MEAN-FIELD MODEL FOR THE DYNAMICS OF ACTIN FILAMENTS IN THE

CONTRACTILE RING

In the following, we will give the mathematical details of the physical model used in the
main text.

Model definition

With our theory, we try to capture essential features of the ring dynamics, such as,
filament polarity, rules of interaction between filaments through molecular motors. Still, in
the following, we use the words actin filaments and myosin for their relevance in cytokinetic
rings.
Consider a ring of actin filaments such that the filaments align with the ring perimeter.
We denote the co-ordinate along the ring perimeter by x and describe the distribution
of (polar) actin filaments along x by the densities c+ for filaments with their plus-end
pointing clockwise and c for filaments of the opposite orientation.

Two filaments of

opposite orientation can join their plus-ends forming a bipolar filament (Fig. 4a, top).
In cells, these joints might be established by protein complexes containing, for example,
formins and myosins (Figs. 2c, 3b). The distribution of bipolar filaments is denoted by
cbp giving the density of their centres. Actin filaments are assumed to have a length
`, bipolar filaments a length 2`. Consequently, the total actin density at a point x
R`
is 0 d (c+ (x + ) + cbp (x + ) + c (x ) + cbp (x )) . Bipolar filaments form at rate
c c+ c , bipolar filaments can split into two filaments of opposite orientations at rate d
(Fig. 4a, top).
Let us now turn to the processes of filament dynamics we account for. Filaments are
assumed to grow at their plus-ends and to shrink at their minus-ends at the same rate .
This treadmilling dynamics leads to movement of actin filaments at velocity vtr = a into the
direction of their plus-ends, where a is the size of an actin monomer. Bipolar filaments are
not displaced by treadmilling. Myosin mini-filaments can induce relative sliding between
filaments. The corresponding velocities are between filaments of the same orientation
(Fig. 4a, middle) and for filaments of opposite orientations (Fig. 4a, bottom). We use these
parameters to quantify the strength of the motor-mediated filament-filament interactions.
1

We assume that myosins are located at the filaments plus-ends, such that c+ + c + cbp is
the distribution of myosin. Finally, fluctuations are accounted for by diffusion terms with
an effective diffusion constant D. The corresponding dynamic equations read:
+

t c (x)

=Dx2 c+ (x)

d c+ (x + ) c+ (x ) c+ (x)
0
Z `
Z `


+
d cbp (x + )c (x) + x
d c (x ) + cbp (x ) c+ (x)
x
x

x vtr c+ (x) c c+ (x)c (x) + d cbp (x)

(1)
Z `


t c (x) =Dx2 c (x) x
d c (x + ) c (x ) c (x)
0
Z `
Z `

+ x
d cbp (x )c (x) x
d c+ (x + ) + cbp (x + ) c (x)
0

0
+

+ x vtr c (x) c c (x)c (x) + d cbp (x)

Z `
2
t cbp (x) =Dx cbp (x) x
d (cbp (x + ) cbp (x )) cbp (x)
0
Z `


d c (x + ) c+ (x ) cbp (x)
x
0
Z `


x
d c+ (x + ) c (x ) cbp (x) + c c+ (x)c (x) d cbp (x)

(2)

(3)

For numerical solution of the dynamic equations, we used a first-order upwind scheme
with adaptive time stepping.

Calculation of the stress in the bundle

The stress in the bundle is defined as the sum of the stresses in the individual filaments.
Stresses in a filament are generated by motors that pull on the filaments and by friction
with the surrounding medium. Explicitly, force balance on a single filament gives
s =

1
v + fmot .

(4)

In this expression, s is the co-ordinate along the filament, the stress in the filament,
a mobility, v the filaments velocity, and fmot the force density exerted by motors on the
filament. Only the effects of motors cross-linking two filaments are accounted for. The stress
along a filament is thus piece-wise linear in s with slope v/, where v = or v =
depending on the orientation and the relative position of the partner filament, the motor is
2

filament (length l)
motor position smot
stress profile

v l/

FIG. 1. Illustration of the stress distribution along a stiff slender rod of length l (the filament, red)
that is drawn by a point force at s0 into the direction of the arrow. The filament velocity is v, its
mobility . The black line indicates the stress profile that results from the applied force and the
filament friction with the environment.

connected to. If there is no motor at a filament end, then the stress vanishes at this point,
and the stress jumps by an amount |v|`/ at the positions smot , where motors are bound
to the filament, see below. The total stress profile along the bundle is then obtained by
summing the stress profiles along all filaments in the bundle. Since the expressions are quite
involved, we refrain from giving them here explicitly.

Results and Observations

Model considering actin polar filaments

This odel has already ee dis ussed i A ti ely Co tra ti g Bu dles of Polar Fila e ts ,PRL
(Vol. 85 No. 8).However the model that we have used to describe dynamics includes treadmmilling
urre t: x(vtrc+- .o I this odel as it's straightfor ard to see fro the graph that c >0 for all
parameters (when vtr=0), which means unstability is due to interaction between parallel filaments
ith c
Also it a e easily see that c increases with increasing D, diffusion coeff.(As
ag itude o ly i graph is a tually */D here is le gth of fila e t oth filaments were
assu ed of sa e le gth is de reasi g , c i reases .As soo as e de rease size of syste L, c
i reases shorte i g of u dles gi es sta ility .Also c decreases with increasing c22 w.r.t. i.e.
increasing filaments of same orientation.(graph Also Whe tread illi g is a ou ted,c
decreases which means treadmilling is also responsible for destabilization of system.Now it might
e said for that regio he c decreases with increase in D is due to dominance of treadmilling in
those region.(see graph 2).

Graph 1 (Here coordinates on X-axis is and coordinates on Y-axis is

.Curves
with green color indicate more fraction of c22 compared to c11 and and red color indicate lesser fraction
c22 compared to c11.)

Results and Observations

Graph 2(Here in this case c11=.3 and c22=.7,Here coordinates on X-axis is

and coordinates on Y-axis is

Model considering actin polar filaments and Bipolar filaments

This model is an extension of previous model with consideration of Bipolar filaments which has
been mentioned in "Self-organization and mechanical properties of active filament bundles",
Physical Review E 67 051913. However the model that we have used to describe dynamics includes
tread
illi g urre t: x(vtrc+-) ).
Also it a e easily see that c decreases with increasing wd ,rate of splitting of bipolar
filaments to result in polar ones.(As
ag itude o ly i graph is a tually / d* here is
le gth of fila e t oth fila e ts ere assu ed of sa e le gth is de reasi g , c decreases).As
soon as we decrease size of syste L, c first decreases then increases(shortening of bundles for
sta iltiy o has arrier .Also c ay i rease ith i rease i ot al ays! I gree ur e,for
egat ie part , c has very random patterns) and sometimes function is multivalued, So no
ertai ity for all <c is stable region , while in previous model (only polar filaments) , c
de reases ith i rease i a d ith ertai ity. o I this odel, ore a ti parallel fila e t
interactions causes stability to system(as i reases,more stabiltiy to system .Also c decreases
ith i reasi g
.r.t. i.e. i reasi g fila e ts of sa e orie tatio .It sta ilizes c increases)
with increase in D,diffusion coefficient.

Results and Observations

10

8.33

alpha

6.67

3.33

1.67

L=5 with c0+=.3,c0-=.7

L=5 with c+0=.3,c0-=1.5
L=10 with c+0=.3,c0-=1.5

-13.33

-6.67

0
beta

Constant

6.67

Prameters:

13.33

20

D=2,wd=1.5

,wc=0.5,vtr=1.5 ,=1)
Here Also Whe tread illi g is a ou ted,c initially increases rapidly with increase in vtr then
increases very slowly (remains almost constant).

Results and Observations

Constant Prameters: (C0+ =.3 , C0- =.7, D=2, =.5 ,

d=.7

,wc=.3 ,=1)

Also c decreases with increase in wc ,So combination of polar filaments(antiparallel) resulting in

bipolar filament causes unstability to system( wc is rate of combination of polar filaments).If this
result is in consistence with other observations, then it can be understood physically that "As
antiparallel filaments may cause stability to system,so making of bipolar filaments by
combination will bring unstability to system."(however this argument seems to be
mathematically wrong when wc<wd ,as for these values of wc ,we get polar filaments efectively as
wc>wd which is not happening as you can see from graph curve with lesser has lesser stability for
all times.By looking at graph ,one thing that is needed to be observed,for system to have stability
with lesser L (contraction) is not so straightforward.as it can be seen that for w c lesser than certain
value,system has more stability than system of lesser size(really ???).

Results and Observations

10
L=10with beta=1.0
L=5 with beta=1.0
L=5 with beta=.5(green)

alpha

2.5

7.5
wc

10

12.5

15

Constant Prameters: (C0+ =.3 , C0- =.7, D=1,wd=2.5 ,vtr=.5 ,=1)

First Order Upwind Scheme for solving Actin Dynamics Equation
Till now we have been trying to understand it's behaviour using linear stability analysis or loosely speaking
geometric intuition.Now we use first order upwind scheme to get numerical solutions of these equations.
Results obtained for stability of system were in consistence with previous results.Also this method extends
it's analysis to initial states of any kind unlike steady state analysis in previous methods.Graphs can be seen
on my previous report describing analysis behind this method.