CINXE.COM
PLOS Biology
<!DOCTYPE html> <html xmlns="http://www.w3.org/1999/xhtml" lang="en" xml:lang="en" itemscope itemtype="http://schema.org/Article" class="no-js"> <head prefix="og: http://ogp.me/ns#"> <link rel="stylesheet" href="/resource/css/home.css?76e559307c04719bc34c8788c3343923"/> <!-- allows for extra head tags --> <!-- hello --> <link rel="stylesheet" type="text/css" href="https://fonts.googleapis.com/css?family=Open+Sans:400,400i,600"> <link media="print" rel="stylesheet" type="text/css" href="/resource/css/print.css"/> <script type="text/javascript"> var siteUrlPrefix = "/plosbiology/"; </script> <script src="/resource/js/vendor/modernizr-v2.7.1.js" type="text/javascript"></script> <script src="/resource/js/vendor/detectizr.min.js" type="text/javascript"></script> <link rel="shortcut icon" href="/resource/img/favicon.ico" type="image/x-icon"/> <meta http-equiv="Content-Type" content="text/html; charset=utf-8"/> <!-- DoubleClick overall ad setup script --> <script type='text/javascript'> var googletag = googletag || {}; googletag.cmd = googletag.cmd || []; (function() { var gads = document.createElement('script'); gads.async = true; gads.type = 'text/javascript'; var useSSL = 'https:' == document.location.protocol; gads.src = (useSSL ? 'https:' : 'http:') + '//www.googletagservices.com/tag/js/gpt.js'; var node = document.getElementsByTagName('script')[0]; node.parentNode.insertBefore(gads, node); })(); </script> <!-- DoubleClick ad slot setup script --> <script id="doubleClickSetupScript" type='text/javascript'> googletag.cmd.push(function() { googletag.defineSlot('/75507958/PBIO_728x90_ATF', [728, 90], 'div-gpt-ad-1458247671871-0').addService(googletag.pubads()); googletag.defineSlot('/75507958/PBIO_160x600_BTF', [160, 600], 'div-gpt-ad-1458247671871-1').addService(googletag.pubads()); googletag.defineSlot('/75507958/PBIO_300x250_ITB1', [300, 250], 'div-gpt-ad-1458247671871-2').addService(googletag.pubads()); googletag.defineSlot('/75507958/PBIO_300x250_ITB2', [300, 250], 'div-gpt-ad-1458247671871-3').addService(googletag.pubads()); googletag.defineSlot('/75507958/PBIO_300x250_ITB3', [300, 250], 'div-gpt-ad-1458247671871-4').addService(googletag.pubads()); var personalizedAds = window.plosCookieConsent && window.plosCookieConsent.hasConsented('advertising'); googletag.pubads().setRequestNonPersonalizedAds(personalizedAds ? 0 : 1); googletag.pubads().enableSingleRequest(); googletag.enableServices(); }); </script> <script type="text/javascript"> var WombatConfig = WombatConfig || {}; WombatConfig.journalKey = "PLoSBiology"; WombatConfig.journalName = "PLOS Biology"; WombatConfig.figurePath = "/plosbiology/article/figure/image"; WombatConfig.figShareInstitutionString = "plos"; WombatConfig.doiResolverPrefix = "https://dx.plos.org/"; </script> <script type="text/javascript"> var WombatConfig = WombatConfig || {}; WombatConfig.metrics = WombatConfig.metrics || {}; WombatConfig.metrics.referenceUrl = "http://lagotto.io/plos"; WombatConfig.metrics.googleScholarUrl = "https://scholar.google.com/scholar"; WombatConfig.metrics.googleScholarCitationUrl = WombatConfig.metrics.googleScholarUrl + "?hl=en&lr=&q="; WombatConfig.metrics.crossrefUrl = "https://www.crossref.org"; </script> <script defer="defer" src="/resource/js/defer.js?94d25648e7eaf3e1771c"></script><script src="/resource/js/sync.js?94d25648e7eaf3e1771c"></script> <script src="/resource/js/vendor/jquery.min.js" type="text/javascript"></script> <script src="/resource/js/vendor/fastclick/lib/fastclick.js" type="text/javascript"></script> <script src="/resource/js/vendor/foundation/foundation.js" type="text/javascript"></script> <script src="/resource/js/vendor/underscore-min.js" type="text/javascript"></script> <script src="/resource/js/vendor/underscore.string.min.js" type="text/javascript"></script> <script src="/resource/js/vendor/moment.js" type="text/javascript"></script> <script src="/resource/js/vendor/jquery-ui-effects.min.js" type="text/javascript"></script> <script src="/resource/js/vendor/foundation/foundation.tooltip.js" type="text/javascript"></script> <script src="/resource/js/vendor/foundation/foundation.dropdown.js" type="text/javascript"></script> <script src="/resource/js/vendor/foundation/foundation.tab.js" type="text/javascript"></script> <script src="/resource/js/vendor/foundation/foundation.reveal.js" type="text/javascript"></script> <script src="/resource/js/vendor/foundation/foundation.slider.js" type="text/javascript"></script> <script src="/resource/js/util/utils.js" type="text/javascript"></script> <script src="/resource/js/components/toggle.js" type="text/javascript"></script> <script src="/resource/js/components/truncate_elem.js" type="text/javascript"></script> <script src="/resource/js/components/tooltip_hover.js" type="text/javascript"></script> <script src="/resource/js/vendor/jquery.dotdotdot.js" type="text/javascript"></script> <!--For Google Tag manager to be able to track site information --> <script> dataLayer = [{ 'mobileSite': 'false', 'desktopSite': 'true' }]; </script> <title>PLOS Biology</title> <!-- for lemur --> <script> $(document).ready(()=> { let lemurTitle = $('article > h1').text(); if (lemurTitle && lemurTitle != '') { $('title').text(lemurTitle + ' | ' + 'PLOS Biology'); } }); </script> </head> <body class="home plosbiology"> <div id="test"></div> <!-- Google Tag Manager --> <noscript><iframe src="//www.googletagmanager.com/ns.html?id=GTM-TP26BH" height="0" width="0" style="display:none;visibility:hidden"></iframe></noscript> <script> (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start': new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0], j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src= '//www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f); })(window,document,'script','dataLayer','GTM-TP26BH'); </script> <noscript><iframe src="//www.googletagmanager.com/ns.html?id=GTM-MQQMGF" height="0" width="0" style="display:none;visibility:hidden"></iframe></noscript> <script>(function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start': new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0], j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src= '//www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f); })(window,document,'script','dataLayer','GTM-MQQMGF');</script> <!-- End Google Tag Manager --> <!-- Start of HubSpot Embed Code --> <script> // from https://developers.hubspot.com/docs/reference/api/analytics-and-events/cookie-banner/cookie-banner-api#example-using-a-third-party-cookie-banner-to-control-hubspot-cookies window.disableHubSpotCookieBanner = true; window._hsp = window._hsp || []; window._hsp.push(['setHubSpotConsent', { analytics: window.plosCookieConsent?.hasConsented('analytics'), advertisement: window.plosCookieConsent?.hasConsented('advertising'), functionality: window.plosCookieConsent?.hasConsented('functionality') } ]); </script> <script type="text/javascript" id="hs-script-loader" async defer src="//js.hs-scripts.com/44092021.js"></script> <!-- End of HubSpot Embed Code --> <!-- New Relic --> <script type="text/javascript"> ;window.NREUM||(NREUM={});NREUM.init={distributed_tracing:{enabled:true},privacy:{cookies_enabled:true},ajax:{deny_list:["bam.nr-data.net"]}}; window.NREUM||(NREUM={}),__nr_require=function(t,e,n){function r(n){if(!e[n]){var o=e[n]={exports:{}};t[n][0].call(o.exports,function(e){var o=t[n][1][e];return r(o||e)},o,o.exports)}return e[n].exports}if("function"==typeof __nr_require)return __nr_require;for(var o=0;o<n.length;o++)r(n[o]);return r}({1:[function(t,e,n){function r(t){try{s.console&&console.log(t)}catch(e){}}var o,i=t("ee"),a=t(32),s={};try{o=localStorage.getItem("__nr_flags").split(","),console&&"function"==typeof console.log&&(s.console=!0,o.indexOf("dev")!==-1&&(s.dev=!0),o.indexOf("nr_dev")!==-1&&(s.nrDev=!0))}catch(c){}s.nrDev&&i.on("internal-error",function(t){r(t.stack)}),s.dev&&i.on("fn-err",function(t,e,n){r(n.stack)}),s.dev&&(r("NR AGENT IN DEVELOPMENT MODE"),r("flags: "+a(s,function(t,e){return t}).join(", ")))},{}],2:[function(t,e,n){function r(t,e,n,r,s){try{l?l-=1:o(s||new UncaughtException(t,e,n),!0)}catch(f){try{i("ierr",[f,c.now(),!0])}catch(d){}}return"function"==typeof u&&u.apply(this,a(arguments))}function UncaughtException(t,e,n){this.message=t||"Uncaught error with no additional information",this.sourceURL=e,this.line=n}function o(t,e){var n=e?null:c.now();i("err",[t,n])}var i=t("handle"),a=t(33),s=t("ee"),c=t("loader"),f=t("gos"),u=window.onerror,d=!1,p="nr@seenError";if(!c.disabled){var l=0;c.features.err=!0,t(1),window.onerror=r;try{throw new Error}catch(h){"stack"in h&&(t(14),t(13),"addEventListener"in window&&t(7),c.xhrWrappable&&t(15),d=!0)}s.on("fn-start",function(t,e,n){d&&(l+=1)}),s.on("fn-err",function(t,e,n){d&&!n[p]&&(f(n,p,function(){return!0}),this.thrown=!0,o(n))}),s.on("fn-end",function(){d&&!this.thrown&&l>0&&(l-=1)}),s.on("internal-error",function(t){i("ierr",[t,c.now(),!0])})}},{}],3:[function(t,e,n){var r=t("loader");r.disabled||(r.features.ins=!0)},{}],4:[function(t,e,n){function r(){U++,L=g.hash,this[u]=y.now()}function o(){U--,g.hash!==L&&i(0,!0);var t=y.now();this[h]=~~this[h]+t-this[u],this[d]=t}function i(t,e){E.emit("newURL",[""+g,e])}function a(t,e){t.on(e,function(){this[e]=y.now()})}var s="-start",c="-end",f="-body",u="fn"+s,d="fn"+c,p="cb"+s,l="cb"+c,h="jsTime",m="fetch",v="addEventListener",w=window,g=w.location,y=t("loader");if(w[v]&&y.xhrWrappable&&!y.disabled){var x=t(11),b=t(12),E=t(9),R=t(7),O=t(14),T=t(8),P=t(15),S=t(10),M=t("ee"),N=M.get("tracer"),C=t(23);t(17),y.features.spa=!0;var L,U=0;M.on(u,r),b.on(p,r),S.on(p,r),M.on(d,o),b.on(l,o),S.on(l,o),M.buffer([u,d,"xhr-resolved"]),R.buffer([u]),O.buffer(["setTimeout"+c,"clearTimeout"+s,u]),P.buffer([u,"new-xhr","send-xhr"+s]),T.buffer([m+s,m+"-done",m+f+s,m+f+c]),E.buffer(["newURL"]),x.buffer([u]),b.buffer(["propagate",p,l,"executor-err","resolve"+s]),N.buffer([u,"no-"+u]),S.buffer(["new-jsonp","cb-start","jsonp-error","jsonp-end"]),a(T,m+s),a(T,m+"-done"),a(S,"new-jsonp"),a(S,"jsonp-end"),a(S,"cb-start"),E.on("pushState-end",i),E.on("replaceState-end",i),w[v]("hashchange",i,C(!0)),w[v]("load",i,C(!0)),w[v]("popstate",function(){i(0,U>1)},C(!0))}},{}],5:[function(t,e,n){function r(){var t=new PerformanceObserver(function(t,e){var n=t.getEntries();s(v,[n])});try{t.observe({entryTypes:["resource"]})}catch(e){}}function o(t){if(s(v,[window.performance.getEntriesByType(w)]),window.performance["c"+p])try{window.performance[h](m,o,!1)}catch(t){}else try{window.performance[h]("webkit"+m,o,!1)}catch(t){}}function i(t){}if(window.performance&&window.performance.timing&&window.performance.getEntriesByType){var a=t("ee"),s=t("handle"),c=t(14),f=t(13),u=t(6),d=t(23),p="learResourceTimings",l="addEventListener",h="removeEventListener",m="resourcetimingbufferfull",v="bstResource",w="resource",g="-start",y="-end",x="fn"+g,b="fn"+y,E="bstTimer",R="pushState",O=t("loader");if(!O.disabled){O.features.stn=!0,t(9),"addEventListener"in window&&t(7);var T=NREUM.o.EV;a.on(x,function(t,e){var n=t[0];n instanceof T&&(this.bstStart=O.now())}),a.on(b,function(t,e){var n=t[0];n instanceof T&&s("bst",[n,e,this.bstStart,O.now()])}),c.on(x,function(t,e,n){this.bstStart=O.now(),this.bstType=n}),c.on(b,function(t,e){s(E,[e,this.bstStart,O.now(),this.bstType])}),f.on(x,function(){this.bstStart=O.now()}),f.on(b,function(t,e){s(E,[e,this.bstStart,O.now(),"requestAnimationFrame"])}),a.on(R+g,function(t){this.time=O.now(),this.startPath=location.pathname+location.hash}),a.on(R+y,function(t){s("bstHist",[location.pathname+location.hash,this.startPath,this.time])}),u()?(s(v,[window.performance.getEntriesByType("resource")]),r()):l in window.performance&&(window.performance["c"+p]?window.performance[l](m,o,d(!1)):window.performance[l]("webkit"+m,o,d(!1))),document[l]("scroll",i,d(!1)),document[l]("keypress",i,d(!1)),document[l]("click",i,d(!1))}}},{}],6:[function(t,e,n){e.exports=function(){return"PerformanceObserver"in window&&"function"==typeof window.PerformanceObserver}},{}],7:[function(t,e,n){function r(t){for(var e=t;e&&!e.hasOwnProperty(u);)e=Object.getPrototypeOf(e);e&&o(e)}function o(t){s.inPlace(t,[u,d],"-",i)}function i(t,e){return t[1]}var a=t("ee").get("events"),s=t("wrap-function")(a,!0),c=t("gos"),f=XMLHttpRequest,u="addEventListener",d="removeEventListener";e.exports=a,"getPrototypeOf"in Object?(r(document),r(window),r(f.prototype)):f.prototype.hasOwnProperty(u)&&(o(window),o(f.prototype)),a.on(u+"-start",function(t,e){var n=t[1];if(null!==n&&("function"==typeof n||"object"==typeof n)){var r=c(n,"nr@wrapped",function(){function t(){if("function"==typeof n.handleEvent)return n.handleEvent.apply(n,arguments)}var e={object:t,"function":n}[typeof n];return e?s(e,"fn-",null,e.name||"anonymous"):n});this.wrapped=t[1]=r}}),a.on(d+"-start",function(t){t[1]=this.wrapped||t[1]})},{}],8:[function(t,e,n){function r(t,e,n){var r=t[e];"function"==typeof r&&(t[e]=function(){var t=i(arguments),e={};o.emit(n+"before-start",[t],e);var a;e[m]&&e[m].dt&&(a=e[m].dt);var s=r.apply(this,t);return o.emit(n+"start",[t,a],s),s.then(function(t){return o.emit(n+"end",[null,t],s),t},function(t){throw o.emit(n+"end",[t],s),t})})}var o=t("ee").get("fetch"),i=t(33),a=t(32);e.exports=o;var s=window,c="fetch-",f=c+"body-",u=["arrayBuffer","blob","json","text","formData"],d=s.Request,p=s.Response,l=s.fetch,h="prototype",m="nr@context";d&&p&&l&&(a(u,function(t,e){r(d[h],e,f),r(p[h],e,f)}),r(s,"fetch",c),o.on(c+"end",function(t,e){var n=this;if(e){var r=e.headers.get("content-length");null!==r&&(n.rxSize=r),o.emit(c+"done",[null,e],n)}else o.emit(c+"done",[t],n)}))},{}],9:[function(t,e,n){var r=t("ee").get("history"),o=t("wrap-function")(r);e.exports=r;var i=window.history&&window.history.constructor&&window.history.constructor.prototype,a=window.history;i&&i.pushState&&i.replaceState&&(a=i),o.inPlace(a,["pushState","replaceState"],"-")},{}],10:[function(t,e,n){function r(t){function e(){f.emit("jsonp-end",[],l),t.removeEventListener("load",e,c(!1)),t.removeEventListener("error",n,c(!1))}function n(){f.emit("jsonp-error",[],l),f.emit("jsonp-end",[],l),t.removeEventListener("load",e,c(!1)),t.removeEventListener("error",n,c(!1))}var r=t&&"string"==typeof t.nodeName&&"script"===t.nodeName.toLowerCase();if(r){var o="function"==typeof t.addEventListener;if(o){var a=i(t.src);if(a){var d=s(a),p="function"==typeof d.parent[d.key];if(p){var l={};u.inPlace(d.parent,[d.key],"cb-",l),t.addEventListener("load",e,c(!1)),t.addEventListener("error",n,c(!1)),f.emit("new-jsonp",[t.src],l)}}}}}function o(){return"addEventListener"in window}function i(t){var e=t.match(d);return e?e[1]:null}function a(t,e){var n=t.match(l),r=n[1],o=n[3];return o?a(o,e[r]):e[r]}function s(t){var e=t.match(p);return e&&e.length>=3?{key:e[2],parent:a(e[1],window)}:{key:t,parent:window}}var c=t(23),f=t("ee").get("jsonp"),u=t("wrap-function")(f);if(e.exports=f,o()){var d=/[?&](?:callback|cb)=([^&#]+)/,p=/(.*)\.([^.]+)/,l=/^(\w+)(\.|$)(.*)$/,h=["appendChild","insertBefore","replaceChild"];Node&&Node.prototype&&Node.prototype.appendChild?u.inPlace(Node.prototype,h,"dom-"):(u.inPlace(HTMLElement.prototype,h,"dom-"),u.inPlace(HTMLHeadElement.prototype,h,"dom-"),u.inPlace(HTMLBodyElement.prototype,h,"dom-")),f.on("dom-start",function(t){r(t[0])})}},{}],11:[function(t,e,n){var r=t("ee").get("mutation"),o=t("wrap-function")(r),i=NREUM.o.MO;e.exports=r,i&&(window.MutationObserver=function(t){return this instanceof i?new i(o(t,"fn-")):i.apply(this,arguments)},MutationObserver.prototype=i.prototype)},{}],12:[function(t,e,n){function r(t){var e=i.context(),n=s(t,"executor-",e,null,!1),r=new f(n);return i.context(r).getCtx=function(){return e},r}var o=t("wrap-function"),i=t("ee").get("promise"),a=t("ee").getOrSetContext,s=o(i),c=t(32),f=NREUM.o.PR;e.exports=i,f&&(window.Promise=r,["all","race"].forEach(function(t){var e=f[t];f[t]=function(n){function r(t){return function(){i.emit("propagate",[null,!o],a,!1,!1),o=o||!t}}var o=!1;c(n,function(e,n){Promise.resolve(n).then(r("all"===t),r(!1))});var a=e.apply(f,arguments),s=f.resolve(a);return s}}),["resolve","reject"].forEach(function(t){var e=f[t];f[t]=function(t){var n=e.apply(f,arguments);return t!==n&&i.emit("propagate",[t,!0],n,!1,!1),n}}),f.prototype["catch"]=function(t){return this.then(null,t)},f.prototype=Object.create(f.prototype,{constructor:{value:r}}),c(Object.getOwnPropertyNames(f),function(t,e){try{r[e]=f[e]}catch(n){}}),o.wrapInPlace(f.prototype,"then",function(t){return function(){var e=this,n=o.argsToArray.apply(this,arguments),r=a(e);r.promise=e,n[0]=s(n[0],"cb-",r,null,!1),n[1]=s(n[1],"cb-",r,null,!1);var c=t.apply(this,n);return r.nextPromise=c,i.emit("propagate",[e,!0],c,!1,!1),c}}),i.on("executor-start",function(t){t[0]=s(t[0],"resolve-",this,null,!1),t[1]=s(t[1],"resolve-",this,null,!1)}),i.on("executor-err",function(t,e,n){t[1](n)}),i.on("cb-end",function(t,e,n){i.emit("propagate",[n,!0],this.nextPromise,!1,!1)}),i.on("propagate",function(t,e,n){this.getCtx&&!e||(this.getCtx=function(){if(t instanceof Promise)var e=i.context(t);return e&&e.getCtx?e.getCtx():this})}),r.toString=function(){return""+f})},{}],13:[function(t,e,n){var r=t("ee").get("raf"),o=t("wrap-function")(r),i="equestAnimationFrame";e.exports=r,o.inPlace(window,["r"+i,"mozR"+i,"webkitR"+i,"msR"+i],"raf-"),r.on("raf-start",function(t){t[0]=o(t[0],"fn-")})},{}],14:[function(t,e,n){function r(t,e,n){t[0]=a(t[0],"fn-",null,n)}function o(t,e,n){this.method=n,this.timerDuration=isNaN(t[1])?0:+t[1],t[0]=a(t[0],"fn-",this,n)}var i=t("ee").get("timer"),a=t("wrap-function")(i),s="setTimeout",c="setInterval",f="clearTimeout",u="-start",d="-";e.exports=i,a.inPlace(window,[s,"setImmediate"],s+d),a.inPlace(window,[c],c+d),a.inPlace(window,[f,"clearImmediate"],f+d),i.on(c+u,r),i.on(s+u,o)},{}],15:[function(t,e,n){function r(t,e){d.inPlace(e,["onreadystatechange"],"fn-",s)}function o(){var t=this,e=u.context(t);t.readyState>3&&!e.resolved&&(e.resolved=!0,u.emit("xhr-resolved",[],t)),d.inPlace(t,y,"fn-",s)}function i(t){x.push(t),m&&(E?E.then(a):w?w(a):(R=-R,O.data=R))}function a(){for(var t=0;t<x.length;t++)r([],x[t]);x.length&&(x=[])}function s(t,e){return e}function c(t,e){for(var n in t)e[n]=t[n];return e}t(7);var f=t("ee"),u=f.get("xhr"),d=t("wrap-function")(u),p=t(23),l=NREUM.o,h=l.XHR,m=l.MO,v=l.PR,w=l.SI,g="readystatechange",y=["onload","onerror","onabort","onloadstart","onloadend","onprogress","ontimeout"],x=[];e.exports=u;var b=window.XMLHttpRequest=function(t){var e=new h(t);try{u.emit("new-xhr",[e],e),e.addEventListener(g,o,p(!1))}catch(n){try{u.emit("internal-error",[n])}catch(r){}}return e};if(c(h,b),b.prototype=h.prototype,d.inPlace(b.prototype,["open","send"],"-xhr-",s),u.on("send-xhr-start",function(t,e){r(t,e),i(e)}),u.on("open-xhr-start",r),m){var E=v&&v.resolve();if(!w&&!v){var R=1,O=document.createTextNode(R);new m(a).observe(O,{characterData:!0})}}else f.on("fn-end",function(t){t[0]&&t[0].type===g||a()})},{}],16:[function(t,e,n){function r(t){if(!s(t))return null;var e=window.NREUM;if(!e.loader_config)return null;var n=(e.loader_config.accountID||"").toString()||null,r=(e.loader_config.agentID||"").toString()||null,f=(e.loader_config.trustKey||"").toString()||null;if(!n||!r)return null;var h=l.generateSpanId(),m=l.generateTraceId(),v=Date.now(),w={spanId:h,traceId:m,timestamp:v};return(t.sameOrigin||c(t)&&p())&&(w.traceContextParentHeader=o(h,m),w.traceContextStateHeader=i(h,v,n,r,f)),(t.sameOrigin&&!u()||!t.sameOrigin&&c(t)&&d())&&(w.newrelicHeader=a(h,m,v,n,r,f)),w}function o(t,e){return"00-"+e+"-"+t+"-01"}function i(t,e,n,r,o){var i=0,a="",s=1,c="",f="";return o+"@nr="+i+"-"+s+"-"+n+"-"+r+"-"+t+"-"+a+"-"+c+"-"+f+"-"+e}function a(t,e,n,r,o,i){var a="btoa"in window&&"function"==typeof window.btoa;if(!a)return null;var s={v:[0,1],d:{ty:"Browser",ac:r,ap:o,id:t,tr:e,ti:n}};return i&&r!==i&&(s.d.tk=i),btoa(JSON.stringify(s))}function s(t){return f()&&c(t)}function c(t){var e=!1,n={};if("init"in NREUM&&"distributed_tracing"in NREUM.init&&(n=NREUM.init.distributed_tracing),t.sameOrigin)e=!0;else if(n.allowed_origins instanceof Array)for(var r=0;r<n.allowed_origins.length;r++){var o=h(n.allowed_origins[r]);if(t.hostname===o.hostname&&t.protocol===o.protocol&&t.port===o.port){e=!0;break}}return e}function f(){return"init"in NREUM&&"distributed_tracing"in NREUM.init&&!!NREUM.init.distributed_tracing.enabled}function u(){return"init"in NREUM&&"distributed_tracing"in NREUM.init&&!!NREUM.init.distributed_tracing.exclude_newrelic_header}function d(){return"init"in NREUM&&"distributed_tracing"in NREUM.init&&NREUM.init.distributed_tracing.cors_use_newrelic_header!==!1}function p(){return"init"in NREUM&&"distributed_tracing"in NREUM.init&&!!NREUM.init.distributed_tracing.cors_use_tracecontext_headers}var l=t(29),h=t(18);e.exports={generateTracePayload:r,shouldGenerateTrace:s}},{}],17:[function(t,e,n){function r(t){var e=this.params,n=this.metrics;if(!this.ended){this.ended=!0;for(var r=0;r<p;r++)t.removeEventListener(d[r],this.listener,!1);e.aborted||(n.duration=a.now()-this.startTime,this.loadCaptureCalled||4!==t.readyState?null==e.status&&(e.status=0):i(this,t),n.cbTime=this.cbTime,s("xhr",[e,n,this.startTime,this.endTime,"xhr"],this))}}function o(t,e){var n=c(e),r=t.params;r.hostname=n.hostname,r.port=n.port,r.protocol=n.protocol,r.host=n.hostname+":"+n.port,r.pathname=n.pathname,t.parsedOrigin=n,t.sameOrigin=n.sameOrigin}function i(t,e){t.params.status=e.status;var n=v(e,t.lastSize);if(n&&(t.metrics.rxSize=n),t.sameOrigin){var r=e.getResponseHeader("X-NewRelic-App-Data");r&&(t.params.cat=r.split(", ").pop())}t.loadCaptureCalled=!0}var a=t("loader");if(a.xhrWrappable&&!a.disabled){var s=t("handle"),c=t(18),f=t(16).generateTracePayload,u=t("ee"),d=["load","error","abort","timeout"],p=d.length,l=t("id"),h=t(24),m=t(22),v=t(19),w=t(23),g=NREUM.o.REQ,y=window.XMLHttpRequest;a.features.xhr=!0,t(15),t(8),u.on("new-xhr",function(t){var e=this;e.totalCbs=0,e.called=0,e.cbTime=0,e.end=r,e.ended=!1,e.xhrGuids={},e.lastSize=null,e.loadCaptureCalled=!1,e.params=this.params||{},e.metrics=this.metrics||{},t.addEventListener("load",function(n){i(e,t)},w(!1)),h&&(h>34||h<10)||t.addEventListener("progress",function(t){e.lastSize=t.loaded},w(!1))}),u.on("open-xhr-start",function(t){this.params={method:t[0]},o(this,t[1]),this.metrics={}}),u.on("open-xhr-end",function(t,e){"loader_config"in NREUM&&"xpid"in NREUM.loader_config&&this.sameOrigin&&e.setRequestHeader("X-NewRelic-ID",NREUM.loader_config.xpid);var n=f(this.parsedOrigin);if(n){var r=!1;n.newrelicHeader&&(e.setRequestHeader("newrelic",n.newrelicHeader),r=!0),n.traceContextParentHeader&&(e.setRequestHeader("traceparent",n.traceContextParentHeader),n.traceContextStateHeader&&e.setRequestHeader("tracestate",n.traceContextStateHeader),r=!0),r&&(this.dt=n)}}),u.on("send-xhr-start",function(t,e){var n=this.metrics,r=t[0],o=this;if(n&&r){var i=m(r);i&&(n.txSize=i)}this.startTime=a.now(),this.listener=function(t){try{"abort"!==t.type||o.loadCaptureCalled||(o.params.aborted=!0),("load"!==t.type||o.called===o.totalCbs&&(o.onloadCalled||"function"!=typeof e.onload))&&o.end(e)}catch(n){try{u.emit("internal-error",[n])}catch(r){}}};for(var s=0;s<p;s++)e.addEventListener(d[s],this.listener,w(!1))}),u.on("xhr-cb-time",function(t,e,n){this.cbTime+=t,e?this.onloadCalled=!0:this.called+=1,this.called!==this.totalCbs||!this.onloadCalled&&"function"==typeof n.onload||this.end(n)}),u.on("xhr-load-added",function(t,e){var n=""+l(t)+!!e;this.xhrGuids&&!this.xhrGuids[n]&&(this.xhrGuids[n]=!0,this.totalCbs+=1)}),u.on("xhr-load-removed",function(t,e){var n=""+l(t)+!!e;this.xhrGuids&&this.xhrGuids[n]&&(delete this.xhrGuids[n],this.totalCbs-=1)}),u.on("xhr-resolved",function(){this.endTime=a.now()}),u.on("addEventListener-end",function(t,e){e instanceof y&&"load"===t[0]&&u.emit("xhr-load-added",[t[1],t[2]],e)}),u.on("removeEventListener-end",function(t,e){e instanceof y&&"load"===t[0]&&u.emit("xhr-load-removed",[t[1],t[2]],e)}),u.on("fn-start",function(t,e,n){e instanceof y&&("onload"===n&&(this.onload=!0),("load"===(t[0]&&t[0].type)||this.onload)&&(this.xhrCbStart=a.now()))}),u.on("fn-end",function(t,e){this.xhrCbStart&&u.emit("xhr-cb-time",[a.now()-this.xhrCbStart,this.onload,e],e)}),u.on("fetch-before-start",function(t){function e(t,e){var n=!1;return e.newrelicHeader&&(t.set("newrelic",e.newrelicHeader),n=!0),e.traceContextParentHeader&&(t.set("traceparent",e.traceContextParentHeader),e.traceContextStateHeader&&t.set("tracestate",e.traceContextStateHeader),n=!0),n}var n,r=t[1]||{};"string"==typeof t[0]?n=t[0]:t[0]&&t[0].url?n=t[0].url:window.URL&&t[0]&&t[0]instanceof URL&&(n=t[0].href),n&&(this.parsedOrigin=c(n),this.sameOrigin=this.parsedOrigin.sameOrigin);var o=f(this.parsedOrigin);if(o&&(o.newrelicHeader||o.traceContextParentHeader))if("string"==typeof t[0]||window.URL&&t[0]&&t[0]instanceof URL){var i={};for(var a in r)i[a]=r[a];i.headers=new Headers(r.headers||{}),e(i.headers,o)&&(this.dt=o),t.length>1?t[1]=i:t.push(i)}else t[0]&&t[0].headers&&e(t[0].headers,o)&&(this.dt=o)}),u.on("fetch-start",function(t,e){this.params={},this.metrics={},this.startTime=a.now(),this.dt=e,t.length>=1&&(this.target=t[0]),t.length>=2&&(this.opts=t[1]);var n,r=this.opts||{},i=this.target;"string"==typeof i?n=i:"object"==typeof i&&i instanceof g?n=i.url:window.URL&&"object"==typeof i&&i instanceof URL&&(n=i.href),o(this,n);var s=(""+(i&&i instanceof g&&i.method||r.method||"GET")).toUpperCase();this.params.method=s,this.txSize=m(r.body)||0}),u.on("fetch-done",function(t,e){this.endTime=a.now(),this.params||(this.params={}),this.params.status=e?e.status:0;var n;"string"==typeof this.rxSize&&this.rxSize.length>0&&(n=+this.rxSize);var r={txSize:this.txSize,rxSize:n,duration:a.now()-this.startTime};s("xhr",[this.params,r,this.startTime,this.endTime,"fetch"],this)})}},{}],18:[function(t,e,n){var r={};e.exports=function(t){if(t in r)return r[t];var e=document.createElement("a"),n=window.location,o={};e.href=t,o.port=e.port;var i=e.href.split("://");!o.port&&i[1]&&(o.port=i[1].split("/")[0].split("@").pop().split(":")[1]),o.port&&"0"!==o.port||(o.port="https"===i[0]?"443":"80"),o.hostname=e.hostname||n.hostname,o.pathname=e.pathname,o.protocol=i[0],"/"!==o.pathname.charAt(0)&&(o.pathname="/"+o.pathname);var a=!e.protocol||":"===e.protocol||e.protocol===n.protocol,s=e.hostname===document.domain&&e.port===n.port;return o.sameOrigin=a&&(!e.hostname||s),"/"===o.pathname&&(r[t]=o),o}},{}],19:[function(t,e,n){function r(t,e){var n=t.responseType;return"json"===n&&null!==e?e:"arraybuffer"===n||"blob"===n||"json"===n?o(t.response):"text"===n||""===n||void 0===n?o(t.responseText):void 0}var o=t(22);e.exports=r},{}],20:[function(t,e,n){function r(){}function o(t,e,n,r){return function(){return u.recordSupportability("API/"+e+"/called"),i(t+e,[f.now()].concat(s(arguments)),n?null:this,r),n?void 0:this}}var i=t("handle"),a=t(32),s=t(33),c=t("ee").get("tracer"),f=t("loader"),u=t(25),d=NREUM;"undefined"==typeof window.newrelic&&(newrelic=d);var p=["setPageViewName","setCustomAttribute","setErrorHandler","finished","addToTrace","inlineHit","addRelease"],l="api-",h=l+"ixn-";a(p,function(t,e){d[e]=o(l,e,!0,"api")}),d.addPageAction=o(l,"addPageAction",!0),d.setCurrentRouteName=o(l,"routeName",!0),e.exports=newrelic,d.interaction=function(){return(new r).get()};var m=r.prototype={createTracer:function(t,e){var n={},r=this,o="function"==typeof e;return i(h+"tracer",[f.now(),t,n],r),function(){if(c.emit((o?"":"no-")+"fn-start",[f.now(),r,o],n),o)try{return e.apply(this,arguments)}catch(t){throw c.emit("fn-err",[arguments,this,t],n),t}finally{c.emit("fn-end",[f.now()],n)}}}};a("actionText,setName,setAttribute,save,ignore,onEnd,getContext,end,get".split(","),function(t,e){m[e]=o(h,e)}),newrelic.noticeError=function(t,e){"string"==typeof t&&(t=new Error(t)),u.recordSupportability("API/noticeError/called"),i("err",[t,f.now(),!1,e])}},{}],21:[function(t,e,n){function r(t){if(NREUM.init){for(var e=NREUM.init,n=t.split("."),r=0;r<n.length-1;r++)if(e=e[n[r]],"object"!=typeof e)return;return e=e[n[n.length-1]]}}e.exports={getConfiguration:r}},{}],22:[function(t,e,n){e.exports=function(t){if("string"==typeof t&&t.length)return t.length;if("object"==typeof t){if("undefined"!=typeof ArrayBuffer&&t instanceof ArrayBuffer&&t.byteLength)return t.byteLength;if("undefined"!=typeof Blob&&t instanceof Blob&&t.size)return t.size;if(!("undefined"!=typeof FormData&&t instanceof FormData))try{return JSON.stringify(t).length}catch(e){return}}}},{}],23:[function(t,e,n){var r=!1;try{var o=Object.defineProperty({},"passive",{get:function(){r=!0}});window.addEventListener("testPassive",null,o),window.removeEventListener("testPassive",null,o)}catch(i){}e.exports=function(t){return r?{passive:!0,capture:!!t}:!!t}},{}],24:[function(t,e,n){var r=0,o=navigator.userAgent.match(/Firefox[\/\s](\d+\.\d+)/);o&&(r=+o[1]),e.exports=r},{}],25:[function(t,e,n){function r(t,e){var n=[a,t,{name:t},e];return i("storeMetric",n,null,"api"),n}function o(t,e){var n=[s,t,{name:t},e];return i("storeEventMetrics",n,null,"api"),n}var i=t("handle"),a="sm",s="cm";e.exports={constants:{SUPPORTABILITY_METRIC:a,CUSTOM_METRIC:s},recordSupportability:r,recordCustom:o}},{}],26:[function(t,e,n){function r(){return s.exists&&performance.now?Math.round(performance.now()):(i=Math.max((new Date).getTime(),i))-a}function o(){return i}var i=(new Date).getTime(),a=i,s=t(34);e.exports=r,e.exports.offset=a,e.exports.getLastTimestamp=o},{}],27:[function(t,e,n){function r(t){return!(!t||!t.protocol||"file:"===t.protocol)}e.exports=r},{}],28:[function(t,e,n){function r(t,e){var n=t.getEntries();n.forEach(function(t){"first-paint"===t.name?p("timing",["fp",Math.floor(t.startTime)]):"first-contentful-paint"===t.name&&p("timing",["fcp",Math.floor(t.startTime)])})}function o(t,e){var n=t.getEntries();if(n.length>0){var r=n[n.length-1];if(c&&c<r.startTime)return;p("lcp",[r])}}function i(t){t.getEntries().forEach(function(t){t.hadRecentInput||p("cls",[t])})}function a(t){if(t instanceof v&&!g){var e=Math.round(t.timeStamp),n={type:t.type};e<=l.now()?n.fid=l.now()-e:e>l.offset&&e<=Date.now()?(e-=l.offset,n.fid=l.now()-e):e=l.now(),g=!0,p("timing",["fi",e,n])}}function s(t){"hidden"===t&&(c=l.now(),p("pageHide",[c]))}if(!("init"in NREUM&&"page_view_timing"in NREUM.init&&"enabled"in NREUM.init.page_view_timing&&NREUM.init.page_view_timing.enabled===!1)){var c,f,u,d,p=t("handle"),l=t("loader"),h=t(31),m=t(23),v=NREUM.o.EV;if("PerformanceObserver"in window&&"function"==typeof window.PerformanceObserver){f=new PerformanceObserver(r);try{f.observe({entryTypes:["paint"]})}catch(w){}u=new PerformanceObserver(o);try{u.observe({entryTypes:["largest-contentful-paint"]})}catch(w){}d=new PerformanceObserver(i);try{d.observe({type:"layout-shift",buffered:!0})}catch(w){}}if("addEventListener"in document){var g=!1,y=["click","keydown","mousedown","pointerdown","touchstart"];y.forEach(function(t){document.addEventListener(t,a,m(!1))})}h(s)}},{}],29:[function(t,e,n){function r(){function t(){return e?15&e[n++]:16*Math.random()|0}var e=null,n=0,r=window.crypto||window.msCrypto;r&&r.getRandomValues&&(e=r.getRandomValues(new Uint8Array(31)));for(var o,i="xxxxxxxx-xxxx-4xxx-yxxx-xxxxxxxxxxxx",a="",s=0;s<i.length;s++)o=i[s],"x"===o?a+=t().toString(16):"y"===o?(o=3&t()|8,a+=o.toString(16)):a+=o;return a}function o(){return a(16)}function i(){return a(32)}function a(t){function e(){return n?15&n[r++]:16*Math.random()|0}var n=null,r=0,o=window.crypto||window.msCrypto;o&&o.getRandomValues&&Uint8Array&&(n=o.getRandomValues(new Uint8Array(31)));for(var i=[],a=0;a<t;a++)i.push(e().toString(16));return i.join("")}e.exports={generateUuid:r,generateSpanId:o,generateTraceId:i}},{}],30:[function(t,e,n){function r(t,e){if(!o)return!1;if(t!==o)return!1;if(!e)return!0;if(!i)return!1;for(var n=i.split("."),r=e.split("."),a=0;a<r.length;a++)if(r[a]!==n[a])return!1;return!0}var o=null,i=null,a=/Version\/(\S+)\s+Safari/;if(navigator.userAgent){var s=navigator.userAgent,c=s.match(a);c&&s.indexOf("Chrome")===-1&&s.indexOf("Chromium")===-1&&(o="Safari",i=c[1])}e.exports={agent:o,version:i,match:r}},{}],31:[function(t,e,n){function r(t){function e(){t(s&&document[s]?document[s]:document[i]?"hidden":"visible")}"addEventListener"in document&&a&&document.addEventListener(a,e,o(!1))}var o=t(23);e.exports=r;var i,a,s;"undefined"!=typeof document.hidden?(i="hidden",a="visibilitychange",s="visibilityState"):"undefined"!=typeof document.msHidden?(i="msHidden",a="msvisibilitychange"):"undefined"!=typeof document.webkitHidden&&(i="webkitHidden",a="webkitvisibilitychange",s="webkitVisibilityState")},{}],32:[function(t,e,n){function r(t,e){var n=[],r="",i=0;for(r in t)o.call(t,r)&&(n[i]=e(r,t[r]),i+=1);return n}var o=Object.prototype.hasOwnProperty;e.exports=r},{}],33:[function(t,e,n){function r(t,e,n){e||(e=0),"undefined"==typeof n&&(n=t?t.length:0);for(var r=-1,o=n-e||0,i=Array(o<0?0:o);++r<o;)i[r]=t[e+r];return i}e.exports=r},{}],34:[function(t,e,n){e.exports={exists:"undefined"!=typeof window.performance&&window.performance.timing&&"undefined"!=typeof window.performance.timing.navigationStart}},{}],ee:[function(t,e,n){function r(){}function o(t){function e(t){return t&&t instanceof r?t:t?f(t,c,a):a()}function n(n,r,o,i,a){if(a!==!1&&(a=!0),!l.aborted||i){t&&a&&t(n,r,o);for(var s=e(o),c=m(n),f=c.length,u=0;u<f;u++)c[u].apply(s,r);var p=d[y[n]];return p&&p.push([x,n,r,s]),s}}function i(t,e){g[t]=m(t).concat(e)}function h(t,e){var n=g[t];if(n)for(var r=0;r<n.length;r++)n[r]===e&&n.splice(r,1)}function m(t){return g[t]||[]}function v(t){return p[t]=p[t]||o(n)}function w(t,e){l.aborted||u(t,function(t,n){e=e||"feature",y[n]=e,e in d||(d[e]=[])})}var g={},y={},x={on:i,addEventListener:i,removeEventListener:h,emit:n,get:v,listeners:m,context:e,buffer:w,abort:s,aborted:!1};return x}function i(t){return f(t,c,a)}function a(){return new r}function s(){(d.api||d.feature)&&(l.aborted=!0,d=l.backlog={})}var c="nr@context",f=t("gos"),u=t(32),d={},p={},l=e.exports=o();e.exports.getOrSetContext=i,l.backlog=d},{}],gos:[function(t,e,n){function r(t,e,n){if(o.call(t,e))return t[e];var r=n();if(Object.defineProperty&&Object.keys)try{return Object.defineProperty(t,e,{value:r,writable:!0,enumerable:!1}),r}catch(i){}return t[e]=r,r}var o=Object.prototype.hasOwnProperty;e.exports=r},{}],handle:[function(t,e,n){function r(t,e,n,r){o.buffer([t],r),o.emit(t,e,n)}var o=t("ee").get("handle");e.exports=r,r.ee=o},{}],id:[function(t,e,n){function r(t){var e=typeof t;return!t||"object"!==e&&"function"!==e?-1:t===window?0:a(t,i,function(){return o++})}var o=1,i="nr@id",a=t("gos");e.exports=r},{}],loader:[function(t,e,n){function r(){if(!P++){var t=T.info=NREUM.info,e=v.getElementsByTagName("script")[0];if(setTimeout(f.abort,3e4),!(t&&t.licenseKey&&t.applicationID&&e))return f.abort();c(R,function(e,n){t[e]||(t[e]=n)});var n=a();s("mark",["onload",n+T.offset],null,"api"),s("timing",["load",n]);var r=v.createElement("script");0===t.agent.indexOf("http://")||0===t.agent.indexOf("https://")?r.src=t.agent:r.src=h+"://"+t.agent,e.parentNode.insertBefore(r,e)}}function o(){"complete"===v.readyState&&i()}function i(){s("mark",["domContent",a()+T.offset],null,"api")}var a=t(26),s=t("handle"),c=t(32),f=t("ee"),u=t(30),d=t(27),p=t(21),l=t(23),h=p.getConfiguration("ssl")===!1?"http":"https",m=window,v=m.document,w="addEventListener",g="attachEvent",y=m.XMLHttpRequest,x=y&&y.prototype,b=!d(m.location);NREUM.o={ST:setTimeout,SI:m.setImmediate,CT:clearTimeout,XHR:y,REQ:m.Request,EV:m.Event,PR:m.Promise,MO:m.MutationObserver};var E=""+location,R={beacon:"bam.nr-data.net",errorBeacon:"bam.nr-data.net",agent:"js-agent.newrelic.com/nr-spa-1212.min.js"},O=y&&x&&x[w]&&!/CriOS/.test(navigator.userAgent),T=e.exports={offset:a.getLastTimestamp(),now:a,origin:E,features:{},xhrWrappable:O,userAgent:u,disabled:b};if(!b){t(20),t(28),v[w]?(v[w]("DOMContentLoaded",i,l(!1)),m[w]("load",r,l(!1))):(v[g]("onreadystatechange",o),m[g]("onload",r)),s("mark",["firstbyte",a.getLastTimestamp()],null,"api");var P=0}},{}],"wrap-function":[function(t,e,n){function r(t,e){function n(e,n,r,c,f){function nrWrapper(){var i,a,u,p;try{a=this,i=d(arguments),u="function"==typeof r?r(i,a):r||{}}catch(l){o([l,"",[i,a,c],u],t)}s(n+"start",[i,a,c],u,f);try{return p=e.apply(a,i)}catch(h){throw s(n+"err",[i,a,h],u,f),h}finally{s(n+"end",[i,a,p],u,f)}}return a(e)?e:(n||(n=""),nrWrapper[p]=e,i(e,nrWrapper,t),nrWrapper)}function r(t,e,r,o,i){r||(r="");var s,c,f,u="-"===r.charAt(0);for(f=0;f<e.length;f++)c=e[f],s=t[c],a(s)||(t[c]=n(s,u?c+r:r,o,c,i))}function s(n,r,i,a){if(!h||e){var s=h;h=!0;try{t.emit(n,r,i,e,a)}catch(c){o([c,n,r,i],t)}h=s}}return t||(t=u),n.inPlace=r,n.flag=p,n}function o(t,e){e||(e=u);try{e.emit("internal-error",t)}catch(n){}}function i(t,e,n){if(Object.defineProperty&&Object.keys)try{var r=Object.keys(t);return r.forEach(function(n){Object.defineProperty(e,n,{get:function(){return t[n]},set:function(e){return t[n]=e,e}})}),e}catch(i){o([i],n)}for(var a in t)l.call(t,a)&&(e[a]=t[a]);return e}function a(t){return!(t&&t instanceof Function&&t.apply&&!t[p])}function s(t,e){var n=e(t);return n[p]=t,i(t,n,u),n}function c(t,e,n){var r=t[e];t[e]=s(r,n)}function f(){for(var t=arguments.length,e=new Array(t),n=0;n<t;++n)e[n]=arguments[n];return e}var u=t("ee"),d=t(33),p="nr@original",l=Object.prototype.hasOwnProperty,h=!1;e.exports=r,e.exports.wrapFunction=s,e.exports.wrapInPlace=c,e.exports.argsToArray=f},{}]},{},["loader",2,17,5,3,4]); ;NREUM.loader_config={accountID:"804283",trustKey:"804283",agentID:"402703674",licenseKey:"cf99e8d2a3",applicationID:"402703674"} ;NREUM.info={beacon:"bam.nr-data.net",errorBeacon:"bam.nr-data.net",licenseKey:"cf99e8d2a3", // Modified this value from the generated script, to pass prod vs dev applicationID: window.location.hostname.includes('journals.plos.org') ? "402703674" : "402694889", sa:1} </script> <!-- End New Relic --> <header> <div id="topslot" class="head-top"> <a id="skip-to-content" tabindex="0" class="button" href="#main-content"> Skip to main content </a> <div class="center"> <div class="title">Advertisement</div> <!-- DoubleClick Ad Zone --> <div class='advertisement' id='div-gpt-ad-1458247671871-0' style='width:728px; height:90px;'> <script type='text/javascript'> googletag.cmd.push(function() { googletag.display('div-gpt-ad-1458247671871-0'); }); </script> </div> </div> </div> <div id="user" class="nav" data-user-management-url="https://community.plos.org"> </div> <div id="pagehdr"> <nav class="nav-main"> <h1 class="logo"> <a href="/plosbiology/.">PLOS Biology</a> </h1> <section class="top-bar-section"> <ul class="nav-elements"> <li class="menu-section-header has-dropdown " id="browse"> <span class="menu-section-header-title"> Browse </span> <ul class="menu-section dropdown " id="browse-dropdown-list"> <li> <a href="/plosbiology/issue" >Current Issue</a> </li> <li> <a href="/plosbiology/volume" >Journal Archive</a> </li> <li> <a href="/plosbiology/s/collections" >Collections</a> </li> <li> <a href="/plosbiology/s/find-and-read-articles" >Find and Read Articles</a> </li> </ul> </li> <li class="multi-col-parent menu-section-header has-dropdown" id="publish"> Publish <div class="dropdown mega "> <ul class="multi-col" id="publish-dropdown-list"> <li class="menu-section-header " id="submissions"> <span class="menu-section-header-title"> Submissions </span> <ul class="menu-section " id="submissions-dropdown-list"> <li> <a href="/plosbiology/s/submission-guidelines" >Submission Guidelines</a> </li> <li> <a href="/plosbiology/s/figures" >Figures</a> </li> <li> <a href="/plosbiology/s/tables" >Tables</a> </li> <li> <a href="/plosbiology/s/supporting-information" >Supporting Information</a> </li> <li> <a href="/plosbiology/s/latex" >LaTeX</a> </li> <li> <a href="/plosbiology/s/what-we-publish" >What We Publish</a> </li> <li> <a href="/plosbiology/s/preprints" >Preprints</a> </li> <li> <a href="/plosbiology/s/revising-your-manuscript" >Revising Your Manuscript</a> </li> <li> <a href="/plosbiology/s/submit-now" >Submit Now</a> </li> </ul> </li> <li class="menu-section-header " id="policies"> <span class="menu-section-header-title"> Policies </span> <ul class="menu-section " id="policies-dropdown-list"> <li> <a href="/plosbiology/s/best-practices-in-research-reporting" >Best Practices in Research Reporting</a> </li> <li> <a href="/plosbiology/s/human-subjects-research" >Human Subjects Research</a> </li> <li> <a href="/plosbiology/s/animal-research" >Animal Research</a> </li> <li> <a href="/plosbiology/s/competing-interests" >Competing Interests</a> </li> <li> <a href="/plosbiology/s/disclosure-of-funding-sources" >Disclosure of Funding Sources</a> </li> <li> <a href="/plosbiology/s/licenses-and-copyright" >Licenses and Copyright</a> </li> <li> <a href="/plosbiology/s/data-availability" >Data Availability</a> </li> <li> <a href="/plosbiology/s/complementary-research" >Complementary Research</a> </li> <li> <a href="/plosbiology/s/materials-software-and-code-sharing" >Materials, Software and Code Sharing</a> </li> <li> <a href="/plosbiology/s/ethical-publishing-practice" >Ethical Publishing Practice</a> </li> <li> <a href="/plosbiology/s/authorship" >Authorship</a> </li> <li> <a href="/plosbiology/s/corrections-expressions-of-concern-and-retractions" >Corrections, Expressions of Concern, and Retractions</a> </li> </ul> </li> <li class="menu-section-header " id="manuscript-review-and-publication"> <span class="menu-section-header-title"> Manuscript Review and Publication </span> <ul class="menu-section " id="manuscript-review-and-publication-dropdown-list"> <li> <a href="/plosbiology/s/editorial-and-peer-review-process" >Editorial and Peer Review Process</a> </li> <li> <a href="/plosbiology/s/reviewer-guidelines" >Guidelines for Reviewers</a> </li> <li> <a href="/plosbiology/s/accepted-manuscripts" >Accepted Manuscripts</a> </li> <li> <a href="/plosbiology/s/comments" >Comments</a> </li> </ul> </li> </ul> <div class="calloutcontainer"> <h3 class="callout-headline">Submit Your Manuscript</h3> <div class="action-contain"> <p class="callout-content"> <em>PLOS Biology</em> provides an Open Access platform to showcase your best research and commentary across all areas of biological science. </p> <p class="button-contain special"> <a class="button button-default" href="/plosbiology/s/submit-now"> Submit Now </a> </p> </div> <!-- opens in siteMenuCalloutDescription --> </div> </div> </li> <li class="menu-section-header has-dropdown " id="about"> <span class="menu-section-header-title"> About </span> <ul class="menu-section dropdown " id="about-dropdown-list"> <li> <a href="/plosbiology/static/publish" >Why Publish with PLOS Biology</a> </li> <li> <a href="/plosbiology/s/journal-information" >Journal Information</a> </li> <li> <a href="/plosbiology/s/staff-editors" >Staff Editors</a> </li> <li> <a href="/plosbiology/s/editorial-board" >Editorial Board</a> </li> <li> <a href="/plosbiology/s/publishing-information" >Publishing Information</a> </li> <li> <a href="https://plos.org/publication-fees" >Publication Fees</a> </li> <li> <a href="https://plos.org/press-and-media" >Press and Media</a> </li> <li> <a href="/plosbiology/s/contact" >Contact</a> </li> </ul> </li> <script src="/resource/js/vendor/jquery.hoverIntent.js" type="text/javascript"></script> <script src="/resource/js/components/menu_drop.js" type="text/javascript"></script> <script src="/resource/js/components/hover_delay.js" type="text/javascript"></script> <li id="navsearch" class="head-search"> <form name="searchForm" action="/plosbiology/search" method="get"> <fieldset> <legend>Search</legend> <label for="search">Search</label> <div class="search-contain"> <input id="search" type="text" name="q" placeholder="SEARCH" required/> <button id="headerSearchButton" type="submit" aria-label="Submit search"> <i title="Submit search" class="search-icon"></i> </button> </div> </fieldset> <input type="hidden" name="filterJournals" value="PLoSBiology"/> </form> <a id="advSearch" href="/plosbiology/search"> advanced search </a> <script src="/resource/js/components/placeholder_style.js" type="text/javascript"></script> </li> </ul> </section> </nav> </div> </header> <main id="main-content"> <div id="home-content"> <div class="row top"> <div class="hero light-text overlay-dark" id="hero"> <a alt="Sonic hedgehog and feather branching" href="/plosbiology/article?id=10.1371/journal.pbio.3003061"><img alt="Sonic hedgehog and feather branching" data-file-extension="jpg" src="/plosbiology/indirect/a184cc3b099781cd56a974c587bba63f66414f5ab2c277c3874a362a5e7114d6"> </a> <div class="hero-content"> <p class="published_at">March 20, 2025</p> <h1 class="title"><a alt="Sonic hedgehog and feather branching" href="/plosbiology/article?id=10.1371/journal.pbio.3003061">Sonic hedgehog and feather branching</a></h1> <div class="blurb authors"> <p>The Sonic Hedgehog (Shh) pathway is a key regulator of feather development. Rory Cooper and Michel Milinkovitch show that <i>in vivo</i> Shh inhibition during early chicken embryogenesis temporarily results in unbranched and non-invaginated embryonic feather buds, similar to those seen in some early dinosaur ancestors of birds.</p> </div> <p class="credit">Image credit: Rory Cooper & Michel Milinkovitch</p> </div> </div> <div id="billboard"> <h3>PLOS Biologue</h3> <h4>Community blog for PLOS Biology, PLOS Genetics and PLOS Computational Biology.</h4><a href="https://biologue.plos.org/">PLOS BIOLOGUE</a> </div> </div> <div class="row one"> <article class="block medium image" id="cellcat-1-1"> <div class="block-content"> <p class="date">03/21/2025</p> <p class="category">Research Article</p> <h2 class="title"><a alt="Navigation strategy learning in worms" href="/plosbiology/article?id=10.1371/journal.pbio.3003005">Navigation strategy learning in worms</a></h2> <div class="blurb"> <p>How do worms navigate based on learned experience? Kevin Chen, Anuj Sharma, Jonathan Pillow and Andrew Leifer reveal the flexible behavioral strategies and distributed neural computations that underlie learning-dependent odor navigation in <i>Caenorhabditis elegans</i>.</p> </div> </div> <p class="credits">Image credit: pbio.3003005</p><a class="block-image" alt="Navigation strategy learning in worms" href="/plosbiology/article?id=10.1371/journal.pbio.3003005"><img alt="Navigation strategy learning in worms" data-file-extension="jpg" src="/plosbiology/indirect/6f678c07837e451383f5c77204715df99bad2ffbc62bc2bd791e32fbceefed13"></a> </article> <div id="article-list"> <h3>Recently Published Articles</h3> <section> <ul id="article-results" class="list-plain results"> <li><a href="article?id=10.1371/journal.pbio.3003089" data-tooltip class="truncated-tooltip" title="Navigating your US bioscience career into the 2030s">Navigating your US bioscience career into the 2030s</a></li> <li><a href="article?id=10.1371/journal.pbio.3003107" data-tooltip class="truncated-tooltip" title="Editorial Note: How sex chromosomes get trapped into nonrecombination">Editorial Note: How sex chromosomes get trapped into nonrecombination</a></li> <li><a href="article?id=10.1371/journal.pbio.3003018" data-tooltip class="truncated-tooltip" title="Neural mechanisms for the attention-mediated propagation of conceptual information in the human brain">Neural mechanisms for the attention-mediated propagation of conceptual information in the human brain</a></li> </ul> </section> <div class="more-link"> <a href="search?sortOrder=DATE_NEWEST_FIRST&filterStartDate=2025-02-27&filterEndDate=2025-03-29&filterJournals=PLoSBiology&q=">See all articles</a> </div> <script src="/resource/js/vendor/jquery.dotdotdot.js" type="text/javascript"></script> </div> <div id="issue"> <a href="/plosbiology/issue"> <img src="/plosbiology/article/figure/image?size=medium&id=10.1371/image.pbio.v23.i02.g001" class="current-img" alt="Current Issue"> </a> <p class="boxtitle"><a href="/plosbiology/issue">Current Issue</a> <span class="subhead">February 2025</span></p> </div> </div> <div class="row two"> <article class="block medium image" id="cellcat-1-2"> <div class="block-content"> <p class="date">03/20/2025</p> <p class="category">Research Article</p> <h2 class="title"><a alt="Enhanced memory and sleep" href="/plosbiology/article?id=10.1371/journal.pbio.3003076">Enhanced memory and sleep</a></h2> <div class="blurb"> <p>Sleep is known to support memory; however, Sheng Huang, Chengji Piao, Stephan Sigrist and colleagues identify an instance where memory hyperfunction may actually provoke sleep deficits. They find that the short-sleep <i>Drosophila </i>mutant, <i>insomniac</i>, has enhanced memory and that PKA signaling balances sleep and memory in these flies.</p> </div> </div> <p class="credits">Image credit: pbio.3003076</p><a class="block-image" alt="Enhanced memory and sleep" href="/plosbiology/article?id=10.1371/journal.pbio.3003076"><img alt="Enhanced memory and sleep" data-file-extension="jpg" src="/plosbiology/indirect/7bfd0d905bf082b5a276474756d5451eebc9452489a8cae77153d463d3e8a63a"></a> </article> <article class="block medium image" id="cellcat-1-3"> <div class="block-content"> <p class="date">03/18/2025</p> <p class="category">Short Reports</p> <h2 class="title"><a alt="Outomes of past conservation efforts" href="/plosbiology/article?id=10.1371/journal.pbio.3003051">Outomes of past conservation efforts</a></h2> <div class="blurb"> <p>Understanding the consequences of past conservation efforts is essential to inform how we maintain and restore species. An analysis of IUCN Red List data on 67,217 animal species, by Ashley Simkins, Silviu Petrovan and co-workers, reveals that a range of different conservation actions have successfully conserved species that are at greatest risk of extinction, but have rarely resulted in full recovery.</p> </div> </div> <p class="credits">Image credit: BR Ansil</p><a class="block-image" alt="Outomes of past conservation efforts" href="/plosbiology/article?id=10.1371/journal.pbio.3003051"><img alt="Outomes of past conservation efforts" data-file-extension="jpg" src="/plosbiology/indirect/c5b351fe56d0e92aba2873d14d09010e75092dd4b46dad002c1f9603961e11dd"></a> </article> </div> <div class="row three"> <div class="column left"> <article class="block large image" id="cellcat2-1"> <div class="block-content"> <p class="date">03/17/2025</p> <p class="category">Short Reports</p> <h2 class="title"><a alt="Double-bladed bacterial toxins" href="/plosbiology/article?id=10.1371/journal.pbio.3003053">Double-bladed bacterial toxins</a></h2> <div class="blurb"> <p>The secretion mechanisms of many type VI secretion system (T6SS) effectors in Gram-negative bacteria remain unclear. Chaya Mushka Fridman, Eran Bosis, Dor Salomon and colleagues identify a new class of T6SS effectors which can harbor either one or two toxic domains and use WHIX as a secretion motif.</p> </div> </div><a class="block-image" alt="Double-bladed bacterial toxins" href="/plosbiology/article?id=10.1371/journal.pbio.3003053"><img alt="Double-bladed bacterial toxins" data-file-extension="jpg" src="/plosbiology/indirect/e09f5a36e2707504ee1c0dda6b5a3f7fdcb2d17aea119e76220c1829bd235de5"></a> <p class="credits">Image credit: pbio.3003053</p> </article> <article class="block large image" id="cellcat2-2"> <div class="block-content"> <p class="date">03/12/2025</p> <p class="category">Primer</p> <h2 class="title"><a alt="Is beauty beyond the eye of the butterfly?" href="/plosbiology/article?id=10.1371/journal.pbio.3003054">Is beauty beyond the eye of the butterfly?</a></h2> <div class="blurb"> <p>The diversity of bright colors observed across the animal world are often used during mate choice. Richard Merrill explores <a href="https://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.3002989">a new study</a> in PLOS Biology which reveals genetic and neural mechanisms contributing to the evolution of visual mating decisions in <i>Heliconius</i> butterflies.</p> </div> </div><a class="block-image" alt="Is beauty beyond the eye of the butterfly?" href="/plosbiology/article?id=10.1371/journal.pbio.3003054"><img alt="Is beauty beyond the eye of the butterfly?" data-file-extension="jpg" src="/plosbiology/indirect/1e501612c9fc045c198e25eae5af2281a33bec477b013b8db02431d1ae313fcf"></a> <p class="credits">Image credit: pbio.3003054</p> </article> <article class="block large image" id="cellcat2-3"> <div class="block-content"> <p class="date">03/11/2025</p> <p class="category">Research Article</p> <h2 class="title"><a alt="Signals from the niche..." href="/plosbiology/article?id=10.1371/journal.pbio.3003049">Signals from the niche...</a></h2> <div class="blurb"> <p>Transcription is known to tightly regulate gene expression profiles in stem and differentiating cells, but what about translational control? Ruoxu Wang, Marc Amoyel anc co-workers reveal that signals from the stem cell niche induce changes in translational rates, regulated via eIF3d1 phosphorylation, to control stem cell fate in the <i>Drosophila </i>testis.</p> </div> </div><a class="block-image" alt="Signals from the niche..." href="/plosbiology/article?id=10.1371/journal.pbio.3003049"><img alt="Signals from the niche..." data-file-extension="jpg" src="/plosbiology/indirect/160212d6be9c3387853a4a8cddf5d654a02cf82c5a0c5a611d47ae13470efa40"></a> <p class="credits">Image credit: pbio.3003049</p> </article> <div class="row-small-blocks"> <article class="block small image" id="cellcat3-1"> <div class="block-content"> <p class="date">03/19/2025</p> <p class="category">Community Page</p> <h2 class="title"><a alt="The IBEX Knowledge-Base" href="/plosbiology/article?id=10.1371/journal.pbio.3003070">The IBEX Knowledge-Base</a></h2> <div class="blurb"> <p>Multiplexed imaging is a powerful approach in spatial biology. This Community Page presents the IBEX Knowledge-Base, a central resource for reagents, protocols and more, to enhance knowledge sharing, optimization and innovation of spatial proteomics techniques.</p> </div> </div><a class="block-image" alt="The IBEX Knowledge-Base" href="/plosbiology/article?id=10.1371/journal.pbio.3003070"><img alt="The IBEX Knowledge-Base" data-file-extension="jpg" src="/plosbiology/indirect/6f8fcabe132214790a787e0737b05ff010d7f6c287a89bd992c2914b5de1f449"></a> <p class="credits">Image credit: pbio.3003070</p> </article> <article class="block small image" id="cellcat3-2"> <div class="block-content"> <p class="date">03/18/2025</p> <p class="category">Perspective</p> <h2 class="title"><a alt="Vector microbiota collections" href="/plosbiology/article?id=10.1371/journal.pbio.3003078">Vector microbiota collections</a></h2> <div class="blurb"> <p>This article outlines how collections of microbial isolate from insect vectors aid in identifying microbial targets, and why their implementation must rely on coordinated international efforts that are ethical and ensure equitable benefit sharing.</p> </div> </div><a class="block-image" alt="Vector microbiota collections" href="/plosbiology/article?id=10.1371/journal.pbio.3003078"><img alt="Vector microbiota collections" data-file-extension="jpg" src="/plosbiology/indirect/8f8e8f8ab61a8812b0e591e0c98493e5b110767b57a860245875c205316099ea"></a> <p class="credits">Image credit: pbio.3003078</p> </article> <article class="block small image" id="cellcat3-3"> <div class="block-content"> <p class="date">03/18/2025</p> <p class="category">Essay</p> <h2 class="title"><a alt="The end of the genetic paradigm of cancer" href="/plosbiology/article?id=10.1371/journal.pbio.3003052">The end of the genetic paradigm of cancer</a></h2> <div class="blurb"> <p>Sui Huang, Ana Soto and Carlos Sonnenschein explore cancer omics data that challenge the somatic mutation theory, and present theories based on gene regulatory networks and tissue fields that help explain inconsistencies in the data.</p> </div> </div><a class="block-image" alt="The end of the genetic paradigm of cancer" href="/plosbiology/article?id=10.1371/journal.pbio.3003052"><img alt="The end of the genetic paradigm of cancer" data-file-extension="jpg" src="/plosbiology/indirect/1f10a17924384139c675bdb4e01f3f7019ba1da420203b422e290fa55cd58c1b"></a> <p class="credits">Image credit: pbio.3003052</p> </article> <article class="block small image" id="cellcat3-4"> <div class="block-content"> <p class="date">02/28/2025</p> <p class="category">Perspective</p> <h2 class="title"><a alt="New solutions for antibiotic discovery" href="/plosbiology/article?id=10.1371/journal.pbio.3003058">New solutions for antibiotic discovery</a></h2> <div class="blurb"> <p>A major challenge in natural-product-based drug discovery is the identification of gene clusters most likely to specify new chemistry and bioactivities. This Perspective discusses the state of the art of antibiotic discovery.</p> </div> </div><a class="block-image" alt="New solutions for antibiotic discovery" href="/plosbiology/article?id=10.1371/journal.pbio.3003058"><img alt="New solutions for antibiotic discovery" data-file-extension="jpg" src="/plosbiology/indirect/53c5dd4bec7d5becf390595c1adbf6c0cb7735997e5afe14813250258386926e"></a> <p class="credits">Image credit: pbio.3001026</p> </article> </div> <div class="row-small-blocks"> <article class="carousel-wrap jcarousel-wrapper small" id="carousel-1"> <div class="block-header"> Collections </div> <div class="jcarousel carousel"> <ul> <li> <h3><a href="https://collections.plos.org/collection/neurotechnology-human-health-cognition/" alt="The promises and challenges of neurotechnology">The promises and challenges of neurotechnology</a></h3> <p></p> <p>Neurotechnology offers unprecedented opportunities to treat neural disorders, restore brain function and enhance cognitive abilities. This collection explores the present and possible futures of neurotechnology to improve human health and cognition.</p> <p></p> <p></p> <p></p></li> <li> <h3><a href="https://collections.plos.org/collection/conservation-and-biodiversity/" alt="Decision making for conservation and biodiversity">Decision making for conservation and biodiversity</a></h3> <p></p> <p>Translating conservation and biodiversity research from the field into the real world is a complex problem. This collection discusses issues around economics, policy, and how to do research that answers questions that decision makers have.</p> <p></p></li> <li> <h3><a href="https://collections.plos.org/collection/symbiosis-across-the-tree-of-life/" alt="Symbiosis across the tree of life">Symbiosis across the tree of life</a></h3> <p></p> <p>Symbiosis research has become a holistic and pervasive field with a mature theoretical basis. This collection showcases symbiotic relationships across the tree of life, exploring their evolutionary basis and underlying mechanisms.</p> <p></p> <p></p></li> <li> <h3><a href="https://collections.plos.org/collection/plos-biology-20th-anniversary/" alt="PLOS Biology 20th Anniversary">PLOS Biology 20th Anniversary<br></a></h3> <p></p> <p>PLOS Biology is 20 and we are celebrating with a collection that contains articles that look back at landmark studies that we published, others that look past and future, and others discussing how publishing and open science have evolved and what is to come.</p> <p>.</p> <p><br></p> <p></p></li> <li> <h3><a href="https://collections.plos.org/collection/engineering-plants-for-a-changing-climate/" alt="Engineering plants for a changing climate">Engineering plants for a changing climate</a></h3> <p></p> <p>This collection explores engineering strategies to help us adapt plants to a changing climate, including breeding techniques, genome engineering, synthetic biology and microbiome engineering.<br></p> <p></p></li> </ul> </div> <div class="carousel-control"> <a class="jcarousel-prev" href="#"></a> <div class="numbers"> <span class="number" data-js="carousel-current-item"></span> <span> of </span> <span class="number" data-js="carousel-total-index"></span> </div><a class="jcarousel-next" href="#"></a> </div> </article> <article class="carousel-wrap jcarousel-wrapper small" id="carousel-2"> <div class="block-header"> Meet our staff at conferences </div> <div class="jcarousel carousel"> <ul> <li> <h3><a href="https://100yearsconference.biologists.com/" alt="Biologists @ 100 conference">Biologists @ 100 conference<br></a></h3> <p></p> <p><br></p> <p>March 24 - 27</p> <p><br></p> <p>Meet Senior Magazine Editor Daniel Routledge (droutledge@plos.org)</p> <p></p></li> <li> <h3><a href="https://www.cosyne.org/" alt="COSYNE 2025">COSYNE 2025</a></h3> <p><br></p> <p>March 27 - 30</p> <p><br></p> <p>Meet Senior Editor Christian Schnell (cschneill@plos.org)</p> <p><br></p> <p></p></li> <li> <h3><a href="https://coursesandconferences.wellcomeconnectingscience.org/event/crispr-and-beyond-perturbations-at-scale-to-understand-genomes-20250402/" alt="Wellcome: CRISPR and Beyond: Perturbations at Scale to Understand Genomes">Wellcome: CRISPR and Beyond: Perturbations at Scale to Understand Genomes</a></h3> <p><br></p> <p>April 2 - 4</p> <p><br></p> <p>Meet Senior Editor Richard Hodge (rhodge@plos.org)</p> <p><br></p> <p></p></li> <li> <h3><a href="https://www.nationalacademies.org/event/44297_02-2025_navigating-the-benefits-and-risks-of-publishing-studies-of-in-silico-modeling-and-computational-approaches-of-biological-agents-and-organisms-a-workshop" alt="Navigating the benefits and risks of publishing studies of in silico modeling and computational approaches...">Navigating the benefits and risks of publishing studies of in silico modeling and computational approaches...</a></h3> <p></p> <p><br></p> <p>April 3 - 4</p> <p><br></p> <p>Meet Senior Editor Luke Smith (lsmith@plos.org)</p> <p></p></li> <li> <h3><a href="https://www.sisb2025.conferences-pasteur.org/" alt="EMBO: The Immune System of Bacteria">EMBO: The Immune System of Bacteria</a></h3> <p><br></p> <p>April 6 - 11</p> <p><br></p> <p>Meet Associate Editor Melissa Vazquez Hernandez (mvazquezhernandez@plos.org).</p> <p></p></li> </ul> </div> <div class="carousel-control"> <a class="jcarousel-prev" href="#"></a> <div class="numbers"> <span class="number" data-js="carousel-current-item"></span> <span> of </span> <span class="number" data-js="carousel-total-index"></span> </div><a class="jcarousel-next" href="#"></a> </div> </article> </div> </div> <div class="column right"> <div id="external-content-top"> <div id="blogs" data-host-name="https://blogs.plos.org/" data-blog-title="PLOS Biologue" data-blog-url="https://blogs.plos.org/biologue" data-feed-url="https://blogs.plos.org/biologue/?feed=json" data-post-count="4"> </div> <div id="submission-links"> <h3>Publish with PLOS</h3> <p><a href="/plosbiology/s/submit-now" id="submissionManu">Submit Your Manuscript</a></p> </div> <div id="social-links"> <h3>Connect with Us</h3> <ul class="social-media"> <li id="social-link-email"><a href="/plosbiology/s/contact" class="social" title="Contact Us"> <span class="icon-email" icon-hidden="true"></span> <span class="icon-text">Contact Us</span> </a></li> <li id="social-link-RSS"><a href="/plosbiology/feed/atom" class="social" title="RSS"> <span class="icon-rss" icon-hidden="true"></span> <span class="icon-text">RSS</span> </a></li> <li id="social-link-bluesky"><a href="https://bsky.app/profile/plosbiology.org" class="social" title="PLOS Biology on Bluesky" target="_blank"> <span class="icon-bluesky" icon-hidden="true"></span> <span class="icon-text">PLOS Biology on Bluesky</span> </a></li> <li id="social-link-facebook"><a href="https://www.facebook.com/plos.org" class="social" title="PLOS on Facebook" target="_blank"> <span class="icon-facebook" icon-hidden="true"></span> <span class="icon-text">PLOS on Facebook</span> </a></li> <li id="social-link-blogs"><a href="https://blogs.plos.org/" class="social-blogs" title="PLOS Blogs">PLOS Blogs</a></li> </ul> </div> </div> <div id="external-content-bottom"> </div> </div> </div> </div><script src="/resource/js/vendor/jquery.carousel.js" type="text/javascript"></script> <script src="/resource/js/components/carousel.js" type="text/javascript"></script> <div class="plos-row"> <div class="home-email-subscription" id="etocs-signup"> <h2>Get new content from PLOS Biology in your inbox</h2> <div class="form-container"> <iframe src="https://cdn.forms-content.sg-form.com/3d84b365-f12d-11ee-80a7-822ee8ce2407" frameborder="0"></iframe> </div> <div class="consent"> <div class="email_consent"> PLOS will use your email address to provide content from <span style="font-style: italic;">PLOS Biology</span>. You can find out more about how PLOS processes your data by reading our <a href="https://plos.org/privacy-policy" target="_blank" title="Page opens in new window">Privacy Policy</a>. You can unsubscribe at any time by clicking the unsubscribe link in our emails or by contacting us at privacy@plos.org. </div> </div> </div> </div> <div class="spotlight"> <section id="ad-slot-bottom"> <div class="ad-item"> <!-- DoubleClick Ad Zone --> <div class='advertisement' id='div-gpt-ad-1458247671871-2' style='width:300px; height:250px;'> <script type='text/javascript'> googletag.cmd.push(function() { googletag.display('div-gpt-ad-1458247671871-2'); }); </script> </div> </div> <div class="ad-item"> <!-- DoubleClick Ad Zone --> <div class='advertisement' id='div-gpt-ad-1458247671871-3' style='width:300px; height:250px;'> <script type='text/javascript'> googletag.cmd.push(function() { googletag.display('div-gpt-ad-1458247671871-3'); }); </script> </div> </div> <div class="ad-item"> <!-- DoubleClick Ad Zone --> <div class='advertisement' id='div-gpt-ad-1458247671871-4' style='width:300px; height:250px;'> <script type='text/javascript'> googletag.cmd.push(function() { googletag.display('div-gpt-ad-1458247671871-4'); }); </script> </div> </div> </section> </div> <div class="journal-issn">PLOS Biology | ISSN: 1545-7885 (online)</div> </main> <footer id="pageftr"> <div class="row"> <div class="block x-small"> <ul class="nav nav-secondary"> <li class="ftr-header"><a href="https://plos.org/our-journals/">Publications</a></li> <li><a href="/plosbiology/" id="ftr-bio">PLOS Biology</a></li> <li><a href="/climate/" id="ftr-climate">PLOS Climate</a></li> <li><a href="/complexsystems/" id="ftr-complex-systems">PLOS Complex Systems</a></li> <li><a href="/ploscompbiol/" id="ftr-compbio">PLOS Computational Biology</a></li> <li><a href="/digitalhealth/" id="ftr-digitalhealth">PLOS Digital Health</a></li> <li><a href="/plosgenetics/" id="ftr-gen">PLOS Genetics</a></li> <li><a href="/globalpublichealth/" id="ftr-globalpublichealth">PLOS Global Public Health</a></li> </ul> </div> <div class="block x-small"> <ul class="nav nav-secondary"> <li class="ftr-header"> </li> <li><a href="/plosmedicine/" id="ftr-med">PLOS Medicine</a></li> <li><a href="/mentalhealth/" id="ftr-mental-health">PLOS Mental Health</a></li> <li><a href="/plosntds/" id="ftr-ntds">PLOS Neglected Tropical Diseases</a></li> <li><a href="/plosone/" id="ftr-one">PLOS One</a></li> <li><a href="/plospathogens/" id="ftr-path">PLOS Pathogens</a></li> <li><a href="/sustainabilitytransformation/" id="ftr-sustainabilitytransformation">PLOS Sustainability and Transformation</a></li> <li><a href="/water/" id="ftr-water">PLOS Water</a></li> </ul> </div> <div class="block xx-small"> <ul class="nav nav-tertiary"> <li> <a href="https://plos.org" id="ftr-home">Home</a> </li> <li> <a href="https://blogs.plos.org" id="ftr-blog">Blogs</a> </li> <li> <a href="https://collections.plos.org/" id="ftr-collections">Collections</a> </li> <li> <a href="mailto:webmaster@plos.org" id="ftr-feedback">Give feedback</a> </li> <li> <a href="/plosbiology/lockss-manifest" id="ftr-lockss">LOCKSS</a> </li> </ul> </div> <div class="block xx-small"> <ul class="nav nav-primary"> <li><a href="https://plos.org/privacy-policy" id="ftr-privacy">Privacy Policy</a></li> <li><a href="https://plos.org/terms-of-use" id="ftr-terms">Terms of Use</a></li> <li><a href="https://plos.org/advertise/" id="ftr-advertise">Advertise</a></li> <li><a href="https://plos.org/media-inquiries" id="ftr-media">Media Inquiries</a></li> <li><a href="https://plos.org/contact" id="ftr-contact">Contact</a></li> </ul> </div> </div> <div class="row"> <p> <img src="/resource/img/logo-plos-footer.png" alt="PLOS" class="logo-footer"/> <span class="footer-non-profit-statement">PLOS is a nonprofit 501(c)(3) corporation, #C2354500, based in California, US</span> </p> <div class="block"> </div> </div> <script src="/resource/js/global.js" type="text/javascript"></script> </footer> <script src="/resource/js/pages/home.js" type="text/javascript"></script> </body> </html>