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Genetic engineering | Definition, Process, Uses, Examples, Techniques, & Facts | Britannica

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data-level="h1"><li data-target="#ref1"><div class="pl-25"><a class="link-gray-900 w-100" href="/science/genetic-engineering">Introduction</a></div><div class="ml-40 toc-drawer sub-toc-drawer"></div></li><li data-target="#ref337079"><div class="d-flex align-items-center"><div class="ml-25"></div><a class="w-100 link-gray-900" href="/science/genetic-engineering#ref337079">Historical developments</a></div><div class="ml-40 toc-drawer sub-toc-drawer"></div></li><li data-target="#ref337080"><div class="d-flex align-items-center"><div class="ml-25"></div><a class="w-100 link-gray-900" href="/science/genetic-engineering/Process-and-techniques">Process and techniques</a></div><div class="ml-40 toc-drawer sub-toc-drawer"></div></li><li data-target="#ref337081"><div class="d-flex align-items-center"><div class="ml-25"></div><a class="w-100 link-gray-900" href="/science/genetic-engineering/Process-and-techniques#ref337081">Applications</a></div><div class="ml-40 toc-drawer sub-toc-drawer"></div></li><li data-target="#ref337082"><div class="d-flex align-items-center"><div class="ml-25"></div><a class="w-100 link-gray-900" href="/science/genetic-engineering/Process-and-techniques#ref337082">Controversy and ethical issues</a></div><div class="ml-40 toc-drawer sub-toc-drawer"></div></li></ul> <a class="toc-extra-link link-gray-900" href="https://www.britannica.com/science/genetic-engineering/additional-info">References &amp; Edit History</a> <a class="toc-extra-link link-gray-900" href="/facts/genetic-engineering">Related Topics</a> </div> <div class="tlr-media-slider pb-10 mb-30"> <a class="section-header link-gray-900 font-serif font-14 font-weight-bold mb-10 mx-10" href="https://www.britannica.com/science/genetic-engineering/images-videos">Images & Videos</a> <div class="slider js-slider position-relative d-inline-flex align-items-center mw-100 "> <div class="slider-container js-slider-container overflow-hidden d-flex overflow-hidden text-nowrap ml-15"> 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</a> <a href="https://cdn.britannica.com/61/131361-050-1A1D205E/corn.jpg" data-href="/media/1/228897/128155" class="media-overlay-link d-inline-block mr-5"> <img loading="lazy" src="https://cdn.britannica.com/61/131361-004-07890150/corn.jpg" alt="Genetically engineered corn (maize)" height="50" /> </a> <a href="https://cdn.britannica.com/41/246541-050-8D5BA567/Golden-rice-GMO-Vitamin-A.jpg" data-href="/media/1/228897/292785" class="media-overlay-link d-inline-block mr-5"> <img loading="lazy" src="https://cdn.britannica.com/41/246541-004-B3F359D3/Golden-rice-GMO-Vitamin-A.jpg" alt="golden rice" height="50" /> </a> <a href="https://cdn.britannica.com/87/22487-050-739115AC/DNA-fragments-genomics-research-cells-vector-replication.jpg" data-href="/media/1/228897/17930" class="media-overlay-link d-inline-block mr-5"> <img loading="lazy" src="https://cdn.britannica.com/87/22487-004-16BC016E/DNA-fragments-genomics-research-cells-vector-replication.jpg" alt="Recombinant DNA" height="50" /> </a> 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School of Medicine - Gene Therapy and Genetic Engineering</a></li> <li><a class="external" href="https://www.eolss.net/sample-chapters/c17/E6-58-03-04.pdf" target="_blank" rel="noopener ">UNESCO-EOLSS - Genetic Engineering of Plants (PDF)</a></li> <li><a class="external" href="https://pmc.ncbi.nlm.nih.gov/articles/PMC3078015/" target="_blank" rel="noopener ">National Center for Biotechnology Information - PubMed Central - Genetic engineering of animals: Ethical issues, including welfare concerns</a></li> <li><a class="external" href="https://www.livescience.com/32648-whats-genetic-engineering.html" target="_blank" rel="noopener ">LiveScience - What's Genetic Engineering?</a></li> <li><a class="external" href="https://iastate.pressbooks.pub/genagbiotech/chapter/genetic-engineering/" target="_blank" rel="noopener ">Iowa State University Digital Press - Genetic Engineering</a></li> <li><a class="external" href="https://bio.libretexts.org/Bookshelves/Introductory_and_General_Biology/Principles_of_Biology/02%3A_Chapter_2/20%3A_Biotechnology/20.03%3A_Genetic_Engineering" target="_blank" rel="noopener ">Biology LibreTexts - Genetic Engineering</a></li> <li><a class="external" href="https://english.umd.edu/research-innovation/journals/interpolations/interpolations-spring-2011/genetic-engineering-serious" target="_blank" rel="noopener ">University of Marland - Department of English - Interpolations - Genetic Engineering: A Serious Threat to Human Society</a></li> <li><a class="external" href="https://agricultureandfoodsecurity.biomedcentral.com/articles/10.1186/2048-7010-2-15" target="_blank" rel="noopener ">BMC - Agriculture and Food Security - Genetic engineering for improving quality and productivity of crops</a></li> <li><a class="external" href="https://www.khanacademy.org/science/high-school-biology/hs-molecular-genetics/hs-biotechnology/v/introduction-to-genetic-engineering" target="_blank" rel="noopener ">Khan Academy - Introduction to genetic engineering</a></li> <li><a class="external" href="https://openbooks.lib.msu.edu/isb202/chapter/biotechnology/" target="_blank" rel="noopener ">OER at MSU Libraries - An Interactive Introduction to Organismal and Molecular Biology, 2nd ed. - Genetic Engineering</a></li> </ul> </div> <div class="md-websites-ebk-title">Britannica Websites</div> <div class="md-websites-ebk-subtitle">Articles from Britannica Encyclopedias for elementary and high school students.</div> <ul class="list-unstyled bps-topic-web-sites lh-lg"> <li><a class="external" href="https://kids.britannica.com/kids/article/genetic-engineering/600760" target="_blank" rel="noopener">genetic engineering - Children's Encyclopedia (Ages 8-11)</a></li> <li><a class="external" href="https://kids.britannica.com/students/article/genetic-engineering/274518" target="_blank" rel="noopener">genetic engineering - Student Encyclopedia (Ages 11 and up)</a></li> </ul> </div> </div> </div> </div> <div class="toc-header-marker"></div> <button class="ai-ask-button btn border-2 js-header-ai-ask-button d-none btn-sm btn-outline-red-400 border-red-400 mr-0 mr-lg-10 ml-5 ml-sm-10 ml-lg-0 p-10"> Ask the Chatbot a Question </button> <div class="md-byline module-spacing "> <div class="font-serif font-12"> <span class="written-by text-gray-700"> Written and fact-checked by </span> <div class="editor-popover popover p-0"> <a class="d-block p-20 qa-editor-popup gtm-byline font-12 byline-contributor" href="/editor/The-Editors-of-Encyclopaedia-Britannica/4419" > <div class="editor-title font-16 font-weight-bold">The Editors of Encyclopaedia Britannica</div> <div class="editor-description font-12 font-serif mt-5 text-black">Encyclopaedia Britannica's editors oversee subject areas in which they have extensive knowledge, whether from years of experience gained by working on that content or via study for an advanced degree. They write new content and verify and edit content received from contributors.</div> </a> <div data-popper-arrow></div> </div> <span class="btn btn-link editor-link p-0 qa-byline-link gtm-byline font-12 byline-contributor text-decoration-underline"> The Editors of Encyclopaedia Britannica</span></div> <div class="last-updated font-12 font-serif"> <span class="text-gray-700"> Last Updated: <time datetime="2025-02-15T00:00:00CST" >Feb 15, 2025</time> •</span> <a class="byline-edit-history" href="https://www.britannica.com/science/genetic-engineering/additional-info#history" rel="nofollow">Article History</a> </div></div> </div> <button class="d-flex d-lg-none btn btn-outline-blue border rounded-sm shadow-sm mobile-toc-button gtm-mobile-toc-inline-button d-none d-sm-block js-sections-inline-button module-spacing btn d-lg-none"> <em class="material-icons mr-5 ml-n10 my-n5 md-icon" data-icon="toc"></em> Table of Contents </button> <div class="d-flex d-sm-none flex-row"> <button class="d-flex d-lg-none btn btn-outline-blue border rounded-sm shadow-sm mobile-toc-button gtm-mobile-toc-inline-button js-sections-inline-button module-spacing"> <em class="material-icons mr-5 ml-n10 my-n5 md-icon" data-icon="toc"></em> Table of Contents </button> <button class="ai-ask-button btn border-2 ai-ask-button btn border-2 module-spacing btn-sm js-inline-ai-ask-button btn-outline-red-400 border-red-400 p-10 ml-5"> Ask the Chatbot a Question </button> </div> <div class="js-qf-module qf-module px-40 px-sm-20 py-15 mx-auto module-spacing font-14 bg-gray-50 rounded"> <div class="facts-list mt-10"> <div class=""> <div class="js-fact mb-10 line-clamp clamp-3"> <dl> <dt>Key People: </dt> <dd><a href="/biography/Ian-Wilmut" topicid="644616">Ian Wilmut</a></dd> <dd><a href="/biography/George-Ledyard-Stebbins-Jr" topicid="564612">George Ledyard Stebbins, Jr.</a></dd> </dl> <button class="js-more-btn d-none btn btn-unstyled font-12 bg-gray-50" aria-label="Toggle more/less fact data"> <em class="js-content link-blue">(Show&nbsp;more)</em> </button> </div> </div> <div class=""> <div class="js-fact mb-10 line-clamp clamp-3"> <dl> <dt>Related Topics: </dt> <dd><a href="/science/recombinant-DNA-technology" topicid="493667">recombinant DNA</a></dd> <dd><a href="/science/genetically-modified-organism" topicid="897705">genetically modified organism</a></dd> <dd><a href="/science/gene-editing" topicid="1998073">gene editing</a></dd> <dd><a href="/science/three-parent-baby" topicid="2002944">three-parent baby</a></dd> <dd><a href="/science/whole-genome-sequencing" topicid="1866136">whole genome sequencing</a></dd> </dl> <button class="js-more-btn d-none btn btn-unstyled font-12 bg-gray-50" aria-label="Toggle more/less fact data"> <em class="js-content link-blue">(Show&nbsp;more)</em> </button> </div> </div> <div class=""> <div class="js-fact mb-10 line-clamp clamp-3"> <dl> <dt>On the Web: </dt> <dd><a href="https://www.eolss.net/sample-chapters/c17/E6-58-03-04.pdf" target="_blank">UNESCO-EOLSS - Genetic Engineering of Plants (PDF)</a> (Feb. 15, 2025)</dd> </dl> <button class="js-more-btn d-none btn btn-unstyled font-12 bg-gray-50" aria-label="Toggle more/less fact data"> <em class="js-content link-blue">(Show&nbsp;more)</em> </button> </div> <div class="text-center"> <a class="btn btn-sm btn-link p-0" href="/facts/genetic-engineering"> See all related content </a> </div> </div> </div> </div><div class="ai-top-questions-module module-spacing ai-tq collapsed font-sans-serif"> <div class="font-weight-bold font-14"> Top Questions </div> <ul class="mt-5"> <li class="ai-qa-item mb-5 bg-gray-50 rounded "> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="0"> <div>What is genetic engineering?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded "> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="1"> <div>How does genetic engineering differ from traditional breeding methods?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded "> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="2"> <div>What are the main techniques used in genetic engineering?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded af3"> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="3"> <div>How do scientists alter the DNA of an organism?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded af3"> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="4"> <div>What are some of the goals of genetic engineering in agriculture?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded af3"> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="5"> <div>What role does genetic engineering play in medicine?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded af3"> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="6"> <div>What are the ethical concerns associated with genetic engineering?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded af3"> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="7"> <div>How is genetic engineering regulated by governments and organizations?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> <li class="ai-qa-item mb-5 bg-gray-50 rounded af3"> <div class="ai-top-question justify-content-between align-items-center font-14 px-10 py-5 rounded " data-qn="8"> <div>What are some real-world examples where genetic engineering has been applied successfully?</div> </div> <div class="answer p-10 ui-hidden" data-loaded="false"></div> <div class="ai-loader ui-hidden" /> </li> </ul> <button class="show-more btn btn-unstyled font-14 p-5 pr-10 mb-10 rounded-sm d-none"><em class="material-icons " data-icon="expand_more"></em> Show more</button> <button class="show-less btn btn-unstyled font-14 p-5 pr-10 mb-10 rounded-sm"><em class="material-icons " data-icon="expand_less"></em> Show less</button> </div><!--[BEFORE-ARTICLE]--><span class="marker before-article"></span><section data-level="1" id="ref1"><!--[PREMOD1]--><span class="marker PREMOD1 mod-inline"></span><p class="topic-paragraph"><strong><span id="ref233967"></span>genetic engineering</strong>, the artificial manipulation, modification, and recombination of <a href="https://www.britannica.com/science/DNA" class="md-crosslink " data-show-preview="true">DNA</a> or other <a href="https://www.britannica.com/science/nucleic-acid" class="md-crosslink " data-show-preview="true">nucleic acid</a> <a href="https://www.britannica.com/science/molecule" class="md-crosslink " data-show-preview="true">molecules</a> in order to modify an <a href="https://www.britannica.com/science/multicellular-organism" class="md-crosslink " data-show-preview="true">organism</a> or <a href="https://www.britannica.com/science/population-biology-and-anthropology" class="md-crosslink " data-show-preview="true">population</a> of organisms. The term <em>genetic engineering</em> is generally used to refer to methods of <a href="https://www.britannica.com/science/recombinant-DNA-technology" class="md-crosslink " data-show-preview="true">recombinant DNA technology</a>, which emerged from basic research in microbial <a href="https://www.britannica.com/science/genetics" class="md-crosslink autoxref " data-show-preview="true">genetics</a>. The techniques employed in genetic engineering have led to the production of medically important products, including human <a href="https://www.britannica.com/science/insulin" class="md-crosslink " data-show-preview="true">insulin</a>, human <a href="https://www.britannica.com/science/growth-hormone" class="md-crosslink " data-show-preview="true">growth hormone</a>, and <a href="https://www.britannica.com/science/hepatitis-B" class="md-crosslink " data-show-preview="true">hepatitis B</a> <a href="https://www.britannica.com/science/vaccine" class="md-crosslink autoxref " data-show-preview="true">vaccine</a>, as well as to the development of <a href="https://www.britannica.com/science/genetically-modified-organism" class="md-crosslink " data-show-preview="true">genetically modified organisms</a> such as disease-resistant plants.</p><!--[MOD1]--><span class="marker MOD1 mod-inline"></span></section> <!--[H2]--><span class="marker h2"></span><section data-level="1" id="ref337079"> <h2 class="h1">Historical developments</h2> <!--[PREMOD2]--><span class="marker PREMOD2 mod-inline"></span><div class="assemblies"><div class="w-100"><figure class="md-assembly m-0 mb-md-0 card card-borderless print-false" data-assembly-id="280930" data-asm-type="video"><div class="md-assembly-wrapper card-media" data-type="video" video-id="240889"><a data-id="240889" class="gtm-assembly-link d-flex justify-content-center" style="--aspect-ratio: 16/9" href="/video/Genetically-Modified-Humans-CRISPR-Cas-9-Explained/-280930"><img src="https://cdn.britannica.com/89/240889-138-BE17294D/Genetically-Modified-Humans-CRISPR-Cas-9-Explained.jpg?w=800&h=450&c=crop" alt="genetically modified humans" loading="lazy"><script type="application/json"> { "sources": [ { "file" : "//content.jwplatform.com/manifests/cUmKL29W.m3u8" } ], "image": "https://cdn.britannica.com/89/240889-138-BE17294D/Genetically-Modified-Humans-CRISPR-Cas-9-Explained.jpg" ,"tracks": [ { "file" : "//assets-jpcust.jwpsrv.com/tracks/xTEcqdjD", "label": "English" } ] ,"adfile": "//content.jwplatform.com/manifests/yfUjoC1u.m3u8" } </script><div class="btn btn-xl btn-white btn-circle position-absolute shadow" style="top: 50%; transform: translateY(-50%)"><em class="material-icons" data-icon="play_arrow"></em></div></a></div><figcaption class="card-body"><div class="md-assembly-caption text-muted font-14 font-serif line-clamp"><span><span class="md-assembly-title font-weight-bold mr-5 d-inline font-sans-serif md-video-caption" video-control="240889">genetically modified humans</span><span>CRISPR and genetically modified humans.</span><button class="js-more-btn d-none btn btn-unstyled font-12 bg-white js-content" aria-label="Toggle more/less fact data"><span class="link-blue">(more)</span></button></span></div><a class="font-14 mt-10 d-inline-block" href="/science/genetic-engineering/images-videos">See all videos for this article</a></figcaption></figure></div></div><p class="topic-paragraph">The term <em>genetic engineering</em> initially referred to various techniques used for the modification or manipulation of organisms through the processes of <a href="https://www.britannica.com/science/heredity-genetics" class="md-crosslink " data-show-preview="true">heredity</a> and <a href="https://www.britannica.com/science/reproduction-biology" class="md-crosslink " data-show-preview="true">reproduction</a>. As such, the term embraced both <a href="https://www.britannica.com/science/selective-breeding" class="md-crosslink autoxref " data-show-preview="true">artificial selection</a> and all the interventions of biomedical techniques, among them <a href="https://www.britannica.com/science/artificial-insemination" class="md-crosslink autoxref " data-show-preview="true">artificial insemination</a>, <a href="https://www.britannica.com/science/in-vitro-fertilization" class="md-crosslink " data-show-preview="true">in vitro fertilization</a> (e.g., “test-tube” babies), <a href="https://www.britannica.com/science/cloning" class="md-crosslink " data-show-preview="true">cloning</a>, and <a href="https://www.britannica.com/science/gene" class="md-crosslink " data-show-preview="true">gene</a> manipulation. In the latter part of the 20th century, however, the term came to refer more specifically to methods of <span id="ref233969"></span><a href="https://www.britannica.com/science/recombinant-DNA-technology" class="md-crosslink " data-show-preview="true">recombinant DNA technology</a> (or <span id="ref233968"></span><a href="https://www.britannica.com/science/recombination-genetics" class="md-crosslink " data-show-preview="true">gene cloning</a>), in which DNA molecules from two or more sources are combined either within <a href="https://www.britannica.com/science/cell-biology" class="md-crosslink " data-show-preview="true">cells</a> or in vitro and are then inserted into host organisms in which they are able to <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="propagate" href="https://www.merriam-webster.com/dictionary/propagate" data-type="MW">propagate</a>.</p><div class="one-good-fact-module"> </div><!--[MOD2]--><span class="marker MOD2 mod-inline"></span> <!--[PREMOD3]--><span class="marker PREMOD3 mod-inline"></span><p class="topic-paragraph">The possibility for <span id="ref1292909"></span><a href="https://www.britannica.com/science/recombinant-DNA-technology" class="md-crosslink " data-show-preview="true">recombinant DNA</a> <a href="https://www.britannica.com/technology/technology" class="md-crosslink autoxref " data-show-preview="true">technology</a> emerged with the discovery of <span id="ref661169"></span><a href="https://www.britannica.com/science/restriction-enzyme" class="md-crosslink " data-show-preview="true">restriction enzymes</a> in 1968 by Swiss microbiologist <a href="https://www.britannica.com/biography/Werner-Arber" class="md-crosslink " data-show-preview="true">Werner Arber</a>. The following year American microbiologist <a href="https://www.britannica.com/biography/Hamilton-O-Smith" class="md-crosslink " data-show-preview="true">Hamilton O. Smith</a> purified so-called <span id="ref661170"></span><a href="https://www.britannica.com/science/type-II-restriction-enzyme" class="md-crosslink ">type II restriction enzymes</a>, which were found to be essential to genetic engineering for their ability to <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="cleave" href="https://www.merriam-webster.com/dictionary/cleave" data-type="MW">cleave</a> a specific site within the DNA (as opposed to type I restriction enzymes, which cleave DNA at random sites). Drawing on Smith’s work, American molecular biologist <a href="https://www.britannica.com/biography/Daniel-Nathans" class="md-crosslink " data-show-preview="true">Daniel Nathans</a> helped advance the technique of DNA recombination in 1970–71 and demonstrated that type II enzymes could be useful in genetic studies. Genetic engineering based on recombination was pioneered in 1973 by American biochemists <a href="https://www.britannica.com/biography/Stanley-Cohen" class="md-crosslink " data-show-preview="true">Stanley N. Cohen</a> and Herbert W. Boyer, who were among the first to cut DNA into fragments, rejoin different fragments, and insert the new genes into <a href="https://www.britannica.com/science/E-coli" class="md-crosslink " data-show-preview="true"><em>E. coli</em></a> <a href="https://www.britannica.com/science/bacteria" class="md-crosslink autoxref " data-show-preview="true">bacteria</a>, which then reproduced.</p><!--[MOD3]--><span class="marker MOD3 mod-inline"></span> </section> <!--[END-OF-CONTENT]--><span class="marker end-of-content"></span><!--[AFTER-ARTICLE]--><span class="marker after-article"></span></div> <div id="chatbot-root"></div> </div> </div> </div> <div class="ai-dialog-placeholder"></div> </div> </div> <aside class="col-md-da-320"></aside> </div> </div> </div> </div> </article> </div> </div> </div> </div> </main> <div id="md-footer"></div> <noscript><iframe src="//www.googletagmanager.com/ns.html?id=GTM-5W6NC8" height="0" width="0" style="display:none;visibility:hidden"></iframe></noscript> <script type="text/javascript" id="_informizely_script_tag"> var IzWidget = IzWidget || {}; (function (d) { var scriptElement = d.createElement('script'); scriptElement.type = 'text/javascript'; scriptElement.async = true; scriptElement.src = "https://insitez.blob.core.windows.net/site/f780f33e-a610-4ac2-af81-3eb184037547.js"; var node = d.getElementById('_informizely_script_tag'); node.parentNode.insertBefore(scriptElement, node); } )(document); </script> <!-- Ortto ebmwprod capture code --> <script> window.ap3c = window.ap3c || {}; var ap3c = window.ap3c; ap3c.cmd = ap3c.cmd || []; ap3c.cmd.push(function() { ap3c.init('ZO4siT4cLwnykPnzZWJtd3Byb2Q', 'https://engage.email.britannica.com/'); ap3c.track({v: 0}); }); ap3c.activity = function(act) { ap3c.act = (ap3c.act || []); ap3c.act.push(act); }; var s, t; s = document.createElement('script'); s.type = 'text/javascript'; s.src = "https://engage.email.britannica.com/app.js"; t = document.getElementsByTagName('script')[0]; t.parentNode.insertBefore(s, t); </script> <script class="marketing-page-info" type="application/json"> {"pageType":"Topic","templateName":"DESKTOP","pageNumber":1,"pagesTotal":2,"pageId":228897,"pageLength":374,"initialLoad":true,"lastPageOfScroll":false} </script> <script class="marketing-content-info" type="application/json"> [] </script> <script src="https://cdn.britannica.com/mendel-resources/3-133/js/libs/jquery-3.5.0.min.js?v=3.133.9"></script> <script type="text/javascript" data-type="Init Mendel Code Splitting"> (function() { $.ajax({ dataType: 'script', cache: true, url: 'https://cdn.britannica.com/mendel-resources/3-133/dist/topic-page.js?v=3.133.9' }); })(); </script> <script class="analytics-metadata" type="application/json"> {"leg":"D","adLeg":"D","userType":"ANONYMOUS","pageType":"Topic","pageSubtype":null,"articleTemplateType":"PAGINATED","gisted":false,"pageNumber":1,"hasSummarizeButton":false,"hasAskButton":true} </script> <script type="text/javascript"> EBStat={accountId:-1,hostnameOverride:'webstats.eb.com',domain:'www.britannica.com', json:''}; </script> <script type="text/javascript"> ( function() { $.ajax( { dataType: 'script', cache: true, url: '//www.britannica.com/webstats/mendelstats.js?v=1' } ) .done( function() { try {writeStat(null,EBStat);} catch(err){} } ); })(); </script> <div id="bc-fixed-dialogue"></div> </body> </html>

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