Increasing Evolvability Considered as a Large-Scale Trend in Evolution

Evolvability is the capacity to evolve. This paper introduces a simple computational model of evolvability and demonstrates that, under certain conditions, evolvability can increase indefinitely, even when there is no direct selection for evolvability. The model shows that increasing evolvability implies an accelerating evolutionary pace. It is suggested that the conditions for indefinitely increasing evolvability are satisfied in biological and cultural evolution. We claim that increasing evolvability is a large-scale trend in evolution. This hypothesis leads to testable predictions about biological and cultural evolution

The Emergence of Canalization and Evolvability in an Open-Ended, Interactive Evolutionary System

Natural evolution has produced a tremendous diversity of functional organisms. Many believe an essential component of this process was the evolution of evolvability, whereby evolution speeds up its ability to innovate by generating a more adaptive pool of offspring. One hypothesized mechanism for evolvability is developmental canalization, wherein certain dimensions of variation become more likely to be traversed and others are prevented from being explored (e.g. offspring tend to have similarly sized legs, and mutations affect the length of both legs, not each leg individually). While ubiquitous in nature, canalization almost never evolves in computational simulations of evolution. Not only does that deprive us of in silico models in which to study the evolution of evolvability, but it also raises the question of which conditions give rise to this form of evolvability. Answering this question would shed light on why such evolvability emerged naturally and could accelerate engineering efforts to harness evolution to solve important engineering challenges. In this paper we reveal a unique system in which canalization did emerge in computational evolution. We document that genomes entrench certain dimensions of variation that were frequently explored during their evolutionary history. The genetic representation of these organisms also evolved to be highly modular and hierarchical, and we show that these organizational properties correlate with increased fitness. Interestingly, the type of computational evolutionary experiment that produced this evolvability was very different from traditional digital evolution in that there was no objective, suggesting that open-ended, divergent evolutionary processes may be necessary for the evolution of evolvability.Comment: SI can be found at: http://www.evolvingai.org/files/SI_0.zi

Degeneracy: a design principle for achieving robustness and evolvability

Robustness, the insensitivity of some of a biological system's functionalities to a set of distinct conditions, is intimately linked to fitness. Recent studies suggest that it may also play a vital role in enabling the evolution of species. Increasing robustness, so is proposed, can lead to the emergence of evolvability if evolution proceeds over a neutral network that extends far throughout the fitness landscape. Here, we show that the design principles used to achieve robustness dramatically influence whether robustness leads to evolvability. In simulation experiments, we find that purely redundant systems have remarkably low evolvability while degenerate, i.e. partially redundant, systems tend to be orders of magnitude more evolvable. Surprisingly, the magnitude of observed variation in evolvability can neither be explained by differences in the size nor the topology of the neutral networks. This suggests that degeneracy, a ubiquitous characteristic in biological systems, may be an important enabler of natural evolution. More generally, our study provides valuable new clues about the origin of innovations in complex adaptive systems.Comment: Accepted in the Journal of Theoretical Biology (Nov 2009

Adaptive evolvability through direct selection instead of indirect, second‐order selection

Can evolvability itself be the product of adaptive evolution? To answer this question is challenging, because any DNA mutation that alters only evolvability is subject to indirect, “second order” selection on the future effects of this mutation. Such indirect selection is weaker than “first-order” selection on mutations that alter fitness, in the sense that it can operate only under restrictive conditions. Here I discuss a route to adaptive evolvability that overcomes this challenge. Specifically, a recent evolution experiment showed that some mutations can enhance both fitness and evolvability through a combination of direct and indirect selection. Unrelated evidence from gene duplication and the evolution of gene regulation suggests that mutations with such dual effects may not be rare. Through such mutations, evolvability may increase at least in part because it provides an adaptive advantage. These observations suggest a research program on the adaptive evolution of evolvability, which aims to identify such mutations and to disentangle their direct fitness effects from their indirect effects on evolvability. If evolvability is itself adaptive, Darwinian evolution may have created more than life's diversity. It may also have helped create the very conditions that made the success of Darwinian evolution possible

Degeneracy: a link between evolvability, robustness and complexity in biological systems

A full accounting of biological robustness remains elusive; both in terms of the mechanisms by which robustness is achieved and the forces that have caused robustness to grow over evolutionary time. Although its importance to topics such as ecosystem services and resilience is well recognized, the broader relationship between robustness and evolution is only starting to be fully appreciated. A renewed interest in this relationship has been prompted by evidence that mutational robustness can play a positive role in the discovery of adaptive innovations (evolvability) and evidence of an intimate relationship between robustness and complexity in biology. This paper offers a new perspective on the mechanics of evolution and the origins of complexity, robustness, and evolvability. Here we explore the hypothesis that degeneracy, a partial overlap in the functioning of multi-functional components, plays a central role in the evolution and robustness of complex forms. In support of this hypothesis, we present evidence that degeneracy is a fundamental source of robustness, it is intimately tied to multi-scaled complexity, and it establishes conditions that are necessary for system evolvability

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

Scuba Search : when selection meets innovation

We proposed a new search heuristic using the scuba diving metaphor. This approach is based on the concept of evolvability and tends to exploit neutrality in fitness landscape. Despite the fact that natural evolution does not directly select for evolvability, the basic idea behind the scuba search heuristic is to explicitly push the evolvability to increase. The search process switches between two phases: Conquest-of-the-Waters and Invasion-of-the-Land. A comparative study of the new algorithm and standard local search heuristics on the NKq-landscapes has shown advantage and limit of the scuba search. To enlighten qualitative differences between neutral search processes, the space is changed into a connected graph to visualize the pathways that the search is likely to follow

Evolution favors protein mutational robustness in sufficiently large populations

BACKGROUND: An important question is whether evolution favors properties such as mutational robustness or evolvability that do not directly benefit any individual, but can influence the course of future evolution. Functionally similar proteins can differ substantially in their robustness to mutations and capacity to evolve new functions, but it has remained unclear whether any of these differences might be due to evolutionary selection for these properties. RESULTS: Here we use laboratory experiments to demonstrate that evolution favors protein mutational robustness if the evolving population is sufficiently large. We neutrally evolve cytochrome P450 proteins under identical selection pressures and mutation rates in populations of different sizes, and show that proteins from the larger and thus more polymorphic population tend towards higher mutational robustness. Proteins from the larger population also evolve greater stability, a biophysical property that is known to enhance both mutational robustness and evolvability. The excess mutational robustness and stability is well described by existing mathematical theories, and can be quantitatively related to the way that the proteins occupy their neutral network. CONCLUSIONS: Our work is the first experimental demonstration of the general tendency of evolution to favor mutational robustness and protein stability in highly polymorphic populations. We suggest that this phenomenon may contribute to the mutational robustness and evolvability of viruses and bacteria that exist in large populations