Lecture 10: Non-linear Dynamical Systems - Part...

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Principles of stability and bifurcation in simple non-linear dynamical systems Lecture 10: Non-linear Dynamical Systems - Part 2 R. Ranganathan Green Center for Systems Biology, ND11.120E Albert Libchaber 1934 - Edward Lorenz 1917-2008 Henri Poincare 1854 - 1912 Robert May 1936 - Mitchell Feigenbaum 1944 - Joseph-Louis LaGrange 1736 - 1813

Transcript of Lecture 10: Non-linear Dynamical Systems - Part...

Page 1: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Principles of stability and bifurcation in simple non-linear dynamical systems

Lecture 10: Non-linear Dynamical Systems - Part 2

R. Ranganathan Green Center for Systems Biology, ND11.120E

Albert Libchaber 1934 -

Edward Lorenz 1917-2008

Henri Poincare 1854 - 1912

Robert May

1936 -

Mitchell Feigenbaum 1944 -

Joseph-Louis LaGrange 1736 - 1813

Page 2: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

So, today we explore the truly astounding emergent complexity inherent in even simple non-linear dynamical systems.

Linear

Nonlinear

n = 1

n = 2 or 3

n >> 1

continuum

exponential growth and decay

single step conformational change

fluorescence emission

pseudo first order kinetics

fixed points

bifurcations, multi stability

irreversible hysteresis

overdamped oscillators

second order reaction kinetics

linear harmonic oscillators

simple feedback control

sequences of conformational change

anharmomic oscillators

relaxation oscillations

predator-prey models

van der Pol systems

Chaotic systems

electrical circuits

molecular dynamics

systems of coupled harmonic oscillators

equilibrium thermodynamics

diffraction, Fourier transforms

systems of non-linear oscillators

non-equilibrium thermodynamics

protein structure/function

neural networks

the cell

ecosystems

Diffusion

Wave propagation

quantum mechanics

viscoelastic systems

Nonlinear wave propagation

Reaction-diffusion in dissipative systems

Turbulent/chaotic flows

adapted from S. Strogatz

Page 3: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

So, today:

(1) A reminder of the van der Pol oscillator - a small non-linear system that illustrates analysis of stability, limit cycle oscillations, and bifurcation.

(2) Concepts of local linearization and formal analyses of stability and bifurcation. Examples in the classic van der Pol oscillator, the Fitzhigh-Nagumo model, and perhaps a higher order system.

Page 4: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

The general solution to second-order linear system…

…a vector of initial conditionsgiven

….where A is the characteristic matrix. It’s properties control all behaviors of the system

Page 5: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Properties of the characteristic matrix…

…the eigenvalues of A are completely determined by the trace and determinant…

Page 6: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

…stability is determined by the real part of the eigenvalue…

System behaviors: the second order linear case

Page 7: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

We know how to analyze the behavior….right?

Seeing behaviors: the linear harmonic oscillator

Page 8: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Eigenvalues are pure imaginary….so centers!

Seeing behaviors: the linear harmonic oscillator

Page 9: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Eigenvalues are pure imaginary….so centers!

And…remember the system nullclines, which help us sketch the behavior in the phase space…

Seeing behaviors: the linear harmonic oscillator

Page 10: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

Here is the non-linearity….with mu controlling the degree of non-linearity.

Page 11: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Re-writing the equations in a more intuitive way....

A non-linear oscillator...

Page 12: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where

Now, we compute fixed points and nullclines and sketch the behavior in the x, y space….

Page 13: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where

Fixed point at origin. At any non-zero (x, y) the system has a limit cycle oscillation…

Page 14: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where

Fixed point at origin. At any non-zero (x, y) the system has a limit cycle oscillation…

A closed orbit that another trajectory spirals into as time goes to infinity…

Page 15: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where

The limit cycle shows the property of a large divergence of time scales….

Page 16: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where

Page 17: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

Page 18: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

a network of elementary passive and active components

Page 19: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

a network of elementary passive and active components

things that use but do not produce energy

(resistor)

(capacitor)

(inductor)

Page 20: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

a network of elementary passive and active components

voltage and current sources

Page 21: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

here, there is also a non-linear resistor…with a cubic response function

Page 22: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Based on Kirchhoff’s voltage law…

A non-linear oscillator...

, or….

Page 23: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Based on Kirchhoff’s voltage law…

A non-linear oscillator...

, or….

Page 24: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Based on Kirchhoff’s voltage law…

A non-linear oscillator...

, or….

because of the chain rule…

Page 25: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Based on Kirchhoff’s voltage law…

A non-linear oscillator...

, or….

we can simplify this using our basic voltage equation for the non-linear resistor…

Page 26: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

if we choose a =1 and b = 1/3….

Page 27: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

if we choose a =1 and b = 1/3….

starting to look like our usual van der Pol system…

Page 28: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

An electrical circuit implementation of the van der Pol oscillator…

A non-linear oscillator...

if we choose a =1 and b = 1/3….

and, taking the derivative….

Page 29: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Based on Kirchhoff’s voltage law…

A non-linear oscillator...

, or….

this is the differential equation that controls our system…

Page 30: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Based on Kirchhoff’s voltage law…

A non-linear oscillator...

, or….

And now for a little magic. I will re-scale time so that

Page 31: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Based on Kirchhoff’s voltage law…

A non-linear oscillator...

, or….

, where…

Page 32: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

Remember the basic van der Pol oscillator equation?

Page 33: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

Remember the basic van der Pol oscillator equation?

Page 34: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

…the actual implementation of our non-linear resistor element, with appropriate choices of R1-R5 to get a=1, b=1/3, and the effective net resistance to be R

Page 35: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

now…let’s carry out stability and bifurcation analysis of this system

Page 36: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

again, a re-writing of our equation to represent the phase space…and we know the fixed point:

To study the stability of the fixed point, we carry out a local linearization…and then look at the flow.

Page 37: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Page 38: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

that is, (x*,y*) is a fixed point

Page 39: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Now, we introduce a disturbance around the fixed point….

Page 40: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Now, we introduce a disturbance around the fixed point….

To see if the disturbance grows or not, we look at the derivatives…

Similar thing for the disturbance v….

Page 41: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Now, we introduce a disturbance around the fixed point….

To see if the disturbance grows or not, we look at the derivatives…

…and in matrix form,

Page 42: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Now, we introduce a disturbance around the fixed point….

To see if the disturbance grows or not, we look at the derivatives…

…and ignoring the quadratic and higher order terms, since they are tiny….

Page 43: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Now, we introduce a disturbance around the fixed point….

To see if the disturbance grows or not, we look at the derivatives…

…this is the locally linearized form of our general non-linear system…

Page 44: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Now, we introduce a disturbance around the fixed point….

To see if the disturbance grows or not, we look at the derivatives…

this matrix is called the Jacobian, and evaluated at the fixed point (x*,y*), tells us about the flow of the system in the local environment…

Page 45: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A general non-linear system…

and let’s say

Now, we introduce a disturbance around the fixed point….

To see if the disturbance grows or not, we look at the derivatives…

the Jacobian. In this form, this matrix is just like the characteristic matrix for a linear system, right? So, we know how to analyze its behavior…

Page 46: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

At the fixed point….

And what are the eigenvalues? Remember that stability is about the sign of the real part of the system eigenvalues…

Page 47: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

At the fixed point….

The trace of the Jacobian is epsilon, and so the system is stable for negative values and unstable for positive…

Page 48: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

At the fixed point….

So, for a van der Pol oscillator, the system loses stability at the place where the real part of the system eigenvalues go from negative to positive….such an event is called a Hopf bifurcation

Page 49: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

, where…

Page 50: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

The neuronal action potential…a slight variation on the van der Pol oscillator…

A non-linear oscillator...

Page 51: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

this is essentially the van der Pol oscillator, with one difference….

Fitzhugh-Nagumo (1962)

membrane pot

slow K+ flux

Page 52: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

the linear term to the w nullcline provides for thresholded oscillation….

membrane pot

slow K+ flux

Page 53: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

A non-linear oscillator...

the linear term to the w nullcline provides for thresholded oscillation….a stable fixed point destabilized to produce relaxation oscillations. We will look at this more closely next time…

membrane pot

slow K+ flux

Page 54: Lecture 10: Non-linear Dynamical Systems - Part 2ranganathanlab.org/.../QB_lecture10_NonLin_part2v2.pdf · Principles of stability and bifurcation in simple non-linear dynamical systems

Next, we will further analyze the simple non-linear oscillator systems…

Linear

Nonlinear

n = 1

n = 2 or 3

n >> 1

continuum

exponential growth and decay

single step conformational change

fluorescence emission

pseudo first order kinetics

fixed points

bifurcations, multi stability

irreversible hysteresis

overdamped oscillators

second order reaction kinetics

linear harmonic oscillators

simple feedback control

sequences of conformational change

anharmomic oscillators

relaxation oscillations

predator-prey models

van der Pol systems

Chaotic systems

electrical circuits

molecular dynamics

systems of coupled harmonic oscillators

equilibrium thermodynamics

diffraction, Fourier transforms

systems of non-linear oscillators

non-equilibrium thermodynamics

protein structure/function

neural networks

the cell

ecosystems

Diffusion

Wave propagation

quantum mechanics

viscoelastic systems

Nonlinear wave propagation

Reaction-diffusion in dissipative systems

Turbulent/chaotic flows

adapted from S. Strogatz