Repeatability of evolution on epistatic landscapes

Evolution is a dynamic process. The two classical forces of evolution are mutation and selection. Assuming small mutation rates, evolution can be predicted based solely on the fitness differences between phenotypes. Predicting an evolutionary process under varying mutation rates as well as varying fitness is still an open question. Experimental procedures, however, do include these complexities along with fluctuating population sizes and stochastic events such as extinctions. We investigate the mutational path probabilities of systems having epistatic effects on both fitness and mutation rates using a theoretical and computational framework. In contrast to previous models, we do not limit ourselves to the typical strong selection, weak mutation (SSWM)-regime or to fixed population sizes. Rather we allow epistatic interactions to also affect mutation rates. This can lead to qualitatively non-trivial dynamics. Pathways, that are negligible in the SSWM-regime, can overcome fitness valleys and become accessible. This finding has the potential to extend the traditional predictions based on the SSWM foundation and bring us closer to what is observed in experimental systems

Eco-evolutionary dynamics of social dilemmas

Social dilemmas are an integral part of social interactions. Cooperative actions, ranging from secreting extra-cellular products in microbial populations to donating blood in humans, are costly to the actor and hence create an incentive to shirk and avoid the costs. Nevertheless, cooperation is ubiquitous in nature. Both costs and benefits often depend non-linearly on the number and types of individuals involved -- as captured by idioms such as `too many cooks spoil the broth' where additional contributions are discounted, or `two heads are better than one' where cooperators synergistically enhance the group benefit. Interaction group sizes may depend on the size of the population and hence on ecological processes. This results in feedback mechanisms between ecological and evolutionary processes, which jointly affect and determine the evolutionary trajectory. Only recently combined eco-evolutionary processes became experimentally tractable in microbial social dilemmas. Here we analyse the evolutionary dynamics of non-linear social dilemmas in settings where the population fluctuates in size and the environment changes over time. In particular, cooperation is often supported and maintained at high densities through ecological fluctuations. Moreover, we find that the combination of the two processes routinely reveals highly complex dynamics, which suggests common occurrence in nature.Comment: 26 pages, 11 figure

Evolutionary Multiplayer Games

Evolutionary game theory has become one of the most diverse and far reaching theories in biology. Applications of this theory range from cell dynamics to social evolution. However, many applications make it clear that inherent non-linearities of natural systems need to be taken into account. One way of introducing such non-linearities into evolutionary games is by the inclusion of multiple players. An example is of social dilemmas, where group benefits could e.g.\ increase less than linear with the number of cooperators. Such multiplayer games can be introduced in all the fields where evolutionary game theory is already well established. However, the inclusion of non-linearities can help to advance the analysis of systems which are known to be complex, e.g. in the case of non-Mendelian inheritance. We review the diachronic theory and applications of multiplayer evolutionary games and present the current state of the field. Our aim is a summary of the theoretical results from well-mixed populations in infinite as well as finite populations. We also discuss examples from three fields where the theory has been successfully applied, ecology, social sciences and population genetics. In closing, we probe certain future directions which can be explored using the complexity of multiplayer games while preserving the promise of simplicity of evolutionary games.Comment: 14 pages, 2 figures, review pape

Mutualism and evolutionary multiplayer games: revisiting the Red King

Coevolution of two species is typically thought to favour the evolution of faster evolutionary rates helping a species keep ahead in the Red Queen race, where `it takes all the running you can do to stay where you are'. In contrast, if species are in a mutualistic relationship, it was proposed that the Red King effect may act, where it can be beneficial to evolve slower than the mutualistic species. The Red King hypothesis proposes that the species which evolves slower can gain a larger share of the benefits. However, the interactions between the two species may involve multiple individuals. To analyse such a situation, we resort to evolutionary multiplayer games. Even in situations where evolving slower is beneficial in a two-player setting, faster evolution may be favoured in a multiplayer setting. The underlying features of multiplayer games can be crucial for the distribution of benefits. They also suggest a link between the evolution of the rate of evolution and group size

Chaotic provinces in the kingdom of the Red Queen

The interplay between parasites and their hosts is found in all kinds of species and plays an important role in understanding the principles of evolution and coevolution. Usually, the different genotypes of hosts and parasites oscillate in their abundances. The well-established theory of oscillatory Red Queen dynamics proposes an ongoing change in frequencies of the different types within each species. So far, it is unclear in which way Red Queen dynamics persists with more than two types of hosts and parasites. In our analysis, an arbitrary number of types within two species are examined in a deterministic framework with constant or changing population size. This general framework allows for analytical solutions for internal fixed points and their stability. For more than two species, apparently chaotic dynamics has been reported. Here we show that even for two species, once more than two types are considered per species, irregular dynamics in their frequencies can be observed in the long run. The nature of the dynamics depends strongly on the initial configuration of the system; the usual regular Red Queen oscillations are only observed in some parts of the parameter region

The pace of evolution across fitness valleys

How fast does a population evolve from one fitness peak to another? We study the dynamics of evolving, asexually reproducing populations in which a certain number of mutations jointly confer a fitness advantage. We consider the time until a population has evolved from one fitness peak to another one with a higher fitness. The order of mutations can either be fixed or random. If the order of mutations is fixed, then the population follows a metaphorical ridge, a single path. If the order of mutations is arbitrary, then there are many ways to evolve to the higher fitness state. We address the time required for fixation in such scenarios and study how it is affected by the order of mutations, the population size, the fitness values and the mutation rate

Implementing the Flipped Classroom in an Undergraduate Corporate Finance Course

This study analyzes flipped mode of instruction in Corporate Finance. In the current environment, with many students in quarantine and greater emphasis on self-study, it is even more relevant to understand how students understand and retain concepts derived from online environment. In this study, performance of students in flipped mode is compared with the performance in traditional lecture-style. In flipped class, students watched a brief video-lecture, took an online assessment quiz prior to attending an interactive discussion-based class session, unlike the traditional lecture style. The results of this study suggest that with flipped mode, most students take greater responsibility of their learning, prefer hands-on learning, achieve more and feel more satisfied with their performance. However, there is no statistical difference in the test scores of students in the flipped classroom as compared to the traditional classroom

Implementing the Flipped Classroom in an Undergraduate Corporate Finance Course

This study analyzes flipped mode of instruction in Corporate Finance. In the current environment, with many students in quarantine and greater emphasis on self-study, it is even more relevant to understand how students understand and retain concepts derived from online environment. In this study, performance of students in flipped mode is compared with the performance in traditional lecture-style. In flipped class, students watched a brief video-lecture, took an online assessment quiz prior to attending an interactive discussion-based class session, unlike the traditional lecture style. The results of this study suggest that with flipped mode, most students take greater responsibility of their learning, prefer hands-on learning, achieve more and feel more satisfied with their performance. However, there is no statistical difference in the test scores of students in the flipped classroom as compared to the traditional classroom

Numerical Modelling of Scramjet Combustor

Numerical modelling of turbulent-reacting flow field of supersonic combustion ramjet(scramjet) combustors are presented. The developed numerical procedure is based on the implicittreatment of chemical source terms by preconditioning and solved along with unstedy turbulentNavier-Stokes equations explicitly. Reaction is modelled using an eight-step hydrogen-airchemistry. Code is validated against a standard wall jet experimental data and is successfullyused to model the turbulent-reacting flow field resulting due to the combustion of hydrogeninjected from diamond-shaped strut and also in the wake region of wedge-shaped strut placedin the heated supersonic airstream. The analysis could demonstrate the effect of interaction ofoblique shock wave with a supersonic stream of hydrogen  in its (fuel-air) mixing and reactionfor strut-based scramjet combustors