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</style><table class="sidebar sidebar-collapse nomobile nowraplinks" style="border:2px solid #90ee90"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#90ee90;padding:0.2em;font-size:175%;font-weight:bold"><a href="Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></th></tr><tr><td class="sidebar-image"><span class="notpageimage" typeof="mw:File"></span><div class="sidebar-caption"><a href="Darwin's_finches" title="Darwin's finches">Darwin's finches</a> by <a href="John_Gould" title="John Gould">John Gould</a></div></td></tr><tr><td class="sidebar-above" style="display:block; margin-bottom:0.55em; background-color: transparent;">
<div class="hlist">
<ul><li><a href="Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li>
<li><a href="Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li>
<li><a href="Evolution" title="Evolution">Main</a></li>
<li><a href="Outline_of_evolution" title="Outline of evolution">Outline</a></li></ul>
<div class="hlist">
</div>
<ul><li><a href="Glossary_of_evolutionary_biology" class="mw-redirect" title="Glossary of evolutionary biology">Glossary</a></li>
<li><a href="Evidence_of_common_descent" title="Evidence of common descent">Evidence</a></li>
<li><a href="History_of_life" title="History of life">History</a></li></ul>
</div></td></tr><tr><td class="sidebar-content" style="background:transparent;">
<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Processes and outcomes</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist">
<ul><li><a href="Population_genetics" title="Population genetics">Population genetics</a></li>
<li><a href="Genetic_variation" title="Genetic variation">Variation</a></li>
<li><a href="Genetic_diversity" title="Genetic diversity">Diversity</a></li>
<li><a href="Mutation" title="Mutation">Mutation</a></li>
<li><a href="Natural_selection" title="Natural selection">Natural selection</a></li>
<li><a href="Adaptation" title="Adaptation">Adaptation</a></li>
<li><a href="Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li>
<li><a href="Genetic_drift" title="Genetic drift">Genetic drift</a></li>
<li><a href="Gene_flow" title="Gene flow">Gene flow</a></li>
<li><a href="Speciation" title="Speciation">Speciation</a></li>
<li><a href="Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li>
<li><a href="Cooperation_(evolution)" title="Cooperation (evolution)">Co-operation</a></li>
<li><a href="Coevolution" title="Coevolution">Coevolution</a></li>
<li><a href="Coextinction" title="Coextinction">Coextinction</a></li>
<li><a href="Contingency_(evolutionary_biology)" title="Contingency (evolutionary biology)">Contingency</a></li>
<li><a href="Divergent_evolution" title="Divergent evolution">Divergence</a></li>
<li><a href="Convergent_evolution" title="Convergent evolution">Convergence</a></li>
<li><a href="Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li>
<li><a href="Extinction" title="Extinction">Extinction</a></li></ul>
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<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Natural history</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist">
<ul><li><a href="Abiogenesis" title="Abiogenesis">Origin of life</a></li>
<li><a href="Common_descent" title="Common descent">Common descent</a></li>
<li><a href="History_of_life" title="History of life">History of life</a></li>
<li><a href="Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li>
<li><a href="Human_evolution" title="Human evolution">Human evolution</a>
<ul><li><a href="Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li></ul></li>
<li><a href="Phylogenetic_tree" title="Phylogenetic tree">Phylogeny</a></li>
<li><a href="Biodiversity" title="Biodiversity">Biodiversity</a></li>
<li><a href="Biogeography" title="Biogeography">Biogeography</a></li>
<li><a href="Taxonomy_(biology)" title="Taxonomy (biology)">Classification</a></li>
<li><a href="Evolutionary_taxonomy" title="Evolutionary taxonomy">Evolutionary taxonomy</a></li>
<li><a href="Cladistics" title="Cladistics">Cladistics</a></li>
<li><a href="Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li>
<li><a href="Extinction_event" title="Extinction event">Extinction event</a></li></ul>
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<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">History of evolutionary theory</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist">
<ul><li><a href="History_of_evolutionary_thought" title="History of evolutionary thought">Overview</a></li>
<li><a href="Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance</a></li>
<li><a href="Transmutation_of_species" title="Transmutation of species">Before Darwin</a></li>
<li><a href="Charles_Darwin" title="Charles Darwin">Darwin</a></li>
<li><i><a href="On_the_Origin_of_Species" title="On the Origin of Species">Origin of Species</a></i></li>
<li><a href="The_eclipse_of_Darwinism" title="The eclipse of Darwinism">Before synthesis</a></li>
<li><a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li>
<li><a href="History_of_molecular_evolution" title="History of molecular evolution">Molecular evolution</a></li>
<li><a href="Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evo-devo</a></li>
<li><a href="Current_research_in_evolutionary_biology" class="mw-redirect" title="Current research in evolutionary biology">Current research</a></li>
<li><a href="History_of_speciation" title="History of speciation">History of speciation</a></li>
<li><a href="History_of_paleontology" title="History of paleontology">History of paleontology</a> (<a href="Timeline_of_paleontology" title="Timeline of paleontology">timeline</a>)</li></ul>
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</tr><tr><td class="sidebar-content" style="background:transparent;">
<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Fields and applications</div><div class="sidebar-list-content mw-collapsible-content plainlist">
<ul><li><a href="Applications_of_evolution" title="Applications of evolution">Applications of evolution</a></li>
<li><a href="Biosocial_criminology" title="Biosocial criminology">Biosocial criminology</a></li>
<li><a href="Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li>
<li><a href="Evolutionary_aesthetics" title="Evolutionary aesthetics">Evolutionary aesthetics</a></li>
<li><a href="Evolutionary_anthropology" title="Evolutionary anthropology">Evolutionary anthropology</a></li>
<li><a href="Evolutionary_computation" title="Evolutionary computation">Evolutionary computation</a></li>
<li><a href="Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li>
<li><a href="Evolutionary_economics" title="Evolutionary economics">Evolutionary economics</a></li>
<li><a href="Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li>
<li><a href="Evolutionary_ethics" title="Evolutionary ethics">Evolutionary ethics</a></li>
<li><a href="Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li>
<li><a href="Evolutionary_linguistics" title="Evolutionary linguistics">Evolutionary linguistics</a></li>
<li><a href="Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li>
<li><a href="Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li>
<li><a href="Evolutionary_physiology" title="Evolutionary physiology">Evolutionary physiology</a></li>
<li><a href="Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></li>
<li><a href="Invasion_genetics" title="Invasion genetics">Invasion genetics</a></li>
<li><a href="Island_biogeography" class="mw-redirect" title="Island biogeography">Island biogeography</a></li>
<li><a href="Phylogenetics" title="Phylogenetics">Phylogenetics</a></li>
<li><a href="Paleontology" title="Paleontology">Paleontology</a></li>
<li><a href="Selective_breeding" title="Selective breeding">Selective breeding</a></li>
<li><a href="Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Speciation experiments</a></li>
<li><a href="Sociobiology" title="Sociobiology">Sociobiology</a></li>
<li><a href="Systematics" title="Systematics">Systematics</a></li>
<li><a href="Universal_Darwinism" title="Universal Darwinism">Universal Darwinism</a></li></ul></div></div></td>
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<div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Social implications</div><div class="sidebar-list-content mw-collapsible-content plainlist">
<ul><li><a href="Eugenics" title="Eugenics">Eugenics</a></li>
<li><a href="Evolution_as_fact_and_theory" title="Evolution as fact and theory">Evolution as fact and theory</a></li>
<li><a href="Dysgenics" title="Dysgenics">Dysgenics</a></li>
<li><a href="Social_effects_of_evolutionary_theory" title="Social effects of evolutionary theory">Social effects</a></li>
<li><a href="Rejection_of_evolution_by_religious_groups" title="Rejection of evolution by religious groups">Creation–evolution controversy</a></li>
<li><a href="Theistic_evolution" title="Theistic evolution">Theistic evolution</a></li>
<li><a href="Objections_to_evolution" title="Objections to evolution">Objections to evolution</a></li>
<li><a href="Level_of_support_for_evolution" title="Level of support for evolution">Level of support</a></li>
<li><a href="Nature_versus_nurture" title="Nature versus nurture">Nature-nurture controversy</a></li></ul></div></div></td>
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<ul><li><span class="nowrap"><span class="nowrap"><span class="noviewer" typeof="mw:File"></span> </span><a href="Portal%3AEvolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></span></li>
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<p><b>Experimental evolution</b> is the use of laboratory experiments or controlled field manipulations to explore evolutionary dynamics.<sup id="cite_ref-Nature_1-0" class="reference"><a href="#cite_note-Nature-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> <a href="Evolution" title="Evolution">Evolution</a> may be observed in the laboratory as populations adapt to new environmental conditions by <a href="Natural_selection" title="Natural selection">natural selection</a>.
</p><p><a href="Adaptation" title="Adaptation">Adaptation</a> can arise in experimental evolution in two different ways. One is via an individual organism gaining a novel beneficial <a href="Mutation" title="Mutation">mutation</a>.<sup id="cite_ref-:0_2-0" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The other is from <a href="Allele" title="Allele">allele</a> frequency change in standing genetic variation already present in a population of organisms.<sup id="cite_ref-:0_2-1" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Other evolutionary forces outside of mutation and natural selection can also play a role or be incorporated into experimental evolution studies, such as <a href="Genetic_drift" title="Genetic drift">genetic drift</a> and <a href="Gene_flow" title="Gene flow">gene flow</a>.<sup id="cite_ref-:1_3-0" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
</p><p>The organism used is decided by the experimenter, based on the hypothesis to be tested. Many <a href="Generation" title="Generation">generations</a> are required for adaptive mutation to occur, and experimental evolution via mutation is carried out in <a href="Virus" title="Virus">viruses</a> or <a href="Unicellular_organism" title="Unicellular organism">unicellular</a> organisms with rapid generation times, such as <a href="Bacterium" class="mw-redirect" title="Bacterium">bacteria</a> and asexual clonal <a href="Yeast" title="Yeast">yeast</a>.<sup id="cite_ref-Nature_1-1" class="reference"><a href="#cite_note-Nature-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Buckling_4-0" class="reference"><a href="#cite_note-Buckling-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> <a href="Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphic</a> populations of asexual or sexual <a href="Yeast" title="Yeast">yeast</a>,<sup id="cite_ref-:0_2-2" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> and <a href="Multicellular_organism" title="Multicellular organism">multicellular</a> <a href="Eukaryote" title="Eukaryote">eukaryotes</a> like <a href="Drosophila" title="Drosophila">Drosophila</a>, can adapt to new environments through allele frequency change in standing genetic variation.<sup id="cite_ref-:1_3-1" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Organisms with longer generations times, although costly, can be used in experimental evolution. Laboratory studies with foxes<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> and with <a href="Rodents" class="mw-redirect" title="Rodents">rodents</a> (see below) have shown that notable adaptations can occur within as few as 10–20 generations and experiments with wild <a href="Guppy" title="Guppy">guppies</a> have observed adaptations within comparable numbers of generations.<sup id="cite_ref-Reznicketal1997_7-0" class="reference"><a href="#cite_note-Reznicketal1997-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>
</p><p>More recently, experimentally evolved individuals or populations are often analyzed using <a href="Whole_genome_sequencing" title="Whole genome sequencing">whole genome sequencing</a>,<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Jha_AR,_Miles_CM,_Lippert_NR,_Brown_CD,_White_KP,_Kreitman_M_9-0" class="reference"><a href="#cite_note-Jha_AR,_Miles_CM,_Lippert_NR,_Brown_CD,_White_KP,_Kreitman_M-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> an approach known as Evolve and Resequence (E&R).<sup id="cite_ref-Turner_TL,_Stewart_AD,_et_al_10-0" class="reference"><a href="#cite_note-Turner_TL,_Stewart_AD,_et_al-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> E&R can identify mutations that lead to adaptation in clonal individuals or identify alleles that changed in frequency in polymorphic populations, by comparing the sequences of individuals/populations before and after adaptation.<sup id="cite_ref-:0_2-3" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The sequence data makes it possible to pinpoint the site in a <a href="DNA" title="DNA">DNA</a> sequence that a mutation/allele frequency change occurred to bring about adaptation.<sup id="cite_ref-Turner_TL,_Stewart_AD,_et_al_10-1" class="reference"><a href="#cite_note-Turner_TL,_Stewart_AD,_et_al-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Jha_AR,_Miles_CM,_Lippert_NR,_Brown_CD,_White_KP,_Kreitman_M_9-1" class="reference"><a href="#cite_note-Jha_AR,_Miles_CM,_Lippert_NR,_Brown_CD,_White_KP,_Kreitman_M-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_2-4" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The nature of the adaptation and functional follow up studies can shed insight into what effect the mutation/allele has on <a href="Phenotype" title="Phenotype">phenotype</a>.
</p>
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<div class="mw-heading mw-heading2"><h2 id="History">History</h2></div>
<div class="mw-heading mw-heading3"><h3 id="Domestication_and_breeding">Domestication and breeding</h3></div>
<p>Unwittingly, humans have carried out evolution experiments for as long as they have been <a href="Domestication" title="Domestication">domesticating</a> plants and animals. <a href="Selective_breeding" title="Selective breeding">Selective breeding</a> of plants and animals has led to varieties that differ dramatically from their original wild-type ancestors. Examples are the <a href="Cabbage" title="Cabbage">cabbage</a> varieties, <a href="Maize" title="Maize">maize</a>, or the large number of different <a href="Dog" title="Dog">dog</a> breeds. The power of human breeding to create varieties with extreme differences from a single species was already recognized by <a href="Charles_Darwin" title="Charles Darwin">Charles Darwin</a>. In fact, he started out his book <i><a href="The_Origin_of_Species" class="mw-redirect" title="The Origin of Species">The Origin of Species</a></i> with a chapter on variation in domestic animals. In this chapter, Darwin discussed in particular the pigeon.
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</style><blockquote class="templatequote"><p>Altogether at least a score of pigeons might be chosen, which if shown to an ornithologist, and he were told that they were wild birds, would certainly, I think, be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would place the English carrier, the short-faced tumbler, the runt, the barb, pouter, and fantail in the same genus; more especially as in each of these breeds several truly-inherited sub-breeds, or species as he might have called them, could be shown him.
(...) I am fully convinced that the common opinion of naturalists is correct, namely, that all have descended from the rock-pigeon (<i>Columba livia</i>), including under this term several geographical races or sub-species, which differ from each other in the most trifling respects.</p></blockquote><div class="templatequotecite"><p style="display: inline; padding-left: 2.3em;">— <a href="Charles_Darwin" title="Charles Darwin">Charles Darwin</a>, The Origin of Species</p></div>
<div class="mw-heading mw-heading3"><h3 id="Early">Early</h3></div>
<p>One of the first to carry out a controlled evolution experiment was <a href="William_Dallinger" title="William Dallinger">William Dallinger</a>. In the late 19th century, he cultivated small <a href="Unicellular_organism" title="Unicellular organism">unicellular organisms</a> in a custom-built incubator over a time period of seven years (1880–1886). Dallinger slowly increased the temperature of the incubator from an initial 60 °F up to 158 °F. The early cultures had shown clear signs of distress at a temperature of 73 °F, and were certainly not capable of surviving at 158 °F. The organisms Dallinger had in his incubator at the end of the experiment, on the other hand, were perfectly fine at 158 °F. However, these organisms would no longer grow at the initial 60 °F. Dallinger concluded that he had found evidence for Darwinian adaptation in his incubator, and that the organisms had adapted to live in a high-temperature environment. Dallinger's incubator was accidentally destroyed in 1886, and Dallinger could not continue this line of research.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup>
</p><p>From the 1880s to 1980, experimental evolution was intermittently practiced by a variety of evolutionary biologists, including the highly influential <a href="Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a>. Like other experimental research in evolutionary biology during this period, much of this work lacked extensive replication and was carried out only for relatively short periods of evolutionary time.<sup id="cite_ref-Dobzhansky_1957_13-0" class="reference"><a href="#cite_note-Dobzhansky_1957-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Modern">Modern</h2></div>
<p>Experimental evolution has been used in various formats to understand underlying evolutionary processes in a controlled system. Experimental evolution has been performed on multicellular<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> and unicellular<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> eukaryotes, prokaryotes,<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> and viruses.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Similar works have also been performed by <a href="Directed_evolution" title="Directed evolution">directed evolution</a> of individual <a href="Enzyme" title="Enzyme">enzyme</a>,<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> <a href="Ribozyme" title="Ribozyme">ribozyme</a><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> and <a href="Self-replication" title="Self-replication">replicator</a><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> genes.
</p>
<div class="mw-heading mw-heading3"><h3 id="Aphids">Aphids</h3></div>
<p>In the 1950s, the Soviet biologist Georgy Shaposhnikov conducted <a href="https://ru.wikipedia.org/wiki/%D0%9E%D0%BF%D1%8B%D1%82%D1%8B_%D0%93%D0%B5%D0%BE%D1%80%D0%B3%D0%B8%D1%8F_%D0%A8%D0%B0%D0%BF%D0%BE%D1%88%D0%BD%D0%B8%D0%BA%D0%BE%D0%B2%D0%B0_%D0%BF%D0%BE_%D0%B8%D1%81%D0%BA%D1%83%D1%81%D1%81%D1%82%D0%B2%D0%B5%D0%BD%D0%BD%D0%BE%D0%B9_%D1%8D%D0%B2%D0%BE%D0%BB%D1%8E%D1%86%D0%B8%D0%B8" class="extiw external" title="ru:Опыты Георгия Шапошникова по искусственной эволюции">experiments</a> on aphids of the <a href="https://species.wikimedia.org/wiki/Dysaphis" class="extiw external" title="species:Dysaphis">Dysaphis</a> genus. By transferring them to plants normally nearly or completely unsuitable for them, he had forced populations of parthenogenetic descendants to adapt to the new food source to the point of reproductive isolation from the regular populations of the same species.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Fruit_flies">Fruit flies</h3></div>
<p>One of the first of a new wave of experiments using this strategy was the laboratory "evolutionary radiation" of <i><a href="Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i> populations that Michael R. Rose started in February, 1980.<sup id="cite_ref-Rose1984a_24-0" class="reference"><a href="#cite_note-Rose1984a-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> This system started with ten populations, five cultured at later ages, and five cultured at early ages. Since then more than 200 different populations have been created in this laboratory radiation, with selection targeting multiple characters. Some of these highly differentiated populations have also been selected "backward" or "in reverse," by returning experimental populations to their ancestral culture regime. Hundreds of people have worked with these populations over the better part of three decades. Much of this work is summarized in the papers collected in the book <i>Methuselah Flies</i>.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup>
</p><p>The early experiments in flies were limited to studying phenotypes but the molecular mechanisms, i.e., changes in DNA that facilitated such changes, could not be identified. This changed with genomics technology.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Subsequently, Thomas Turner coined the term Evolve and Resequence (E&R)<sup id="cite_ref-Turner_TL,_Stewart_AD,_et_al_10-2" class="reference"><a href="#cite_note-Turner_TL,_Stewart_AD,_et_al-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> and several studies used E&R approach with mixed success.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> One of the more interesting experimental evolution studies was conducted by Gabriel Haddad's group at UC San Diego, where Haddad and colleagues evolved flies to adapt to low oxygen environments, also known as hypoxia.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> After 200 generations, they used E&R approach to identify genomic regions that were selected by natural selection in the hypoxia adapted flies.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> More recent experiments are following up E&R predictions with RNAseq<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> and genetic crosses.<sup id="cite_ref-Jha_AR,_Miles_CM,_Lippert_NR,_Brown_CD,_White_KP,_Kreitman_M_9-2" class="reference"><a href="#cite_note-Jha_AR,_Miles_CM,_Lippert_NR,_Brown_CD,_White_KP,_Kreitman_M-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Such efforts in combining E&R with experimental validations should be powerful in identifying genes that regulate adaptation in flies.
</p><p>Much recently the experimental evolution in flies have taken the course to address the molecular mechanisms<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> and in doing so it might pave way to understand physiology of an organism better and thus redefine disease therapeutics.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Microbes">Microbes</h3></div>
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</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="Serial_passage" title="Serial passage">Serial passage</a></div>
<p>Many microbial species have short <a href="Generation_time" title="Generation time">generation times</a>, easily sequenced genomes, and well-understood biology. They are therefore commonly used for experimental evolution studies. The bacterial species most commonly used for experimental evolution include <i>P. fluorescens</i>,<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> <i>Pseudomonas aeruginosa</i>,<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> <i>Enterococcus faecalis</i> <sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> and <i>E. coli</i> (see below), while the Yeast <i>S. cerevisiae</i> has been used as a model for the study of eukaryotic evolution.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading4"><h4 id="Lenski's_E._coli_experiment">Lenski's <i>E. coli</i> experiment</h4></div>
<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="E._coli_long-term_evolution_experiment" title="E. coli long-term evolution experiment">E. coli long-term evolution experiment</a></div>
<p>One of the most widely known examples of laboratory bacterial evolution is the <a href="E._coli_long-term_evolution_experiment" title="E. coli long-term evolution experiment">long-term <i>E.coli</i> experiment</a> of <a href="Richard_Lenski" title="Richard Lenski">Richard Lenski</a>. On February 24, 1988, Lenski started growing twelve lineages of <i><a href="E._coli" class="mw-redirect" title="E. coli">E. coli</a></i> under identical growth conditions.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> When one of the populations evolved the ability to aerobically metabolize citrate from the growth medium and showed greatly increased growth,<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> this provided a dramatic observation of evolution in action. The experiment continues to this day, and is now the longest-running (in terms of generations) controlled evolution experiment ever undertaken. Since the inception of the experiment, the bacteria have grown for more than 60,000 generations. Lenski and colleagues regularly publish updates on the status of the experiments.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Leishmania_donovani"><i>Leishmania donovani</i></h3></div>
<p>Bussotti and collaborators isolated amastigotes from <i><a href="Leishmania_donovani" title="Leishmania donovani">Leishmania donovani</a></i> and cultured them in vitro for 3800 generations (36 weeks). The culture of these parasites showed how they adapted to in vitro conditions by compensating for the loss of a <a href="NIMA-related_kinase_1" title="NIMA-related kinase 1">NIMA-related kinase</a>, important for the correct progression of mitosis, by increasing the expression of another orthologous kinase as the culture generations progressed. Furthermore, it was observed how <i>L. donovani</i> has been adapted to in vitro culture by reducing the expression of 23 transcripts related to <a href="Flagellum" title="Flagellum">flagellar biogenesis</a> and increasing the expression of ribosomal protein clusters and non-coding RNAs such as <a href="Small_nucleolar_RNA" title="Small nucleolar RNA">nucleolar small RNAs</a>. Flagella are considered less necessary by the parasite in in vitro culture and therefore the progression of generations leads to their elimination, causing an energy saving due to lower motility so that proliferation and growth rate in culture is higher. The amplified snoRNAs also lead to increased ribosomal biosynthesis, increased protein biosynthesis and thus increased growth rate of the culture. These adaptations observed over generations of parasites are governed by <a href="Copy_number_variation" title="Copy number variation">copy number variations (CNV)</a> and <a href="Epistasis" title="Epistasis">epistatic interactions</a> between affected genes, and allow us to justify <i>Leishmania</i> <a href="Genome_instability" title="Genome instability">genomic instability</a> through its post-transcriptional regulation of gene expression.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="High_Runner_laboratory_house_mice">High Runner laboratory house mice</h3></div>
<p>In 1993, <a href="Theodore_Garland%2C_Jr." class="mw-redirect" title="Theodore Garland, Jr.">Theodore Garland, Jr.</a> and colleagues started a long-term experiment that involves selective breeding of <a href="Laboratory_mouse" title="Laboratory mouse">mice</a> for high voluntary activity levels on running wheels.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> This experiment also continues to this day (> 105 <a href="Generation" title="Generation">generations</a>). Mice from the four replicate "High Runner" lines evolved to run almost three times as many running-wheel revolutions per day compared with the four unselected control lines of mice, mainly by running faster than the control mice rather than running for more minutes/day. However, the High Runner lines have evolved in somewhat different ways, with some emphasizing running speed versus duration or vice versa, thus demonstrating "multiple solutions"
<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup>
that seem to be based partly in evolved muscle characteristics.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup>
</p>
<p>The HR mice have an elevated endurance running ability
<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup>
and <a href="VO2max" class="mw-redirect" title="VO2max">maximal aerobic capacity</a>
<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> when tested on a motorized treadmill. They also exhibit alterations in <a href="Motivation" title="Motivation">motivation</a> and the <a href="Reward_system" title="Reward system">reward system</a> of the <a href="Brain" title="Brain">brain</a>. <a href="Pharmacological" class="mw-redirect" title="Pharmacological">Pharmacological</a> studies point to alterations in <a href="Dopamine" title="Dopamine">dopamine</a> function and the <a href="Endocannabinoid_system" title="Endocannabinoid system">endocannabinoid system</a>.<sup id="cite_ref-Keeneyetal2008_49-0" class="reference"><a href="#cite_note-Keeneyetal2008-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> The High Runner lines have been proposed as a model to study human attention-deficit hyperactivity disorder (<a href="ADHD" class="mw-redirect" title="ADHD">ADHD</a>), and administration of <a href="Ritalin" class="mw-redirect" title="Ritalin">Ritalin</a> reduces their wheel running approximately to the levels of control mice.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Multidirectional_selection_on_bank_voles">Multidirectional selection on bank voles</h3></div>
<p>In 2005 Paweł Koteja with Edyta Sadowska and colleagues from the <a href="Jagiellonian_University" title="Jagiellonian University">Jagiellonian University</a> (Poland) started a multidirectional selection on a non-laboratory rodent, the <a href="Bank_vole" title="Bank vole">bank vole</a> <i>Myodes (= Clethrionomys) glareolus</i>.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> The voles are selected for three distinct traits, which played important roles in the <a href="Adaptive_radiation" title="Adaptive radiation">adaptive radiation</a> of terrestrial vertebrates: high maximum rate of aerobic metabolism, predatory propensity, and herbivorous capability. Aerobic lines are selected for the maximum rate of oxygen consumption achieved during swimming at 38°C; Predatory lines – for a short time to catch live <a href="Crickets" class="mw-redirect" title="Crickets">crickets</a>; <a href="Herbivore" title="Herbivore"> Herbivorous</a> lines – for capability to maintain body mass when fed a low-quality diet “diluted” with dried, powdered grass. Four replicate lines are maintained for each of the three selection directions and another four as unselected Controls.
</p><p>After approximately 20 generations of selective breeding, voles from the Aerobic lines evolved a 60% higher swim-induced metabolic rate than voles from the unselected Control lines. Although the selection protocol does not impose a thermoregulatory burden, both the <a href="Basal_metabolic_rate" title="Basal metabolic rate">basal metabolic rate</a> and <a href="Thermogenesis" title="Thermogenesis"> thermogenic</a> capacity increased in the Aerobic lines.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> Thus, the results have provided some support for the “aerobic capacity model” for the evolution of <a href="Endothermy" class="mw-redirect" title="Endothermy">endothermy</a> in mammals.
</p><p>More than 85% of the Predatory voles capture the crickets, compared to only about 15% of unselected Control voles, and they catch the crickets faster. The increased predatory behavior is associated with a more proactive <a href="Coping" title="Coping"> coping style</a> (“<a href="Personality" title="Personality">personality</a>”).<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup>
</p><p>During the test with low-quality diet, the Herbivorous voles lose approximately 2 grams less mass (approximately 10% of the original body mass) than the Control ones. The Herbivorous voles have an altered composition of the bacterial <a href="Microbiota" title="Microbiota"> microbiome</a> in their <a href="Caecum" class="mw-redirect" title="Caecum">caecum</a>.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Thus, the selection has resulted in evolution of the entire holobiome, and the experiment may offer a laboratory model of <a href="Hologenome_theory_of_evolution" title="Hologenome theory of evolution"> hologenome evolution</a>.
</p>
<div class="mw-heading mw-heading3"><h3 id="Synthetic_biology">Synthetic biology</h3></div>
<p><a href="Synthetic_biology" title="Synthetic biology">Synthetic biology</a> offers unique opportunities for experimental evolution, facilitating the interpretation of evolutionary changes by inserting genetic modules into host genomes and applying selection specifically targeting such modules. <a href="Synthetic_biological_circuits" class="mw-redirect" title="Synthetic biological circuits">Synthetic biological circuits</a> inserted into the genome of <a href="Escherichia_coli" title="Escherichia coli">Escherichia coli</a><sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> or the budding yeast <a href="Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a><sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> degrade (lose function) during laboratory evolution. With appropriate selection, mechanisms underlying the evolutionary regain of lost biological function can be studied.<sup id="cite_ref-KheirGouda_2019_58-0" class="reference"><a href="#cite_note-KheirGouda_2019-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> Experimental evolution of mammalian cells harboring synthetic gene circuits<sup id="cite_ref-Farquhar_2019_59-0" class="reference"><a href="#cite_note-Farquhar_2019-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> reveals the role of cellular heterogeneity in the evolution of drug resistance, with implications for <a href="Chemotherapy" title="Chemotherapy">chemotherapy</a> resistance of cancer cells.
</p>
<div class="mw-heading mw-heading3"><h3 id="Other_examples">Other examples</h3></div>
<p><a href="Stickleback" title="Stickleback">Stickleback</a> fish have both marine and freshwater species, the freshwater species evolving since the last ice age. Freshwater species can survive colder temperatures. Scientists tested to see if they could reproduce this evolution of cold-tolerance by keeping marine sticklebacks in cold freshwater. It took the marine sticklebacks only three generations to evolve to match the 2.5 degree Celsius improvement in cold-tolerance found in wild freshwater sticklebacks.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup>
</p><p>Microbial cells <sup id="cite_ref-DragositsMattanovich2013_61-0" class="reference"><a href="#cite_note-DragositsMattanovich2013-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> and recently mammalian cells <sup id="cite_ref-MaralingannavarParmar2017_62-0" class="reference"><a href="#cite_note-MaralingannavarParmar2017-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> are evolved under nutrient limiting conditions to study their metabolic response and engineer cells for useful characteristics.
</p>
<div class="mw-heading mw-heading2"><h2 id="For_teaching">For teaching</h2></div>
<p>Because of their rapid generation times microbes offer an opportunity to study <a href="Microevolution" title="Microevolution">microevolution</a> in the classroom. A number of exercises involving bacteria and yeast teach concepts ranging from the evolution of resistance<sup id="cite_ref-Hyman2014_63-0" class="reference"><a href="#cite_note-Hyman2014-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> to the evolution of multicellularity.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> With the advent of next-generation sequencing technology it has become possible for students to conduct an evolutionary experiment, sequence the evolved genomes, and to analyze and interpret the results.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="For_industrial_use">For industrial use</h2></div>
<p>Laboratory synthetic evolution, not necessarily directed, is also used to develop new phenotypes in organisms used for <a href="Biomanufacturing" title="Biomanufacturing">biomanufacturing</a>. Natural mutations can be complemented with methods that directly generate phenotypic diversity. With yeast, the generation of new phenotypes can be accelerated by manipulation at the transcription level: by stochastically introducing artificial transcription factors, globally altering the transcription machinery, CRISPR interference and activation. Also with yeast, transposons, loxP sequences, and highly error-prone orthogonal DNAP-DNA plasmid pairs can be used to accelerate the generation of mutations.<sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div>
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<ul><li><a href="Artificial_selection" class="mw-redirect" title="Artificial selection">Artificial selection</a></li>
<li><a href="Bacteriophage_experimental_evolution" title="Bacteriophage experimental evolution">Bacteriophage experimental evolution</a></li>
<li><a href="Directed_evolution" title="Directed evolution">Directed evolution</a></li>
<li><a href="Domestication" title="Domestication">Domestication</a></li>
<li><a href="Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li>
<li><a href="Evolutionary_physiology" title="Evolutionary physiology">Evolutionary physiology</a></li>
<li><a href="Genetics" title="Genetics">Genetics</a></li>
<li><a href="Genomics_of_domestication" class="mw-redirect" title="Genomics of domestication">Genomics of domestication</a></li>
<li><a href="Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Laboratory experiments of speciation</a></li>
<li><a href="Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li>
<li><a href="Selection_limits" title="Selection limits">Selection limits</a></li>
<li><a href="Selective_breeding" title="Selective breeding">Selective breeding</a></li>
<li><a href="Tame_Silver_Fox" class="mw-redirect" title="Tame Silver Fox">Tame Silver Fox</a></li></ul>
</div>
<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>
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<li id="cite_note-Hyman2014-63"><span class="mw-cite-backlink"><b><a href="#cite_ref-Hyman2014_63-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFHyman2014" class="citation journal cs1">Hyman P (January 2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3895413">"Bacteriophage as instructional organisms in introductory biology labs"</a>. <i>Bacteriophage</i>. <b>4</b> (1): e27336. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.4161%2Fbact.27336">10.4161/bact.27336</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3895413">3895413</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24478938">24478938</a>.</cite></span>
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<li id="cite_note-64"><span class="mw-cite-backlink"><b><a href="#cite_ref-64">^</a></b></span> <span class="reference-text"><cite id="CITEREFRatcliffRaneyWestreichCotner2014" class="citation journal cs1">Ratcliff WC, Raney A, Westreich S, Cotner S (2014). "A Novel Laboratory Activity for Teaching about the Evolution of Multicellularity". <i>The American Biology Teacher</i>. <b>76</b> (2): <span class="nowrap">81–</span>87. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1525%2Fabt.2014.76.2.3">10.1525/abt.2014.76.2.3</a>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0002-7685">0002-7685</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:86079463">86079463</a>.</cite></span>
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<li id="cite_note-65"><span class="mw-cite-backlink"><b><a href="#cite_ref-65">^</a></b></span> <span class="reference-text"><cite id="CITEREFMikheyevArora2015" class="citation report cs1">Mikheyev, Alexander S; Arora, Jigyasa (9 September 2015). Using experimental evolution and next-generation sequencing to teach bench and bioinformatic skills (Preprint). <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.7287%2Fpeerj.preprints.1356v1">10.7287/peerj.preprints.1356v1</a></span>.</cite></span>
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<li id="cite_note-66"><span class="mw-cite-backlink"><b><a href="#cite_ref-66">^</a></b></span> <span class="reference-text"><cite id="CITEREFWangQiRenLin2025" class="citation journal cs1">Wang, Zhen; Qi, Xianni; Ren, Xinru; Lin, Yuping; Zeng, Fanli; Wang, Qinhong (February 2025). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC11868838">"Synthetic evolution of Saccharomyces cerevisiae for biomanufacturing: Approaches and applications"</a>. <i>mLife</i>. <b>4</b> (1): <span class="nowrap">1–</span>16. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fmlf2.12167">10.1002/mlf2.12167</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC11868838">11868838</a></span>.</cite></span>
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<div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2></div>
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<ul><li><cite id="CITEREFBennett2003" class="citation journal cs1">Bennett AF (2003). <a rel="nofollow" class="external text" href="https://zenodo.org/record/1059074">"Experimental evolution and the Krogh principle: generating biological novelty for functional and genetic analyses"</a>. <i>Physiological and Biochemical Zoology</i>. <b>76</b> (1): <span class="nowrap">1–</span>11. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1086%2F374275">10.1086/374275</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12695982">12695982</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:9032244">9032244</a>.</cite></li>
<li><cite id="CITEREFDallinger1887" class="citation journal cs1">Dallinger WH (April 1887). "The president's address". <i>Journal of the Royal Microscopical Society</i>. <b>7</b> (2): <span class="nowrap">185–</span>99. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1365-2818.1887.tb01566.x">10.1111/j.1365-2818.1887.tb01566.x</a>.</cite></li>
<li><cite id="CITEREFGarland_Jr2003" class="citation book cs1"><a href="Theodore_Garland%2C_Jr." class="mw-redirect" title="Theodore Garland, Jr.">Garland Jr T</a> (2003). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20150923190230/http://www.biology.ucr.edu/people/faculty/Garland/Garland_2003.pdf">"Selection experiments: an under-utilized tool in biomechanics and organismal biology."</a> <span class="cs1-format">(PDF)</span>. In Bels VL, Gasc JP, Casinos A (eds.). <i>Vertebrate biomechanics and evolution</i>. Oxford, UK: <a href="BIOS_Scientific_Publishers" title="BIOS Scientific Publishers">BIOS Scientific Publishers</a>. pp. <span class="nowrap">23–</span>56. Archived from <a rel="nofollow" class="external text" href="http://www.biology.ucr.edu/people/faculty/Garland/Garland_2003.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 2015-09-23<span class="reference-accessdate">. Retrieved <span class="nowrap">2007-02-10</span></span>.</cite></li>
<li><cite id="CITEREFGarland_JrRose2009" class="citation book cs1">Garland Jr T, Rose MR, eds. (2009). <a rel="nofollow" class="external text" href="https://www.ucpress.edu/book/9780520261808/experimental-evolution"><i>Experimental evolution: concepts, methods, and applications of selection experiments</i></a>. Berkeley, California: University of California Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-520-26180-8</bdi>.</cite></li>
<li><cite id="CITEREFGibbs1999" class="citation journal cs1">Gibbs AG (October 1999). "Laboratory selection for the comparative physiologist". <i>The Journal of Experimental Biology</i>. <b>202</b> (Pt 20): <span class="nowrap">2709–</span>2718. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1242%2Fjeb.202.20.2709">10.1242/jeb.202.20.2709</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/10504307">10504307</a>.</cite></li>
<li><cite id="CITEREFLenski2003" class="citation book cs1">Lenski, Richard E. (2003). "Phenotypic and Genomic Evolution during a 20,000-Generation Experiment with the Bacterium <i>Escherichia coli</i>". <i>Plant Breeding Reviews</i>. pp. <span class="nowrap">225–</span>265. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2F9780470650288.ch8">10.1002/9780470650288.ch8</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-471-46892-9</bdi>.</cite></li>
<li><cite id="CITEREFLenskiRoseSimpsonTadler1991" class="citation journal cs1">Lenski RE, Rose MR, Simpson SC, Tadler SC (1991). "Long-term experimental evolution in <i>Escherichia coli</i>. I. Adaptation and divergence during 2,000 generations". <i>American Naturalist</i>. <b>138</b> (6): <span class="nowrap">1315–</span>1341. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1086%2F285289">10.1086/285289</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:83996233">83996233</a>.</cite></li>
<li><cite id="CITEREFMcKenzieBatterham1994" class="citation journal cs1">McKenzie JA, Batterham P (May 1994). "The genetic, molecular and phenotypic consequences of selection for insecticide resistance". <i>Trends in Ecology & Evolution</i>. <b>9</b> (5): <span class="nowrap">166–</span>169. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1994TEcoE...9..166M">1994TEcoE...9..166M</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2F0169-5347%2894%2990079-5">10.1016/0169-5347(94)90079-5</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/21236810">21236810</a>.</cite></li>
<li><cite id="CITEREFReznickBryantRoffGhalambor2004" class="citation journal cs1">Reznick DN, Bryant MJ, Roff D, Ghalambor CK, Ghalambor DE (October 2004). "Effect of extrinsic mortality on the evolution of senescence in guppies". <i>Nature</i>. <b>431</b> (7012): <span class="nowrap">1095–</span>1099. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2004Natur.431.1095R">2004Natur.431.1095R</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnature02936">10.1038/nature02936</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15510147">15510147</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:205210169">205210169</a>.</cite></li>
<li><cite id="CITEREFRosePassanantiMatos2004" class="citation book cs1">Rose MR, Passananti HB, Matos M, eds. (2004). <i>Methuselah flies: A case study in the evolution of aging</i>. Singapore: World Scientific Publishing.</cite></li>
<li><cite id="CITEREFSwallowGarland2005" class="citation journal cs1">Swallow JG, <a href="Theodore_Garland%2C_Jr." class="mw-redirect" title="Theodore Garland, Jr.">Garland T</a> (June 2005). <a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Ficb%2F45.3.387">"Selection Experiments as a Tool in Evolutionary and Comparative Physiology: Insights into Complex Traits--an Introduction to the Symposium"</a>. <i>Integrative and Comparative Biology</i>. <b>45</b> (3): <span class="nowrap">387–</span>390. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Ficb%2F45.3.387">10.1093/icb/45.3.387</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/21676784">21676784</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:2305227">2305227</a>.</cite></li></ul>
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<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>
<ul><li><a rel="nofollow" class="external text" href="http://myxo.css.msu.edu/ecoli/">E. coli Long-term Experimental Evolution Project Site</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170727225642/http://myxo.css.msu.edu/ecoli/">Archived</a> 2017-07-27 at the <a href="Wayback_Machine" title="Wayback Machine">Wayback Machine</a>, Lenski lab, <a href="Michigan_State_University" title="Michigan State University">Michigan State University</a></li>
<li>A <a rel="nofollow" class="external text" href="http://www.biology.ucr.edu/people/faculty/Garland/Girard01.mov">movie</a> illustrating the dramatic differences in wheel-running behavior.</li>
<li><a rel="nofollow" class="external text" href="https://sites.google.com/ucr.edu/hrmice/publications">Experimental Evolution Publications by Ted Garland: Artificial Selection for High Voluntary Wheel-Running Behavior in House Mice</a> — a detailed list of publications.</li>
<li><a rel="nofollow" class="external text" href="http://biology.ucr.edu/people/faculty/Garland/ExperimentalEvolution.html">Experimental Evolution</a> — a list of laboratories that study experimental evolution.</li>
<li><a rel="nofollow" class="external text" href="http://nere.bio.uci.edu/">Network for Experimental Research on Evolution</a>, <a href="University_of_California" title="University of California">University of California</a>.</li>
<li><cite id="CITEREFNicholls2009" class="citation news cs1">Nicholls, Henry (30 September 2009). <a rel="nofollow" class="external text" href="https://www.newscientist.com/article/mg20427281-500-my-little-zebra-the-secrets-of-domestication/">"My little zebra: The secrets of domestication"</a>. <i>New Scientist</i>.</cite></li>
<li><a href="Inquiry-based_learning" title="Inquiry-based learning">Inquiry-based</a> <a rel="nofollow" class="external text" href="https://biol.ucr.edu/idea/born_to_run/Born_to_Run.html">middle school lesson plan: "Born to Run: Artificial Selection Lab"</a></li>
<li><a rel="nofollow" class="external text" href="http://avida-ed.msu.edu/">Digital Evolution for Education software</a></li></ul></div><!--htdig_noindex--><div><div class="zim-footer">
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