Generation and annotation of the DNA sequences of human chromosomes 2 and 4

<!DOCTYPE html>     <html class="no-js" lang="en">  <head> <title>Generation and annotation of the DNA sequences of human chromosomes 2 and 4 - NASA/ADS</title>  <link rel="apple-touch-icon" sizes="180x180" href="//styles/favicon/apple-touch-icon.png" /> <link rel="icon" type="image/png" sizes="32x32" href="//styles/favicon/favicon-32x32.png" /> <link rel="icon" type="image/png" sizes="16x16" href="//styles/favicon/favicon-16x16.png" /> <link rel="manifest" href="//styles/favicon/site.webmanifest" /> <link rel="mask-icon" href="//styles/favicon/safari-pinned-tab.svg" color="#5bbad5" /> <meta name="apple-mobile-web-app-title" content="NASA ADS" /> <meta name="application-name" content="NASA ADS" /> <meta name="msapplication-TileColor" content="#ffc40d" /> <meta name="theme-color" content="#ffffff" />  <link rel="stylesheet" href="/styles/css/styles.css"> <meta name="robots" content="noarchive"> <link rel="canonical" href="http://ui.adsabs.harvard.edu/abs/2005Natur.434..724H/abstract"/> <meta name="description" content="Human chromosome 2 is unique to the human lineage in being the product of a head-to-head fusion of two intermediate-sized ancestral chromosomes. Chromosome 4 has received attention primarily related to the search for the Huntington's disease gene, but also for genes associated with Wolf-Hirschhorn syndrome, polycystic kidney disease and a form of muscular dystrophy. Here we present approximately 237 million base pairs of sequence for chromosome 2, and 186 million base pairs for chromosome 4, representing more than 99.6% of their euchromatic sequences. Our initial analyses have identified 1,346 protein-coding genes and 1,239 pseudogenes on chromosome 2, and 796 protein-coding genes and 778 pseudogenes on chromosome 4. Extensive analyses confirm the underlying construction of the sequence, and expand our understanding of the structure and evolution of mammalian chromosomes, including gene deserts, segmental duplications and highly variant regions.">  <meta property="og:type" content="article"> <meta property="og:title" content="Generation and annotation of the DNA sequences of human chromosomes 2 and 4"> <meta property="og:site_name" content="NASA/ADS"> <meta property="og:description" content="Human chromosome 2 is unique to the human lineage in being the product of a head-to-head fusion of two intermediate-sized ancestral chromosomes. Chromosome 4 has received attention primarily related to the search for the Huntington's disease gene, but also for genes associated with Wolf-Hirschhorn syndrome, polycystic kidney disease and a form of muscular dystrophy. Here we present approximately 237 million base pairs of sequence for chromosome 2, and 186 million base pairs for chromosome 4, representing more than 99.6% of their euchromatic sequences. Our initial analyses have identified 1,346 protein-coding genes and 1,239 pseudogenes on chromosome 2, and 796 protein-coding genes and 778 pseudogenes on chromosome 4. Extensive analyses confirm the underlying construction of the sequence, and expand our understanding of the structure and evolution of mammalian chromosomes, including gene deserts, segmental duplications and highly variant regions."> <meta property="og:url" content="https://ui.adsabs.harvard.edu/abs/2005Natur.434..724H/abstract"> <meta property="og:image" content="https://ui.adsabs.harvard.edu/styles/img/transparent_logo.svg"> <meta property="article:published_time" content="04/2005"> <meta property="article:author" content="Hillier, LaDeana W."> <meta property="article:author" content="Graves, Tina A."> <meta property="article:author" content="Fulton, Robert S."> <meta property="article:author" content="Fulton, Lucinda A."> <meta property="article:author" content="Pepin, Kymberlie H."> <meta property="article:author" content="Minx, Patrick"> <meta property="article:author" content="Wagner-McPherson, Caryn"> <meta property="article:author" content="Layman, Dan"> <meta 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Chromosome 4 has received attention primarily related to the search for the Huntington's disease gene, but also for genes associated with Wolf-Hirschhorn syndrome, polycystic kidney disease and a form of muscular dystrophy. Here we present approximately 237 million base pairs of sequence for chromosome 2, and 186 million base pairs for chromosome 4, representing more than 99.6% of their euchromatic sequences. Our initial analyses have identified 1,346 protein-coding genes and 1,239 pseudogenes on chromosome 2, and 796 protein-coding genes and 778 pseudogenes on chromosome 4. 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<li class="author"><a href="/search/?q=author%3A%22Graves%2C+Tina+A.%22">Graves, Tina A.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Fulton%2C+Robert+S.%22">Fulton, Robert S.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Fulton%2C+Lucinda+A.%22">Fulton, Lucinda A.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Pepin%2C+Kymberlie+H.%22">Pepin, Kymberlie H.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Minx%2C+Patrick%22">Minx, Patrick</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Wagner-McPherson%2C+Caryn%22">Wagner-McPherson, Caryn</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Layman%2C+Dan%22">Layman, Dan</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Wylie%2C+Kristine%22">Wylie, Kristine</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Sekhon%2C+Mandeep%22">Sekhon, Mandeep</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Becker%2C+Michael+C.%22">Becker, Michael C.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Fewell%2C+Ginger+A.%22">Fewell, Ginger A.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Delehaunty%2C+Kimberly+D.%22">Delehaunty, Kimberly D.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Miner%2C+Tracie+L.%22">Miner, Tracie L.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Nash%2C+William+E.%22">Nash, William E.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Kremitzki%2C+Colin%22">Kremitzki, Colin</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Oddy%2C+Lachlan%22">Oddy, Lachlan</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Du%2C+Hui%22">Du, Hui</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Sun%2C+Hui%22">Sun, Hui</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Bradshaw-Cordum%2C+Holland%22">Bradshaw-Cordum, Holland</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Ali%2C+Johar%22">Ali, Johar</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Carter%2C+Jason%22">Carter, Jason</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Cordes%2C+Matt%22">Cordes, Matt</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Harris%2C+Anthony%22">Harris, Anthony</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Isak%2C+Amber%22">Isak, Amber</a> </li>; <li class="author"><a href="/search/?q=author%3A%22van+Brunt%2C+Andrew%22">van Brunt, Andrew</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Nguyen%2C+Christine%22">Nguyen, Christine</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Du%2C+Feiyu%22">Du, Feiyu</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Courtney%2C+Laura%22">Courtney, Laura</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Kalicki%2C+Joelle%22">Kalicki, Joelle</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Ozersky%2C+Philip%22">Ozersky, Philip</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Abbott%2C+Scott%22">Abbott, Scott</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Armstrong%2C+Jon%22">Armstrong, Jon</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Belter%2C+Edward+A.%22">Belter, Edward A.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Caruso%2C+Lauren%22">Caruso, Lauren</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Cedroni%2C+Maria%22">Cedroni, Maria</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Cotton%2C+Marc%22">Cotton, Marc</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Davidson%2C+Teresa%22">Davidson, Teresa</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Desai%2C+Anu%22">Desai, Anu</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Elliott%2C+Glendoria%22">Elliott, Glendoria</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Erb%2C+Thomas%22">Erb, Thomas</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Fronick%2C+Catrina%22">Fronick, Catrina</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Gaige%2C+Tony%22">Gaige, Tony</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Haakenson%2C+William%22">Haakenson, William</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Haglund%2C+Krista%22">Haglund, Krista</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Holmes%2C+Andrea%22">Holmes, Andrea</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Harkins%2C+Richard%22">Harkins, Richard</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Kim%2C+Kyung%22">Kim, Kyung</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Kruchowski%2C+Scott+S.%22">Kruchowski, Scott S.</a> </li>; <li class="author"><a href="/search/?q=author%3A%22Strong%2C+Cynthia+Madsen%22">Strong, Cynthia Madsen</a> </li> <li> <em>and 72 more</em> </li> </ul> </div> <div class="s-abstract-text"> <h4 class="sr-only">Abstract</h4> <p> Human chromosome 2 is unique to the human lineage in being the product of a head-to-head fusion of two intermediate-sized ancestral chromosomes. Chromosome 4 has received attention primarily related to the search for the Huntington's disease gene, but also for genes associated with Wolf-Hirschhorn syndrome, polycystic kidney disease and a form of muscular dystrophy. Here we present approximately 237 million base pairs of sequence for chromosome 2, and 186 million base pairs for chromosome 4, representing more than 99.6% of their euchromatic sequences. Our initial analyses have identified 1,346 protein-coding genes and 1,239 pseudogenes on chromosome 2, and 796 protein-coding genes and 778 pseudogenes on chromosome 4. Extensive analyses confirm the underlying construction of the sequence, and expand our understanding of the structure and evolution of mammalian chromosomes, including gene deserts, segmental duplications and highly variant regions. </p> </div> <br> <dl class="s-abstract-dl-horizontal"> <dt>Publication:</dt> <dd> <div id="article-publication">Nature</div> </dd> <dt>Pub Date:</dt> <dd>April 2005</dd> <dt>DOI:</dt> <dd> <span> <a href="/link_gateway/2005Natur.434..724H/doi:10.1038/nature03466" target="_blank" rel="noopener">10.1038/nature03466</a> <i class="fa fa-external-link"></i> </span> </dd> <dt>Bibcode:</dt> <dd> <a href="/abs/2005Natur.434..724H/abstract"> 2005Natur.434..724H </a> <i class="icon-help" title="The bibcode is assigned by the ADS as a unique identifier for the paper."></i> </dd> </dl> </article> </div> <div data-widget="ShowCitations"></div> <div data-widget="ShowReferences"></div> <div data-widget="ShowCoreads"></div> <div data-widget="ShowSimilar"></div> <div data-widget="ShowTableofcontents"></div> <div data-widget="ShowGraphics"></div> <div data-widget="ShowExportcitation" data-origin="abstract"></div> <div data-widget="ShowMetrics" data-allow-redirect="false"></div> <div data-widget="MetaTagsWidget"></div> </div> </div> </div> <div class="s-right-col-container col-xs-12 col-sm-12 col-md-3 col-lg-2 s-right-column" id="right-col-container" > <div data-widget="ShowResources"> <div data-reactroot="" class="s-right-col-widget-container" style="padding: 10px" > <div> <div class="resources__container"> <div class="resources__full__list"> <div class="resources__header__row"> <i class="fa fa-file-text-o" aria-hidden="true"> </i> <div class="resources__header__title">full text sources</div> </div> <div class="resources__content"> <div class="resources__content__title">Publisher</div> <div class="resources__content__links"> <span> <div class="resources__content__link__separator">|</div> </span> <span> <a href="/link_gateway/2005Natur.434..724H/PUB_HTML" rel="noopener" class="resources__content__link " > <i class="fa fa-file-text" aria-hidden="true"> </i> </a> </span> </div> </div> </div> </div> <div data-widget="ShowAssociated"> </div> </div> </div> </div> <div data-widget="ShowGraphicsSidebar"> </div> </div> </div> </div> </div> </div> </div> <div id="footer-container"> <div data-widget="FooterWidget"> <div class="footer s-footer"> <footer> <div class="__footer_wrapper"> <div class="__footer_brand"> 漏 The SAO/NASA Astrophysics Data System <div class="__footer_brand_extra"> <p> <i class="fa fa-envelope"></i> adshelp[at]cfa.harvard.edu </p> <p> The ADS is operated by the Smithsonian Astrophysical Observatory under NASA Cooperative Agreement <em>NNX16AC86A</em> </p> </div> <div class="__footer_brand_logos"> <a href="http://www.nasa.gov" target="_blank" rel="noopener"> <img src="/styles/img/nasa.svg" alt="NASA logo" id="nasa-logo"> </a> <a href="http://www.si.edu" target="_blank" rel="noopener"> <img id="smithsonian-logo" src="/styles/img/smithsonian.svg" alt="Smithsonian logo"> </a> <a href="https://www.cfa.harvard.edu/" target="_blank" rel="noopener"> <img src="/styles/img/cfa.png" title="Harvard Center for Astrophysics logo" id="cfa-logo"> </a> </div> </div> <div class="__footer_list"> <div class="__footer_list_title"> Resources </div> <ul class="__footer_links"> <li> <a href="/about/" target="_blank" rel="noopener"> <i class="fa fa-question-circle"></i> About ADS </a> </li> <li> <a href="//ui.adsabs.harvard.edu/help/" target="_blank" rel="noopener"> <i class="fa fa-info-circle"></i> ADS Help </a> </li> <li> <a href="//ui.adsabs.harvard.edu/help/whats_new/" target="_blank" rel="noopener"> <i class="fa fa-bullhorn"></i> What's New </a> </li> <li> <a href="/about/careers/" target="_blank" rel="noopener"> <i class="fa fa-group"></i> Careers@ADS </a> </li> </ul> </div> <div class="__footer_list"> <div class="__footer_list_title"> Social </div> <ul class="__footer_links"> <li> <a href="//twitter.com/adsabs" target="_blank" rel="noopener"> <i class="fa fa-twitter"></i> @adsabs </a> </li> <li> <a href="//ui.adsabs.harvard.edu/blog/" target="_blank" rel="noopener"> <i class="fa fa-newspaper-o"></i> ADS Blog </a> </li> </ul> </div> <div class="__footer_list"> <div class="__footer_list_title"> Project </div> <ul class="__footer_links"> <li> <a href="/core/never">Switch to full ADS</a> </li> <li> <a href="https://adsisdownorjustme.herokuapp.com/" target="_blank" rel="noopener">Is ADS down? 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// Add class "autocomplete-active": x[currentFocus].classList.add("autocomplete-active"); } function removeActive(x) { // Remove the "active" class from all autocomplete items: for (var i = 0; i < x.length; i++) { x[i].classList.remove("autocomplete-active"); } } function closeAllLists(elmnt) { // Close all autocomplete lists in the document, except the one passed as an argument: var x = document.getElementsByClassName("autocomplete-items"); for (var i = 0; i < x.length; i++) { if (elmnt != x[i] && elmnt != searchBox) { x[i].parentNode.removeChild(x[i]); } } } // Any other clicks in the document: document.addEventListener("click", function (e) { closeAllLists(e.target); }); } var autoList = [ { value: 'author:""', label: 'Author', match: 'author:"' }, { value: 'author:"^"', label: 'First Author', match: 'first author' }, { value: 'author:"^"', label: 'First Author', match: 'author:"^' }, { value: 'bibcode:""', label: 'Bibcode', desc: 'e.g. bibcode:1989ApJ...342L..71R', match: 'bibcode:"' }, { value: 'bibstem:""', label: 'Publication', desc: 'e.g. bibstem:ApJ', match: 'bibstem:"' }, { value: 'bibstem:""', label: 'Publication', desc: 'e.g. bibstem:ApJ', match: 'publication (bibstem)' }, { value: 'arXiv:', label: 'arXiv ID', match: 'arxiv:' }, { value: 'doi:', label: 'DOI', match: 'doi:' }, { value: 'full:""', label: 'Full text search', desc: 'title, abstract, and body', match: 'full:' }, { value: 'full:""', label: 'Full text search', desc: 'title, abstract, and body', match: 'fulltext' }, { value: 'full:""', label: 'Full text search', desc: 'title, abstract, and body', match: 'text' }, { value: 'year:', label: 'Year', match: 'year' }, { value: 'year:1999-2005', label: 'Year Range', desc: 'e.g. 1999-2005', match: 'year range' }, { value: 'aff:""', label: 'Affiliation', match: 'aff:' }, { value: 'abs:""', label: 'Search abstract + title + keywords', match: 'abs:' }, { value: 'database:astronomy', label: 'Limit to papers in the astronomy database', match: 'database:astronomy' }, { value: 'database:physics', label: 'Limit to papers in the physics database', match: 'database:physics' }, { value: 'title:""', label: 'Title', match: 'title:"' }, { value: 'orcid:', label: 'ORCiD identifier', match: 'orcid:' }, { value: 'object:', label: 'SIMBAD object (e.g. object:LMC)', match: 'object:' }, { value: 'property:refereed', label: 'Limit to refereed', desc: '(property:refereed)', match: 'refereed' }, { value: 'property:refereed', label: 'Limit to refereed', desc: '(property:refereed)', match: 'property:refereed' }, { value: 'property:notrefereed', label: 'Limit to non-refereed', desc: '(property:notrefereed)', match: 'property:notrefereed' }, { value: 'property:notrefereed', label: 'Limit to non-refereed', desc: '(property:notrefereed)', match: 'notrefereed' }, { value: 'property:eprint', label: 'Limit to eprints', desc: '(property:eprint)', match: 'eprint' }, { value: 'property:eprint', label: 'Limit to eprints', desc: '(property:eprint)', match: 'property:eprint' }, { value: 'property:openaccess', label: 'Limit to open access', desc: '(property:openaccess)', match: 'property:openaccess' }, { value: 'property:openaccess', label: 'Limit to open access', desc: '(property:openaccess)', match: 'openaccess' }, { value: 'doctype:software', label: 'Limit to software', desc: '(doctype:software)', match: 'software' }, { value: 'doctype:software', label: 'Limit to software', desc: '(doctype:software)', match: 'doctype:software' }, { value: 'property:inproceedings', label: 'Limit to papers in conference proceedings', desc: '(property:inproceedings)', match: 'proceedings' }, { value: 'property:inproceedings', label: 'Limit to papers in conference proceedings', desc: '(property:inproceedings)', match: 'property:inproceedings' }, { value: 'citations()', label: 'Citations', desc: 'Get papers citing your search result set', match: 'citations(' }, { value: 'references()', label: 'References', desc: 'Get papers referenced by your search result set', match: 'references(' }, { value: 'trending()', label: 'Trending', desc: 'Get papers most read by users who recently read your search result set', match: 'trending(' }, { value: 'reviews()', label: 'Review Articles', desc: 'Get most relevant papers that cite your search result set', match: 'reviews(' }, { value: 'useful()', label: 'Useful', desc: 'Get papers most frequently cited by your search result set', match: 'useful(' }, { value: 'similar()', label: 'Similar', desc: 'Get papers that have similar full text to your search result set', match: 'similar(' }, ]; // initiate the autocomplete function on the "q" element, and pass along the operators array as possible autocomplete values: inputBox = document.getElementById("q") if (inputBox) { inputBox.focus() // autofucs inputBox.setSelectionRange(inputBox.value.length, inputBox.value.length); // bring cursor to the end autocomplete(inputBox, autoList); } </script> <script> (function() { // turn off no-js if we have javascript document.documentElement.className = document.documentElement.className.replace("no-js", "js"); function getCookie(cname) { var name = cname + "="; var decodedCookie = decodeURIComponent(document.cookie); var ca = decodedCookie.split(';'); for (var i = 0; i < ca.length; i++) { var c = ca[i]; while (c.charAt(0) == ' ') { c = c.substring(1); } if (c.indexOf(name) == 0) { return c.substring(name.length, c.length); } } return ""; } (function() { // looks for the cookie, and sets true if its 'always' const coreCookie = getCookie('core') === 'always'; // only load bumblebee if we detect the core cookie and we are on abstract page if (coreCookie || (!(/^\/abs\//.test(document.location.pathname)) && !coreCookie)) { return; 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Generation and annotation of the DNA sequences of human chromosomes 2 and 4 - NASA/ADS