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Stochastic Equations and Processes in physics and biology Andrey Pototsky Swinburne University AMSI 2017 Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 1 / 34

Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

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Page 1: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Stochastic Equations and Processes in physics and

biology

Andrey Pototsky

Swinburne University

AMSI 2017

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 1 / 34

Page 2: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Lecture 2:

Stochastic Processes

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 2 / 34

Page 3: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Stochastic Process

Definition

Any stochastic process is a probabilistic time series x(t), where x(t) is atime-dependent random variable

Coordinate of a Brownian particle vs time

0 200 400 600 800 1000time

-20

0

20

40

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 3 / 34

Page 4: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Stochastic Process

Electrocardiogram (ECG)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 4 / 34

Page 5: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Continuous stochastic process x(t)

t0 time

x0

x(t)

t3

x1

t2

x2x

3

t1

Choose a series of points on the time line

(t1 > t2 > t3 . . . ) with (x1 = x(t1), x2 = x(t2), x3 = x(t3) . . . )

x(t) is completely described by the joint distribution

p(x1, t1;x2, t2;x3, t3; . . . )

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 5 / 34

Page 6: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Ensemble of realizations

Role of initial conditions

For identical initial conditions x(t = 0) = x0, each realization (trajectory)of x(t) is different!

t0 time

x0

x0

x0

x0

x(t)

t3

(3)

t2

(2)

(1)

(4)

t1

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 6 / 34

Page 7: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Ensemble average

Averaging (integrating) over x

p(x1, t1) =

Ωdx2 p(x1, t1;x2, t2),

where Ω is the space of all possible values of x.For t1 > t2 we define transitional probability to go from (2) to (1)

p(x1, t1|x2, t2) =p(x1, t1;x2, t2)

p(x2, t2)

Time-dependent ensemble mean

〈x(t)|x0, t0〉 =

dxx p(x, t|x0, t0).

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 7 / 34

Page 8: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Autocorrelation function (ACF)

Definition

For process x(t), defined on (−∞,∞), we introduce theautocorrelation function

G(τ) = limT→∞

1

T

∫ T

0dtx(t)x(t+ τ),

ACF is an even function

G(−τ) = G(τ)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 8 / 34

Page 9: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Autocorrelation function (ACF)

Proof of G(−τ) = G(τ)

G(−τ) = limT→∞

1

T

∫ T

0dtx(t)x(t− τ) =

t− τ = y∣

T−τ

−τ, dt = dy

= limT→∞

1

T

∫ T−τ

−τ

dyx(τ + y)x(y)

= limT→∞

1

T

[∫ 0

−τ

(. . . ) +

∫ T

0(. . . )−

∫ T

T−τ

(. . . )

]

= G(τ)

The last equality holds in the limit T → ∞

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 9 / 34

Page 10: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

ACF and the ensemble average

Non-stationary ACF

The conditional (non-stationary) ACF is determined as

〈x(t)x(t′)|x0, t0〉 =

dx dx′ xx′p(x, t;x′, t′|x0, t0)

Average over time vs ensemble average

For a general process x(t)

〈x(t)x(t′)|x0, t0〉 6= limT→∞

1

T

∫ T

0dtx(t)x(t+ τ).

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 10 / 34

Page 11: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Markov processes

Absence of memory

For any t1 > t2 > t3, the probability at time t1 only conditionally dependson the state at time t3, i.e.

p(x1, t1|x2, t2;x3, t3) = p(x1, t1|x3, t3).

Consequence

p(x1, t1;x2, t2|x3, t3) = p(x1, t1|x2, t2)p(x2, t2|x3, t3).

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 11 / 34

Page 12: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Proof

Using the Markov property

p(x1, t1|x2, t2)p(x2, t2|x3, t3) = p(x1, t1|x2, t2;x3, t3)p(x2, t2|x3, t3)

=p(x1, t1;x2, t2;x3, t3)

p(x2, t2;x3, t3)

p(x2, t2;x3, t3)

p(x3, t3)

=p(x1, t1;x2, t2;x3, t3)

p(x3, t3)

= p(x1, t1;x2, t2|x3, t3).

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 12 / 34

Page 13: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

The Chapman-Kolmogorov equation

Consider all possible ways to go from (3) to (1) over (2)

p(x1, t1|x3, t3) =

Ωdx2 p(x1, t1;x2, t2|x3, t3)

=

Ωdx2

p(x1, t1;x2, t2;x3, t3)

p(x3, t3)

For Markovian process x(t)∫

Ωdx2

p(x1, t1;x2, t2;x3, t3)

p(x3, t3)=

Ωdx2 p(x1, t1|x2, t2)p(x2, t2|x3, t3)

Chapman-Kolmogorov equation

p(x1, t1|x3, t3) =

Ωdx2 p(x1, t1|x2, t2)p(x2, t2|x3, t3)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 13 / 34

Page 14: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random
Page 15: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Stationary processes

Definition

Process x(t) is called stationary if for any ǫ > 0, x(t+ ǫ) has the samestatistics as x(t). Stationary process corresponds to a remote past:

t0 → −∞.

Properties of a stationary process

limt0→−∞

p(x, t|x0, t0) = ps(x)

〈x(t)|x0, t0〉 =

x p(x, t|x0, t0) dx =

xps(x) dx

= constant

〈x(t)x(t′)|x0, t0〉 = f(t− t′).

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 14 / 34

Page 16: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

ACF of a stationary processes

In the limit t0 → −∞

ACF(t, t′)s = limt0→−∞

〈x(t)x(t′)|x0, t0〉

= limt0→−∞

xx′dx dx′ p(x, t;x′, t′|x0, t0)

= limt0→−∞

xx′dx dx′ p(x, t|x′, t′)p(x′, t′|x0, t0)

=

xx′dx dx′ p(x, t|x′, t′)ps(x′)

=

x′dx′ 〈x(t)|x′, t′〉ps(x′)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 15 / 34

Page 17: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Ergodic processes

Definition

For an ergodic process x(t), the averaging over time is equivalent to theaveraging over the ensemble.

Ergodicity vs Stationarity

Note that ergodicity is stronger than stationarity

Example:

x(t) = A, A is uniformly distributed in [0; 1]

Any realization is a straight line x(t) = Ai

Ai are different for different realizations so that 〈x(t)〉 = 0.5Average over time for a single realization

dt x(t) = Ai

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 16 / 34

Page 18: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Kramers theory of chemical reactions: (H. A. Kramers, 1940)Two reacting chemicals: X1 and X2

X1 X2

Associated bistable system x(t) = (a, b) with transition probabilities(reaction rates):

λ = lim∆t→0

1

∆tP (x = b, t+∆t|x = a, t)

µ = lim∆t→0

1

∆tP (x = a, t+∆t|x = b, t)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 17 / 34

Page 19: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Schematic representation and bistable systems

0 time

a

b

x(t)

a bx

U(x

)Other applications:

in physics: spin systems and magnetism

in finance: stoch market prices

in biology: bistable neurons

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 18 / 34

Page 20: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Master equation (Chapman-Kolmogorov)

∂tP (a, t|x, t0) = −λP (a, t|x, t0) + µP (b, t|x, t0)

∂tP (b, t|x, t0) = λP (a, t|x, t0)− µP (b, t|x, t0)

Conservation of probability at all times

∂t(P (a, t|x, t0) + P (b, t|x, t0)) = 0 ⇒ P (a, t|x, t0) + P (b, t|x, t0) = 1

Initial conditions

P (x′, t0|x0, t0) = δx′,x0=

0, x′ 6= x0,

1, x′ = x0

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 19 / 34

Page 21: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Characteristic eigenvalues γ

−λ− γ µ

λ −µ− γ

∣= 0

γ2 + γ(λ+ µ) = 0

γ1 = 0, γ2 = −(λ+ µ)

Corresponding eigenvectors

v(γ=0) =

(

C1,λ

µC1

)

, v(γ=−(λ+µ)) = (C2,−C2) ,

General solution

P (a, t|x0, t0) = C1 + C2e−(λ+µ)t,

P (b, t|x0, t0) =λ

µC1 − C2e

−(λ+µ)t

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 20 / 34

Page 22: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Solution in the compact form

P (a, t|x0, t0) =µ

λ+ µ+ e−(λ+µ)(t−t0)

(

λ

λ+ µδa,x0

−µ

λ+ µδb,x0

)

,

P (b, t|x0, t0) =λ

λ+ µ− e−(λ+µ)(t−t0)

(

λ

λ+ µδa,x0

−µ

λ+ µδb,x0

)

Stationary distribution: t0 → −∞

Ps(a) =µ

λ+ µ, Ps(b) =

λ

λ+ µ.

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 21 / 34

Page 23: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Time-dependent ensemble average

〈x(t)|x0, t0〉 = aP (a, t|x0, t0) + b P (b, t|x0, t0)

=aµ+ bλ

µ+ λ+ exp [−(λ+ µ)(t− t0)]

(a− b)(λδa,x0− µδb,x0

)

µ+ λ

Note that(

x0 −aµ+ bλ

µ+ λ

)

=(a− b)(λδa,x0

− µδb,x0)

µ+ λ

Final result

〈x(t)|x0, t0〉 =aµ+ bλ

µ+ λ+ exp [−(λ+ µ)(t− t0)]

(

x0 −aµ+ bλ

µ+ λ

)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 22 / 34

Page 24: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Stationary average xs

xs = limt0→−∞

〈x(t)|x0, t0〉 =aµ+ bλ

µ+ λ.

Stationary variance Var(x)s = 〈x2〉s − x2s

Var(x)s =a2µ

λ+ µ+

b2λ

λ+ µ−

(aµ+ bλ)2

(λ+ µ)2

=a2µ(λ+ µ) + b2λ(λ+ µ)− (a2µ2 + b2λ2 − 2abµλ)

(λ+ µ)2

=(a− b)2µλ

(λ+ µ)2

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 23 / 34

Page 25: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Stationary ACF

ACF(t, t′)s =∑

(x,x′=a,b)

xx′P (x, t|x′, t′)Ps(x′)

=∑

(x′=a,b)

x′Ps(x′)

(x=a,b)

xP (x, t|x′, t′)

=∑

(x′=a,b)

x′〈x(t)|x′, t′〉Ps(x′)

= a〈x(t)|a, t′〉Ps(a) + b〈x(t)|b, t′〉Ps(b)

=

(

aµ+ bλ

µ+ λ

)2

+(a− b)2µλ

(µ+ λ)2exp [−(λ+ µ)(t− t′)]

= x2s +Var(x)s exp [−(λ+ µ)(t− t′)]

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 24 / 34

Page 26: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Process centered about mean value x(t)− xs

〈x(t)− xs|x0, t0〉 = exp [−(λ+ µ)(t− t0)] (x0 − xs)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 25 / 34

Page 27: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Stationary ACF of the centered process

G(t, t′) = 〈(x(t)− xs)(x(t′)− xs)|x0, t0〉

=

dx dx′(x− xs)(x′ − xs)p(x, t;x

′, t′|x0, t0)

=

dx dx′(xx′ − xsx− xsx′ + x2s)p(x, t;x

′, t′|x0, t0)

= 〈x(t)x(t′)|x0, t0〉

− xs

dx dx′ x′p(x, t;x′, t′|x0, t0)

− xs

dx dx′ xp(x, t;x′, t′|x0, t0)

+ x2s

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 26 / 34

Page 28: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Note that

dx dx′ x′p(x, t;x′, t′|x0, t0) =

dx′ x′p(x′, t′|x0, t0)

= 〈x(t′)|x0, t0〉∫

dx dx′ xp(x, t;x′, t′|x0, t0) =

dxxp(x, t|x0, t0)

= 〈x(t)|x0, t0〉

in the limit t0 → −∞

〈x(t)|x0, t0〉 = 〈x(t′)|x0, t0〉 = xs

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 27 / 34

Page 29: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Stationary ACF of the centered process

G(t, t′) = ACF(t, t′)s − 〈x〉2s

= Var(x)s exp [−(λ+ µ)(t− t′)]

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 28 / 34

Page 30: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Random telegraph process

Survival probability

Given that at time t = t0 the system was in state x = a, what is theprobability P (a, t|a, t0) for the system to stay in the same state at timet > t0?

∂tP (a, t|a, t0) = −λP (a, t|a, t0), P (a, t = t0|a, t0) = 1

∂tP (b, t|b, t0) = −µP (b, t|b, t0), P (b, t = t0|b, t0) = 1

Survival probabilities for a random telegraph process

P (a, t|a, t0) = e−λ(t−t0), P (b, t|b, t0) = e−µ(t−t0).

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 29 / 34

Page 31: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Residence times

ta . . . time spent in state a

tb . . . time spent in state b

cdf of the residence times

cdfa(t) = Pr(ta ≤ t) = 1− Pr(ta ≥ t) = 1− P (a, t|a, t0) = 1− e−λ(t−t0)

cdfb(t) = Pr(tb ≤ t) = 1− Pr(tb ≥ t) = 1− P (b, t|b, t0) = 1− e−µ(t−t0)

pdf and the average residence times

pdfa(t) = cdfa(t)′ = λe−λ(t−t0), 〈ta〉 = λ−1

pdfb(t) = cdfb(t)′ = µe−µ(t−t0), 〈tb〉 = µ−1

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 30 / 34

Page 32: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Relation of the random telegraph process to thePoisson distribution

Exponential vs Poisson

If the distribution of the residence times is exponential with the parameterλ, then that distribution of the number of switches N in any interval τfollows the Poisson distribution

P (N = k) =(τλ)k

k!e−τλ.

Expected value and variance

E(N) = τλ, Var(N) = τλ

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 31 / 34

Page 33: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Examples of the Poisson distribution

Radioactivity (number of decays in a given time interval)

Retail markets (number of customers arriving at a shop in a giventime interval, or the number of purchases per day, per hour or perminute)

Telecommunication (number of telephone calls arriving per given timeinterval)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 32 / 34

Page 34: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Renewal process

Sequence of random variables si with identical distribution

(s1, s2, s3, . . . )

General theory

David R. Cox Renewal Theory

Igor Goychuk and Peter Hanggi, Theory of non-Markovian stochastic

resonance, Phys Rev. E 69, 021104 (2004)

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 33 / 34

Page 35: Stochastic Equations and Processes in physics and …...Stochastic Process Definition Any stochastic process is a probabilistic time series x(t), where x(t) is a time-dependent random

Firing neuron as a renewal process

t

-1

1

x

-2

0

2

4x

-1

1x

(a)

(b)

(c)

TW

TRTE

TE TR

Excitable (FitzHugh−Nagumo) system

Representing excitable system as a two−state system

Noisy bistable system with a single delay

=

Andrey Pototsky (Swinburne University) Stochastic Equations and Processes in physics and biology AMSI 2017 34 / 34