#copy-dynamic{padding:0 20px}.highwire_clipboard_form_ajax_shareit{display:none}#mailit .item-list ul li{margin:0}.clipboard-copy{width:100%}.copy-button{float:right}.highwire_clipboard_form_ajax_shareit{display:none}.ctools-collapsible-container .ctools-toggle{float:left;width:21px;height:21px;cursor:pointer;background-position:7px 7px;background-repeat:no-repeat;background-image:url(https://science.sciencemag.org/sites/all/modules/contrib/ctools/images/collapsible-expanded.png)}.ctools-collapsible-container .ctools-collapsible-handle{display:none}html.js .ctools-collapsible-container .ctools-collapsible-handle{display:block}.ctools-collapsible-container .ctools-collapsible-handle{cursor:pointer}.ctools-collapsible-container .ctools-toggle-collapsed{background-image:url(https://science.sciencemag.org/sites/all/modules/contrib/ctools/images/collapsible-collapsed.png)}.hw-icon-large:before{vertical-align:-10%;font-size:1.3333333333333333em}a [class^="hw-icon-"],a [class*=" hw-icon-"]{display:inline}[class^="hw-icon-"].hw-icon-fixed-width,[class*=" hw-icon-"].hw-icon-fixed-width{display:inline-block;width:1.1428571428571428em;text-align:right;padding-right:.2857142857142857em}[class^="hw-icon-"].hw-icon-fixed-width.hw-icon-large,[class*=" hw-icon-"].hw-icon-fixed-width.hw-icon-large{width:1.4285714285714286em}.icons-ul .hw-icon-li{position:absolute;left:-2.142857142857143em;width:2.142857142857143em;text-align:center;line-height:inherit}[class^="hw-icon-"].hide,[class*=" hw-icon-"].hide{display:none}.hw-icon-muted{color:#eee}.hw-icon-light{color:#fff}.hw-icon-dark{color:#333}.hw-icon-border{border:solid 1px #eee;padding:.2em .25em .15em;-webkit-border-radius:3px;-moz-border-radius:3px;border-radius:3px}.hw-icon-2x{font-size:2em}.hw-icon-2x.hw-icon-border{border-width:2px;-webkit-border-radius:4px;-moz-border-radius:4px;border-radius:4px}.hw-icon-3x{font-size:3em}.hw-icon-3x.hw-icon-border{border-width:3px;-webkit-border-radius:5px;-moz-border-radius:5px;border-radius:5px}.hw-icon-4x{font-size:4em}.hw-icon-4x.hw-icon-border{border-width:4px;-webkit-border-radius:6px;-moz-border-radius:6px;border-radius:6px}.hw-icon-5x{font-size:5em}.hw-icon-5x.hw-icon-border{border-width:5px;-webkit-border-radius:7px;-moz-border-radius:7px;border-radius:7px}.pull-right{float:right}.pull-left{float:left}[class^="hw-icon-"].pull-left,[class*=" hw-icon-"].pull-left{margin-right:.3em}[class^="hw-icon-"].pull-right,[class*=" hw-icon-"].pull-right{margin-left:.3em}[class^="hw-icon-"],[class*=" hw-icon-"]{display:inline;width:auto;height:auto;line-height:normal;vertical-align:baseline;background-image:none;background-position:0 0;background-repeat:repeat;margin-top:0}.hw-icon-white,.nav-pills>.active>a>[class^="hw-icon-"],.nav-pills>.active>a>[class*=" hw-icon-"],.nav-list>.active>a>[class^="hw-icon-"],.nav-list>.active>a>[class*=" hw-icon-"],.navbar-inverse .nav>.active>a>[class^="hw-icon-"],.navbar-inverse .nav>.active>a>[class*=" hw-icon-"],.dropdown-menu>li>a:hover>[class^="hw-icon-"],.dropdown-menu>li>a:hover>[class*=" hw-icon-"],.dropdown-menu>.active>a>[class^="hw-icon-"],.dropdown-menu>.active>a>[class*=" hw-icon-"],.dropdown-submenu:hover>a>[class^="hw-icon-"],.dropdown-submenu:hover>a>[class*=" hw-icon-"]{background-image:none}.btn [class^="hw-icon-"].hw-icon-large,.nav [class^="hw-icon-"].hw-icon-large,.btn [class*=" hw-icon-"].hw-icon-large,.nav [class*=" hw-icon-"].hw-icon-large{line-height:.9em}
.btn [class^="hw-icon-"].hw-icon-spin,.nav [class^="hw-icon-"].hw-icon-spin,.btn [class*=" hw-icon-"].hw-icon-spin,.nav [class*=" hw-icon-"].hw-icon-spin{display:inline-block}.nav-tabs [class^="hw-icon-"],.nav-pills [class^="hw-icon-"],.nav-tabs [class*=" hw-icon-"],.nav-pills [class*=" hw-icon-"],.nav-tabs [class^="hw-icon-"].hw-icon-large,.nav-pills [class^="hw-icon-"].hw-icon-large,.nav-tabs [class*=" hw-icon-"].hw-icon-large,.nav-pills [class*=" hw-icon-"].hw-icon-large{line-height:.9em}.btn [class^="hw-icon-"].pull-left.hw-icon-2x,.btn [class*=" hw-icon-"].pull-left.hw-icon-2x,.btn [class^="hw-icon-"].pull-right.hw-icon-2x,.btn [class*=" hw-icon-"].pull-right.hw-icon-2x{margin-top:.18em}.btn [class^="hw-icon-"].hw-icon-spin.hw-icon-large,.btn [class*=" hw-icon-"].hw-icon-spin.hw-icon-large{line-height:.8em}.btn.btn-small [class^="hw-icon-"].pull-left.hw-icon-2x,.btn.btn-small [class*=" hw-icon-"].pull-left.hw-icon-2x,.btn.btn-small [class^="hw-icon-"].pull-right.hw-icon-2x,.btn.btn-small [class*=" hw-icon-"].pull-right.hw-icon-2x{margin-top:.25em}.btn.btn-large [class^="hw-icon-"],.btn.btn-large [class*=" hw-icon-"]{margin-top:0}.btn.btn-large [class^="hw-icon-"].pull-left.hw-icon-2x,.btn.btn-large [class*=" hw-icon-"].pull-left.hw-icon-2x,.btn.btn-large [class^="hw-icon-"].pull-right.hw-icon-2x,.btn.btn-large [class*=" hw-icon-"].pull-right.hw-icon-2x{margin-top:.05em}.btn.btn-large [class^="hw-icon-"].pull-left.hw-icon-2x,.btn.btn-large [class*=" hw-icon-"].pull-left.hw-icon-2x{margin-right:.2em}.btn.btn-large [class^="hw-icon-"].pull-right.hw-icon-2x,.btn.btn-large [class*=" hw-icon-"].pull-right.hw-icon-2x{margin-left:.2em}.nav-list [class^="hw-icon-"],.nav-list [class*=" hw-icon-"]{line-height:inherit}.icon-stack{position:relative;display:inline-block;width:2em;height:2em;line-height:2em;vertical-align:-35%}.icon-stack [class^="hw-icon-"],.icon-stack [class*=" hw-icon-"]{display:block;text-align:center;position:absolute;width:100%;height:100%;font-size:1em;line-height:inherit;*line-height:2em;}.hw-icon-stack .hw-icon-stack-base{font-size:2em;*line-height:1em;}.hw-icon-spin{display:inline-block;-moz-animation:spin 2s infinite linear;-o-animation:spin 2s infinite linear;-webkit-animation:spin 2s infinite linear;animation:spin 2s infinite linear}a .hw-icon-stack,a .hw-icon-spin{display:inline-block;text-decoration:none}@-moz-keyframes spin{0%{-moz-transform:rotate(0deg)}100%{-moz-transform:rotate(359deg)}}@-webkit-keyframes spin{0%{-webkit-transform:rotate(0deg)}100%{-webkit-transform:rotate(359deg)}}@-o-keyframes spin{0%{-o-transform:rotate(0deg)}100%{-o-transform:rotate(359deg)}}@-ms-keyframes spin{0%{-ms-transform:rotate(0deg)}100%{-ms-transform:rotate(359deg)}}@keyframes spin{0%{transform:rotate(0deg)}100%{transform:rotate(359deg)}}.hw-icon-rotate-90:before{-webkit-transform:rotate(90deg);-moz-transform:rotate(90deg);-ms-transform:rotate(90deg);-o-transform:rotate(90deg);transform:rotate(90deg);filter:progid:DXImageTransform.Microsoft.BasicImage(rotation=1)}.hw-icon-rotate-180:before{-webkit-transform:rotate(180deg);-moz-transform:rotate(180deg);-ms-transform:rotate(180deg);-o-transform:rotate(180deg);transform:rotate(180deg);filter:progid:DXImageTransform.Microsoft.BasicImage(rotation=2)}
.hw-icon-rotate-270:before{-webkit-transform:rotate(270deg);-moz-transform:rotate(270deg);-ms-transform:rotate(270deg);-o-transform:rotate(270deg);transform:rotate(270deg);filter:progid:DXImageTransform.Microsoft.BasicImage(rotation=3)}.hw-icon-flip-horizontal:before{-webkit-transform:scale(-1,1);-moz-transform:scale(-1,1);-ms-transform:scale(-1,1);-o-transform:scale(-1,1);transform:scale(-1,1)}.hw-icon-flip-vertical:before{-webkit-transform:scale(1,-1);-moz-transform:scale(1,-1);-ms-transform:scale(1,-1);-o-transform:scale(1,-1);transform:scale(1,-1)}a .hw-icon-rotate-90:before,a .hw-icon-rotate-180:before,a .hw-icon-rotate-270:before,a .hw-icon-flip-horizontal:before,a .hw-icon-flip-vertical:before{display:inline-block}.hw-icon-color-highwire{color:#a90533}.hw-icon-color-pdf{color:#dd1f12}.hw-icon-color-epub{color:#90bf2d}.hw-icon-color-orcid{color:#a6ce39}.hw-icon-readcube,a .hw-icon-readcube{display:inline-block}.hw-icon-readcube{background:url(https://science.sciencemag.org/sites/all/modules/highwire/highwire/highwire_theme_tools/img/readcube.svg) 0 0 no-repeat transparent;background-size:100%;height:1.2em;width:1.2em;text-indent:-9999em}@font-face{font-family:'hwicons';src:url(https://science.sciencemag.org/sites/all/modules/highwire/highwire/highwire_theme_tools/fonts/hwicons.eot?sc9idg);src:url(https://science.sciencemag.org/sites/all/modules/highwire/highwire/highwire_theme_tools/fonts/hwicons.eot?sc9idg#iefix) format('embedded-opentype'),url(https://science.sciencemag.org/sites/all/modules/highwire/highwire/highwire_theme_tools/fonts/hwicons.ttf?sc9idg) format('truetype'),url(https://science.sciencemag.org/sites/all/modules/highwire/highwire/highwire_theme_tools/fonts/hwicons.woff?sc9idg) format('woff'),url(https://science.sciencemag.org/sites/all/modules/highwire/highwire/highwire_theme_tools/fonts/hwicons.svg?sc9idg#hwicons) format('svg');font-weight:normal;font-style:normal}.hw-icon,[class^="hw-icon-"],[class*=" hw-icon-"]{font-family:'hwicons' !important;speak:none;font-style:normal;font-weight:normal;font-variant:normal;text-transform:none;line-height:1;-webkit-font-smoothing:antialiased;-moz-osx-font-smoothing:grayscale}.hw-icon-highwire:before{content:"\e903"}.hw-icon-readcube:before{content:"\e902"}.hw-icon-orcid:before{content:"\e901"}.hw-icon-epub:before{content:"\e900"}.hw-icon-free:before{content:"\e601"}.hw-icon-closed-access-alt:before{content:"\e602"}.hw-icon-closed-access:before{content:"\e603"}.hw-icon-open-access:before{content:"\e604"}.hw-icon-pdf:before{content:"\e600"}.highwire-markup .section .list-simple li,.highwire-markup div.contributors ol.contributor-list,.highwire-markup div.contributors ol.affiliation-list,.highwire-markup div.contributors ol.corresp-list,.highwire-markup div.contributors .fn-track,.highwire-markup div.fn-group ul,.highwire-markup ol.ref-use-labels,.highwire-markup #content-block-markup ul.history-list,.highwire-markup ul.copyright-statement{list-style:none;list-style-type:none;margin-left:0;padding:0}q.attrib{quotes:none}.highwire-markup .contributors,.highwire-markup .contributors .contributor-list{text-align:left}.highwire-markup .contributors .contributor-list li{display:inline;padding:0;margin:0;text-align:left}.highwire-markup .contributors ol li a{border:0;font-weight:normal;text-decoration:none}.highwire-markup .affiliation-list-reveal a.view-more{margin:0;padding:0 5px;font-size:14px;line-height:1;text-align:center;display:inline-block;vertical-align:middle;width:25px}.highwire-markup .hideaffil{left:-9999px;position:absolute;width:5000px}.highwire-markup div.contributors ol.affiliation-list li{display:list-item;padding-left:0}
.highwire-markup div.contributors ol.affiliation-list li address{border:0;display:inline;font-style:italic;padding:0;margin:0}.highwire-markup div.article ul.list-unord{list-style-type:disc;margin:15px 0 15px 22.5px}.highwire-markup .section.intro{font-family:"Benton Sans WGL","Helvetica Neue",Helvetica,Arial,sans-serif;font-size:1em;line-height:1.5;padding:0;margin:0;border:0}.highwire-markup #content-block-markup ul.history-list{padding:0}.publication-history li{margin-top:0}.highwire-markup div.fig a.in-nw-vis{display:inline}#colorbox #cboxTitle{background:rgba(83,83,83,0.9);color:#FFF}.highwire-markup div.caption-clear{clear:both}.highwire-markup ul.copyright-statement{list-style-type:none}.highwire-markup div.fn-group ul{margin-left:0}.highwire-markup div.bio{margin-top:1em}.highwire-markup div.fn-group ul li p{margin-bottom:0}.highwire-markup #content-block-markup div.fn-group ul li p{margin:.5em 0;padding:0}.highwire-markup ol.cit-list>li>.rev-xref-ref+span{display:block}.highwire-markup div.ref-list .rev-xref-ref,.highwire-markup div.ref-list .rev-xref-ref:link{border:0;text-decoration:none}.highwire-markup .ref-cit div,.highwire-markup .ref-cit cite,.highwire-markup .ref-cit ol,.highwire-markup .ref-cit li{display:inline;margin-left:0;margin-right:0;padding-left:0;padding-right:0}.highwire-markup div.ref-cit .cit-metadata li{display:inline}.highwire-markup ol.cit-list li{display:list-item}.highwire-markup div.ref-cit cite{font-style:normal}.highwire-markup div.ref-cit .cit-vol{font-weight:bolder}.highwire-markup div.ref-cit div.cit-metadata{margin-right:.5rem}.highwire-markup div.ref-cit div.cit-extra a{display:inline-block;font-size:88%;margin-left:0;margin-right:.75rem;text-decoration:underline;vertical-align:middle;white-space:nowrap}.highwire-markup div.ref-cit cite span.cit-source,.highwire-markup div.ref-cit cite abbr.cit-jnl-abbrev{margin-right:.1em;font-style:italic}.hw-ref-link-tooltip{max-width:33%}.hw-ref-link-tooltip.qtip-pos-bl .qtip-tip,.hw-ref-link-tooltip.qtip-pos-bc .qtip-tip,.hw-ref-link-tooltip.qtip-pos-br .qtip-tip{-moz-transform:scaleY(-1);-o-transform:scaleY(-1);-webkit-transform:scaleY(-1);transform:scaleY(-1);filter:FlipV;-ms-filter:"FlipV"}.article__body .highwire-markup .abstract>.highwire-quick-nav:first-child+*,.article__body .highwire-markup .boxed-text>.highwire-quick-nav:first-child+*,.article__body .highwire-markup .structured>.highwire-quick-nav:first-child+*,.article__body .highwire-markup .editor-summary>.highwire-quick-nav:first-child+*{margin-top:0}.highwire-quick-nav{float:right}.highwire-quick-nav a{font-size:1em;line-height:1;padding:0}.icon-caret-down:after{content:"\25BC"}.icon-caret-up:after{content:"\25B2"}.orig-source{text-align:center}.table-expansion{overflow:auto;width:100%}.table-expansion table,.table-expansion table *{border-color:#e0dbd5}.table-center,.table-char{text-align:center}.nlm-sup,.nlm-sub,.contributor-list>li>*[class^="xref-"],.corresp-list .corresp-label{font-size:75%;line-height:0;position:relative;top:-.5em;vertical-align:baseline}.contributor-list>li>*[class^="xref-"] sup,.corresp-list .corresp-label sup{font-size:100%;top:auto}.nlm-sub{top:auto;bottom:-.5em}.cit-ref-sprinkles{margin-left:10px}ol.duplicate{display:none}.highwire-markup .sc{font-size:.85em;text-transform:uppercase}.underline{text-decoration:underline}.highwire-markup{word-wrap:break-word}.highwire-markup .figure.nonresearch-content{min-width:auto}.highwire-markup .figure.nonresearch-content img{width:auto}html.js .highwire-markup .lazyload.fragment-image,html.js .highwire-markup .lazyloading.fragment-image{height:auto;width:auto}.highwire-markup span.disp-formula-label{padding-left:0}figcaption.fig-caption.attrib{font-size:14px;margin:.5em 0 0;text-align:left;text-transform:inherit;color:#666}