A minimal scenario for the origin of non-equilibrium order

Extant life contains numerous non-equilibrium mechanisms to create order not achievable at equilibrium; it is generally assumed that these mechanisms evolved because the resulting order was sufficiently beneficial to overcome associated costs of time and energy. Here, we identify a broad range of conditions under which non-equilibrium order-creating mechanisms will evolve as an inevitable consequence of self-replication, even if the order is not directly functional. We show that models of polymerases, when expanded to include known stalling effects, will evolve kinetic proofreading mechanisms through selection for fast replication alone, consistent with data from recent mutational screens. Similarly, replication contingent on self-assembly will select for non-equilibrium mechanisms like dynamic instability and result in more ordered structures despite the absence of direct selection for such ordered structures. We abstract these results into a framework that predicts that self-replication intrinsically amplifies dissipative order-enhancing mechanisms if the distribution of replication times is wide enough. Our work suggests the intriguing possibility that non-equilibrium order can arise more easily than assumed, even before that order is directly functional, with consequences impacting the evolution of mutation rates and kinetic traps in self-assembly to the origin of life.