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Saturday, June 13, 2020

R. classicum ...? More like R. catalanii

The R. classicum reproductive behavior leads to some familiar combinatorial behavior involving the Catalan numbers. I'm not totally sure how to get the made up scientific nomenclature of a non-existent bacteria changed ... but I think we should look into it.

First the setup:
R. classicum either splits into two copies (with probability p) or dies (with probability 1p). If you start with a single R. classicum bacterium, what is the probability that it will lead to an everlasting colony (i.e., the colony will theoretically persist for an infinite amount of time)?

Theorem. The probability that a single R. classicum bacteria leads to an everlasting colony is E(p)=max{21p,0}. In addition to the colony population at time t, Nt, let's additionally catalog the history of a bacteria colony by tracking two quantities: Rt and Dt, the number of reproductions/doublings and deaths that have occurred up until time t. For consistency, let's define R0=D0=0.

Proposition. For each t0, Nt=RtDt+1.
Proof. We will proceed by induction. The base case of t=0 satisfies the equation.
Assume that for some time t=τ0 that the formula holds, that is, Nτ=RτDτ+1.
Mechanically we see that Nτ+1 is equal to twice the number of bacteria that reproduced at time t=τ+1, so in particular Rτ+1=Nτ+12+Rτ. Additionally, the remainder of the population at time t=τ that did not reproduce must have died, we have Dτ+1=(NτNτ+12)+Dτ. If we subtract the deaths equation from the reproductions, we get Rτ+1Dτ+1=Nτ+1Nτ+RτDτ. From the inductive hypothesis we have, Nτ=RτDτ+1, so we can substitute above and get Rτ+1Dτ+1=Nτ+1(RτDτ+1)+RτDτ=Nτ+11, or equivalently Nτ+1=Rτ+1Dτ+1+1. Thus, by simple induction, the proposition holds for all time steps t0.

The corollary follows immediately.

Corollary. The colony population collapses if and only if there is some step t>0 for which Dt=Rt+1.

Using the corollary, we can then proceed with the proof of the main Theorem.

Proof. Let's calculate 1E(p), which is the the probability that the R. classicum colony eventually collapses. From the corollary, this means that there is some t>0 for which Dt=Rt+1. We see that the probability of a colony history with, say k reproductions and k+1 deaths is pk(1p)k+1. Let Ck denote the number of distinct colony histories that end in collapse after k reproductions and k+1 deaths.
Then the probability of that the colony eventually collapses is 1E(p)=k=0Ckpk(1p)k+1. Now the only thing that remains to show is to find out what Ck is (though I imagine you've already gathered where I'm going with this...). Since a population collapse needs to end it at least one death, the remaining k reproductions and k deaths can be ordered in any way such that the cumulative number of deaths is never more than the cumulative number of reproductions. Thus there is a simple bijection between colony histories with k reproductions and k+1 deaths and nonnegative Dyck paths of length 2k (for instance set reproductions to ``up'' steps and deaths to ``down'' steps), of which there are Ck=1k+1(2kk), that is, the kth Catalan number, such paths.
Using the generating function for the Catalan numbers, c(x)=n=0Cnxn=114x2x we get that the probability of eventual colony collapse is 1E(p)=k=0Ckpk(1p)k+1=(1p)c(p(1p))=(1p)114p(1p)2p(1p)=114p+4p22p=1|12p|2p. So, for p>0.5, we have E(p)=11|12p|2p=11(2p1)2p=12(1p)2p=21p; whereas for p0.5, we have E(p)=11|12p|2p=11(12p)2p=12p2p=0. Altogether, the formula E(p)=max{21p,0} is proved.

Editor's Note: I actually solved this brute force in Excel by setting up several steps of the process, seeing the when p=0.8 that the probability of having a zero population quickly to 0.25. Varying the input value of p quickly showed that the probability of having a zero population was min{1,(1p)/p}. However, I was convinced that there was a theoretical solution and was pleasantly surprised that it led solidly back to my undergraduate research in enumerative combinatorics with Prof. Ira Gessel at Brandeis University (cf. this sloppy arXiv article).

Editor's Note 2: This seems like a really convoluted way of proving that if the expected population of the colony decreases to zero over time (that is, when p<0.5), that population collapse is almost certain. I suppose it is a roughly serviceable level of convolution for showing almost sure population collapse when the expected value is constant (that is, when p=0.5).

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