Weak selection and stochastic evolutionary stability in a stochastic replicator dynamics.

It is a very challenging problem whether natural selection is able to effectively resist the continuous disturbance of environmental noise such that the direction or outcome of evolution determined by the deterministic selection pressure will not be changed. By analyzing the impact of weak selection on the evolutionary stability of a stochastic replicator dynamics with n possible pure strategies, we found that the weak selection is able to enhance the evolutionary stability, that is, under weak selection, the stochastic evolutionary stability of the system is determined by the mean payoff matrix. This finding strongly implies that the weak selection should be regarded as an important mechanism to ensure evolutionary stability in stochastic environments.

Evolutionary game dynamics with non-uniform interaction rates in finite population.

In this study, we extend evolutionary game dynamics with non-uniform interaction rates to the situation with finite population. Our main goal is to show how the fixation probability is influenced by the non-uniform interaction rates under weak selection. Based on the diffusion approximation of the Moran process and assumption of weak selection, the stochastic dynamic properties of a two-phenotype game with non-uniform interaction rates in a finite population are investigated. By the analysis of some cases, we show that the non-uniform interaction rates may result in the potential evolutionary complexity of game dynamics in finite population.

Inclusive fitness and Hamilton's rule in a stochastic environment.

The evolution of cooperation in Prisoner's Dilemmas with additive random cost and benefit for cooperation cannot be accounted for by Hamilton's rule based on mean effects transferred from recipients to donors weighted by coefficients of relatedness, which defines inclusive fitness in a constant environment. Extensions that involve higher moments of stochastic effects are possible, however, and these are connected to a concept of random inclusive fitness that is frequency-dependent. This is shown in the setting of pairwise interactions in a haploid population with the same coefficient of relatedness between interacting players. In an infinite population, fixation of cooperation is stochastically stable if a mean geometric inclusive fitness of defection when rare is negative, while fixation of defection is stochastically unstable if a mean geometric inclusive fitness of cooperation when rare is positive, and these conditions are generally not equivalent. In a finite population, the probability for cooperation to ultimately fix when represented once exceeds the probability under neutrality or the corresponding probability for defection if the mean inclusive fitness of cooperation when its frequency is 1/3 or 1/2, respectively, exceeds 1. All these results rely on the simplifying assumption of a linear fitness function. It is argued that meaningful applications of random inclusive fitness in complex settings (multi-player game, diploidy, population structure) would generally require conditions of weak selection and additive gene action.

Hamilton's rule and kin competition in a finite kin population.

Kin selection means that individuals can increase their own inclusive fitness through displaying more altruistically toward their relatives. So, Hamilton's rule says kin selection will work if the coefficient of relatedness exceeds the cost-to-benefit ratio of the altruistic act. However, some studies have shown that the kin competition due to the altruism among relatives can reduce, and even totally negate, the kin-selected benefits of altruism toward relatives. In order to understand how the evolution of cooperation is influenced by both kin selection and kin competition under a general theoretical framework, we here consider the evolutionary dynamics of cooperation in a finite kin population, where kin competition is incorporated into a simple Prisoner's Dilemma game between relatives. Differently from the previous studies, we emphasize that the difference between the effects of mutually and unilaterally altruistic acts on kin competition may play an important role for the evolution of cooperation. The main results not only show the conditions that Hamilton's rule still works under the kin competition but also reveal the evolutionary biological mechanism driving the evolution of cooperation in a finite kin population.

Transitions in the cell-fate induction induced by colored noise associated with the inductive stimulus.

The cell-fate induction based on the saddle-node bifurcation is undoubtedly a very important concept in developmental biology, which provides a possible mechanism to explain the intrinsic irreversibility in the developmental process. In this paper, the effect of a colored noise, which is associated with the inductive stimulus, on the saddle-node landscape of cell-fate induction is investigated, especially, the effect of the change of correlation time of colored noise on cell-fate induction. The main results show clearly that the change of correlation time of colored noise could induce the transitions of the system. This implies that the colored noise associated with inductive stimulus may have a profound effect on the saddle-node bifurcation landscape of cell-fate induction. This will also help us to understand more deeply the role of cell-fate induction in developmental biology.

Dynamic transcriptional symmetry-breaking in pre-implantation mammalian embryo development revealed by single-cell RNA-seq.

During mammalian pre-implantation embryo development, when the first asymmetry emerges and how it develops to direct distinct cell fates remain longstanding questions. Here, by analyzing single-blastomere transcriptome data from mouse and human pre-implantation embryos, we revealed that the initial blastomere-to-blastomere biases emerge as early as the first embryonic cleavage division, following a binomial distribution pattern. The subsequent zygotic transcriptional activation further elevated overall blastomere-to-blastomere biases during the two- to 16-cell embryo stages. The trends of transcriptional asymmetry fell into two distinct patterns: for some genes, the extent of asymmetry was minimized between blastomeres (monostable pattern), whereas other genes, including those known to be lineage specifiers, showed ever-increasing asymmetry between blastomeres (bistable pattern), supposedly controlled by negative or positive feedbacks. Moreover, our analysis supports a scenario in which opposing lineage specifiers within an early blastomere constantly compete with each other based on their relative ratio, forming an inclined 'lineage strength' that pushes the blastomere onto a predisposed, yet flexible, lineage track before morphological distinction.

Environmental Noise Could Promote Stochastic Local Stability of Behavioral Diversity Evolution.

In this Letter, we investigate stochastic stability in a two-phenotype evolutionary game model for an infinite, well-mixed population undergoing discrete, nonoverlapping generations. We assume that the fitness of a phenotype is an exponential function of its expected payoff following random pairwise interactions whose outcomes randomly fluctuate with time. We show that the stochastic local stability of a constant interior equilibrium can be promoted by the random environmental noise even if the system may display a complicated nonlinear dynamics. This result provides a new perspective for a better understanding of how environmental fluctuations may contribute to the evolution of behavioral diversity.

A simple rule of direct reciprocity leads to the stable coexistence of cooperation and defection in the Prisoner's Dilemma game.

The long-term coexistence of cooperation and defection is a common phenomenon in nature and human society. However, none of the theoretical models based on the Prisoner's Dilemma (PD) game can provide a concise theoretical model to explain what leads to the stable coexistence of cooperation and defection in the long-term even though some rules for promoting cooperation have been summarized (Nowak, 2006, Science 314, 1560-1563). Here, based on the concept of direct reciprocity, we develop an elementary model to show why stable coexistence of cooperation and defection in the PD game is possible. The basic idea behind our theoretical model is that all players in a PD game prefer a cooperator as an opponent, and our results show that considering strategies allowing opting out against defection provide a general and concise way of understanding the fundamental importance of direct reciprocity in driving the evolution of cooperation.

Activation of Smurf E3 ligase promoted by smoothened regulates hedgehog signaling through targeting patched turnover.

Hedgehog signaling plays conserved roles in controlling embryonic development; its dysregulation has been implicated in many human diseases including cancers. Hedgehog signaling has an unusual reception system consisting of two transmembrane proteins, Patched receptor and Smoothened signal transducer. Although activation of Smoothened and its downstream signal transduction have been intensively studied, less is known about how Patched receptor is regulated, and particularly how this regulation contributes to appropriate Hedgehog signal transduction. Here we identified a novel role of Smurf E3 ligase in regulating Hedgehog signaling by controlling Patched ubiquitination and turnover. Moreover, we showed that Smurf-mediated Patched ubiquitination depends on Smo activity in wing discs. Mechanistically, we found that Smo interacts with Smurf and promotes it to mediate Patched ubiquitination by targeting the K1261 site in Ptc. The further mathematic modeling analysis reveals that a bidirectional control of activation of Smo involving Smurf and Patched is important for signal-receiving cells to precisely interpret external signals, thereby maintaining Hedgehog signaling reliability. Finally, our data revealed an evolutionarily conserved role of Smurf proteins in controlling Hh signaling by targeting Ptc during development.

Impact of environmental stochastic fluctuations on the evolutionary stability of imitation dynamics.

To show the impact of environmental noise on imitation dynamics, the stochastic stability and stochastic evolutionary stability of a discrete-time imitation dynamics with random payoffs are studied in this paper. Based on the stochastic local stability of fixation states and constant interior equilibria in a two-phenotype model, we extend the concept of stochastic evolutionary stability to the stochastic imitation dynamics, which is defined as a strategy such that, if all the members of the population adopt it, then the probability for any mutant strategy to invade the population successfully under the influence of natural selection is arbitrarily low. Our main results show clearly that the stochastic evolutionary stability of the system depends only on the properties of the mean matrix of the random payoff matrix and is independent of the randomness of the random payoff matrix. Moreover, as two examples, we show also that under the framework of stochastic imitation dynamics, the noise intensity affects the evolution of cooperative behavior in a stochastic prisoner's dilemma game and the system's nonlinear dynamic behavior.

Evolutionary dynamics of cooperation coupled with ecological feedback compensation.

A simple theoretical model (or a demonstrative example) was developed to illustrate how the evolution of cooperation can be affected by the density-dependent survival competition, in which we assume that the fertility of an individual depends only on the pairwise interaction between him and other individuals based on Prisoner's Dilemma game, while its viability is only related to the density-dependent survival competitiveness. Our results show that not only cooperation could be evolutionarily stable if the advantage of cooperators in viability can compensate for the cost they pay for their fertility, but also the long-term stable coexistence of cooperation and defection is possible if none of cooperation and defection is evolutionarily stable. Moreover, for the stochastic evolutionary dynamics in a finite population, our analysis shows that the increase (or decrease) of the survival competitiveness of cooperators (or defectors) should be conductive to the evolutionary emergence of cooperation.

Costly punishment does not always increase cooperation.

In a pairwise interaction, an individual who uses costly punishment must pay a cost in order that the opponent incurs a cost. It has been argued that individuals will behave more cooperatively if they know that their opponent has the option of using costly punishment. We examined this hypothesis by conducting two repeated two-player Prisoner's Dilemma experiments, that differed in their payoffs associated to cooperation, with university students from Beijing as participants. In these experiments, the level of cooperation either stayed the same or actually decreased when compared with the control experiments in which costly punishment was not an option. We argue that this result is likely due to differences in cultural attitudes to cooperation and punishment based on similar experiments with university students from Boston that found cooperation did increase with costly punishment.

Stochastic replicator dynamics and evolutionary stability.

To develop the concept of evolutionary stability in a stochastic environment, we investigate the continuous-time dynamics of a two-phenotype linear evolutionary game with generally correlated random payoffs in pairwise interactions. By using the Gram-Schmidt orthogonalization procedure and Itô's formula, we deduce a stochastic differential equation for the phenotype frequencies that extends the replicator equation, called the stochastic replicator equation. We give conditions for stochastic stability of a fixation state or a constant interior equilibrium point with respect to the stochastic dynamics of the two phenotypes. We show that, if a fixation state is stochastically stable, then the pure strategy corresponding to this fixation state must be stochastically evolutionarily stable with respect to mixed strategies. However, this is not the case for a mixed strategy that corresponds to a stochastically stable constant interior equilibrium point with respect to the two phenotypes.

Stochastic evolutionary stability in matrix games with random payoffs.

Evolutionary game theory and the concept of an evolutionarily stable strategy have been not only extensively developed and successfully applied to explain the evolution of animal behavior, but also widely used in economics and social sciences. Recently, in order to reveal the stochastic dynamical properties of evolutionary games in randomly fluctuating environments, the concept of stochastic evolutionary stability based on conditions for stochastic local stability for a fixation state was developed in the context of a symmetric matrix game with two phenotypes and random payoffs in pairwise interactions [Zheng et al., Phys. Rev. E 96, 032414 (2017)2470-004510.1103/PhysRevE.96.032414]. In this paper, we extend this study to more general situations, namely, multiphenotype symmetric as well as asymmetric matrix games with random payoffs. Conditions for stochastic local stability and stochastic evolutionary stability are established. Conditions for a fixation state to be stochastically unstable and almost everywhere stochastically unstable are distinguished in a multiphenotype setting according to the initial population state. Our results provide some alternative perspective and a more general theoretical framework for a better understanding of the evolution of animal behavior in a stochastic environment.

Effects of bidirectional regulation on noises in gene networks.

To investigate the effects of bidirectional regulation on the noise in protein concentration, a theoretical and simple three-gene network model is considered. The basic idea behind this model is from Paulsson's proposition (J. Paulsson, Phys. Life Rev. 2005, 2, 157-175), where the synthesis and degradation of a mRNA species corresponding to a target protein are regulated directly and indirectly by a certain sigma-factor, and a random increase in the concentration of the sigma-factor should increase both the synthesis and degradation rates of the mRNA species (bidirectional regulation). Using the standard Omega-expansion technique (linear noise approximation) and Monte Carlo simulation, our main results show clearly that for the steady-state statistics the effects of the noise of the sigma-factor on the stochastic fluctuation of the target protein could partially cancel out.

Weak selection can filter environmental noise in the evolution of animal behavior.

Weak selection is an important assumption in theoretical evolutionary biology, but its biological significance remains unclear. In this study, we investigate the effect of weak selection on stochastic evolutionary stability in a two-phenotype evolutionary game dynamics with a random payoff matrix assuming an infinite, well-mixed population undergoing discrete, nonoverlapping generations. We show that, under weak selection, both stochastic local stability and stochastic evolutionary stability in this system depend on the means of the random payoffs but not on their variances. Moreover, although stochastic local stability or instability of an equilibrium may not depend on environmental noise if selection is weak enough, the growth rate near an equilibrium not only depends on environmental noise, but can even be enhanced by environmental noise if selection is weak. This is the case, for instance, when the variances of the random payoffs as well as the covariances are equal. These results suggest that natural selection could be able to filter (or resist) the effect of environmental noise on the evolution of animal behavior if selection is weak.

6mA-DNA-binding factor Jumu controls maternal-to-zygotic transition upstream of Zelda.

A long-standing question in the field of embryogenesis is how the zygotic genome is precisely activated by maternal factors, allowing normal early embryonic development. We have previously shown that N6-methyladenine (6mA) DNA modification is highly dynamic in early Drosophila embryos and forms an epigenetic mark. However, little is known about how 6mA-formed epigenetic information is decoded. Here we report that the Fox-family protein Jumu binds 6mA-marked DNA and acts as a maternal factor to regulate the maternal-to-zygotic transition. We find that zelda encoding the pioneer factor Zelda is marked by 6mA. Our genetic assays suggest that Jumu controls the proper zygotic genome activation (ZGA) in early embryos, at least in part, by regulating zelda expression. Thus, our findings not only support that the 6mA-formed epigenetic marks can be read by specific transcription factors, but also uncover a mechanism by which the Jumu regulates ZGA partially through Zelda in early embryos.

Stochastic fluctuations in frequency-dependent selection: a one-locus, two-allele and two-phenotype model.

Stochastic fluctuations in a simple frequency-dependent selection model with one-locus, two-alleles and two-phenotypes are investigated. The steady-state statistics of allele frequencies for an interior stable phenotypic equilibrium are shown to be similar to the stochastic fluctuations in standard evolutionary game dynamics [Tao, Y., Cressman, R., 2007. Stochastic fluctuations through intrinsic noise in evolutionary game dynamics. Bull. Math. Biol. 69, 1377-1399]. On the other hand, for an interior stable phenotypic or genotypic equilibrium, our main results show that the deterministic model cannot be used to predict the expectation of phenotypic frequency. The variance of phenotypic frequency for an interior stable genotypic equilibrium is more sensitive to the expected population size than for an interior stable phenotypic equilibrium. Furthermore, the stochastic fluctuations of allele frequency and phenotypic frequency can be considered approximately independent of each other for these genotypic equilibria, but not for phenotypic.

Evolutionary stability concepts in a stochastic environment.

Over the past 30 years, evolutionary game theory and the concept of an evolutionarily stable strategy have been not only extensively developed and successfully applied to explain the evolution of animal behaviors, but also widely used in economics and social sciences. Nonetheless, the stochastic dynamical properties of evolutionary games in randomly fluctuating environments are still unclear. In this study, we investigate conditions for stochastic local stability of fixation states and constant interior equilibria in a two-phenotype model with random payoffs following pairwise interactions. Based on this model, we develop the concepts of stochastic evolutionary stability (SES) and stochastic convergence stability (SCS). We show that the condition for a pure strategy to be SES and SCS is more stringent than in a constant environment, while the condition for a constant mixed strategy to be SES is less stringent than the condition to be SCS, which is less stringent than the condition in a constant environment.

Limiting similarity of competitive species and demographic stochasticity.

The limiting similarity of competitive species and its relationship with the competitive exclusion principle is still one of the most important concepts in ecology. In the 1970s, May [R. M. May, Stability and Complexity in Model Ecosystems (Princeton University, Princeton, NJ, 1973)] developed a concise theoretical framework to investigate the limiting similarity of competitive species. His theoretical results show that no limiting similarity threshold of competitive species can be identified in the deterministic model system whereby species more similar than this threshold never coexist. Theoretically, for competitive species coexisting in an unvarying environment, deterministic interspecific interactions and demographic stochasticity can be considered two sides of a coin. To investigate how the "tension" between these two forces affects the coexistence of competing species, a simple two-species competitive system based only on May's model system is transformed into an equivalent replicator equation. The effect of demographic stochasticity on the system stability is measured by the expected drift of the Lyapunov function. Our main results show that the limiting similarity of competitive species should not be considered to be an absolute measure. Specifically, very similar competitive species should be able to coexist in an environment with a high productivity level but big differences between competitive species should be necessary in an ecosystem with a low productivity level.