Charles Ofria

PERSPECTIVE:EVOLUTION AND DETECTION OF GENETIC ROBUSTNESS

Abstract Robustness is the invariance of phenotypes in the face of perturbation. The robustness of phenotypes appears at various levels of biological organization, including gene expression, protein folding, metabolic flux, physiological homeostasis, development, and even organismal fitness. The mechanisms underlying robustness are diverse, ranging from thermodynamic stability at the RNA and protein level to behavior at the organismal level. Phenotypes can be robust either against heritable perturbations (e.g., mutations) or nonheritable perturbations (e.g., the weather). Here we primarily focus on the first kind of robustness—genetic robustness—and survey three growing avenues of research: (1) measuring genetic robustness in nature and in the laboratory; (2) understanding the evolution of genetic robustness; and (3) exploring the implications of genetic robustness for future evolution.

The evolutionary origin of complex features

A long-standing challenge to evolutionary theory has been whether it can explain the origin of complex organismal features. We examined this issue using digital organisms—computer programs that self-replicate, mutate, compete and evolve. Populations of digital organisms often evolved the ability to perform complex logic functions requiring the coordinated execution of many genomic instructions. Complex functions evolved by building on simpler functions that had evolved earlier, provided that these were also selectively favoured. However, no particular intermediate stage was essential for evolving complex functions. The first genotypes able to perform complex functions differed from their non-performing parents by only one or two mutations, but differed from the ancestor by many mutations that were also crucial to the new functions. In some cases, mutations that were deleterious when they appeared served as stepping-stones in the evolution of complex features. These findings show how complex functions can originate by random mutation and natural selection.

Avida: a software platform for research in computational evolutionary biology

Avida is a software platform for experiments with self-replicating and evolving computer programs. It provides detailed control over experimental settings and protocols, a large array of measurement tools, and sophisticated methods to analyze and post-process experimental data. We explain the general principles on which Avida is built, as well as its main components and their interactions. We also explain how experiments are set up, carried out, and analyzed.

Genome complexity, robustness and genetic interactions in digital organisms

Digital organisms are computer programs that self-replicate, mutate and adapt by natural selection. They offer an opportunity to test generalizations about living systems that may extend beyond the organic life that biologists usually study. Here we have generated two classes of digital organism: simple programs selected solely for rapid replication, and complex programs selected to perform mathematical operations that accelerate replication through a set of defined ‘metabolic’ rewards. To examine the differences in their genetic architecture, we introduced millions of single and multiple mutations into each organism and measured the effects on the organisms fitness. The complex organisms are more robust than the simple ones with respect to the average effects of single mutations. Interactions among mutations are common and usually yield higher fitness than predicted from the component mutations assuming multiplicative effects; such interactions are especially important in the complex organisms. Frequent interactions among mutations have also been seen in bacteria, fungi and fruitflies. Our findings support the view that interactions are a general feature of genetic systems.

Balancing robustness and evolvability.

Can a single unifying mathematical framework help to explain robustness - the ability of organisms to persist in the face of changing conditions - at all biological scales, from biochemical to ecological?

Evolving coordinated quadruped gaits with the HyperNEAT generative encoding

Legged robots show promise for complex mobility tasks, such as navigating rough terrain, but the design of their control software is both challenging and laborious. Traditional evolutionary algorithms can produce these controllers, but require manual decomposition or other problem simplification because conventionally-used direct encodings have trouble taking advantage of a problems regularities and symmetries. Such active intervention is time consuming, limits the range of potential solutions, and requires the user to possess a deep understanding of the problems structure. This paper demonstrates that HyperNEAT, a new and promising generative encoding for evolving neural networks, can evolve quadruped gaits without an engineer manually decomposing the problem. Analyses suggest that HyperNEAT is successful because it employs a generative encoding that can more easily reuse phenotypic modules. It is also one of the first neuroevolutionary algorithms that exploits a problems geometric symmetries, which may aid its performance. We compare HyperNEAT to FT-NEAT, a direct encoding control, and find that HyperNEAT is able to evolve impressive quadruped gaits and vastly outperforms FT-NEAT. Comparative analyses reveal that HyperNEAT individuals are more holistically affected by genetic operators, resulting in better leg coordination. Overall, the results suggest that HyperNEAT is a powerful algorithm for evolving control systems for complex, yet regular, devices, such as robots.

Sexual reproduction reshapes the genetic architecture of digital organisms

Modularity and epistasis, as well as other aspects of genetic architecture, have emerged as central themes in evolutionary biology. Theory suggests that modularity promotes evolvability, and that aggravating (synergistic) epistasis among deleterious mutations facilitates the evolution of sex. Here, by contrast, we investigate the evolution of different genetic architectures using digital organisms, which are computer programs that self-replicate, mutate, compete and evolve. Specifically, we investigate how genetic architecture is shaped by reproductive mode. We allowed 200 populations of digital organisms to evolve for over 10 000 generations while reproducing either asexually or sexually. For 10 randomly chosen organisms from each population, we constructed and analysed all possible single mutants as well as one million mutants at each mutational distance from 2 to 10. The genomes of sexual organisms were more modular than asexual ones; sites encoding different functional traits had less overlap and sites encoding a particular trait were more tightly clustered. Net directional epistasis was alleviating (antagonistic) in both groups, although the overall strength of this epistasis was weaker in sexual than in asexual organisms. Our results show that sexual reproduction profoundly influences the evolution of the genetic architecture.

On the Performance of Indirect Encoding Across the Continuum of Regularity

This paper investigates how an evolutionary algorithm with an indirect encoding exploits the property of phenotypic regularity, an important design principle found in natural organisms and engineered designs. We present the first comprehensive study showing that such phenotypic regularity enables an indirect encoding to outperform direct encoding controls as problem regularity increases. Such an ability to produce regular solutions that can exploit the regularity of problems is an important prerequisite if evolutionary algorithms are to scale to high-dimensional real-world problems, which typically contain many regularities, both known and unrecognized. The indirect encoding in this case study is HyperNEAT, which evolves artificial neural networks (ANNs) in a manner inspired by concepts from biological development. We demonstrate that, in contrast to two direct encoding controls, HyperNEAT produces both regular behaviors and regular ANNs, which enables HyperNEAT to significantly outperform the direct encodings as regularity increases in three problem domains. We also show that the types of regularities HyperNEAT produces can be biased, allowing domain knowledge and preferences to be injected into the search. Finally, we examine the downside of a bias toward regularity. Even when a solution is mainly regular, some irregularity may be needed to perfect its functionality. This insight is illustrated by a new algorithm called HybrID that hybridizes indirect and direct encodings, which matched HyperNEATs performance on regular problems yet outperformed it on problems with some irregularity. HybrIDs ability to improve upon the performance of HyperNEAT raises the question of whether indirect encodings may ultimately excel not as stand-alone algorithms, but by being hybridized with a further process of refinement, wherein the indirect encoding produces patterns that exploit problem regularity and the refining process modifies that pattern to capture irregularities. This paper thus paints a more complete picture of indirect encodings than prior studies because it analyzes the impact of the continuum between irregularity and regularity on the performance of such encodings, and ultimately suggests a path forward that combines indirect encodings with a separate process of refinement.

EFFECTS OF POPULATION SIZE AND MUTATION RATE ON THE EVOLUTION OF MUTATIONAL ROBUSTNESS

Abstract It is often assumed that the efficiency of selection for mutational robustness would be proportional to mutation rate and population size, thus being inefficient in small populations. However, Krakauer and Plotkin (2002) hypothesized that selection in small populations would favor robustness mechanisms, such as redundancy, that mask the effect of deleterious mutations. In large populations, by contrast, selection is more effective at removing deleterious mutants and fitness would be improved by eliminating mechanisms that mask the effect of deleterious mutations and thus impede their removal. Here, we test whether these predictions are supported in experiments with evolving populations of digital organisms. Digital organisms are self-replicating programs that inhabit a virtual world inside a computer. Like their organic counterparts, digital organisms mutate, compete, evolve, and adapt by natural selection to their environment. In this study, 160 populations evolved at different combinations of mutation rate and population size. After 104 generations, we measured the mutational robustness of the most abundant genotype in each population. Mutational robustness tended to increase with mutation rate and to decline with population size, although the dependence with population size was in part mediated by a negative relationship between fitness and robustness. These results are independent of whether genomes were constrained to their original length or allowed to change in size.

Distributed Cooperative Caching in Social Wireless Networks

This paper introduces cooperative caching policies for minimizing electronic content provisioning cost in Social Wireless Networks (SWNET). SWNETs are formed by mobile devices, such as data enabled phones, electronic book readers etc., sharing common interests in electronic content, and physically gathering together in public places. Electronic object caching in such SWNETs are shown to be able to reduce the content provisioning cost which depends heavily on the service and pricing dependences among various stakeholders including content providers (CP), network service providers, and End Consumers (EC). Drawing motivation from Amazons Kindle electronic book delivery business, this paper develops practical network, service, and pricing models which are then used for creating two object caching strategies for minimizing content provisioning costs in networks with homogenous and heterogeneous object demands. The paper constructs analytical and simulation models for analyzing the proposed caching strategies in the presence of selfish users that deviate from network-wide cost-optimal policies. It also reports results from an Android phone-based prototype SWNET, validating the presented analytical and simulation results.