Abstract
Genotype–phenotype mapping plays an essential role in the design of an evolutionary algorithm. Variation occurs at the genotypic level but fitness is evaluated at the phenotypic level, therefore, this mapping determines if and how variations are effectively translated into quality improvements. In evolutionary algorithms, this mapping has often been observed as highly redundant, i.e., multiple genotypes can map to the same phenotype, as well as heterogeneous, i.e., some phenotypes are represented by a large number of genotypes while some phenotypes only have few. We numerically study the redundant genotype–phenotype mapping of a simple Boolean linear genetic programming system and quantify the mutational connections among phenotypes using tools of complex network analysis. The analysis yields several interesting statistics of the phenotype network. We show the evidence and provide explanations for the observation that some phenotypes are much more difficult to find as the _target of a search than others. Our study provides a quantitative analysis framework to better understand the genotype–phenotype map, and the results may be utilized to inspire algorithm design that allows the search of a difficult _target to be more effective.
Similar content being viewed by others
Explore related subjects
Discover the latest articles, news and stories from top researchers in related subjects.References
P. Alberch, From genes to phenotype: dynamical systems and evolvability. Genetica 84, 5–11 (1991)
L. Altenberg, The evolution of evolvability in genetic programming, in Advances in Genetic Programming, (MIT Press, Cambridge, MA, 1994), pp. 47–74
L. Altenberg. Genome growth and the evolution of the genotype-phenotype map, in W. Banzhaf and F. Eeckman, eds., Evolution and Biocomputation, volume 899 of Lecture Notes in Computer Science. (Springer, 1995), pp. 205–259
W. Banzhaf, Genotype–phenotype mapping and neutral variation—a case study in genetic programming, in Parallel Problem Solving from Nature, volume of 866 Lecture Notes in Computer Science, ed. by Y. Davidor, H.-P. Schwefel, R. Manner (Springer, Berlin, 1994), pp. 322–332
A.-L. Barábasi, Network Science (Cambridge University Press, Cambridge, 2016)
J.D. Bloom, S.T. Labthavikul, C.R. Otey, F.H. Arnold, Protein stability promotes evolvability. Proc. Nat. Acad. Sci. 103(15), 5869–5874 (2006)
M.F. Brameier, W. Banzhaf, Linear Genetic Programming (Springer, Berlin, 2007)
P. Catalan, A. Wagner, S. Manrubia, J.A. Cuesta, Adding levels of complexity enhances robustness and evolvability in a multilevel genotype–phenotype map. J. R. Soc. Interface. 15(138), 20170516 (2018)
J. Clune, K.O. Stanley, R.T. Pennock, C. Ofria, On the performance of indirect encoding across the continuum of regularity. IEEE Trans. Evolut. Comput. 15(3), 346–367 (2011)
M.C. Cowperthwaite, E.P. Economo, W.R. Harcombe, E.L. Miller, L.A. Meyers, The ascent of the abundant: how mutational networks constrain evolution. PLoS Comput. Biol. 4(7), e1000110 (2008)
M.C. Cowperthwaite, L.A. Meyers, How mutational networks shape evolution: lessons from RNA models. Annu. Rev. Ecol. Evol. Syst. 38, 203–230 (2007)
G. Csardi, T. Nepusz, The igraph software package for complex network research. InterJ. Complex Syst. 1695, 1–9 (2006)
S. Cussat-Blanc, K. Harrington, W. Banzhaf, Artificial gene regulatory networks—a review. Artif. lLfe 24(4), 296–328 (2019)
E.H. Davidson, The Regulatory Genome: Gene Regulatory Networks in Development and Evolution (Elsevier, Amsterdam, 2010)
J.A.G.M. de Visser, J. Krug, Empirical fitness landscapes and the predictability of evolution. Nat. Rev. Genet. 15, 480–490 (2014)
M. Ebner, M. Shackleton, R. Shipman, How neutral networks influence evolvability. Complexity 7(2), 19–33 (2002)
A. Fontana, Epigenetic tracking: biological implications, in European Conference on Artificial Life, (Springer, 2009), pp. 10–17
E. Galvan-Lopez, R. Poli, An empirical investigation of how and why neutrality affects evolutionary search, in M. Cattolico, ed., Proceedings of the Genetic and Evolutionary Computation Conference, (2006), pp. 1149–1156
T. Hu, W. Banzhaf, Neutrality and variability: Two sides of evolvability in linear genetic programming, in Proceedings of the 18th Genetic and Evolutionary Computation Conference (GECCO), (2009), pp. 963–970
T. Hu, W. Banzhaf, Quantitative analysis of evolvability using vertex centralities in phenotype network, in Proceedings of the 25th Genetic and Evolutionary Computation Conference (GECCO), (2016), pp. 733–740
T. Hu, W. Banzhaf, Neutrality, robustness, and evolvability in genetic programming, in R. Riolo, B. Worzel, B. Goldman, B. Tozier, eds., Genetic Programming Theory and Practice XIV, chapter 7, (Springer, 2018), pp. 101–117
T. Hu, W. Banzhaf, J.H. Moore, The effect of recombination on phenotypic exploration and robustness in evolution. Artif. Life 20(4), 457–470 (2014)
T. Hu, J. Payne, W. Banzhaf, J.H. Moore, Evolutionary dynamics on multiple scales: a quantitative analysis of the interplay between genotype, phenotype, and fitness in linear genetic programming. Genet. Program. Evolv. Mach. 13(3), 305–337 (2012)
T. Hu, M. Tomassini, W. Banzhaf, Complex network analysis of a genetic programming phenotype network, in Proceedings of the 22nd European Conference on Genetic Programming (EuroGP), volume 11451 of Lecture Notes in Computer Science, (2019), pp. 49–63
S. Kauffman, S. Levin, Towards a general theory of adaptive walks on rugged landscapes. J. Theor. Biol. 128(1), 11–45 (1987)
D.B. Kell, Genotype-phenotype mapping: genes as computer programs. Trends Genet. 18(11), 555–559 (2002)
M. Kirschner, J. Gerhart, Evolvability. Proc. Natl. Acad. Sci. 95, 8420–8427 (1998)
M. Kirschner, J.C. Gerhart, The Plausibility of Life: Resolving Darwin’s Dilemma (Yale University Press, New Haven, 2006)
J. D. Knowles, R. A. Watson, On the utility of redundant encodings in mutation-based evolutionary search, in Parallel Problem Solving from Nature—PPSN VII, volume 2439 of Lecture Notes in Computer Science, (2002), pp. 88–98
J.R. Koza, D. Andre, M.A. Keane, F.H. Bennett III, Genetic Programming III: Darwinian Invention and Problem Solving, vol. 3 (Morgan Kaufmann, Burlington, 1999)
R.E. Lenski, J.E. Barrick, C. Ofria, Balancing robustness and evolvability. PLoS Biol. 4(12), e428 (2006)
N. Masuda, M.A. Porter, R. Lambiotte, Random walk and diffusion in networks. Phys. Rep. 716, 1–58 (2017)
R.C. McBride, C.B. Ogbunugafor, P.E. Turner, Robustness promotes evolvability of thermotolerance in an RNA virus. BMC Evolut. Biol. 8, 231 (2008)
J.F. Miller, W. Banzhaf, Evolving the program for a cell: From French flags to Boolean circuits, in On Growth, Form and Computers, ed. by S. Kumar, P. Bentley (Academic, New York, 2003), pp. 278–301
M.E.J. Newman, Networks: An Introduction (Oxford University Press, Oxford, 2018)
M.E.J. Newman, R. Engelhardt, Effects of selective neutrality on the evolution of molecular species. Proc. R. Soc. B 265(1403), 1333–1338 (1998)
M.E.J. Newman, M. Girvan, Finding and evaluating community structure in networks. Phys. Rev. E 69, 026113 (2004)
K. L. Nickerson, Y. Chen, F. Wang, T. Hu, Measuring evolvability and accessibility using the Hyperlink-Induced Topic Search algorithm, in Proceedings of the 27th Genetic and Evolutionary Computation Conference (GECCO), (2018), pp. 1175–1182
L. Page, S. Brin, R. Motwani, T. Winograd, The pagerank citation ranking: Bringing order to the web Technical report, Stanford InfoLab (1999)
J.L. Payne, A. Wagner, The causes of evolvability and their evolution. Nat. Rev. Genet. 20, 24–38 (2019)
R. Rezazadegan, C. Barrett, C. Reidys, Multiplicity of phenotypes and RNA evolution. J. Theoret. Biol. 447, 139–146 (2018)
F. Rothlauf, D.E. Goldberg, Redundant representations in evolutionary computation. Evolut. Comput. 11(4), 381–415 (2003)
S. Schaper, A.A. Louis, The arrival of the frequent: how bias in genotype–phenotype maps can steer populations to local optima. PLoS One 9(2), e86635 (2014)
P. Schuster, W. Fontana, P .F. Stadler, I .L. Hofacker, From sequences to shapes and back: a case study in RNA secondary structures. Proc. R. Soc. Lond. Ser. B Biol. Sci. 255(1344), 279–284 (1994)
P. Shannon, A. Markiel, O. Ozier, N.S. Baliga, J.T. Wang, D. Ramage, N. Amin, B. Schwikowski, T. Ideker, Cytoscape: a software environment for integrated models of biomolecular interaction networks. Genome Res. 13, 2498–2504 (2003)
T. Smith, P. Husbands, M. O’Shea, Neutral networks and evolvability with complex genotype-phenotype mapping, in J. Kelemen, P. Sosik, eds., Proceedings of the European Conference on Artificial Life, volume 2159 of Lecture Notes in Artificial Intelligence, (Springer-Verlag, 2001), pp. 272–281
E. van Nimwegen, J.P. Crutchfield, M.A. Huynen, Neutral evolution of mutational robustness. Proc. Natl. Acad. Sci. 96(17), 9716–9720 (1999)
A. Wagner, Robustness, evolvability, and neutrality. Fed. Eur. Biochem. Soc. Lett. 579(8), 1772–1778 (2005)
A. Wagner, Robustness and evolvability: a paradox resolved. Proc. R. Soc. B 275(1630), 91–100 (2008)
G.P. Wagner, L. Altenberg, Perspective: Complex adaptations and the evolution of evolvability. Evolution 50(3), 967–976 (1996)
Acknowledgements
This research was supported by the Natural Sciences and Engineering Research Council (NSERC) of Canada Discovery Grant RGPIN-2016-04699 to T.H., and the Koza Endowment fund provided to W.B. by Michigan State University and supported by its BEACON Center for the Study of Evolution in Action.
Author information
Authors and Affiliations
Corresponding author
Additional information
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
About this article
Cite this article
Hu, T., Tomassini, M. & Banzhaf, W. A network perspective on genotype–phenotype mapping in genetic programming. Genet Program Evolvable Mach 21, 375–397 (2020). https://doi.org/10.1007/s10710-020-09379-0
Received:
Revised:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10710-020-09379-0