A system for studying evolution of life-like virtual organisms

BackgroundFitness landscapes, the dependences of fitness on the genotype, are of critical importance for the evolution of living beings. Unfortunately, fitness landscapes that are relevant to the evolution of complex biological functions are very poorly known. As a result, the existing theory of evolution is mostly based on postulated fitness landscapes, which diminishes its usefulness. Attempts to deduce fitness landscapes from models of actual biological processes led, so far, to only limited success.ResultsWe present a model system for studying the evolution of biological function, which makes it possible to attribute fitness to genotypes in a natural way. The system mimics a very simple cell and takes into account the basic properties of gene regulation and enzyme kinetics. A virtual cell contains only two small molecules, an organic nutrient A and an energy carrier X, and proteins of five types – two transcription factors, two enzymes, and a membrane transporter. The metabolism of the cell consists of importing A from the environment and utilizing it in order to produce X and an unspecified end product. The genome may carry an arbitrary number of genes, each one encoding a protein of one of the five types. Both major mutations that affect whole genes and minor mutations that affect individual characteristics of genes are possible. Fitness is determined by the ability of the cell to maintain homeostasis when its environment changes. The system has been implemented as a computer program, and several numerical experiments have been performed on it. Evolution of the virtual cells usually involves a rapid initial increase of fitness, which eventually slows down, until a fitness plateau is reached. The origin of a wide variety of genetic networks is routinely observed in independent experiments performed under the same conditions. These networks can have different, including very high, levels of complexity and often include large numbers of non-essential genes.ConclusionThe described system displays a rich repertoire of biologically sensible behaviors and, thus, can be useful for investigating a number of unresolved issues in evolutionary biology, including evolution of complexity, modularity and redundancy, as well as for studying the general properties of genotype-to-fitness maps.ReviewersThis article was reviewed by Drs. Eugene Koonin, Shamil Sunyaev and Arcady Mushegian.

[1]  J. Tyson,et al.  Regulation of the eukaryotic cell cycle: molecular antagonism, hysteresis, and irreversible transitions. , 2001, Journal of theoretical biology.

[2]  J. Crow,et al.  Efficiency of truncation selection. , 1979, Proceedings of the National Academy of Sciences of the United States of America.

[3]  N. K. Rozov,et al.  Differential Equations with Small Parameters and Relaxation Oscillations , 1980 .

[4]  D. Hartl,et al.  Principles of population genetics , 1981 .

[5]  John R. Koza,et al.  Genetic Programming III: Darwinian Invention & Problem Solving , 1999 .

[6]  J. Ross,et al.  Genetic-algorithm selection of a regulatory structure that directs flux in a simple metabolic model. , 1995, Biophysical journal.

[7]  C. Ofria,et al.  Adaptive Radiation from Resource Competition in Digital Organisms , 2004, Science.

[8]  H. Sauro,et al.  Preliminary Studies on the In Silico Evolution of Biochemical Networks , 2004, Chembiochem : a European journal of chemical biology.

[9]  B. Palsson,et al.  Escherichia coli K-12 undergoes adaptive evolution to achieve in silico predicted optimal growth , 2002, Nature.

[10]  William H. Press,et al.  Numerical recipes in C , 2002 .

[11]  Riccardo Poli,et al.  Foundations of Genetic Programming , 1999, Springer Berlin Heidelberg.

[12]  V. Hakim,et al.  Design of genetic networks with specified functions by evolution in silico. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[13]  F. Cross,et al.  Growth and division. , 1994, Science.

[14]  Heekuck Oh,et al.  Neural Networks for Pattern Recognition , 1993, Adv. Comput..

[15]  John H. Holland,et al.  Adaptation in Natural and Artificial Systems: An Introductory Analysis with Applications to Biology, Control, and Artificial Intelligence , 1992 .

[16]  David E. Goldberg,et al.  The Design of Innovation: Lessons from and for Competent Genetic Algorithms , 2002 .

[17]  W. J. Dickinson,et al.  Marginal fitness contributions of nonessential genes in yeast. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[18]  S. Ehrlich,et al.  Essential Bacillus subtilis genes , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[19]  M. Goebl,et al.  Most of the yeast genomic sequences are not essential for cell growth and division , 1986, Cell.

[20]  D Botstein,et al.  Functional Analysis of the Genes of Yeast Chromosome V by Genetic Footprinting , 1996, Science.

[21]  C. Ofria,et al.  Genome complexity, robustness and genetic interactions in digital organisms , 1999, Nature.

[22]  James M. Roberts,et al.  Living with or without cyclins and cyclin-dependent kinases. , 2004, Genes & development.

[23]  Robert T. Pennock,et al.  The evolutionary origin of complex features , 2003, Nature.

[24]  John H. Holland,et al.  Genetic Algorithms and the Optimal Allocation of Trials , 1973, SIAM J. Comput..

[25]  John R. Koza Genetic Programming III - Darwinian Invention and Problem Solving , 1999, Evolutionary Computation.

[26]  S. Gould,et al.  The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme , 1979, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[27]  C. Ofria,et al.  Evolution of digital organisms at high mutation rates leads to survival of the flattest , 2001, Nature.

[28]  B. Palsson,et al.  Genome-scale models of microbial cells: evaluating the consequences of constraints , 2004, Nature Reviews Microbiology.

[29]  P. Nordin Genetic Programming III - Darwinian Invention and Problem Solving , 1999 .

[30]  C. Adami,et al.  Introduction To Artificial Life , 1997, IEEE Trans. Evol. Comput..

[31]  J. W. Campbell,et al.  Experimental Determination and System Level Analysis of Essential Genes in Escherichia coli MG1655 , 2003, Journal of bacteriology.

[32]  David E. Goldberg,et al.  Genetic Algorithms in Search Optimization and Machine Learning , 1988 .