and
examine how anisotropic they are.
The aim is now to relate these local
measurements to the internal components
of the cell under different conditions e.g
of cell adhesion, motility, stress etc.
We do not yet know if the variation is
real, and we will be able to make these
correlations, or simply due to a spread in
values for statistical reasons.

35