Two Channels of One River: How Chance Shapes Evolution ✎

Original: "How deterministic trajectories and their fluctuations underlie allele-frequency statistics in the strong-selection regime"
· David Waxman
arXiv:2609.14549v1 · 2026-09-13 · arXiv non-exclusive · independent analysis · 1 min
Even when selection is strong, random events can wipe out a beneficial trait, and that has to be factored in.

A river flows downhill, but some of its water slips into a side channel. Evolution works the same way: selection pushes a beneficial gene toward fixation, while chance can drag it toward extinction. For a long time, scientists watched only the main channel, tracking how a gene's frequency rises. The side channel, where the gene is lost, was dismissed as unimportant. That mistake was a big one: when calculating the spread of frequency, it reached 100%.

The new method keeps an eye on both channels. It splits a gene's history into two paths: fixation and loss. Each path gets its own trajectory, and then they are combined with probabilities taken into account. This makes it possible to predict precisely the mean frequency of a gene and its spread at any moment. For example, with selection strength R=50 and an initial frequency of 0.001, the error in the mean drops from 10.6% to 3.2%, and the error in the spread falls from 12.6% to 4.9%.

Even under strong selection, random fluctuations can dominate the spread of a gene's frequency in the early stages.

Now researchers can see both channels of the river. The method helps analyze data from evolution experiments on viruses, bacteria, and artificial selection. It estimates selection strength more accurately and predicts whether a beneficial gene will fix in a small population. The river of evolution has become clearer.

Even under strong selection, when the deterministic force is large, the variance of the allele frequency can be almost entirely determined by random fluctuations for a long time. For example, at R=50 and initial frequency y=0.001, the deterministic contribution to the variance is essentially zero up to time τ≈5, whereas the fluctuation contribution accurately describes the actual variance.

R = 2 N_e |s|
R shows how many times the deterministic change in allele frequency due to selection exceeds random changes due to drift. When R >> 1, selection is strong.
Q(\tau) \simeq Q_0(\tau) + \frac{1}{R} Q_1(\tau)
The statistic Q (e.g., mean or variance) is expanded in the small parameter 1/R. Q0 is the contribution of the two deterministic trajectories, and Q1/R is the contribution of fluctuations.
\pi^{(1)}(y) = \frac{1 - e^{-2 \sigma R y}}{1 - e^{-2 \sigma R}}
The probability that an allele with initial frequency y eventually fixes (reaches frequency 1). This is Kimura's formula, depending on R and the sign of selection σ.
Concepts
Wright–Fisher model Fisher's fundamental theorem of natural selection statistical method trials factor transcription factor Percentile stochastic gene expression regime shift random walk average diagnostic accuracy