← Stochastic Processes for Biology

Extinction and outbreak probabilities

At the beginning of an epidemic, only a few infectious individuals may be present. Even when transmission is strong enough for infection to grow on average, chance can still determine whether the first transmission chains disappear or become established.

Core idea. A deterministic model describes average growth. A stochastic model can additionally ask: what is the probability that infection dies out, and what is the probability that it develops into an outbreak?

Why can infection die out when \(R_0>1\)?

Suppose \(R_0=2\). This does not mean that every infectious person infects exactly two others. It means that the average number of secondary infections is 2 under the assumptions defining \(R_0\).

One person may infect nobody, another one person, and another four people. Therefore an early transmission chain can still disappear by chance.

\(R_0>1\) means growth is possible on average; it does not guarantee that every introduction causes a major outbreak.

Two possible outcomes from the same model

Two trajectories generated from the same early birth–death epidemic model. One reaches zero and becomes extinct; the other grows. The difference comes only from the random event sequence.

What do we mean by extinction?

For a simple closed epidemic without imported infection, extinction means that no infectious individuals remain:

\[\boxed{I(t)=0}.\]

If state 0 is absorbing, infection cannot restart from that state.

For an SEIR model, \(I=0\) alone may not mean extinction if exposed individuals remain. A more appropriate disease-free condition is

\[E=0\quad\text{and}\quad I=0.\]

What do we mean by an outbreak?

“Outbreak” must be defined before its probability is calculated. A useful definition is

\[\boxed{\text{outbreak}=\{I(t)\text{ reaches }K\text{ before }0\}},\]

where \(K\) is a chosen threshold.

The threshold \(K\) is a modelling choice. It should be selected to match the scientific question and reported explicitly.

Early transmission as a branching process

When only a few people are infected, almost everyone is still susceptible. Under suitable assumptions, each infectious individual can be viewed as starting a new transmission branch.

initial infectionone branch stopsanother continues
Some transmission branches terminate quickly while others continue. Extinction occurs only if every branch eventually terminates.

The offspring distribution

Let \(X\) be the number of secondary infections caused by one infectious individual during the early phase:

\[P(X=k)=p_k,\qquad k=0,1,2,\ldots\]

The mean is

\[E[X]=\sum_{k=0}^{\infty}kp_k.\]

In a simple branching approximation, this mean plays the role of the reproduction number.

Why the full distribution matters

Two pathogens can have the same mean number of secondary infections but different variability. If many individuals cause no infections while a few cause many, extinction behaviour can differ markedly from a model in which transmission is much more even.

This is one connection to superspreading. Transmission heterogeneity changes the offspring distribution, and therefore can change early extinction probability.

The probability-generating function

Define

\[\boxed{G(s)=p_0+p_1s+p_2s^2+p_3s^3+\cdots}.\]

Equivalently, \(G(s)=E[s^X]\).

Why does extinction satisfy \(q=G(q)\)?

Let

\[q=P(\text{eventual extinction from one initial infectious individual}).\]

If the first person produces exactly \(k\) new infectious individuals, all \(k\) descendant branches must eventually die out. Under the branching approximation, that has probability

\[q^k.\]

Weighting over all possible values of \(k\) gives

\[q=p_0+p_1q+p_2q^2+p_3q^3+\cdots,\]

so

\[\boxed{q=G(q)}.\]
Interpretation. The epidemic becomes extinct if every transmission branch created by the first case eventually becomes extinct.

Seeing the extinction fixed point

For a Poisson offspring distribution with mean 2, \(G(s)=e^{2(s-1)}\). The smaller intersection with \(y=s\) is the extinction probability; the second intersection is always at 1.

The graph is generated directly from the function \(G(s)=e^{2(s-1)}\), not drawn by eye.

Why do we choose the smaller solution?

The equation \(q=G(q)\) always has \(q=1\) as a solution because \(G(1)=1\). If the mean offspring number is at most 1, extinction occurs with probability 1 under the standard branching assumptions.

If the mean is greater than 1, a second fixed point may appear below 1. The extinction probability is the smallest solution in \([0,1]\).

A simple birth–death epidemic approximation

Suppose each infectious individual generates new infections at rate \(\lambda\) and recovers at rate \(\mu\). The next event for one infectious person is either infection or recovery.

The probabilities are

\[P(\text{recovery first})=\frac{\mu}{\lambda+\mu},\qquad P(\text{infection first})=\frac{\lambda}{\lambda+\mu}.\]

If recovery occurs first, the chain ends. If infection occurs first, there are two infectious branches, and both must eventually die out for extinction. Therefore

\[q=\frac{\mu}{\lambda+\mu}+\frac{\lambda}{\lambda+\mu}q^2.\]

This gives

\[(q-1)(\lambda q-\mu)=0.\]

Hence

\[\boxed{q=\begin{cases}1,&\lambda\le\mu,\\\mu/\lambda,&\lambda>\mu.\end{cases}}\]

Connection with \(R_0\)

For this particular simple birth–death approximation,

\[R_0=\frac{\lambda}{\mu}.\]

Therefore, when \(R_0>1\),

\[\boxed{q=\frac1{R_0}},\qquad\boxed{P(\text{major outbreak})\approx1-\frac1{R_0}}.\]
This formula is not universal. It belongs to this particular birth–death approximation. Different offspring distributions can give different extinction probabilities even when they have the same \(R_0\).

How outbreak probability changes with \(R_0\)

For the simple birth–death approximation, the major-outbreak probability is 0 for \(R_0\le1\) and \(1-1/R_0\) for \(R_0>1\). The curve is calculated directly from that formula.

Near \(R_0=1\), extinction is still very likely. As \(R_0\) increases, establishment becomes more likely, but extinction from a small introduction remains possible.

Starting with several infectious individuals

If the extinction probability from one initial infectious individual is \(q\), and the initial transmission branches can be approximated as independent, then

\[\boxed{P(\text{extinction}\mid I(0)=i_0)\approx q^{i_0}}.\]

Therefore

\[\boxed{P(\text{major outbreak}\mid I(0)=i_0)\approx1-q^{i_0}}.\]
Example. If \(q=0.6\) and \(i_0=3\), then the extinction probability is approximately \(0.6^3=0.216\), so the major-outbreak probability is approximately \(0.784\).

Finite-population hitting probabilities

For a finite birth–death CTMC, define

\[h_i=P(\text{reach }K\text{ before }0\mid I(0)=i).\]

The boundary conditions are

\[h_0=0,\qquad h_K=1.\]

For \(1\le i\le K-1\), conditioning on the next jump gives

\[h_i=\frac{b(i)}{b(i)+d(i)}h_{i+1}+\frac{d(i)}{b(i)+d(i)}h_{i-1}.\]

Equivalently,

\[\boxed{b(i)[h_{i+1}-h_i]+d(i)[h_{i-1}-h_i]=0}.\]
Interpretation. From state \(i\), the outbreak probability is the weighted average of the outbreak probabilities after the two possible next jumps.

Monte Carlo estimation

When an analytical calculation is difficult, simulate the stochastic model many times. For each run define

\[Y_r=\begin{cases}1,&\text{outbreak threshold reached first},\\0,&\text{extinction reached first}.\end{cases}\]

Then

\[\boxed{\widehat P(\text{outbreak})=\frac1n\sum_{r=1}^{n}Y_r}.\]

With enough simulations, this estimate approaches the corresponding model probability, subject to Monte Carlo sampling error.

What each quantity tells us

QuantityQuestion answered
\(R_0\)Is average early growth possible?
extinction probabilityWhat is the chance the chain eventually disappears?
outbreak probabilityWhat is the chance a specified outbreak criterion is reached?
Monte Carlo estimateHow can we estimate these probabilities for a complicated model?
Key idea. \(R_0>1\) is an average-growth threshold, not a guarantee of outbreak. Early random events can still terminate transmission. Branching processes give intuitive analytical approximations, finite CTMCs give hitting probabilities, and Monte Carlo simulation can estimate these risks for more complicated biological models.

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