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The evolution of multicellularity was a major transition in the history of life on earth. Conditions under which multicellularity is favored have been studied theoretically and experimentally. But since the construction of a multicellular organism requires multiple rounds of cell division, a natural question is whether these cell divisions should be synchronous or not. We study a simple population model in which there compete simple multicellular organisms that grow either by synchronous or asynchronous cell divisions. We demonstrate that natural selection can act differently on synchronous and asynchronous cell division, and we offer intuition for why these phenotypes are generally not neutral variants of each other.

A trademark of eusocial insect species is reproductive division of labor, in which workers forego their own reproduction while the queen produces almost all offspring. The presence of the queen is key for maintaining social harmony, but the specific role of the queen in the evolution of eusociality remains unclear. A long-discussed scenario is that a queen either behaviorally or chemically sterilizes her workers. However, the demographic and ecological conditions that enable such manipulation are unknown. Accordingly, we propose a simple model of evolutionary dynamics that is based on haplodiploid genetics. We consider a mutation that acts in a queen, causing her to control the reproductive behavior of her workers. Our mathematical analysis yields precise conditions for the evolutionary emergence and stability of queen-induced worker sterility. These conditions do not depend on the queen's mating frequency. Moreover, we find that queen control is always established if it increases colony reproductive efficiency and can evolve even if it decreases colony efficiency. We further outline the conditions under which queen control is evolutionarily stable against invasion by mutant, reproductive workers.

Evolutionary game dynamics are often studied in the context of different population structures. Here we propose a new population structure that is inspired by simple multicellular life forms. In our model, cells reproduce but can stay together after reproduction. They reach complexes of a certain size, n, before producing single cells again. The cells within a complex derive payoff from an evolutionary game by interacting with each other. The reproductive rate of cells is proportional to their payoff. We consider all two-strategy games. We study deterministic evolutionary dynamics with mutations, and derive exact conditions for selection to favor one strategy over another. Our main result has the same symmetry as the well-known sigma condition, which has been proven for stochastic game dynamics and weak selection. For a maximum complex size of n=2 our result holds for any intensity of selection. For n > 2 it holds for weak selection. As specific examples we study the prisoner's dilemma and hawk-dove games. Our model advances theoretical work on multicellularity by allowing for frequency-dependent interactions within groups., Comment: 44 pages, 11 figues

Workers in insect societies are sometimes observed to kill male eggs of other workers, a phenomenon known as worker policing. We perform a mathematical analysis of the evolutionary dynamics of policing. We investigate the selective forces behind policing for both dominant and recessive mutations for different numbers of matings of the queen. The traditional, relatedness-based argument suggests that policing evolves if the queen mates with more than two males, but does not evolve if the queen mates with a single male. We derive precise conditions for the invasion and stability of policing alleles. We find that the relatedness-based argument is not robust with respect to small changes in colony efficiency caused by policing. We also calculate evolutionarily singular strategies and determine when they are evolutionarily stable. We use a population genetics approach that applies to dominant or recessive mutations of any effect size.

We study the evolution of cooperation in a finite population interacting according to a simple model of like-with-like assortment. Evolution proceeds as a Moran process, and payoffs from the underlying cooperator-defector game are translated to positive fitnesses by an exponential transformation. These evolutionary dynamics can arise, for example, in a nest-structured population with rare migration. The use of the exponential transformation, rather than the usual linear one, is appropriate when interactions have multiplicative fitness effects, and allows for a tractable characterization of the effect of assortment on the evolution of cooperation. We define two senses in which a greater degree of assortment can favour the evolution of cooperation, the first stronger than the second: (i) greater assortment increases, at all population states, the probability that the number of cooperators increases, relative to the probability that the number of defectors increases; and (ii) greater assortment increases the fixation probability of cooperation, relative to that of defection. We show that, by the stronger definition, greater assortment favours the evolution of cooperation for a subset of cooperative dilemmas: prisoners' dilemmas, snowdrift games, stag-hunt games, and some prisoners' delight games. For other cooperative dilemmas, greater assortment favours cooperation by the weak definition, but not by the strong definition. Our results hold for any strength of selection.

We model evolution according to an asymmetric game as occurring in multiple finite populations, one for each role in the game, and study the effect of subjecting individuals to stochastic strategy mutations. We show that, when these mutations occur sufficiently infrequently, the dynamics over all population states simplify to an ergodic Markov chain over just the pure population states (where each population is monomorphic). This makes calculation of the stationary distribution computationally feasible. The transition probabilities of this embedded Markov chain involve fixation probabilities of mutants in single populations. The asymmetry of the underlying game leads to fixation probabilities that are derived from frequency-independent selection, in contrast to the analogous single-population symmetric-game case (Fudenberg and Imhof 2006). This frequency independence is useful in that it allows us to employ results from the population genetics literature to calculate the stationary distribution of the evolutionary process, giving sharper, and sometimes even analytic, results. We demonstrate the utility of this approach by applying it to a battle-of-the-sexes game, a Crawford-Sobel signalling game, and the beer-quiche game of Cho and Kreps (1987)., Comment: Journal of Economic Theory (2015)

We introduce and analyze a purely competitive dynamics for the evolution of an infinite population subject to a 3-strategy game. We argue that this dynamics represents a characterization of how certain systems, both natural and artificial, are governed. In each period, the population is randomly sorted into pairs, which engage in a once-off play of the game; the probability that a member propagates its type to its offspring is proportional only to its payoff within the pair. We show that if a type is dominant (obtains higher payoffs in games with both other types), its 'pure' population state, comprising only members of that type, is globally attracting. If there is no dominant type, there is an unstable 'mixed' fixed point; the population state eventually oscillates between the three near-pure states. We then allow for mutations, where offspring have a non-zero probability of randomly changing their type. In this case, the existence of a dominant type renders a point near its pure state globally attracting. If no dominant type exists, a supercritical Hopf bifurcation occurs at the unique mixed fixed point, and above a critical (typically low) mutation rate, this fixed point becomes globally attracting: the implication is that even very low mutation rates can stabilize a system that would, in the absence of mutations, be unstable., Comment: 13 pages, 6 figures