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Diagrammatic expressions for steady-state distribution and static responses in population dynamics

Koya Katayama1,*, Ryuna Nagayama1, and Sosuke Ito1,2

  • *Contact author: koya.katayama@ubi.s.u-tokyo.ac.jp
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Phys. Rev. Research 8, 013312 – Published 24 March, 2026

DOI: https://doi.org/10.1103/fc35-47fs

Abstract

One of the fundamental questions in population dynamics is how biological populations respond to environmental perturbations. In population dynamics, the mean fitness and the fraction of a trait in the steady state are important because they indicate how well the trait and the population adapt to the environment. In this study, we examine the parallel mutation–reproduction model, which is one of the simplest models of an evolvable population. As an extension of the Markov chain tree theorem, we derive diagrammatic expressions for the static responses of the mean fitness and the steady-state distribution of the population. For the parallel mutation–reproduction model, we consider self-loops, which represent trait reproduction and are excluded from the Markov chain tree theorem for the linear master equation. To generalize the theorem, we introduce the concept of rooted 0/1 loop forests, which generalize spanning trees with loops. We demonstrate that the weights of rooted 0/1 loop forests yield the static responses of the mean fitness and the steady-state distribution. Our results provide exact expressions for the static responses and the steady-state distribution. Additionally, we discuss approximations of these expressions in cases where reproduction or mutation is dominant. We provide numerical examples to illustrate these approximations and exact expressions. We also demonstrate how our results can be used to design control strategies for harmful populations.

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Phys. Rev. Research 8, 013312– Published 24 March, 2026
  • Received 5 June 2025
  • Accepted 20 January 2026
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