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5. DYNAMICAL SYSTEM
5.1 WHAT IS DYNAMICAL SYSTEM
WHAT IS A DYNAMICAL SYSTEM?
Dynamical system is a system that changes over
time according to a set of fixed rules that
determine how one state of the system moves to
another state.
A DYNAMICAL SYSTEM HAS TWO PARTS
a)a State space, which determines possible values of the
state vector. State vector consists of a set of variables
whose values can be within certain interval.
b) a Function, which tells us, given the current state,
what the state of the system will be in the next instant
of time
CLASSIFICATION OF DYNAMICAL SYSTEMS
DYNAMICAL SYSTEM CAN BE
Either Or
Deterministic Stochastic
Discrete Continous
Linear Nonlinear
Autonomous Nonautonomous
WHEN A DYNAMICAL SYSTEM IS
DETERMINISTIC?
• Deterministic model is one whose behavior is
entire predictable. The system is perfectly
understood, then it is possible to predict precisely
what will happen.
• Stochastic model is one whose behavior cannot
be entirely predicted.
• Chaotic model is a deterministic model with a
behavior that cannot be entirely predicted
DISCRETE VS CONTINUOUS MODELS
• Discrete: the state variables change only at a
countable number of points in time. These points in
time are the ones at which the event occurs/change in
state.
• Continuous: the state variables change in a
continuous way, and not abruptly from one state to
another (infinite number of states).
100000*03.1)(
)(03.1)1(
100000)0(
k
kx
kxkx
x



DISCRETE SYSTEM
• The system is typically specified by the set of equations
such as difference equations.
• We denote time by k, and the system can be solved by
iteration calculation.
• Typical example is annual progress of a bank account. If
the initial deposit is 100000 euros and annual interest is
3%, then we can describe the system by
CONTINUOUS SYSTEM
It is described by a differential equation or a set of them.
For example, vertical throw is described by initial conditions
h(0), v(0) and equations
where h is height and v is velocity of a body.
)(´
)0( 0
xfx
xx


gtv
tvth


)´(
)()´(
10
DIFFERENTIAL EQUATIONS
• Even though an analytical treatment of dynamical systems is usually
very complicated, obtaining a numerical solution is (often) straight
forward.
• Solving differential equations numerically can be done by a number
of schemes. The easiest way is by the 1st order Euler’s Method:
,..)()()(
,...)(
)()(
,...)(
xftttxtx
xf
t
ttxtx
xf
dt
dx





t
x
Δt
x(t)
x(t-Δt)
LINEAR VS. NONLINEAR SYSTEM
Linear system : Function which is describing the system
behaviour must satisfy two basic properties
• additivity
• homogeneity
Nonlinear system is described by a nonlinear function. It
does not satisfy previous basic properties
)()()( yfxfyxf 
)()( xfxf  
LINEAR VS. NONLINEAR SYSTEM
Which one is linear ? A: g(x) = 3x; g(y) = 3y or B: f(x) = x2; f(y)
= y2?
• additivity g(x+y) = 3(x+y) = 3x + 3y = g(x) + g(y)
• homogeneity 5 * g(x) = 5* 3x = 15x = g(5x)
222
22222
5)(525)5()5(
)()(2)()(
xxfxxxf
yxyfxfyxyxyxyxf


A differential equation is linear, if the coefficients are constant or functions only of the independent variable
(time).
AUTONOMOUS VS. NONAUTONOMOUS SYSTEM
Autonomous system is a system of ordinary differential
equations, which do not depend on the independent
variable. If the independent variable is time, we call it time-
invariant system.
Condition: If the input signal x(t) produces an output y(t) then
any time shifted input, x(t + δ), results in a time-shifted
output y(t + δ)
Consider these two systems,
System A: System B: )(10)( txty 
AUTONOMOUS VS. NONAUTONOMOUS SYSTEM
Clearly , therefore the system is not time-invariant or is
nonautonomous.
System A:
Start with a delay of the
input
Now we delay the output by
δ
How about System B ?
5.2 DYNAMICAL SYSTEMS TERMINOLOGY
DYNAMICAL SYSTEMS TERMINOLOGY
• A phase space is a space in which all possible states
of a system are represented, with each possible state
of the system corresponding to one unique point in
the phase space.
• The phase space of a two-dimensional system is
called a phase plane.
• A phase curve is a plot of the solution of equations of
motion in a phase space.
• A phase portrait is a plot of single phase curve or
multiple phase curves corresponding to different
initial conditions in the same phase plane.
DIFFERENTIAL EQUATION OF A SYSTEMS
• Predator-Prey Relationships
18
LOTKA-VOLTERRA EQUATIONS
• The Lotka-Volterra equations describe the interaction between two
species, prey vs. predators, e.g. rabbits vs. foxes.
x: number of prey
y: number of predators
α, β, γ, δ: parameters
)(
)(
xy
dt
dy
yx
dt
dx




Can you model of Predator-Prey system using Cellular Automata ?
19
LOTKA-VOLTERRA EQUATIONS
• The Lotka-Volterra equations describe the interaction between two
species, prey vs. predators, e.g. rabbits vs. foxes.
Can you solve predator-prey system in the Mathematica and reproduce these
graphs?
HARMONIC OSCILLATOR
• Look at particle motion, under a force, linear in the displacement (Hooke’s
Law):
• Solutions: x(t) = A cos(ω0t - α)
y(t) = B cos(ω0t - β)
Or
x˝ + (ω0)2 x = 0
y˝ + (ω0)2 y = 0
-kx = mx˝
-ky = my˝
HARMONIC OSCILLATOR
• The amplitudes A, B, & the phases α, β are determined
by initial conditions.
• The motion is simple harmonic in each of the 2
dimensions
• Both oscillations have the same frequency but (in
general) different amplitudes.
x(t) = A cos(ω0t - α) y(t) = B cos(ω0t -
β)
EQUATION FOR THE PATH
• The equation for the path in the xy plane is obtained by
eliminating t in the solution .
• Define δ  α – β  B2x2 -2ABxy cosδ +A2y2 =A2B2sin2δ
• Except for special cases, the general path is an ellipse!
• One of special cases:
• δ = π/2  An ellipse: (x/A)2 + (y/B)2 = 1
• When the path will be a circle?
How about straight line ? Is there any δ which result in straight line ?
5.3 DYNAMICAL SYSTEMS ANALYSIS
DYNAMICAL SYSTEMS ANALYSIS
How many equilibria a coin balanced on a table have
? Heads, Tails, Edge
Start on its edge, it is in equilibrium … is it stable?
… No! Small movement away (perturbation) means that coin will
end up in one of 2 different equilibria: Heads/ Tails.
Both stable as perturbation does not result in a change in state
Similarly, think of a ball at rest in a dark landscape. It’s either on top of a hill or
at the bottom of a valley. To find out which, push it (perturb it), and see if it
comes back. (What about flat bits?)
ATTRACTOR & PERTURBATION
• An attractor is a set towards which a dynamical system
evolves over time. It can be a point, a curve or more
complicated structure
• A perturbation is a small change in a physical system,
most often in a physical system at equilibrium that is
disturbed from the outside
• Each attractor has a basin of attraction which contains
all the initial conditions which will generate trajectories
joining asymptotically this attractor
FIXED POINTS OF 1D SYSTEMS
One dimensional system, dx/dt = f(x, t)
At a fixed point, x doesn’t change as time increases:
dx/dt = 0
So, to find fixed points, f(x, t) = 0 should be solved
Note: same procedure for finding maxima and minima
What are the fixed points of this system dx/dt = 6x(1 – x) ?
so either 6x = 0 => x = 0 or 1 – x = 0 => x = 1
Set:dx/dt = 0 ie: 6x(1-x) = 0
But to perturb them must have a model of the system.
Use difference equation: x(t+h) = x(t) +h dx/dt from various different
initial x’s
Perturb the points and see what happens …
x=0 unstable and x=1 stable.
Start from:
x = 0 + 0.01, x = 0 – 0.01,
x = 1 + 0.01, x = 1 – 0.01
….
0 0.5 1 1.5
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
t
x
CLASSIFICATION OF ONE-DIMENSIONAL
DYNAMICAL SYSTEMS
Derivative at fixed point Fixed point is
f'(a0) <0 Stable
f'(a0) >0 Unstable
f'(a0) =0 Cannot be decided
if f’(a0) = 0 it can be semi-stable from above or below or periodic …
STABILITY AND FIXED POINTS
• A fixed point is a special point of the dynamical system
which does not change in time. It is also called an
equilibrium, steady-state, or singular point of the system.
• If a system is defined by an equation dx/dt = f(x), then the
fixed point 𝑥 can be found by examining of condition f( 𝑥)=0.
We do not need to know analytic solution of x(t).
• A stable fixed point: for all starting values x0 near the 𝑥 the
system converges to the 𝑥 as t→∞.
• A marginally stable fixed point: for all starting values x0 near
𝑥, the system stays near 𝑥 but does not converge to 𝑥 .
• An unstable fixed point: for starting values x0 very near 𝑥 the
system moves far away from 𝑥
BACTERIA GROWTH MODEL
• We leave a nutritive solution and some bacteria in a
dish.
• Let b be relative rate at which the bacteria reproduce
and p be relative rate at which they die. Then the
population is growing at the rate r = b−p.
• If there are x bacteria in the dish then the rate at
which the number of bacteria is increasing is (b − p)x,
that is, dx/dt = rx.
• Solution of this equation for x(0)=x0 is
rt
extx 0)( 
Is this model
realistic?
BACTERIA GROWTH MODEL (REVISITED)
• The model is not realistic, because bacteria population
goes to the infinity for r>0.
• As the number of bacteria rises, they produce more toxic
products.
• Instead of constant relative perish rate p, we will assume
relative perish rate dependent on their number px.
• Now the number of bacteria increases by bx and their
number decreases by px2.
• New differential equation will be 2
pxbx
dt
dx

How many fixed point has this model?
FIXED POINT OF BACTERIA GROWTH MODEL
• To be able to find a fixed point, we have to set the
right-hand side of the differential equation to zero.
• There are two possible solutions, we have two fixed
points:
0)~(~
0~~ 2


xpbx
xpxb
p
b
x
x


2
1
~
0~
What is the analytic solution ?
FIRST FIXED POINT IS UNSTABLE
• There are no bacteria, None can be born, None can
die
• But after small contamination (perturbation) which
is smaller than b/p, the number of bacteria will
increase by dx/dt = bx-px2>0 and will never return
to the zero state.
THE SECOND FIXED POINT IS STABLE AND IS ALSO AN ATTRACTOR
• Second point 𝑥= b/p, at this population level, bacteria are
being born at a rate b 𝑥=b(b/p) = b2/p and are dying at a rate
p 𝑥2 = b2/p, so birth and death rates are exactly in balance.
• If the number of bacteria would be slightly increased, then
dx/dt = bx-px2<0 and would return to equilibrium.
• If the number of bacteria would be slightly decreased, then
dx/dt = bx-px2>0 and would return to equilibrium.
• Small perturbations away from 𝑥= b/p will self-correct back to
b/p.
How does the graphic solution in Mathematica look like?
5.4 2D DYNAMICAL SYSTEMS
2D DYNAMICAL SYSTEMS
1. Find fixed points
2. Examine stability of fixed points
3. Examine the phase plane and trajectories
),(
),(
yxgy
dt
dy
yxfx
dt
dx




Suppose we have the following system :
FIND FIXED POINTS
• Find fixed points as before we need to solve: f(x,y) = 0
and g(x,y) = 0 to get fixed points (x0, y0)
• Predator-prey system :
• If x=0, dy/dt = -0.4y -> y=0 for dy/dt=0 -> fixed point at
(0,0) and similarly, if y=0, dx/dt = 0 for x=0 so again, (0,0)
is fixed point.
• Other fixed point at (80, 12)
yxyy
xyxx
4.0005.0
05.06.0




0 5 10 15 20 25
0
20
40
60
80
100
120
140
160
180
200
t
Populationsize
20 40 60 80 100 120 140 160 180 200
0
5
10
15
20
25
x
y
EXAMINE BEHAVIOR AT/NEAR FIXED POINTS
x and y in
phase-
plane
x and y
over time
Cyclic behaviour is fixed but system not at a fixed point: complications of higher dimensions
JACOBIAN MATRIX







































n
nnn
n
n
x
f
x
f
x
f
x
f
x
f
x
f
x
f
x
f
x
f
...
.........
...
...
21
2
2
2
1
2
1
2
1
1
1
J
Jacobian matrix is the matrix of all first-order partial derivatives of a
vector- or scalar-valued function with respect to another vector. This
matrix is frequently being marked as J, Df or A.
),....,,(
.
.
),....,,(
),....,,(
21
2122
2
2111
1
nnn
n
n
n
xxxfx
dt
dx
xxxfx
dt
dx
xxxfx
dt
dx






),....,,(
.
.
),....,,(
),....,,(
21
2122
2
2111
1
nnn
n
n
n
xxxfx
dt
dx
xxxfx
dt
dx
xxxfx
dt
dx






CLASSIFICATION OF 2-DIMENSIONAL LINEAR
212122
212111
),(
),(
dxcxxxfx
bxaxxxfx
































dc
ba
x
f
x
f
x
f
x
f
2
2
1
2
2
1
1
1
AJ
0)(0))((
0det0)det(
2










bcaddabcda
dc
ba



EA
Set of equations for 2D system
Jacobian matrix for 2D system
Calculation of eigenvalues
CLASSIFY FIXED POINTS
• If eigenvalues have negative real parts, x0 asymptotically
stable
• If at least one has positive real part, x0 unstable
• If eigenvalues are pure imaginary, stable or unstable
Complex numbers are made up of a real part (normal number) and an imaginary
part eg 4 + 3i where1i
BRIEF CLASSIFICATION OF 2D SYSTEMS
BACK TO EXAMPLE
• fx = 0.6 – 0.05y, fy = 0.05x, gx = 0.005y, gy = 0.005x – 0.4
• For J(0,0) eigenvalues 0.6 and –0.4 and eigenvectors (1,0) and (0,1).
Unstable (a saddle point) with main axes coordinate axes
• For J(80,12) eigenvalues are pure imaginary: need more info but will be
like a centre…. to the phase-plane!








4.00
06.0
)0,0(J 






04
6.00
)12,80(J
yxyy
xyxx
4.0005.0
05.06.0




QUALITATIVE BEHAVIOR
• For a dynamical system to be stable:
• The real parts of all eigenvalues must be negative.
• All eigenvalues lie in the left half complex plane.
• Terminology:
• Underdamped = spiral (some complex eigenvalue)
• Overdamped = nodal (all eigenvalues real)
• Critically damped = the boundary between.
Is there any classification for linear 3D systems using Eigen
values?
CLASSIFICATION OF DYNAMICAL SYSTEMS
USING JACOBIAN MATRIX
ATTRACTING NODE OF 2D LINEAR SYSTEMS
22
11
4xx
xx













40
01
A
Equations
Jacobian matrix
Eigenvalues
λ1= -1
λ2= -4
Eigenvectors












1
0
0
1
There is a stable fixed point, the attracting node or sink
SADDLE POINT
22
11
4xx
xx











40
01
A
Equations
Jacobian matrix
Eigenvalues
λ1= -1
λ2= 4
Eigenvectors












1
0
0
1
REPELLING NODE
There is an unstable fixed point for both
systems
22
11
4xx
xx




Equations
Jacobian matrix
Eigenvalues
λ1= 1
λ2= 4







40
01
A
Eigenvectors












1
0
0
1
5.5 THE LORENZ SYSTEM
• Professor of Meteorology at the
Massachusetts Institute of
Technology
• In 1963 derived a three
dimensional system in efforts to
model long range predictions for
the weather
• The weather is complicated! A
theoretical simplification was
necessary
Edward Lorenz
THE LORENZ SYSTEM
D. Gulick, Encounters with Chaos, Mc-Graw Hill, Inc., New York, 1992.
• The Lorenz systems describes the motion of a fluid between two layers
at different temperature. Specifically, the fluid is heated uniformly from
below and cooled uniformly from above.
• By rising the temperature difference between the two surfaces, we
observe initially a linear temperature gradient, and then the formation of
Rayleigh-Bernard convection cells. After convection, turbulent regime is
also observed.
53
LORENZ ATTRACTOR
• The Lorenz attractor defines a 3-dimensional trajectory by the
differential equations:
dx
dt
 (y  x)
dy
dt
 rx  y  xz
dz
dt
 xy  bz
σ, r, b are parameters
The most frequently used values of parameters:
δ= 10, r= 28 and β= 8/3
LORENZ ATTRACTOR
• When calculating this model, Lorenz encountered a
strange phenomenon. After entering slightly different
input values for two successive attempts he obtained
completely different outputs.
• This effect was later named a butterfly effect, which
should express the sensitivity to initial condition by a
metaphor that „a single flap of butterfly wings in South
America can change weather in Texas“.
LORENZ RESULT COMPARISON
Very slight change of initial condition results in large change in solution of th
function.
EQUILIBRIA
S1 = 0,0,0( )
S2 = b r -1( ), b r -1( ),r -1( )
S3 = - b r -1( ),- b r -1( ),r -1( )
LINEAR STABILITY OF S1
Jacobian
Eigenvalues of J l1,2 =
- s +1( )± s -1( )2
+ 4sr
2
l3 = -b
J 0,0,0( ) =
-s s 0
r -1 0
0 0 - b
é
ë
ê
ê
ê
ù
û
ú
ú
ú
What can you say about the Linear stability of S1 and parameter r
CHAOS
• A chaotic system is roughly defined by sensitivity to
initial conditions: infinitesimal differences in the initial
conditions of the system result in large differences in
behavior.
• Chaotic systems do not usually go out of control, but stay
within bounded operating conditions.
• Chaos provides a balance between flexibility and
stability, adaptiveness and dependability.
• Chaos lives on the edge between order and
randomness.
AN ATTRACTOR OF DYNAMICAL SYSTEM
• Fixed point. The system evolves towards a single state and remains
there. An example is damped pendulum or a sphere at the bottom of
a spheric bowl.
• Periodic or quasiperiodic attractor: The system evolves towards a limit
cycle. An example is undamped pendulum or a planet orbiting
around the Sun.
• Chaotic attractor: The system is very sensitive to initial conditions and
we are not able to simply predict its behavior. An example is the
Lorenz attractor.
• Strange attractor: The system is also very sensitive to initial conditions
and we are not able to simply predict its behavior, but in this case the
system has the same properties like fractals. An example is the
Mandelbrot set.
FRACTALS
• The fractal is a geometric shape, that has the following features:
• It is self-similar, which means, that observing the shape in various zooms results in
the same characteristic shapes (or at least approximately the same)
• It has a simple and recursive definition
• It has a fine structure at arbitrarily small scales
• It is too irregular to be easily described in traditional Euclidean geometric
language.
• It has a Hausdorff dimension of its border higher than the topological dimension
of the border.
• Topological dimension – a point has topological dimension 0, a line has topological
dimension 1, a surface has topological dimension 2, etc.
• Hausdorff dimension – if an object contains n copies of itself reduced to one k-th of the
original dimension, the Hausdorff dimension can be calculated as log(n)/log(k)
Example – a Cantor set
DYNAMICAL SYSTEMS
• systems live in phase space
• of low dimension, compact or open
• or of high dimension (infinite in case of PDEs)
• and develop in time
• continuous time: differential equations
• discrete time: difference equations
• the dynamical laws may be
• deterministic (no uncertainty in the laws)
• stochastic (due to fluctuations)
5.6 DISCRETE DYNAMICAL SYSTEM
CONTINUOUS OR DISCRETE
The pendulum is a 2-dimensional continuous dynamical
system.
A Cellular Automaton or Boolean network are a discrete
dynamical system.
If there are 100 cells, then it is a 100-dim DS with a
deterministic trajectory (following all the update rules) --
strictly you need a 100-dim graph to picture it.
BOOLEAN NETWORK
Boolean network is
- a directed graph G(V,E)
characterized by
- the number of nodes : N
- the number of inputs per node: k
A B
C
E
D
F G
BOOLEAN LOGIC
Boolean models are discrete (in state and time) and deterministic.
(George Boole, 1815-1864)
Each node can assume one of two states:
on („1“) or off („0“)
Background:
Not enough information for more detailed description
Increasing complexity and computational effort for more specific models
Replacement of continuous
functions (e.g. Hill function)
by step function
BOOLEAN NETWORK
7
2
2
Boolean networks have
always a finite number of possible states: 2N
and, therefore, a finite number of state transitions:
A B
C
E
D
F G
N=7, 27 states, theoretically possible state transition
N
2
2
DYNAMICS OF BOOLEAN NETWORKS
The dynamics are described by rules:
„if input value/s at time t is/are...., then output value at t+1 is....“
A B
A(t) B(t+1)
BOOLEAN MODELS: TRUTH FUNCTIONS
in output
0 0 0 1 1
1 0 1 0 1
p p not p
rule 0 1 2 3
A B
B(t+1) = not (A(t))
rule 2
A(t) B(t+1)
STATE SPACE OF BOOLEAN
NETWORKS
State of the net: Row listing the present value of all N elements (0 or 1)
Finite number of states (2N)  as system passes along a sequence of states from
an arbitrary initial state, it must eventually re-enter a state previously passed  a
cycle
Cycle length: number of states on a re-enterant cycle of behavior
Cycle of length 1 – equilibrial state
Transient (or run-in) length: number of state between initial states and entering the
cycle
Confluent: set of states leading to or being part of a cycle
DYNAMICS OF BOOLEAN NETWORKS WITH K=1
Linear chainA B C D
A fixed (no input).
A(t)  B(t+1)
B(t+1)  C(t+2)
C(t+2)  D(t+3)
The system reaches a steady state after N-1 time steps.
Rules 0 and 3 not considered (since independence of input).
DYNAMICS OF BOOLEAN NETWORKS WITH K=1
Ring
A B
C D
A
B
Again: Rules 0 and 3 not considered (since independence of
input).
A(t+1)=B(t)
B(t+1)=A(t)
Both rule 1
A B A B A B A B
0 0 1 0 0 1 1 1
0 0 0 1 1 0 1 1
0 0 1 0 0 1 1 1
A(t+1)=not B(t)
B(t+1)=A(t)
Both rule 2
A B A B A B A B
0 0 1 0 0 1 1 1
1 0 1 1 0 0 0 1
1 1 0 1 1 0 0 0
0 1 0 0 1 1 1 0
0 0 1 0 0 1 1 1
Fixed point or
cycle of length 2
depending on
initial conditions
Cycle of length 4
independent of
initial conditions.
5.7 KAUFFMAN MODEL
KAUFFMAN MODEL
• In 1969, Stuart Kauffman proposed using Random
Boolean Networks (RBNs) as an abstract model of
gene regulatory networks
• Each vertex represents a gene
• “on” state of a vertex indicates that the gene is
expressed
• An edge from one vertex to another implies the
the former gene regulates the later
• “0” and “1” values on edges indicate presence/
absence of activating/repressing proteins
• Boolean functions assigned to vertices represent
rules of regulatory interactions between genes
EXAMPLE NETWORK
Three genes X, Y, and Z
X
Y
Z
Rules
X(t+1) = X(t) and Y(t)
Y(t+1) = X(t) or Y(t)
Z(t+1) = X(t) or (not Y(t) and Z(t))
Current Next
state state
000 000
001 001
010 010
011 010
100 011
101 011
110 111
111 111
000 001 010 011
100
101 110 111
EXAMPLE NETWORK
000 001 010 011
100
101 110 111
X
Y
Z
- The number of accessible states is finite, .
- Cyclic trajectories are possible.
- Not every state must be approachable from every other state.
- The successor state is unique, the predecessor state is not unique.
N
2
RANDOM BOOLEAN NETWORKS
If the rules for updating states are unknown
 select rules randomly
N nodes ½ pN (N-1) edges
Rule 2
Rule 0
Rule 1
Rule 2
KAUFFMAN’S NK BOOLEAN NETWORKS
An NK automaton is an autonomous random network of N Boolean logic
elements. Each element has K inputs and one output. The signals at inputs
and outputs take binary (0 or 1) values. The Boolean elements of the
network and the connections between elements are chosen in a random
manner. There are no external inputs to the network. The number of
elements N is assumed to be large.
KAUFFMAN’S NK BOOLEAN NETWORKS
K connections: 22K
Boolean input functions
Nets are free of external inputs.
Once, connections and rules are selected, they remain constant and the
time evolution is deterministic.
Earlier work by Walker and Ashby (1965): same Boolean functions for all
genes:
Choice of Boolean function affects length of cycles:
“and” yields short cycles,
“exclusive or” yields cycles of immense length
DYNAMICS WHEN N
• Number and length of attractors of depend on k and p
kc = ½ p(1-p)
 k < kc frozen phase
 many small attractors, not sensitive to changes in structure
 k > kc chaotic phase
 few large attractors, very sensitive to changes in structure
 k = kc phase transition behavior
 n attractors of length n, not sensitive to most changes, only to
some rare ones
KAUFFMAN MODEL
• Gene regulatory networks of living cells are believed
to exhibit phase transition behavior, on the border
between the frozen and chaotic phases
• Kaufman has shown that if k = 2 and p = 0.5, then the
statistical features of RBNs match the characteristics of
living cells
• Number of attractors  number of cell types
• Length of attractors  cell cycle time
Unit 5: All

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Unit 5: All

  • 2. 5.1 WHAT IS DYNAMICAL SYSTEM
  • 3. WHAT IS A DYNAMICAL SYSTEM? Dynamical system is a system that changes over time according to a set of fixed rules that determine how one state of the system moves to another state.
  • 4. A DYNAMICAL SYSTEM HAS TWO PARTS a)a State space, which determines possible values of the state vector. State vector consists of a set of variables whose values can be within certain interval. b) a Function, which tells us, given the current state, what the state of the system will be in the next instant of time
  • 5. CLASSIFICATION OF DYNAMICAL SYSTEMS DYNAMICAL SYSTEM CAN BE Either Or Deterministic Stochastic Discrete Continous Linear Nonlinear Autonomous Nonautonomous
  • 6. WHEN A DYNAMICAL SYSTEM IS DETERMINISTIC? • Deterministic model is one whose behavior is entire predictable. The system is perfectly understood, then it is possible to predict precisely what will happen. • Stochastic model is one whose behavior cannot be entirely predicted. • Chaotic model is a deterministic model with a behavior that cannot be entirely predicted
  • 7. DISCRETE VS CONTINUOUS MODELS • Discrete: the state variables change only at a countable number of points in time. These points in time are the ones at which the event occurs/change in state. • Continuous: the state variables change in a continuous way, and not abruptly from one state to another (infinite number of states).
  • 8. 100000*03.1)( )(03.1)1( 100000)0( k kx kxkx x    DISCRETE SYSTEM • The system is typically specified by the set of equations such as difference equations. • We denote time by k, and the system can be solved by iteration calculation. • Typical example is annual progress of a bank account. If the initial deposit is 100000 euros and annual interest is 3%, then we can describe the system by
  • 9. CONTINUOUS SYSTEM It is described by a differential equation or a set of them. For example, vertical throw is described by initial conditions h(0), v(0) and equations where h is height and v is velocity of a body. )(´ )0( 0 xfx xx   gtv tvth   )´( )()´(
  • 10. 10 DIFFERENTIAL EQUATIONS • Even though an analytical treatment of dynamical systems is usually very complicated, obtaining a numerical solution is (often) straight forward. • Solving differential equations numerically can be done by a number of schemes. The easiest way is by the 1st order Euler’s Method: ,..)()()( ,...)( )()( ,...)( xftttxtx xf t ttxtx xf dt dx      t x Δt x(t) x(t-Δt)
  • 11. LINEAR VS. NONLINEAR SYSTEM Linear system : Function which is describing the system behaviour must satisfy two basic properties • additivity • homogeneity Nonlinear system is described by a nonlinear function. It does not satisfy previous basic properties )()()( yfxfyxf  )()( xfxf  
  • 12. LINEAR VS. NONLINEAR SYSTEM Which one is linear ? A: g(x) = 3x; g(y) = 3y or B: f(x) = x2; f(y) = y2? • additivity g(x+y) = 3(x+y) = 3x + 3y = g(x) + g(y) • homogeneity 5 * g(x) = 5* 3x = 15x = g(5x) 222 22222 5)(525)5()5( )()(2)()( xxfxxxf yxyfxfyxyxyxyxf   A differential equation is linear, if the coefficients are constant or functions only of the independent variable (time).
  • 13. AUTONOMOUS VS. NONAUTONOMOUS SYSTEM Autonomous system is a system of ordinary differential equations, which do not depend on the independent variable. If the independent variable is time, we call it time- invariant system. Condition: If the input signal x(t) produces an output y(t) then any time shifted input, x(t + δ), results in a time-shifted output y(t + δ) Consider these two systems, System A: System B: )(10)( txty 
  • 14. AUTONOMOUS VS. NONAUTONOMOUS SYSTEM Clearly , therefore the system is not time-invariant or is nonautonomous. System A: Start with a delay of the input Now we delay the output by δ How about System B ?
  • 15. 5.2 DYNAMICAL SYSTEMS TERMINOLOGY
  • 16. DYNAMICAL SYSTEMS TERMINOLOGY • A phase space is a space in which all possible states of a system are represented, with each possible state of the system corresponding to one unique point in the phase space. • The phase space of a two-dimensional system is called a phase plane. • A phase curve is a plot of the solution of equations of motion in a phase space. • A phase portrait is a plot of single phase curve or multiple phase curves corresponding to different initial conditions in the same phase plane.
  • 17. DIFFERENTIAL EQUATION OF A SYSTEMS • Predator-Prey Relationships
  • 18. 18 LOTKA-VOLTERRA EQUATIONS • The Lotka-Volterra equations describe the interaction between two species, prey vs. predators, e.g. rabbits vs. foxes. x: number of prey y: number of predators α, β, γ, δ: parameters )( )( xy dt dy yx dt dx     Can you model of Predator-Prey system using Cellular Automata ?
  • 19. 19 LOTKA-VOLTERRA EQUATIONS • The Lotka-Volterra equations describe the interaction between two species, prey vs. predators, e.g. rabbits vs. foxes. Can you solve predator-prey system in the Mathematica and reproduce these graphs?
  • 20. HARMONIC OSCILLATOR • Look at particle motion, under a force, linear in the displacement (Hooke’s Law): • Solutions: x(t) = A cos(ω0t - α) y(t) = B cos(ω0t - β) Or x˝ + (ω0)2 x = 0 y˝ + (ω0)2 y = 0 -kx = mx˝ -ky = my˝
  • 21. HARMONIC OSCILLATOR • The amplitudes A, B, & the phases α, β are determined by initial conditions. • The motion is simple harmonic in each of the 2 dimensions • Both oscillations have the same frequency but (in general) different amplitudes. x(t) = A cos(ω0t - α) y(t) = B cos(ω0t - β)
  • 22. EQUATION FOR THE PATH • The equation for the path in the xy plane is obtained by eliminating t in the solution . • Define δ  α – β  B2x2 -2ABxy cosδ +A2y2 =A2B2sin2δ • Except for special cases, the general path is an ellipse! • One of special cases: • δ = π/2  An ellipse: (x/A)2 + (y/B)2 = 1 • When the path will be a circle? How about straight line ? Is there any δ which result in straight line ?
  • 24. DYNAMICAL SYSTEMS ANALYSIS How many equilibria a coin balanced on a table have ? Heads, Tails, Edge Start on its edge, it is in equilibrium … is it stable? … No! Small movement away (perturbation) means that coin will end up in one of 2 different equilibria: Heads/ Tails. Both stable as perturbation does not result in a change in state
  • 25. Similarly, think of a ball at rest in a dark landscape. It’s either on top of a hill or at the bottom of a valley. To find out which, push it (perturb it), and see if it comes back. (What about flat bits?)
  • 26. ATTRACTOR & PERTURBATION • An attractor is a set towards which a dynamical system evolves over time. It can be a point, a curve or more complicated structure • A perturbation is a small change in a physical system, most often in a physical system at equilibrium that is disturbed from the outside • Each attractor has a basin of attraction which contains all the initial conditions which will generate trajectories joining asymptotically this attractor
  • 27. FIXED POINTS OF 1D SYSTEMS One dimensional system, dx/dt = f(x, t) At a fixed point, x doesn’t change as time increases: dx/dt = 0 So, to find fixed points, f(x, t) = 0 should be solved Note: same procedure for finding maxima and minima
  • 28. What are the fixed points of this system dx/dt = 6x(1 – x) ? so either 6x = 0 => x = 0 or 1 – x = 0 => x = 1 Set:dx/dt = 0 ie: 6x(1-x) = 0 But to perturb them must have a model of the system. Use difference equation: x(t+h) = x(t) +h dx/dt from various different initial x’s Perturb the points and see what happens …
  • 29. x=0 unstable and x=1 stable. Start from: x = 0 + 0.01, x = 0 – 0.01, x = 1 + 0.01, x = 1 – 0.01 …. 0 0.5 1 1.5 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 t x
  • 30. CLASSIFICATION OF ONE-DIMENSIONAL DYNAMICAL SYSTEMS Derivative at fixed point Fixed point is f'(a0) <0 Stable f'(a0) >0 Unstable f'(a0) =0 Cannot be decided if f’(a0) = 0 it can be semi-stable from above or below or periodic …
  • 31. STABILITY AND FIXED POINTS • A fixed point is a special point of the dynamical system which does not change in time. It is also called an equilibrium, steady-state, or singular point of the system. • If a system is defined by an equation dx/dt = f(x), then the fixed point 𝑥 can be found by examining of condition f( 𝑥)=0. We do not need to know analytic solution of x(t). • A stable fixed point: for all starting values x0 near the 𝑥 the system converges to the 𝑥 as t→∞. • A marginally stable fixed point: for all starting values x0 near 𝑥, the system stays near 𝑥 but does not converge to 𝑥 . • An unstable fixed point: for starting values x0 very near 𝑥 the system moves far away from 𝑥
  • 32. BACTERIA GROWTH MODEL • We leave a nutritive solution and some bacteria in a dish. • Let b be relative rate at which the bacteria reproduce and p be relative rate at which they die. Then the population is growing at the rate r = b−p. • If there are x bacteria in the dish then the rate at which the number of bacteria is increasing is (b − p)x, that is, dx/dt = rx. • Solution of this equation for x(0)=x0 is rt extx 0)(  Is this model realistic?
  • 33. BACTERIA GROWTH MODEL (REVISITED) • The model is not realistic, because bacteria population goes to the infinity for r>0. • As the number of bacteria rises, they produce more toxic products. • Instead of constant relative perish rate p, we will assume relative perish rate dependent on their number px. • Now the number of bacteria increases by bx and their number decreases by px2. • New differential equation will be 2 pxbx dt dx  How many fixed point has this model?
  • 34. FIXED POINT OF BACTERIA GROWTH MODEL • To be able to find a fixed point, we have to set the right-hand side of the differential equation to zero. • There are two possible solutions, we have two fixed points: 0)~(~ 0~~ 2   xpbx xpxb p b x x   2 1 ~ 0~ What is the analytic solution ?
  • 35. FIRST FIXED POINT IS UNSTABLE • There are no bacteria, None can be born, None can die • But after small contamination (perturbation) which is smaller than b/p, the number of bacteria will increase by dx/dt = bx-px2>0 and will never return to the zero state.
  • 36. THE SECOND FIXED POINT IS STABLE AND IS ALSO AN ATTRACTOR • Second point 𝑥= b/p, at this population level, bacteria are being born at a rate b 𝑥=b(b/p) = b2/p and are dying at a rate p 𝑥2 = b2/p, so birth and death rates are exactly in balance. • If the number of bacteria would be slightly increased, then dx/dt = bx-px2<0 and would return to equilibrium. • If the number of bacteria would be slightly decreased, then dx/dt = bx-px2>0 and would return to equilibrium. • Small perturbations away from 𝑥= b/p will self-correct back to b/p. How does the graphic solution in Mathematica look like?
  • 37. 5.4 2D DYNAMICAL SYSTEMS
  • 38. 2D DYNAMICAL SYSTEMS 1. Find fixed points 2. Examine stability of fixed points 3. Examine the phase plane and trajectories ),( ),( yxgy dt dy yxfx dt dx     Suppose we have the following system :
  • 39. FIND FIXED POINTS • Find fixed points as before we need to solve: f(x,y) = 0 and g(x,y) = 0 to get fixed points (x0, y0) • Predator-prey system : • If x=0, dy/dt = -0.4y -> y=0 for dy/dt=0 -> fixed point at (0,0) and similarly, if y=0, dx/dt = 0 for x=0 so again, (0,0) is fixed point. • Other fixed point at (80, 12) yxyy xyxx 4.0005.0 05.06.0    
  • 40. 0 5 10 15 20 25 0 20 40 60 80 100 120 140 160 180 200 t Populationsize 20 40 60 80 100 120 140 160 180 200 0 5 10 15 20 25 x y EXAMINE BEHAVIOR AT/NEAR FIXED POINTS x and y in phase- plane x and y over time Cyclic behaviour is fixed but system not at a fixed point: complications of higher dimensions
  • 41. JACOBIAN MATRIX                                        n nnn n n x f x f x f x f x f x f x f x f x f ... ......... ... ... 21 2 2 2 1 2 1 2 1 1 1 J Jacobian matrix is the matrix of all first-order partial derivatives of a vector- or scalar-valued function with respect to another vector. This matrix is frequently being marked as J, Df or A. ),....,,( . . ),....,,( ),....,,( 21 2122 2 2111 1 nnn n n n xxxfx dt dx xxxfx dt dx xxxfx dt dx       ),....,,( . . ),....,,( ),....,,( 21 2122 2 2111 1 nnn n n n xxxfx dt dx xxxfx dt dx xxxfx dt dx      
  • 42. CLASSIFICATION OF 2-DIMENSIONAL LINEAR 212122 212111 ),( ),( dxcxxxfx bxaxxxfx                                 dc ba x f x f x f x f 2 2 1 2 2 1 1 1 AJ 0)(0))(( 0det0)det( 2           bcaddabcda dc ba    EA Set of equations for 2D system Jacobian matrix for 2D system Calculation of eigenvalues
  • 43. CLASSIFY FIXED POINTS • If eigenvalues have negative real parts, x0 asymptotically stable • If at least one has positive real part, x0 unstable • If eigenvalues are pure imaginary, stable or unstable Complex numbers are made up of a real part (normal number) and an imaginary part eg 4 + 3i where1i
  • 45. BACK TO EXAMPLE • fx = 0.6 – 0.05y, fy = 0.05x, gx = 0.005y, gy = 0.005x – 0.4 • For J(0,0) eigenvalues 0.6 and –0.4 and eigenvectors (1,0) and (0,1). Unstable (a saddle point) with main axes coordinate axes • For J(80,12) eigenvalues are pure imaginary: need more info but will be like a centre…. to the phase-plane!         4.00 06.0 )0,0(J        04 6.00 )12,80(J yxyy xyxx 4.0005.0 05.06.0    
  • 46. QUALITATIVE BEHAVIOR • For a dynamical system to be stable: • The real parts of all eigenvalues must be negative. • All eigenvalues lie in the left half complex plane. • Terminology: • Underdamped = spiral (some complex eigenvalue) • Overdamped = nodal (all eigenvalues real) • Critically damped = the boundary between.
  • 47. Is there any classification for linear 3D systems using Eigen values? CLASSIFICATION OF DYNAMICAL SYSTEMS USING JACOBIAN MATRIX
  • 48. ATTRACTING NODE OF 2D LINEAR SYSTEMS 22 11 4xx xx              40 01 A Equations Jacobian matrix Eigenvalues λ1= -1 λ2= -4 Eigenvectors             1 0 0 1 There is a stable fixed point, the attracting node or sink
  • 49. SADDLE POINT 22 11 4xx xx            40 01 A Equations Jacobian matrix Eigenvalues λ1= -1 λ2= 4 Eigenvectors             1 0 0 1 REPELLING NODE There is an unstable fixed point for both systems 22 11 4xx xx     Equations Jacobian matrix Eigenvalues λ1= 1 λ2= 4        40 01 A Eigenvectors             1 0 0 1
  • 50. 5.5 THE LORENZ SYSTEM
  • 51. • Professor of Meteorology at the Massachusetts Institute of Technology • In 1963 derived a three dimensional system in efforts to model long range predictions for the weather • The weather is complicated! A theoretical simplification was necessary Edward Lorenz
  • 52. THE LORENZ SYSTEM D. Gulick, Encounters with Chaos, Mc-Graw Hill, Inc., New York, 1992. • The Lorenz systems describes the motion of a fluid between two layers at different temperature. Specifically, the fluid is heated uniformly from below and cooled uniformly from above. • By rising the temperature difference between the two surfaces, we observe initially a linear temperature gradient, and then the formation of Rayleigh-Bernard convection cells. After convection, turbulent regime is also observed.
  • 53. 53 LORENZ ATTRACTOR • The Lorenz attractor defines a 3-dimensional trajectory by the differential equations: dx dt  (y  x) dy dt  rx  y  xz dz dt  xy  bz σ, r, b are parameters The most frequently used values of parameters: δ= 10, r= 28 and β= 8/3
  • 54. LORENZ ATTRACTOR • When calculating this model, Lorenz encountered a strange phenomenon. After entering slightly different input values for two successive attempts he obtained completely different outputs. • This effect was later named a butterfly effect, which should express the sensitivity to initial condition by a metaphor that „a single flap of butterfly wings in South America can change weather in Texas“.
  • 55. LORENZ RESULT COMPARISON Very slight change of initial condition results in large change in solution of th function.
  • 56. EQUILIBRIA S1 = 0,0,0( ) S2 = b r -1( ), b r -1( ),r -1( ) S3 = - b r -1( ),- b r -1( ),r -1( )
  • 57. LINEAR STABILITY OF S1 Jacobian Eigenvalues of J l1,2 = - s +1( )± s -1( )2 + 4sr 2 l3 = -b J 0,0,0( ) = -s s 0 r -1 0 0 0 - b é ë ê ê ê ù û ú ú ú What can you say about the Linear stability of S1 and parameter r
  • 58. CHAOS • A chaotic system is roughly defined by sensitivity to initial conditions: infinitesimal differences in the initial conditions of the system result in large differences in behavior. • Chaotic systems do not usually go out of control, but stay within bounded operating conditions. • Chaos provides a balance between flexibility and stability, adaptiveness and dependability. • Chaos lives on the edge between order and randomness.
  • 59. AN ATTRACTOR OF DYNAMICAL SYSTEM • Fixed point. The system evolves towards a single state and remains there. An example is damped pendulum or a sphere at the bottom of a spheric bowl. • Periodic or quasiperiodic attractor: The system evolves towards a limit cycle. An example is undamped pendulum or a planet orbiting around the Sun. • Chaotic attractor: The system is very sensitive to initial conditions and we are not able to simply predict its behavior. An example is the Lorenz attractor. • Strange attractor: The system is also very sensitive to initial conditions and we are not able to simply predict its behavior, but in this case the system has the same properties like fractals. An example is the Mandelbrot set.
  • 60. FRACTALS • The fractal is a geometric shape, that has the following features: • It is self-similar, which means, that observing the shape in various zooms results in the same characteristic shapes (or at least approximately the same) • It has a simple and recursive definition • It has a fine structure at arbitrarily small scales • It is too irregular to be easily described in traditional Euclidean geometric language. • It has a Hausdorff dimension of its border higher than the topological dimension of the border. • Topological dimension – a point has topological dimension 0, a line has topological dimension 1, a surface has topological dimension 2, etc. • Hausdorff dimension – if an object contains n copies of itself reduced to one k-th of the original dimension, the Hausdorff dimension can be calculated as log(n)/log(k) Example – a Cantor set
  • 61. DYNAMICAL SYSTEMS • systems live in phase space • of low dimension, compact or open • or of high dimension (infinite in case of PDEs) • and develop in time • continuous time: differential equations • discrete time: difference equations • the dynamical laws may be • deterministic (no uncertainty in the laws) • stochastic (due to fluctuations)
  • 63. CONTINUOUS OR DISCRETE The pendulum is a 2-dimensional continuous dynamical system. A Cellular Automaton or Boolean network are a discrete dynamical system. If there are 100 cells, then it is a 100-dim DS with a deterministic trajectory (following all the update rules) -- strictly you need a 100-dim graph to picture it.
  • 64. BOOLEAN NETWORK Boolean network is - a directed graph G(V,E) characterized by - the number of nodes : N - the number of inputs per node: k A B C E D F G
  • 65. BOOLEAN LOGIC Boolean models are discrete (in state and time) and deterministic. (George Boole, 1815-1864) Each node can assume one of two states: on („1“) or off („0“) Background: Not enough information for more detailed description Increasing complexity and computational effort for more specific models Replacement of continuous functions (e.g. Hill function) by step function
  • 66. BOOLEAN NETWORK 7 2 2 Boolean networks have always a finite number of possible states: 2N and, therefore, a finite number of state transitions: A B C E D F G N=7, 27 states, theoretically possible state transition N 2 2
  • 67. DYNAMICS OF BOOLEAN NETWORKS The dynamics are described by rules: „if input value/s at time t is/are...., then output value at t+1 is....“ A B A(t) B(t+1)
  • 68. BOOLEAN MODELS: TRUTH FUNCTIONS in output 0 0 0 1 1 1 0 1 0 1 p p not p rule 0 1 2 3 A B B(t+1) = not (A(t)) rule 2 A(t) B(t+1)
  • 69. STATE SPACE OF BOOLEAN NETWORKS State of the net: Row listing the present value of all N elements (0 or 1) Finite number of states (2N)  as system passes along a sequence of states from an arbitrary initial state, it must eventually re-enter a state previously passed  a cycle Cycle length: number of states on a re-enterant cycle of behavior Cycle of length 1 – equilibrial state Transient (or run-in) length: number of state between initial states and entering the cycle Confluent: set of states leading to or being part of a cycle
  • 70. DYNAMICS OF BOOLEAN NETWORKS WITH K=1 Linear chainA B C D A fixed (no input). A(t)  B(t+1) B(t+1)  C(t+2) C(t+2)  D(t+3) The system reaches a steady state after N-1 time steps. Rules 0 and 3 not considered (since independence of input).
  • 71. DYNAMICS OF BOOLEAN NETWORKS WITH K=1 Ring A B C D A B Again: Rules 0 and 3 not considered (since independence of input). A(t+1)=B(t) B(t+1)=A(t) Both rule 1 A B A B A B A B 0 0 1 0 0 1 1 1 0 0 0 1 1 0 1 1 0 0 1 0 0 1 1 1 A(t+1)=not B(t) B(t+1)=A(t) Both rule 2 A B A B A B A B 0 0 1 0 0 1 1 1 1 0 1 1 0 0 0 1 1 1 0 1 1 0 0 0 0 1 0 0 1 1 1 0 0 0 1 0 0 1 1 1 Fixed point or cycle of length 2 depending on initial conditions Cycle of length 4 independent of initial conditions.
  • 73. KAUFFMAN MODEL • In 1969, Stuart Kauffman proposed using Random Boolean Networks (RBNs) as an abstract model of gene regulatory networks • Each vertex represents a gene • “on” state of a vertex indicates that the gene is expressed • An edge from one vertex to another implies the the former gene regulates the later • “0” and “1” values on edges indicate presence/ absence of activating/repressing proteins • Boolean functions assigned to vertices represent rules of regulatory interactions between genes
  • 74. EXAMPLE NETWORK Three genes X, Y, and Z X Y Z Rules X(t+1) = X(t) and Y(t) Y(t+1) = X(t) or Y(t) Z(t+1) = X(t) or (not Y(t) and Z(t)) Current Next state state 000 000 001 001 010 010 011 010 100 011 101 011 110 111 111 111 000 001 010 011 100 101 110 111
  • 75. EXAMPLE NETWORK 000 001 010 011 100 101 110 111 X Y Z - The number of accessible states is finite, . - Cyclic trajectories are possible. - Not every state must be approachable from every other state. - The successor state is unique, the predecessor state is not unique. N 2
  • 76. RANDOM BOOLEAN NETWORKS If the rules for updating states are unknown  select rules randomly N nodes ½ pN (N-1) edges Rule 2 Rule 0 Rule 1 Rule 2
  • 77. KAUFFMAN’S NK BOOLEAN NETWORKS An NK automaton is an autonomous random network of N Boolean logic elements. Each element has K inputs and one output. The signals at inputs and outputs take binary (0 or 1) values. The Boolean elements of the network and the connections between elements are chosen in a random manner. There are no external inputs to the network. The number of elements N is assumed to be large.
  • 78. KAUFFMAN’S NK BOOLEAN NETWORKS K connections: 22K Boolean input functions Nets are free of external inputs. Once, connections and rules are selected, they remain constant and the time evolution is deterministic. Earlier work by Walker and Ashby (1965): same Boolean functions for all genes: Choice of Boolean function affects length of cycles: “and” yields short cycles, “exclusive or” yields cycles of immense length
  • 79. DYNAMICS WHEN N • Number and length of attractors of depend on k and p kc = ½ p(1-p)  k < kc frozen phase  many small attractors, not sensitive to changes in structure  k > kc chaotic phase  few large attractors, very sensitive to changes in structure  k = kc phase transition behavior  n attractors of length n, not sensitive to most changes, only to some rare ones
  • 80. KAUFFMAN MODEL • Gene regulatory networks of living cells are believed to exhibit phase transition behavior, on the border between the frozen and chaotic phases • Kaufman has shown that if k = 2 and p = 0.5, then the statistical features of RBNs match the characteristics of living cells • Number of attractors  number of cell types • Length of attractors  cell cycle time