xmlhttprequesthttp://www.pnas.org/ imagehttp://www.pnas.org/favicon.ico xmlhttprequesthttps://apis.google.com/_/scs/apps-static/_/js/k=oz.plusone.en_GB.aoBzmI3bt8Y.O/m=p1b,p1p/rt=j/sv=1/d=1/ed=1/rs=AGLTcCMTk9Z5YO2RVneK5nvFOoN-IhYS8Q/t=zcms/cb=gapi.loaded_1 xmlhttprequesthttps://apis.google.com/_/scs/apps-static/_/js/k=oz.gapi.en_GB.B7GSEr9a240.O/m=googleapis_client,iframes_styles_bubble_internal,gapi_iframes_style_common,gapi_iframes_iframer/rt=j/sv=1/d=1/ed=1/am=AQ/rs=AGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg/t=zcms/cb=gapi.loaded_0 xmlhttprequesthttps://apis.google.com/_/scs/apps-static/_/js/k=oz.plusone.en_GB.aoBzmI3bt8Y.O/m=p1b,p1p/rt=j/sv=1/d=1/ed=1/rs=AGLTcCMTk9Z5YO2RVneK5nvFOoN-IhYS8Q/t=zcms/cb=gapi.loaded_1 xmlhttprequesthttps://apis.google.com/_/scs/apps-static/_/js/k=oz.gapi.en_GB.B7GSEr9a240.O/m=googleapis_client,iframes_styles_bubble_internal,gapi_iframes_style_common,gapi_iframes_iframer/rt=j/sv=1/d=1/ed=1/am=AQ/rs=AGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg/t=zcms/cb=gapi.loaded_0 stylesheethttps://apis.google.com/_/scs/apps-static/_/ss/k=oz.plusone.-1qnmlzwsnet54.L.W.O/d=0/rs=AGLTcCMbPCnX2rqwIm_midHdbqkfLkaZGQ scripthttps://ssl.gstatic.com/accounts/o/2006856415-postmessagerelay.js scripthttps://oauth.googleusercontent.com/gadgets/js/core:rpc:shindig.random:shindig.sha1.js?c=2 scripthttps://apis.google.com/js/api.js stylesheethttps://apis.google.com/_/scs/apps-static/_/ss/k=oz.plusone.-1qnmlzwsnet54.L.W.O/d=0/rs=AGLTcCMbPCnX2rqwIm_midHdbqkfLkaZGQ inline-scripthttps://apis.google.com/u/0/se/0/_/+1/fastbutton?usegapi=1&size=small&count=false&origin=http%3A%2F%2Fwww.pnas.org&url=http%3A%2F%2Fwww.pnas.org%2Fcontent%2F111%2F45%2F15873&gsrc=3p&jsh=m%3B%2F_%2Fscs%2Fapps-static%2F_%2Fjs%2Fk%3Doz.gapi.en_GB.B7GSEr9a240.O%2Fm%3D__features__%2Fam%3DAQ%2Frt%3Dj%2Fd%3D1%2Ft%3Dzcms%2Frs%3DAGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg#_methods=onPlusOne%2C_ready%2C_close%2C_open%2C_resizeMe%2C_renderstart%2Concircled%2Cdrefresh%2Cerefresh%2Conload&id=I1_1425463531152&parent=http%3A%2F%2Fwww.pnas.org&pfname=&rpctoken=40753148{inline-script} inline-scripthttps://accounts.google.com/o/oauth2/postmessageRelay?parent=http%3A%2F%2Fwww.pnas.org#rpctoken=481024953&forcesecure=1{inline-script} inline-scripthttps://apis.google.com/u/0/se/0/_/+1/fastbutton?usegapi=1&size=small&count=false&origin=http%3A%2F%2Fwww.pnas.org&url=http%3A%2F%2Fwww.pnas.org%2Fcontent%2F111%2F45%2F15873&gsrc=3p&jsh=m%3B%2F_%2Fscs%2Fapps-static%2F_%2Fjs%2Fk%3Doz.gapi.en_GB.B7GSEr9a240.O%2Fm%3D__features__%2Fam%3DAQ%2Frt%3Dj%2Fd%3D1%2Ft%3Dzcms%2Frs%3DAGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg#_methods=onPlusOne%2C_ready%2C_close%2C_open%2C_resizeMe%2C_renderstart%2Concircled%2Cdrefresh%2Cerefresh%2Conload&id=I0_1425463531127&parent=http%3A%2F%2Fwww.pnas.org&pfname=&rpctoken=27338699{inline-script} sub_framehttps://accounts.google.com/o/oauth2/postmessageRelay?parent=http%3A%2F%2Fwww.pnas.org#rpctoken=481024953&forcesecure=1 sub_framehttps://apis.google.com/u/0/se/0/_/+1/fastbutton?usegapi=1&size=small&count=false&origin=http%3A%2F%2Fwww.pnas.org&url=http%3A%2F%2Fwww.pnas.org%2Fcontent%2F111%2F45%2F15873&gsrc=3p&jsh=m%3B%2F_%2Fscs%2Fapps-static%2F_%2Fjs%2Fk%3Doz.gapi.en_GB.B7GSEr9a240.O%2Fm%3D__features__%2Fam%3DAQ%2Frt%3Dj%2Fd%3D1%2Ft%3Dzcms%2Frs%3DAGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg#_methods=onPlusOne%2C_ready%2C_close%2C_open%2C_resizeMe%2C_renderstart%2Concircled%2Cdrefresh%2Cerefresh%2Conload&id=I1_1425463531152&parent=http%3A%2F%2Fwww.pnas.org&pfname=&rpctoken=40753148 sub_framehttps://apis.google.com/u/0/se/0/_/+1/fastbutton?usegapi=1&size=small&count=false&origin=http%3A%2F%2Fwww.pnas.org&url=http%3A%2F%2Fwww.pnas.org%2Fcontent%2F111%2F45%2F15873&gsrc=3p&jsh=m%3B%2F_%2Fscs%2Fapps-static%2F_%2Fjs%2Fk%3Doz.gapi.en_GB.B7GSEr9a240.O%2Fm%3D__features__%2Fam%3DAQ%2Frt%3Dj%2Fd%3D1%2Ft%3Dzcms%2Frs%3DAGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg#_methods=onPlusOne%2C_ready%2C_close%2C_open%2C_resizeMe%2C_renderstart%2Concircled%2Cdrefresh%2Cerefresh%2Conload&id=I0_1425463531127&parent=http%3A%2F%2Fwww.pnas.org&pfname=&rpctoken=27338699 scripthttps://apis.google.com/_/scs/apps-static/_/js/k=oz.gapi.en_GB.B7GSEr9a240.O/m=auth/exm=plusone/rt=j/sv=1/d=1/ed=1/am=AQ/rs=AGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg/t=zcms/cb=gapi.loaded_1 scripthttps://apis.google.com/_/scs/apps-static/_/js/k=oz.gapi.en_GB.B7GSEr9a240.O/m=plusone/rt=j/sv=1/d=1/ed=1/am=AQ/rs=AGLTcCPm3UQfwvh9ZG2zSt9BbS5nrYUrQg/t=zcms/cb=gapi.loaded_0 stylesheethttps://d1bxh8uas1mnw7.cloudfront.net/assets/embed-deec67c8878bf9a094e44df5cac3dc81.css imagehttps://altmetric-badges.a.ssl.fastly.net/?size=240&score=26&types=bttttttg imagehttp://crossmark.crossref.org/images/bg_dialog_footer.png scripthttps://apis.google.com/js/plusone.js?_=1425463530968 xmlhttprequesthttp://www.pnas.org/entrez-links/25326421 xmlhttprequesthttp://www.pnas.org/isi-links//pnas/111/45/15873?rt=yes xmlhttprequesthttp://www.pnas.org/cited-by/pnas/111/45/15873 xmlhttprequesthttp://www.pnas.org/isi-links/has-related/pnas/111/45/15873 scripthttps://api.altmetric.com/v1/doi/10.1073/pnas.1411798111?callback=_altmetric.embed_callback&domain=www.pnas.org&key=3c130976ca2b8f2e88f8377633751ba1&cache_until=10-4 imagehttp://www.pnas.org/local/img/misc/arrow-blue-down.png imagehttp://www.pnas.org/local/img/misc/arrow-blue-side.png imagehttp://www.pnas.org/local/img/nav/btn-search.gif imagehttp://www.pnas.org/local/img/nav/footer-pnas.png imagehttp://www.pnas.org/local/img/nav/glencoe_logo_small.png imagehttp://www.pnas.org/local/img/nav/icon-pnas.gif imagehttp://www.pnas.org/local/img/nav/icon-highwire.png imagehttp://www.pnas.org/local/img/nav/icon-nas.png imagehttp://www.pnas.org/content/111/45/15873/F1.small.gif stylesheethttp://crossmark.crossref.org/stylesheets/crossmark_widget.css imagehttp://www.pnas.org/local/img/misc/icon-mendeley.gif imagehttp://www.pnas.org/local/img/nav/nav-divider-article.png imagehttp://www.pnas.org/local/img/nav/nav-article-bg.png imagehttp://www.pnas.org/local/img/nav/nav-divider2.png imagehttp://www.pnas.org/local/img/nav/background-expand-lg.png xmlhttprequesthttp://www.pnas.org/similar-articles?resource=111/45/15873 scripthttp://movie-usa.glencoesoftware.com/metadata/10.1073/pnas.1411798111?callback=jQuery172040633983444422483_1425463528439&_=1425463529649 fonthttp://fonts.gstatic.com/s/opensans/v10/xjAJXh38I15wypJXxuGMBo4P5ICox8Kq3LLUNMylGO4.woff2 fonthttp://fonts.gstatic.com/s/opensans/v10/cJZKeOuBrn4kERxqtaUH3VtXRa8TVwTICgirnJhmVJw.woff2 fonthttp://fonts.gstatic.com/s/opensans/v10/k3k702ZOKiLJc3WVjuplzOgdm0LZdjqr5-oayXSOefg.woff2 imagehttp://www.pnas.org/shared/img/common/social-bookmarking/digg.gif imagehttp://www.pnas.org/local/img/misc/icon-delicious.gif imagehttp://www.pnas.org/local/img/misc/icon-cite.gif imagehttp://www.pnas.org/local/img/misc/icon-google.gif imagehttp://www.pnas.org/local/img/misc/icon-twitter.gif imagehttps://d1uo4w7k31k5mn.cloudfront.net/donut/0.png scripthttps://d1bxh8uas1mnw7.cloudfront.net/assets/altmetric_badges-d99e7772b00f31495fed09fece0b7237.js imagehttp://www.pnas.org/local/img/misc/icon-facebook.png − /adsystem/imagehttp://www.pnas.org/adsystem/graphics/8109264191440784/pnas/Science-Sessions-224x90-10_22_12.gif?ad=37081&adview=true − /adsystem/imagehttp://www.pnas.org/adsystem/graphics/44400126833185627/pnas/pna-FINAL-bn-physics-portal.gif?ad=41780&adview=true − ||google-analytics.com^scripthttp://www.google-analytics.com/ga.js scripthttp://cdn.mathjax.org/mathjax/latest/config/TeX-AMS-MML_HTMLorMML.js?rev=2.5.1 imagehttp://www.pnas.org/local/img/nav/logo.gif stylesheethttp://www.pnas.org/shared/css/hw-print.css imagehttp://www.pnas.org/local/img/misc/btn-arrow-next.gif imagehttp://www.pnas.org/local/img/misc/btn-arrow-prev.gif imagehttp://www.pnas.org/content/111/45.cover.gif imagehttp://www.pnas.org/local/img/nav/breadcrumb-home.gif imagehttp://www.pnas.org/local/img/nav/headline.png stylesheethttp://www.pnas.org/shared/css/hw-content-int-data-supp.css stylesheethttp://www.pnas.org/shared/css/hw-content-tables.css stylesheethttp://www.pnas.org/shared/css/hw-content-figures.css scripthttps://d1bxh8uas1mnw7.cloudfront.net/assets/embed.js stylesheethttp://www.pnas.org/shared/css/standard-designs/design25/d25-sidebar.css stylesheethttp://www.pnas.org/shared/css/standard-designs/design25/d25-page-home.css stylesheethttp://www.pnas.org/shared/css/standard-designs/design25/ui.css stylesheethttp://www.pnas.org/shared/css/hw-global-dynamic-elements.css stylesheethttp://www.pnas.org/shared/css/hw-global-classes.css stylesheethttp://www.pnas.org/shared/css/hw-global-citation.css stylesheethttp://www.pnas.org/shared/css/hw-global-elements.css stylesheethttp://www.pnas.org/shared/css/hw-global-sidebars.css imagehttp://crossmark.crossref.org/images/crossmark_button.png scripthttp://crossmark.crossref.org/javascripts/crossmark.js scripthttps://ajax.googleapis.com/ajax/libs/jqueryui/1.8.7/jquery-ui.min.js scripthttp://cdn.mathjax.org/mathjax/latest/MathJax.js?config=TeX-AMS-MML_HTMLorMML scripthttp://www.pnas.org/shared/js/util/hw-mathjax.js scripthttp://www.pnas.org/local/js/hw-local-tabbed.js scripthttp://www.pnas.org/shared/js/util/hw-floatabs.js scripthttp://www.pnas.org/local/js/hw-local-content.js scripthttp://www.pnas.org/local/js/article-dynamic-elements.js scripthttp://www.pnas.org/shared/js/util/hw-access-check.js scripthttp://www.pnas.org/shared/js/util/hw-view-specific-access-check.js scripthttp://admin.brightcove.com/js/BrightcoveExperiences_all.js scripthttp://www.pnas.org/shared/js/fancybox/jquery.mousewheel-3.0.4.pack.js scripthttp://www.pnas.org/shared/js/fancybox/jquery.easing-1.3.pack.js scripthttp://www.pnas.org/shared/js/fancybox/jquery.fancybox-1.3.4.js scripthttp://www.pnas.org/shared/js/pages/hw-content.js scripthttp://pnas-movie.glencoesoftware.com/static/video_inline.min.js scripthttp://www.pnas.org/shared/js/design/hw-design1.js scripthttp://www.pnas.org/local/js/script.js scripthttp://www.pnas.org/local/js/expand.js scripthttp://www.pnas.org/local/js/hw-local-global.js scripthttp://www.pnas.org/local/js/jquery.tabify.js scripthttp://www.pnas.org/local/js/hw-local-js-settings.js scripthttp://www.pnas.org/shared/js/hw-shared.js − /fingerprint.jsscripthttp://www.pnas.org/shared/js/fingerprint.js scripthttp://www.pnas.org/local/js/modernizr.js scripthttp://www.pnas.org/local/js/jquery.ui.widget.min.js scripthttp://www.pnas.org/local/js/jquery-1.7.2.min.js stylesheethttp://www.pnas.org/shared/css/standard-designs/design2/leaderboard.css stylesheethttp://www.pnas.org/site/css/global_styles.css stylesheethttp://www.pnas.org/site/css/main.css stylesheethttp://www.pnas.org/local/css/hw-tab-redesign.css stylesheethttp://www.pnas.org/local/css/hw-local-content.css stylesheethttp://www.pnas.org/shared/css/jquery.fancybox-1.3.4.css stylesheethttp://www.pnas.org/shared/css/hw-page-content.css stylesheethttp://pnas-movie.glencoesoftware.com/static/video-js.min.css stylesheethttp://netdna.bootstrapcdn.com/font-awesome/4.0.3/css/font-awesome.css stylesheethttp://www.pnas.org/local/css/hw-local-sidebar.css stylesheethttp://fonts.googleapis.com/css?family=Open+Sans:400italic,700italic,400,700 stylesheethttp://fonts.googleapis.com/css?family=Open+Sans:400,700,600 stylesheethttp://www.pnas.org/local/css/hw-local-global-pages.css stylesheethttp://www.pnas.org/local/css/hw-local-global-overrides.css stylesheethttp://www.pnas.org/local/css/hw-local-global.css stylesheethttp://www.pnas.org/shared/css/standard-designs/design25/main.css stylesheethttp://www.pnas.org/shared/css/hw-global.css inline-scripthttp://www.pnas.org/content/111/45/15873{inline-script} main_framehttp://www.pnas.org/content/111/45/15873
