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William C Ratcliff

Publications and source records attributed to William C Ratcliff.

4 recordsLinked to original sources

Time-dependent fitness effects can drive bet-hedging populations extinct

To survive unpredictable environmental change, many organisms adopt bet-hedging strategies that trade short-term population growth for long-term fitness benefits. Because the benefits of bet-hedging may manifest over long time intervals, bet-hedging strategies may be out-competed by strategies maximizing short-term fitness. Here, we investigate the interplay between two drivers of selection, environmental fluctuations and competition for limited resources, on different bet-hedging strategies. We consider an environment with frequent disasters that switch between which phenotypes they affect in a temporally-correlated fashion. We determine how organisms that stochastically switch between phenotypes at different rates fare in both competition and survival. When disasters are correlated in time, the best strategy for competition is among the worst for survival. Since the time scales over which the two agents of selection act are significantly different, environmental fluctuations and resource competition act in opposition and lead populations to evolve diversification strategies that ultimately drive them extinct.

Evolutionary Biology

Replicator Dynamics with Feedback-Evolving Games: Towards a Co-Evolutionary Game Theory

A tragedy of the commons occurs when individuals take actions to maximize their payoffs even as their combined payoff is less than the global maximum had the players coordinated. The originating example is that of over-grazing of common pasture lands. In game theoretic treatments of this example there is rarely consideration of how individual behavior subsequently modifies the commons and associated payoffs. Here, we generalize evolutionary game theory by proposing a class of replicator dynamics with feedback-evolving games in which environment-dependent payoffs and strategies coevolve. We initially apply our formulation to a system in which the payoffs favor unilateral defection and cooperation, given replete and depleted environments respectively. Using this approach we identify and characterize a new class of dynamics: an oscillatory tragedy of the commons in which the system cycles between deplete and replete environmental states and cooperation and defection behavior states. We generalize the approach to consider outcomes given all possible rational choices of individual behavior in the depleted state when defection is favored in the replete state. In so doing we find that incentivizing cooperation when others defect in the depleted state is necessary to avert the tragedy of the commons. In closing, we propose new directions for the study of control and influence in games in which individual actions exert a substantive effect on the environmental state.

Evolutionary Biology

Trait Heritability in Major Transitions

A crucial component of major transitions theory is that after the transition, adaptation occurs primarily at the level of the new, higher-level unit. For collective-level adaptations to occur, though, collective-level traits must be heritable. Since collective-level trait values are functions of lower-level trait values, collective-level heritability is related to particle-level heritability. However, the nature of this relationship has rarely been explored in the context of major transitions. We examine relationships between particle-level heritability and collective-level heritability for several functions that express collective-level trait value in terms of particle-level trait values. When a collective-level trait value is a linear function of particle-level trait values, the heritability of a collective-level trait is never less than that of the corresponding particle-level trait and is higher under most conditions. For more complicated functions, collective-level heritability is higher under most conditions, but can be lower when the function relating particle to cell-level trait values is sensitive to small fluctuations in the state of the particles within the collective. Rather than being an impediment to major transitions, we show that collective-level heritability superior to that of the lower-level units can often arise for free, simply as a byproduct of collective formation.

Evolutionary Biology

Apoptosis in snowflake yeast: novel trait, or side effect of toxic waste?

Recent experiments evolving de novo multicellularity in yeast have found that large-cluster forming genotypes also exhibit higher rates of programmed cell death (apoptosis). This was previously interpreted as the evolution of a simple form of cellular division of labor: apoptosis results in the scission of cell-cell connections, allowing snowflake yeast to produce proportionally smaller, faster-growing propagules. Through spatial simulations, Duran-Nebreda and Sole (2015) develop the novel null hypothesis that apoptosis is not an adaptation, per se, but is instead caused by the accumulation of toxic metabolites in large clusters. Here we test this hypothesis by synthetically creating unicellular derivatives of snowflake yeast through functional complementation with the ancestral ACE2 allele. We find that multicellular snowflake yeast with elevated apoptosis exhibit a similar rate of apoptosis when cultured as single cells. We also show that larger snowflake yeast clusters tend to contain a greater fraction of older, senescent cells, which may explain why larger clusters of a given genotype are more apoptotic. Our results show that apoptosis is not caused by side effects of spatial structure, such as starvation or waste product accumulation, and are consistent with the hypothesis that elevated apoptosis is a trait which co-evolves with large cluster size.

Evolutionary Biology