Evolvability and organismal architecture:The blind watchmaker and the reminiscent architect

Organisms are constantly faced with the challenge of adapting to new circumstances. In this thesis, I argue that the ability to adapt to new circumstances, “evolvability”, is deeply ingrained in the genetic, developmental, morphological, and physiological architecture of organisms. Using a blend of conceptual research, theoretical modelling, and multidisciplinary studies, I demonstrate how organismal architecture can evolve so that organisms can cope better and better with future environmental challenges. As a first step, I systematically classify the many factors contributing to evolvability. Then I use a simulation approach to show how evolvability-enhancing structures can readily evolve in gene-regulatory networks. This happens via the evolution of "mutational transformers" - structural elements that convert random mutations at the genetic level into adaptation-enhancing mutations at the phenotypic level. In another thesis chapter, I demonstrate that even if selection acts only sporadically, complex adaptations can evolve and persist over long time periods. In other words, complex adaptations do not require constant selection pressure. In an interdisciplinary contribution, I apply biological insights regarding the properties of an evolvability-enhancing mutation structure to the design of algorithms used in Artificial Intelligence. The result is the “Facilitated Mutation” method which enhances the performance of the algorithms in various respects, highlighting the potential for leveraging biological principles in computational sciences. Finally, I embed my research findings in a philosophical context. I emphasise the importance of organismal architecture in retaining evolutionary memories and suggest future research directions to further enhance our understanding of evolvability

Selection, tinkering and emergence in complex networks: crossing the land of tinkering

Complex biological networks have very different origins than technologic ones. The latter involve extensive design and, as engineered structures, include a high level of optimization. The former involve (in principle) contingency and structural constraints, with new structures being incorporated through tinkering with previously evolved modules or units. However, the observation of the topological features of different biological nets suggests that nature can have a limited repertoire of ”attractors” that essentially optimize communication under some basic constraints of cost and architecture or that allow the biological nets to reach a high degree of homeostasis. Conversely, the topological features exhibited by some technology graphs indicate that tinkering and internal constraints play a key role, in spite of the ”designed” nature of these structures. Previous scenarios suggested to explain the overall trends of evolution are re-analyzed in light of topological patterns.Peer ReviewedPostprint (author's final draft

`The frozen accident' as an evolutionary adaptation: A rate distortion theory perspective on the dynamics and symmetries of genetic coding mechanisms

We survey some interpretations and related issues concerning the frozen hypothesis due to F. Crick and how it can be explained in terms of several natural mechanisms involving error correction codes, spin glasses, symmetry breaking and the characteristic robustness of genetic networks. The approach to most of these questions involves using elements of Shannon's rate distortion theory incorporating a semantic system which is meaningful for the relevant alphabets and vocabulary implemented in transmission of the genetic code. We apply the fundamental homology between information source uncertainty with the free energy density of a thermodynamical system with respect to transcriptional regulators and the communication channels of sequence/structure in proteins. This leads to the suggestion that the frozen accident may have been a type of evolutionary adaptation

Artificial Ontogenies: A Computational Model of the Control and Evolution of Development

Understanding the behaviour of biological systems is a challenging task. Gene regulation, development and evolution are each a product of nonlinear interactions between many individual agents: genes, cells or organisms. Moreover, these three processes are not isolated, but interact with one another in an important fashion. The development of an organism involves complex patterns of dynamic behaviour at the genetic level. The gene networks that produce this behaviour are subject to mutations that can alter the course of development, resulting in the production of novel morphologies. Evolution occurs when these novel morphologies are favoured by natural selection and survive to pass on their genes to future generations. Computational models can assist us to understand biological systems by providing a framework within which their behaviour can be explored. Many natural processes, including gene regulation and development, have a computational element to their control. Constructing formal models of these systems enables their behaviour to be simulated, observed and quantified on a scale not otherwise feasible. This thesis uses a computational simulation methodology to explore the relationship between development and evolution. An important question in evolutionary biology is how to explain the direction of evolution. Conventional explanations of evolutionary history have focused on the role of natural selection in orienting evolution. More recently, it has been argued that the nature of development, and the way it changes in response to mutation, may also be a significant factor. A network-lineage model of artificial ontogenies is described that incorporates a developmental mapping between the dynamics of a gene network and a cell lineage representation of a phenotype. Three series of simulation studies are reported, exploring: (a) the relationship between the structure of a gene network and its dynamic behaviour; (b) the characteristic distributions of ontogenies and phenotypes generated by the dynamics of gene networks; (c) the effect of these characteristic distributions on the evolution of ontogeny. The results of these studies indicate that the model networks are capable of generating a diverse range of stable behaviours, and possess a small yet significant sensitivity to perturbation. In the context of developmental control, the intrinsic dynamics of the model networks predispose the production of ontogenies with a modular, quasi-systematic structure. This predisposition is reflected in the structure of variation available for selection in an adaptive search process, resulting in the evolution of ontogenies biased towards simplicity. These results suggest a possible explanation for the levels of ontogenetic complexity observed in biological organisms: that they may be a product of the network architecture of developmental control. By quantifying complexity, variation and bias, the network-lineage model described in this thesis provides a computational method for investigating the effects of development on the direction of evolution. In doing so, it establishes a viable framework for simulating computational aspects of complex biological systems

Artificial evolution with Binary Decision Diagrams: a study in evolvability in neutral spaces

This thesis develops a new approach to evolving Binary Decision Diagrams, and uses it to study evolvability issues. For reasons that are not yet fully understood, current approaches to artificial evolution fail to exhibit the evolvability so readily exhibited in nature. To be able to apply evolvability to artificial evolution the field must first understand and characterise it; this will then lead to systems which are much more capable than they are currently. An experimental approach is taken. Carefully crafted, controlled experiments elucidate the mechanisms and properties that facilitate evolvability, focusing on the roles and interplay between neutrality, modularity, gradualism, robustness and diversity. Evolvability is found to emerge under gradual evolution as a biased distribution of functionality within the genotype-phenotype map, which serves to direct phenotypic variation. Neutrality facilitates fitness-conserving exploration, completely alleviating local optima. Population diversity, in conjunction with neutrality, is shown to facilitate the evolution of evolvability. The search is robust, scalable, and insensitive to the absence of initial diversity. The thesis concludes that gradual evolution in a search space that is free of local optima by way of neutrality can be a viable alternative to problematic evolution on multi-modal landscapes

Landscapes and Effective Fitness

The concept of a fitness landscape arose in theoretical biology, while that of effective fitness has its origin in evolutionary computation. Both have emerged as useful conceptual tools with which to understand the dynamics of evolutionary processes, especially in the presence of complex genotype-phenotype relations. In this contribution we attempt to provide a unified discussion of these two approaches, discussing both their advantages and disadvantages in the context of some simple models. We also discuss how fitness and effective fitness change under various transformations of the configuration space of the underlying genetic model, concentrating on coarse-graining transformations and on a particular coordinate transformation that provides an appropriate basis for illuminating the structure and consequences of recombination