{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,27]],"date-time":"2026-01-27T21:43:36Z","timestamp":1769550216597,"version":"3.49.0"},"reference-count":40,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,6,10]],"date-time":"2021-06-10T00:00:00Z","timestamp":1623283200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/2.zoppoz.workers.dev:443\/https\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Robot. AI"],"abstract":"<jats:p>Given that selection removes genetic variance from evolving populations, thereby reducing exploration opportunities, it is important to find mechanisms that create genetic variation without the disruption of adapted genes and genomes caused by random mutation. Just such an alternative is offered by random epigenetic error, a developmental process that acts on materials and parts expressed by the genome. In this system of embodied computational evolution, simulated within a physics engine, epigenetic error was instantiated in an explicit genotype-to-phenotype map as transcription error at the initiation of gene expression. The hypothesis was that transcription error would create genetic variance by shielding genes from the direct impact of selection, creating, in the process, <jats:italic>masquerading genomes<\/jats:italic>. To test this hypothesis, populations of simulated embodied biorobots and their developmental systems were evolved under steady directional selection as equivalent rates of random mutation and random transcriptional error were covaried systematically in an 11 \u00d7 11 fully factorial experimental design. In each of the 121 different experimental conditions (unique combinations of mutation and transcription error), the same set of 10 randomly created replicate populations of 60 individuals were evolved. Selection for the improved locomotor behavior of individuals led to increased mean fitness of populations over 100 generations at nearly all levels and combinations of mutation and transcription error. When the effects of both types of error were partitioned statistically, increasing transcription error was shown to increase the final genetic variance of populations, incurring a fitness cost but acting on variance independently and differently from genetic mutation. Thus, random epigenetic errors in development feed back through selection of individuals with masquerading genomes to the population\u2019s genetic variance over generational time. Random developmental processes offer an additional mechanism for exploration by increasing genetic variation in the face of steady, directional selection.<\/jats:p>","DOI":"10.3389\/frobt.2021.674823","type":"journal-article","created":{"date-parts":[[2021,6,10]],"date-time":"2021-06-10T05:49:58Z","timestamp":1623304198000},"update-policy":"https:\/\/2.zoppoz.workers.dev:443\/https\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":3,"title":["Embodied Computational Evolution: Feedback Between Development and Evolution in Simulated Biorobots"],"prefix":"10.3389","volume":"8","author":[{"given":"Joshua","family":"Hawthorne-Madell","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eric","family":"Aaron","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ken","family":"Livingston","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"suffix":"Jr.","given":"John H.","family":"Long","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2021,6,10]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1093\/icb\/23.2.347","article-title":"Morphology, Performance and Fitness","volume":"23","author":"Arnold","year":"1983","journal-title":"Am. Zool"},{"key":"B2","doi-asserted-by":"publisher","first-page":"e83438","DOI":"10.1371\/journal.pone.0083438","article-title":"Critical Mutation Rate Has an Exponential Dependence on Population Size in Haploid and Diploid Populations","volume":"8","author":"Aston","year":"2013","journal-title":"PLoS One"},{"key":"B3","doi-asserted-by":"crossref","DOI":"10.1109\/CEC.2002.1004528","article-title":"Evolving Modular Genetic Regulatory Networks","author":"Bongard","year":"2002"},{"key":"B4","doi-asserted-by":"crossref","DOI":"10.7551\/978-0-262-29714-1-ch019","article-title":"Discovering and Maintaining Behaviours Inaccessible to Incremental Genetic Evolution through Transcription Errors and Cultural Transmission","volume-title":"Advances in Artificial Life, ECAL 2011: Proceedings of the Eleventh European Conference on the Synthesis and Simulation of Living Systems","author":"Borg","year":"2011"},{"key":"B5","article-title":"The Emergence and Utility of Social Behaviour and Social Learning in Artificial Evolutionary Systems","author":"Borg","year":"2018"},{"key":"B6","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/frobt.2017.00001","article-title":"Epigenetic Operators and the Evolution of Physically Embodied Robots","volume":"4","author":"Brawer","year":"2017","journal-title":"Front. Robotics AI"},{"key":"B7","doi-asserted-by":"publisher","first-page":"1662","DOI":"10.1073\/pnas.1423275112","article-title":"Causes of Natural Variation in Fitness: Evidence from Studies of Drosophila Populations","volume":"112","author":"Charlesworth","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"B8","doi-asserted-by":"publisher","first-page":"396","DOI":"10.1073\/pnas.76.1.396","article-title":"Efficiency of Truncation Selection","volume":"76","author":"Crow","year":"1979","journal-title":"Proc. Natl. Acad. Sci."},{"key":"B9","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/j.ydbio.2016.06.005","article-title":"Epigenetics: The Origins and Evolution of a Fashionable Topic","volume":"416","author":"Deichmann","year":"2016","journal-title":"Develop. Biol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"4","DOI":"10.3389\/frobt.2015.00004","article-title":"Evolutionary Robotics: what, Why, and where to","volume":"2","author":"Doncieux","year":"2015","journal-title":"Front. Robotics AI"},{"key":"B11","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1055\/s-0029-1237423","article-title":"Epigenetics: Definition, Mechanisms and Clinical Perspective","volume":"27","author":"Dupont","year":"","journal-title":"Semin. Reprod. Med."},{"key":"B12","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1038\/nrmicro3367","article-title":"At the centre: Influenza A Virus Ribonucleoproteins","volume":"13","author":"Eisfeld","year":"2015","journal-title":"Nat. Rev. Microbiol."},{"key":"B13","doi-asserted-by":"publisher","first-page":"250","DOI":"10.1162\/artl_a_00296","article-title":"Mapping the Peaks: Fitness Landscapes of the Fittest and the Flattest","volume":"25","author":"Franklin","year":"2019","journal-title":"Artif. Life"},{"key":"B14","volume-title":"Population Genetics: A Concise Guide","author":"Gillespie","year":"1998"},{"key":"B15","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1080\/03081070108960712","article-title":"The Notion of Stability in Open Dynamical Systems from an Ecological Perspective","volume":"30","author":"Gregorius","year":"2001","journal-title":"Int. J. Gen. Syst."},{"key":"B16","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/978-3-642-23363-0_7","article-title":"Solving Deceptive Tasks in Robot Body-Brain Co-evolution by Searching for Behavioral novelty","volume-title":"Advances in Robotics and Virtual Reality","author":"Kr\u010dah","year":"2012"},{"key":"B17","doi-asserted-by":"publisher","first-page":"318","DOI":"10.1126\/science.aay2727","article-title":"Tales from the Crypt(ic)","volume":"365","author":"Lee","year":"2019","journal-title":"Science"},{"key":"B18","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1162\/EVCO_a_00025","article-title":"Abandoning Objectives: Evolution through the Search for novelty Alone","volume":"19","author":"Lehman","year":"2011","journal-title":"Evol. Comput."},{"key":"B19","first-page":"329","article-title":"Exploiting Open-Endedness to Solve Problems through the Search for novelty","author":"Lehman","year":"2008"},{"key":"B20","volume-title":"Darwin's Devices: What Evolving Robots Can Teach Us About the History of Life and the Future of Technology","author":"Long","year":"2012"},{"key":"B21","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1016\/j.tig.2010.05.003","article-title":"Evolution of the Mutation Rate","volume":"26","author":"Lynch","year":"2010","journal-title":"Trends Genet."},{"key":"B22","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1016\/j.tig.2010.06.002","article-title":"Robustness and Evolvability","volume":"26","author":"Masel","year":"2010","journal-title":"Trends Genet."},{"key":"B23","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1016\/j.robot.2008.09.009","article-title":"Fitness Functions in Evolutionary Robotics: A Survey and Analysis","volume":"57","author":"Nelson","year":"2009","journal-title":"Robotics Autonomous Syst."},{"key":"B24","doi-asserted-by":"publisher","first-page":"743","DOI":"10.1093\/icb\/42.4.743","article-title":"Understanding Vertebrate Brain Evolution","volume":"42","author":"Northcutt","year":"2002","journal-title":"Integr. Comp. Biol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1038\/nrg3688","article-title":"Cryptic Genetic Variation: Evolution's Hidden Substrate","volume":"15","author":"Paaby","year":"2014","journal-title":"Nat. Rev. Genet."},{"key":"B26","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/3585.001.0001","volume-title":"How the Body Shapes the Way We Think: A New View of Intelligence","author":"Pfeifer","year":"2006"},{"key":"B27","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1162\/1064546053279017","article-title":"New Robotics: Design Principles for Intelligent Systems","volume":"11","author":"Pfeifer","year":"2005","journal-title":"Artif. Life"},{"key":"B28","doi-asserted-by":"publisher","first-page":"855","DOI":"10.1038\/nrg2452","article-title":"Epistasis - the Essential Role of Gene Interactions in the Structure and Evolution of Genetic Systems","volume":"9","author":"Phillips","year":"2008","journal-title":"Nat. Rev. Genet."},{"key":"B29","doi-asserted-by":"publisher","first-page":"40","DOI":"10.3389\/frobt.2016.00040","article-title":"Quality Diversity: A New Frontier for Evolutionary Computation","volume":"3","author":"Pugh","year":"2016","journal-title":"Front. Robotics AI"},{"key":"B30","first-page":"967","article-title":"Confronting the challenge of Quality Diversity","author":"Pugh","year":"2015"},{"key":"B31","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1111\/ele.13189","article-title":"From Low to High Gear: There Has Been a Paradigm Shift in Our Understanding of Evolution","volume":"22","author":"Reznick","year":"2019","journal-title":"Ecol. Lett."},{"key":"B32","doi-asserted-by":"publisher","first-page":"e201204006","DOI":"10.5936\/csbj.201204006","article-title":"Genetic Code Redundancy and its Influence on the Encoded Polypeptides","volume":"1","author":"Spencer","year":"2012","journal-title":"Comput. Struct. Biotechnol. J."},{"key":"B33","first-page":"214","article-title":"Stochastic Ontogenesis in Evolutionary Robotics","author":"Stanton","year":"2018"},{"key":"B34","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1126\/science.1257008","article-title":"Rapid Evolution of a Native Species Following Invasion by a Congener","volume":"346","author":"Stuart","year":"2014","journal-title":"Science"},{"key":"B35","volume-title":"Probabilistic Robotics","author":"Thrun","year":"2005"},{"key":"B36","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1016\/j.physrep.2012.03.004","article-title":"Collective Motion","volume":"517","author":"Vicsek","year":"2012","journal-title":"Phys. Rep."},{"key":"B37","volume-title":"Robustness and Evolvability in Living Systems","author":"Wagner","year":"2005"},{"key":"B38","first-page":"116","article-title":"The GENITOR Algorithm and Selection Pressure: Why Rank-Based Allocation of Reproductive Trials Is Best","volume":"89","author":"Whitley","year":"","journal-title":"Icga"},{"key":"B39","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1038\/35085569","article-title":"Evolution of Digital Organisms at High Mutation Rates Leads to Survival of the Flattest","volume":"412","author":"Wilke","year":"2001","journal-title":"Nature"},{"key":"B40","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1126\/science.aax1837","article-title":"Cryptic Genetic Variation Accelerates Evolution by Opening Access to Diverse Adaptive Peaks","volume":"365","author":"Zheng","year":"2019","journal-title":"Science"}],"container-title":["Frontiers in Robotics and AI"],"original-title":[],"link":[{"URL":"https:\/\/2.zoppoz.workers.dev:443\/https\/www.frontiersin.org\/articles\/10.3389\/frobt.2021.674823\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,6,10]],"date-time":"2021-06-10T05:50:13Z","timestamp":1623304213000},"score":1,"resource":{"primary":{"URL":"https:\/\/2.zoppoz.workers.dev:443\/https\/www.frontiersin.org\/articles\/10.3389\/frobt.2021.674823\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,6,10]]},"references-count":40,"alternative-id":["10.3389\/frobt.2021.674823"],"URL":"https:\/\/2.zoppoz.workers.dev:443\/https\/doi.org\/10.3389\/frobt.2021.674823","relation":{},"ISSN":["2296-9144"],"issn-type":[{"value":"2296-9144","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,6,10]]},"article-number":"674823"}}