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Register a new userDiversity and coevolutionary dynamics in high-dimensional phenotype spaces
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We study macroevolutionary dynamics by extending microevolutionary competition models to long time scales. It has been shown that for a general class of competition models, gradual evolutionary change in continuous phenotypes (evolutionary dynamics) can be non-stationary and even chaotic when the dimension of the phenotype space in which the evolutionary dynamics unfold is high. It has also been shown that evolutionary diversification can occur along non-equilibrium trajectories in phenotype space. We combine these lines of thinking by studying long-term coevolutionary dynamics of emerging lineages in multi-dimensional phenotype spaces. We use a statistical approach to investigate the evolutionary dynamics of many different systems. We find: 1) for a given dimension of phenotype space, the coevolutionary dynamics tends to be fast and non-stationary for an intermediate number of coexisting lineages, but tends to stabilize as the evolving communities reach a saturation level of diversity; and 2) the amount of diversity at the saturation level increases rapidly (exponentially) with the dimension of phenotype space. These results have implications for theoretical perspectives on major macroevolutionary patterns such as adaptive radiation, long-term temporal patterns of phenotypic changes, and the evolution of diversity.
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Most theories of evolutionary diversification are based on equilibrium assumptions: they are either based on optimality arguments involving static fitness landscapes, or they assume that populations first evolve to an equilibrium state before diversification occurs, as exemplified by the concept of evolutionary branching points in adaptive dynamics theory. Recent results indicate that adaptive dynamics may often not converge to equilibrium points and instead generate complicated trajectories if evolution takes place in high-dimensional phenotype spaces. Even though some analytical results on diversification in complex phenotype spaces are available, to study this problem in general we need to reconstruct individual-based models from the adaptive dynamics generating the non-equilibrium dynamics. Here we first provide a method to construct individual-based models such that they faithfully reproduce the given adaptive dynamics attractor without diversification. We then show that a propensity to diversify can by introduced by adding Gaussian competition terms that generate frequency dependence while still preserving the same adaptive dynamics. For sufficiently strong competition, the disruptive selection generated by frequency-dependence overcomes the directional evolution along the selection gradient and leads to diversification in phenotypic directions that are orthogonal to the selection gradient.
Living species, ranging from bacteria to animals, exist in environmental conditions that exhibit spatial and temporal heterogeneity which requires them to adapt. Risk-spreading through spontaneous phenotypic variations is a known concept in ecology, which is used to explain how species may survive when faced with the evolutionary risks associated with temporally varying environments. In order to support a deeper understanding of the adaptive role of spontaneous phenotypic variations in fluctuating environments, we consider a system of non-local partial differential equations modelling the evolutionary dynamics of two competing phenotype-structured populations in the presence of periodically oscillating nutrient levels. The two populations undergo spontaneous phenotypic variations at different rates. The phenotypic state of each individual is represented by a continuous variable, and the phenotypic landscape of the populations evolves in time due to variations in the nutrient level. Exploiting the analytical tractability of our model, we study the long-time behaviour of the solutions to obtain a detailed mathematical depiction of evolutionary dynamics. The results suggest that when nutrient levels undergo small and slow oscillations, it is evolutionarily more convenient to rarely undergo spontaneous phenotypic variations. Conversely, under relatively large and fast periodic oscillations in the nutrient levels, which bring about alternating cycles of starvation and nutrient abundance, higher rates of spontaneous phenotypic variations confer a competitive advantage. We discuss the implications of our results in the context of cancer metabolism.
The processes and mechanisms underlying the origin and maintenance of biological diversity have long been of central importance in ecology and evolution. The competitive exclusion principle states that the number of coexisting species is limited by the number of resources, or by the species similarity in resource use. Natural systems such as the extreme diversity of unicellular life in the oceans provide counter examples. It is known that mathematical models incorporating population fluctuations can lead to violations of the exclusion principle. Here we use simple eco-evolutionary models to show that a certain type of population dynamics, boom-bust dynamics, can allow for the evolution of much larger amounts of diversity than would be expected with stable equilibrium dynamics. Boom-bust dynamics are characterized by long periods of almost exponential growth (boom) and a subsequent population crash due to competition (bust). When such ecological dynamics are incorporated into an evolutionary model that allows for adaptive diversification in continuous phenotype spaces, desynchronization of the boom-bust cycles of coexisting species can lead to the maintenance of high levels of diversity.
Mutation and drift play opposite roles in genetics. While mutation creates diversity, drift can cause gene variants to disappear, especially when they are rare. In the absence of natural selection and migration, the balance between the drift and mutation in a well-mixed population defines its diversity. The Moran model captures the effects of these two evolutionary forces and has a counterpart in social dynamics, known as the Voter model with external opinion influencers. Two extreme outcomes of the Voter model dynamics are consensus and polarization, which correspond to low and high diversity in the Moran model. Here we use a Shannons information-theoretic approach to characterize the smooth transition between the ordered and disordered states of consensus and polarization in the Voter model. Mapping the Moran into the Voter model we extend the results to the mutation-drift balance and characterize the transition between low and high diversity in finite populations. Describing the population as a network of connected individuals we show that the transition between the two regimes depends on the geographic structure of the population and on the possible asymmetries in the mutation rates.
We study the coevolutionary dynamics of the diversity of phenotype expression and the evolution of cooperation in the Prisoners Dilemma game. Rather than pre-assigning zero-or-one interaction rate, we diversify the rate of interaction by associating it with the phenotypes shared in common. Individuals each carry a set of potentially expressible phenotypes and expresses a certain number of phenotypes at a cost proportional to the number. The number of expressed phenotypes and thus the rate of interaction is an evolvable trait. Our results show that nonnegligible cost of expressing phenotypes restrains phenotype expression, and the evolutionary race mainly proceeds on between cooperative strains and defective strains who express a very few phenotypes. It pays for cooperative strains to express a very few phenotypes. Though such a low level of expression weakens reciprocity between cooperative strains, it decelerates rate of interaction between cooperative strains and defective strains to a larger degree, leading to the predominance of cooperative strains over defective strains. We also find that evolved diversity of phenotype expression can occasionally destabilize due to the invasion of defective mutants, implying that cooperation and diversity of phenotype expression can mutually reinforce each other. Therefore, our results provide new insights into better understanding the coevolution of cooperation and the diversity of phenotype expression.
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