Generically, suppose that we have a system of particles that can generate or split into other particles of the same type. Here are some typical examples:
The particles are biological organisms that reproduce.
The particles are neutrons in a chain reaction.
The particles are electrons in an electron multiplier.
We assume that the lifetime of each particle is exponentially distributed with parameter , and at the end of its life, is replaced by a random number of new particles that we will refer to as children of the original particle. The number of children of a particle has probability density function on . The particles act independently, so in addition to being identically distributed, the lifetimes and the number of children are independent from particle to particle. Finally, we assume that , so that a particle cannot simply die and be replaced by a single new particle. Let and denote the mean and variance of the number of offspring of a single particle. So
We assume that is finite and so makes sense. In our study of discrete-time Markov chains, we studied branching chains in terms of generational time. Here we want to study the model in real time.
Let denote the number of particles at time . Then is a continuous-time Markov chain on , known as a branching chain. The exponential parameter function and jump transition matrix are given by
for
for and .
Details:
That is a continuous-time Markov chain follows from the assumptions and the basic structure of continuous-time Markov chains. In turns out that the assumption that implies that is regular, so that as , where is the time of the th jump for .
Starting with particles, the time of the first state change is the minimum of independent variables, each exponentially distributed with parameter . As we know, this minimum is also exponentially distributed with parameter .
Starting in state , the next state will be for , if the particle dies and leaves children in her place. This happens with probability .
Of course 0 is an absorbing state, since this state means extinction with no particles. (Note that and so by default, .) So with a branching chain, there are essentially two types of behavior: population extinction or population explosion.
For the branching chain one of the following events occurs with probability 1:
Extinction: for some and hence for all .
Explosion: as .
Details:
If then all states lead to the absorbing state 0 and hence the set of positive staties is transient. With probability 1, the jump chain visits a transient state only finitely many times, so with probability 1 either for some or as . If then is strictly increasing in , since by assumption. Hence with probability 1, as .
Without the assumption that , explosion can actually occur in finite time. On the other hand, the assumption that is for convenience. Without this assumption, would still be a continuous-time Markov chain, but as discussed in the Introduction, the exponential parameter function would be for and the jump transition matrix would be
Because all particles act identically and independently, the branching chain starting with particles is essentially independent copies of the branching chain starting with 1 particle. In many ways, this is the fundamental insight into branching chains, and in particular, means that we can often condition on .
Generator and Transition Matrices
As usual, we will let denote the semigroup of transition matrices of , so that for . Similarly, denotes the infinitesimal generator matrix of .
The infinitesimal generator is given by
Details:
This follows immediately from the exponential parameter function and the jump transition matrix in [1].
The Kolmogorov backward equation is
Details:
The backward equation is , so the result follows from [3].
Unlike some of our other continuous-time models, the jump chain governed by is not the discrete-time version of the model. That is, is not a discrete-time branching chain, since in discrete time, the index represents the th generation, whereas here it represent the th time that a particle reproduces. However, there are lots of discrete-time branching chain embedded in the continuous-time chain.
In general, we know that sampling a (homogeneous) continuous-time Markov chain at multiples of a fixed , results in a (homogeneous) discrete-time Markov chain. For to be a branching chain, we just need to note that
where is the convolution power of of order . This is a consequence of the fundamental fact that given has the same distribution as the sum of independent copies of given . Recall that the PDF of a sum of independent variables is the convolution of the individual PDFs.
For let denote the probability generating function of given
Let denote the probability generating function of The generating functions are defined (the series are absolutely convergent) at least for .
The collection of generating functions gives the same information as the collection of probability density functions . With the fundamental insight that the branching process starting with one particle determines the branching process in general, actually determines the transition semigroup .
For and , the probability generating function of given is :
Details:
Again, given , the number of particles at time has the same distribution as the sum of independent copies of given . Recall that the PGF of a sum of independent variables is the product of the PGFs of the variables.
Note that is the generating function of the offspring distribution for the embedded discrete-time branching chain for . On the other hand, is the generating function of the offspring distribution for the continuous-time chain. So our main goal in this discussion is to see how is built from . Because is a semigroup under matrix multiplication, and because the particles act identically and independently, is a semigroup under composition.
for .
Details:
Using the semigroup property (the Chapman-Kolmogorov equations) and [7] we have
Note also that for all . This also follows from the semigroup property: . The fundamental relationship between the collection of generating functions and the generating function is given in the following theorem:
The mapping satisfies the differential equation
Details:
Using the Kolmogorov backward equation in [4] we have
Using the generator in [2],
Substituting and using [7] gives
This differential equation, along with the initial condition for all determines the collection of generating functions . In fact, an implicit solution for is given by the integral equation
Another relationship is given in the following theorem. Here, refers to the derivative of the generating function with respect to its argument, of course (so , not ).
For ,
Details:
From the semigroup property, we have for . Differentiating with respect to and using the chain rule along with [9] gives
Evaluating at and using the condition we have
Using the [9] once again gives
Solving for gives the result.
Moments
In this discussion, we wil study the mean and variance of the number of particles at time . Let
so that and are the mean and variance, starting with a single particle. As always with a branching process, it suffices to consider a single particle:
For and ,
Details:
Once again, the distribution of given is the same as the distribution of the sum of independent copies of given . Recall that the mean of a sum of variables is the sum of the individual means, and the variance of the sum of independent variables is the sum of the individual variances.
Recall also that and are the the mean and variance of the number of offspring of a particle. Here is the connection between the means:
for .
If then as . This is extinction in the mean.
If then as . This is explosion in the mean.
If then for all . This is stability in the mean.
Details:
From the proof of [10],
Differentiating with respect to , interchanging the order of integration on the left, and using the product rule on the right gives
Now let and recall that . We get
From the basic theory of probability generating functions, and similarly, . Hence we have
Of course we have the initial condition .
This result is intuitively very appealing. As a function of time, the expected number of particles either grows or decays exponentially, depending on whether the expected number of offspring of a particle is greater or less than one. The connection between the variances is more complicated. We assume that .
If then
If then .
If then as
If then as
Details:
Probability generating functions are naturally connected to factorial moments, so it's best to work with these. Thus, let for and let . These are the factorial moments of order 2. In the proof of the last theorem we showed that
Differentiating with respect to again gives
Now substitute . Recall that , , , , and . We get the differential equation
with the initial condition .
Suppose that . Then using standard methods for a linear, first order differential equations with constant coefficients and an exponential forcing function, the solution is
But , and similarly with . Substitution and some algebra then gives the result.
Suppose now that . Then also for all and so and . The differential equation above reduces simply to
with initial condition so trivially . Finally, in the context of part (b), note that if we must have since we have assumed that .
If so that as and we have extinction in the mean, then as also. If so that as and we have explosion in the mean, then as also. We would expect these results. On the other hand, if so that for all and we have stability in the mean, then grows linearly in . This gives some insight into what to expect next when we consider the probability of extinction.
The Probability of Extinction
As shown in [2], there are two types of behavior for a branching process, either population extinction or population explosion. In this discussion, we study the extinction probability, starting as usual with a single particle:
Need we say it? The extinction probability starting with an arbitrary number of particles is easy.
For ,
Details:
Given , extinction has occurred by time if and only if extinction has occurred by time for each of the independent branching chains formed from the descendents of the initial particles.
We can easily relate extinction for the continuous-time branching chain to extinction for any of the embedded discrete-time branching chains.
If extinction occurs for then extinction occurs for for every . Conversely if extinction occurs for for some then extinction occurs for for every and extinction occurs for . Hence is the minimum solution in of the equation for every .
Details:
The statements about the extinction event follow immediately from the fact that is absorbing, so that if for some then for every . The result for the extinction probability follows from the theory of discrete-time branching chains.
So whether or not extinction is certain depends on the critical parameter .
The extinction probability and the mean of the offspring distribution are related as follows:
If then , so extinction is certain.
If then , so there is a positive probability of extinction and a positive probability of explosion.
Details:
These results follow from the corresponding results for discrete-time branching chains. Fix and recall that is the mean of the offspring distribution for the discrete-time chain . From [12],
If then .
If then .
It would be nice to have an equation for in terms of the offspring probability generating function . This is also easy
The probability of extinction is the minimum solution in of the equation .
Details:
From [16], for every . Substituting in exercose [9], we have and hence . As in the theory of discrete branching chains, the equation has only the solution 1 in (0, 1] if or there are two solutions and if . In both cases, is the smaller solution.
Special Models
We now turn our attention to a number of special branching chains that are important in applications or lead to interesting insights. We will use the notation established above, so that is the parameter of the exponential lifetime of a particle, is the transition matrix of the jump chain, is the infinitesimal generator matrix, and is the transition matrix at time . Similarly, , , and are the mean, variance, and generating function of the number of particles at time , starting with a single particle. As always, be sure to try these exercises yourself before looking at the proofs and solutions.
The Pure Death Branching Chain
First we consider the branching chain in which each particle simply dies without offspring. Sadly for these particles, extinction is inevitable, but this case is still a good place to start because the analysis is simple and lead to explicit formulas. Thus, suppose that is a branching process with lifetime parameter and offspring probability density function with .
The transition matrix of the jump chain and the generator matrix are given by
and for
for and for
The time-varying functions are more interesting.
Let . Then
for
Given the distribution of is binomial with trial parameter and success parameter .
Details:
All of these results follow from the general methods above, with and for . But it's helpful to give direct proofs. Given , let be the time until the first transition, which is simply the lifetime of the particle. So has the exponential distribution with parameter . For , is an indicator random variable (taking just values 0 and 1) with
Part (a), (b), and (c) are standard results for an indicator variable. For part (d), given , each of the particles, independently, is still alive at time with probability . Hence the number of particles still alive has the binomial distribution with parameters and .
In particular, note that as . that is, the probability of extinction by time increases to 1 exponentially fast. Since we have an explicit formula for the transition matrices, we can find an explicit formula for the potential matrices as well. The result uses the beta function.
For the potential matrix is given by
For , the potential matrix is given by
for
for and .
Details:
Suppose that and that with . By definition
Substitute so that or equivalently . After some algebra, the result is
By definition, the last integral is .
For ,
For with , the derivation above and properties of the beta function give
We could argue the results for the potential directly. Recall that is the expected time spent in state starting in state . Since 0 is absorbing and all states lead to 0, for . If and , then leads to with probability 1. Once in state the time spent in has an exponential distribution with parameter , and so the mean is . Of course, when the chain leaves , it never returns.
Recall that is a transition probability matrix for , and in fact is the probability density function of given where is independent of has the exponential distribution with parameter . For the next result, recall the ascending power notation
For and , the function is the beta-binomial probability density function with parameters , , and 1.
Details:
From [20] and properties of the beta function.
But from properties of the beta function,
Substituting gives the result
The Yule Process
Next we consider the pure birth branching chain in which each particle, at the end of its life, is replaced by 2 new particles. Equivalently, we can think of particles that never die, but each particle gives birth to a new particle at a constant rate. This chain could serve as the model for an unconstrained nuclear reaction, and is known as the Yule process, named for George Yule. So specifically, let be the branching chain with exponential parameter and offspring probability density function given by . Explosion is inevitable, starting with at least one particle, but other properties of the Yule process are interesting. in particular, there are fascinating parallels with the pure death branching chain in . Since 0 is an isolated, absorbing state, we will sometimes restrict our attention to positive states.
The transition matrix of the jump chain and the generator matrix are given by
and for
for and for
Since the Yule process is a pure birth process and the birth rate in state is , the process is also called the linear birth chain. As with the pure death process, we can give the distribution of specifically.
Proof from the general results: Parts (a) and (b) follow from the general moment results above, with and . For part (c), note that for , so the integral equation for is
From partial fractions, , so the result follows by standard integration and algebra. We recognize as the probability generating function of the geometric distribution on with success parameter , so for part (d) we use our standard argument. Given , has the same distribution as the sum of independent copies of given , and so this is the distribution of the sum of independent variables each with the geometric distribution on with parameter . But this is the negative binomial distribution on with parameters and .
Direct proof: As usual, let and let denote the time of the th transition (birth) for . Given , the population is at time . So the random interval (the time until the next birth) has the exponential distribution with parameter and these intervals are independent as varies. From a result in the section on the exponential distribution, it follows that has distribution function given by
Curiously, this is also the distribution function of the maximum of independent variables, each with the exponential distribution with rate . Hence
and therefore
So given , has the geometric distribution with parameter . The other results then follow easily.
Recall that the negative binomial distribution with parameters and governs the trial number of the th success in a sequence of Bernoulli Trials with success parameter . So the occurrence of this distribution in the Yule process suggests such an interpretation. However this interpretation is not nearly as obvious as with the binomial distribution in the pure death branching chain. Next we give the potential matrices.
For the potential matrix is given by
If , the function is the beta-negative binomial probability density function with parameters , , and 1:
Details:
The proof is very similar to the one in [23]. Suppose that and that with . By definition
Substitute so that or equivalently . After some algebra, the result is
By definition, the last integral is .
If we think of the Yule process in terms of particles that never die, but each particle gives birth to a new particle at rate , then we can study the age of the particles at a given time. As usual, we can start with a single, new particle at time 0. So to set up the notation, let be the Yule branching chain with birth rate , and assume that . Let and for , let denote the time of the th transition (birth).
For , let denote the total age of the particles at time . Then
The random process is the age process.
Details:
Note that there have been births in the interval . For , the age at time of the particle born at time is .
Here is another expression for the age process.
Again, let be the age process for the Yule chain starting with a single particle. Then
Details:
Suppose that where , so that . Note that for and , while for . Hence
From [25],
With the last representation, we can easily find the expected total age at time .
Again, let be the age process for the Yule chain starting with a single particle. Then
Details:
We can interchange the expected value and the integral by Fubini's theorem. So using [23],
The General Birth-Death Branching Chain
Next we consider the continuous-time branching chain in which each particle, at the end of its life, leaves either no children or two children. At each transition, the number of particles either increases by 1 or decreases by 1, and so such a branching chain is also a continuous-time birth-death chain. Specifically, let be a continuous-time branching chain with lifetime parameter and offspring probability density function given by , , where . When we have the pure death chain in [18], and when we have the Yule process in [22]. We have already studied these, so the interesting case is when so that both extinction and explosion are possible.
The transition matrix of the jump chain and the generator matrix are given by
, and , for
for , and , for
As mentioned earlier, is also a continuous-time birth-death chain on , with 0 absorbing. In state , the birth rate is and the death rate is . The moment functions are given next.
For ,
If ,
If , .
Details:
These results follow from the general formulas above for and , since and .
The next result gives the generating function of the offspring distribution and the extinction probability.
For the birth-death branching chain,
for .
if and if .
Details:
By definition, .
The equation factors into . The extinction probability is the smaller solution in .
Graphs of and when Graphs of and when
For , the generating function is given by
Details:
The integral equation for is
The denominator in the integral factors into . If , use partial fractions, standard integration, and some algebra. If the factoring is and partial fractions is not necessary. Again, use standard integration and algebra.