<p>Evolution by natural selection can occur at multiple biological scales. This is particularly the case for host-pathogen systems, where selection occurs both within each infected host as well as through transmission between hosts. Despite there being established mathematical models for understanding evolution at a single biological scale, fewer tractable models exist for multiscale evolutionary dynamics. Here I present mathematical approaches using tools from probability and stochastic processes as well as dynamical systems to handle multiscale evolutionary systems. The first problem I address concerns the antigenic evolution of influenza. Using a combination of ordinary differential equations and inhomogeneous Poisson processes, I study how immune selection pressures at the within-host level impact population-level evolutionary dynamics. The second problem involves the more general question of evolutionary dynamics when selection occurs antagonistically at two biological scales. In addition to host-pathogen systems, such situations arise naturally in the evolution of traits such as the production of a public good and the use of a common resource. I introduce a model for this general phenomenon that is intuitively visualized as a a stochastic ball-and-urn system and can be used to systematically obtain general properties of antagonistic multiscale evolution. Lastly, this ball-and-urn framework is in itself an interesting mathematical object which can studied as either a measure-valued process or an interacting particle system. In this mathematical context, I show that under different scalings, the measure-valued process can have either a propagation of chaos or Fleming-Viot limit.</p> / Dissertation

This dissertation begins with a comprehensive review of the evolutionary biological debate over the evolution of altruism and a discussion of how various models of mechanism have influenced the models of circumstance that paleoanthropologists continue to use in reconstructing details about the level of cooperation displayed by early hominin societies. The remainder of the dissertation concerns itself with systematically exposing previously unquestioned assumptions to potential falsification as well as with exploring some new scenarios concerning the evolution and landscape archaeology of Plio-Pleistocene hominin food sharing, all via a null agent-based model called SHARE. This heuristic model was built to address two major questions: (1) What range of ecological and social conditions facilitates the evolution of food sharing in artificial Plio-Pleistocene hominin populations and (2) Is food sharing at central places necessary for the formation of the so-called "scatter and patches" archaeological landscapes that are characteristic of the Plio-Pleistocene record in East Africa? In answer to the first question, population-level genetic results collected from artificial societies of hominin agents demonstrate that the so-called transitional zone of ecological patchiness can facilitate the evolution of altruistic food sharing in mixed starting populations, even if foragers lack the ability to remember past interactions or to avoid social cheaters. In answer to the second question, ecological patchiness can affect the movements of simple foragers such that the artificial archaeological landscapes they create display the same spatial signature that characterizes observed Oldowan landscapes. That is, in ecological conditions marked by fragmented food resources, archaeological landscapes composed of both concentrated patches and diffuse scatters can form as a result of solitary foragers using simple routes that are in no way tethered to culturally-defined and culturally-maintained central places. In the end, SHARE provides new hypotheses about how ecological patchiness could have influenced both the evolution of altruistic food sharing and the structure of Lower Paleolithic archaeological landscapes. The latter can be tested in the field by looking at the relationship between artifact density and the paleoenvironmental characteristics of locales both in which artifacts are abundant and from which they are conspicuously absent.

abstract: During the 1960s, the long-standing idea that traits or behaviors could be explained by natural selection acting on traits that persisted "for the good of the group" prompted a series of debates about group-level selection and the effectiveness with which natural selection could act at or across multiple levels of biological organization. For some this topic remains contentious, while others consider the debate settled, even while disagreeing about when and how resolution occurred, raising the question: "Why have these debates continued?" Here I explore the biology, history, and philosophy of the possibility of natural selection operating at levels of biological organization other than the organism by focusing on debates about group-level selection that have occurred since the 1960s. In particular, I use experimental, historical, and synthetic methods to review how the debates have changed, and whether different uses of the same words and concepts can lead to different interpretations of the same experimental data. I begin with the results of a group-selection experiment I conducted using the parasitoid wasp Nasonia, and discuss how the interpretation depends on how one conceives of and defines a "group." Then I review the history of the group selection controversy and argue that this history is best interpreted as multiple, interrelated debates rather than a single continuous debate. Furthermore, I show how the aspects of these debates that have changed the most are related to theoretical content and empirical data, while disputes related to methods remain largely unchanged. Synthesizing this material, I distinguish four different "approaches" to the study of multilevel selection based on the questions and methods used by researchers, and I use the results of the Nasonia experiment to discuss how each approach can lead to different interpretations of the same experimental data. I argue that this realization can help to explain why debates about group and multilevel selection have persisted for nearly sixty years. Finally, the conclusions of this dissertation apply beyond evolutionary biology by providing an illustration of how key concepts can change over time, and how failing to appreciate this fact can lead to ongoing controversy within a scientific field. / Dissertation/Thesis / Doctoral Dissertation Biology 2014

Cooperation presents a major challenge for evolutionary theory: how can competition favor a trait that imposes a cost on the individual expressing it while benefitting another? This challenge has been answered by theory that emphasizes the importance of assortment between individuals that tend to cooperate and those who tend to behave selfishly, or `cheat'. Microbial cooperation remains puzzling, given the generally high genetic and taxonomic diversity of most microbial communities. Many microbial populations rely on shared, beneficial extracellular products for an array of functions in nature. However, when these lineages are maintained in liquid cultures, many are invaded and outcompeted by spontaneous `cheater' mutants that forego investments in these products while benefitting from those produced by neighbors. The apparent evolutionary instability of microbial investments in extracellular products in well-mixed laboratory cultures finds a natural parallel in the phenomenon of toxic microalgal blooms. These extremely dense populations of often free-living microalgae destroy populations of competing microalgae and grazing zooplankton that normally control population densities. Bloom populations of planktonic microalgae are unstructured, and seem ill suited for the evolution of cooperation. In this thesis, I have established a new theoretical framework for understanding the evolution of microbial external goods. This framework highlights the importance of cell-level structure in the distribution of these external products, as well as genetic structuring in populations. This perspective informed an investigation into the social niche of a biofilm-dwelling regulatory mutant of the important biocontrol strain Pseudomonas chlororaphis. In the highly self-structured environment of a bacterial biofilm, a surprising mutualistic association between this mutant and the wild type emerged, underscoring the importance of microbial ecology in understanding the evolution of niche construction. Extending these lessons to the evolutionary problem of exotoxins in free-swimming microalgae yields the novel possibility that fluctuations in density of toxic strains shift a cell-level functioning exotoxin into a true public good that may be exploited by cheaters. I show that exotoxicity can serve cell-level functions in Prymnesium parvum. Despite these cell-level benefits, the existence of nontoxic lineages within toxic blooms hints at a complex interaction between rapid evolutionary and ecological changes in toxic blooms.

Many plant species recognize kin and respond with changes in functional traits. Researchers hypothesize that siblings compete less than strangers. However, no study has directly tested whether siblings are less competitive. Measuring natural selection on kin recognition responses in root allocation and other destructively measured traits is challenging, since trait and fitness cannot be measured on the same individual. Here, a methodology using family-level selection is developed, measuring the trait on one individual and measuring its fitness value using another related individual. Three greenhouse pot experiments were conducted using six Brassica oleracea cultivars at two nutrient levels. We investigated whether root allocation and morphological traits were under natural selection. We tested whether or not there was cultivar recognition or resource partitioning in B. oleracea. We found that putative competitive traits (size, height, emergence and root allocation) had expected patterns of individual selection for an increase in each trait and group selection for a reduction of each trait. There was no indication that resource partitioning was occurring or that B. oleracea could recognize cultivars. However, plants were experiencing competitive interactions within pots. In conclusion, we demonstrate that using family-level selection estimates the fitness consequences of root allocation and morphological traits. / Thesis / Master of Science (MSc)