<!DOCTYPE html>
<html lang="en">
<!-- Served by Oscar-EBS -->
<head>
    <title>Nature</title>
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    <meta name="theme-color" content="#940720">
    <style>
        html {
            font-family: sans-serif;
            line-height: 1.15;
            -ms-text-size-adjust: 100%;
            -webkit-text-size-adjust: 100%
        }

        body {
            margin: 0
        }

        article, aside, footer, header, nav, section {
            display: block
        }

        h1 {
            font-size: 2em;
            margin: .67em 0
        }

        figcaption, figure, main {
            display: block
        }

        figure {
            margin: 1em 40px
        }

        hr {
            box-sizing: content-box;
            height: 0;
            overflow: visible
        }

        pre {
            font-family: monospace, monospace;
            font-size: 1em
        }

        a {
            background-color: transparent;
            -webkit-text-decoration-skip: objects
        }

        a:active, a:hover {
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        }

        abbr[title] {
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            text-decoration: underline dotted
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        }

        b, strong {
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        }

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            font-size: 1em
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        dfn {
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        mark {
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        small {
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        }

        sub, sup {
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            position: relative;
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        sub {
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            font-size: 100%;
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        }

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        }

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        }

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            outline: 1px dotted ButtonText
        }

        fieldset {
            border: 1px solid silver;
            margin: 0 2px;
            padding: .35em .625em .75em
        }

        legend {
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            color: inherit;
            display: table;
            max-width: 100%;
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        }

        progress {
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        }

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        }

        [type=number]::-webkit-inner-spin-button, [type=number]::-webkit-outer-spin-button {
            height: auto
        }

        [type=search] {
            -webkit-appearance: textfield;
            outline-offset: -2px
        }

        [type=search]::-webkit-search-cancel-button, [type=search]::-webkit-search-decoration {
            -webkit-appearance: none
        }

        ::-webkit-file-upload-button {
            -webkit-appearance: button;
            font: inherit
        }

        details, menu {
            display: block
        }

        summary {
            display: list-item
        }

        canvas {
            display: inline-block
        }

        template {
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        }

        [hidden] {
            display: none
        }

        body {
            font-size: 13px;
            line-height: 1.5
        }

        .container {
            max-width: 728px;
            margin: 0 auto
        }

        a {
            text-decoration: none;
            color: #069
        }

        a:hover, a:focus {
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        }

        button {
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        }

        .background-gray-medium {
            background-color: #dadada;
        }

        .small-header-icons {
            padding-right: .5em
        }

        .small-space-below {
            margin-bottom: .5em
        }

        img.block {
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        }

        h1, .h1 {
            font-size: 30px;
            font-size: 3rem
        }

        h2, .h2 {
            font-size: 24px;
            font-size: 2.4rem
        }

        h3, .h3 {
            font-size: 17px;
            font-size: 1.7rem
        }

        h4, .h4 {
            font-size: 14px;
            font-size: 1.4rem
        }

        .text-orange {
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        }

        .upper {
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        }

        .no-underline {
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        }

        cite {
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        }

        ins {
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        }

        .hide {
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            visibility: hidden
        }

        .show {
            display: block;
            visibility: visible
        }

        .show-inline {
            display: inline;
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        }

        .clean-list {
            padding: 0;
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        }

        .number {
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        }

        .bullet {
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        }

        .none {
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        }

        .ucletter {
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        }

        .lcletter {
            list-style-type: lower-alpha
        }

        .ucroman {
            list-style-type: upper-roman
        }

        .lcroman {
            list-style-type: lower-roman
        }

        .inside {
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        }

        .tighten-line-height {
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        }

        .credit {
            font-size: 10px
        }

        .inline-list > li, .inline-list > dt, .inline-list > dd {
            display: inline
        }

        .inline-group-item {
            display: inline-block;
            vertical-align: bottom
        }

        img {
            border: 0;
            max-width: 100%;
            height: auto;
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        }

        .header-logo-container h1 {
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            padding: 10px;
            background: #333
        }

        .menu-button {
            border: 1px solid #ccc;
            padding: 10px 20px;
            float: right;
            margin: 5px 10px
        }

        button {
            font-family: Source Sans Pro, arial, helvetica, sans-serif
        }

        #menu {
            display: block;
            visibility: visible
        }

        .content {
            max-width: 728px;
            margin: 0 auto
        }

        .grade-c-hide {
            display: none
        }

        .grade-c-show {
            display: block;
            visibility: visible
        }

        .grade-c-invisible {
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        }

        .js .js-hide {
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        }

        #content {
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        }

        .js-readcube-pdf {
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            top: 0;
            right: 50%
        }

        .box-sizing {
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            -moz-box-sizing: border-box;
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        }

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            top: 20px
        }

        .bottom-negative-20 {
            bottom: -20px
        }

        .flex-box {
            display: inline-block;
            display: -webkit-flex;
            display: flex
        }

        .flex-box-item {
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        }

        .flex-box-column {
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            flex-direction: column
        }

        .flex-box-column-item {
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        }

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            align-items: center;
            justify-content: center
        }

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        }

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                display: flex
            }

            .mq1200-flex-box-item {
                flex-grow: 1
            }
        }

        @media only screen and (max-width: 54.688em) {
            .mq875-flex-box {
                display: inline-block;
                display: flex
            }

            .mq875-flex-box-item {
                flex-grow: 1
            }

            .mq875-kill-flex-space-between {
                justify-content: flex-start
            }
        }

        @media only screen and (max-width: 40em) {
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                display: inline-block;
                display: flex
            }

            .mq640-flex-box-item {
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            }
        }

        @media only screen and (max-width: 30em) {
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                display: inline-block;
                display: flex
            }

            .mq480-flex-box-item {
                flex-grow: 1
            }
        }

        .grid, .grid-ng {
            float: left;
            padding-left: 0 !important;
            padding-right: 0 !important
        }

        .grid-inline-wrapper {
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            margin: 0;
            padding: 0;
            letter-spacing: -.31em
        }

        .grid-inline, .grid-inline-ng {
            display: inline-block;
            letter-spacing: .01em;
            vertical-align: top;
            padding-left: 0 !important;
            padding-right: 0 !important
        }

        .grid-left-2 {
            margin-left: 17.5%;
        }

        .grid-8 {
            width: 65.6%;
        }

        .container {
            max-width: none;
            margin: 0;
            padding: 0
        }

        .pin-right {
            float: right
        }

        .pin-left {
            float: left
        }

        .clear-float {
            float: none
        }

        .position-absolute {
            position: absolute
        }

        .position-relative {
            position: relative
        }

        .position-right {
            right: 0
        }

        .block {
            display: block
        }

        .inline-block {
            display: inline-block
        }

        .cleared {
            zoom: 1
        }

        .cleared:before, .cleared:after {
            content: ' ';
            display: table
        }

        .cleared:after {
            clear: both
        }

        .clear {
            clear: both
        }

        .clear-left {
            clear: left
        }

        .clear-right {
            clear: right
        }

        .hide, .grade-a-hide, .js .js-hide, .svg .js-svg-hide {
            display: none;
            visibility: hidden
        }

        .show, .js .js-show {
            display: block;
            visibility: visible
        }

        .show-inline, .js .js-show-inline {
            display: inline;
            visibility: visible
        }

        .visually-hidden, .js .js-visually-hidden {
            width: 1px;
            height: 1px;
            position: absolute !important;
            clip: rect(1px, 1px, 1px, 1px)
        }

        .search-input-field input, .search-input-field input[type=search] {
            border: 1px solid transparent;
            outline: 0;
            padding: 0;
            width: 100%;
            background: none;
            font-size: 2.8rem;
            line-height: 1.25
        }

        .cleared:before, .cleared:after {
            content: ' ';
            display: table;
        }

        .search-input-btn {
            top: 0;
            right: 0;
            width: 4rem;
            min-width: 40px;
            height: 4rem;
            min-height: 40px;
            padding-left: 1rem;
            padding-bottom: 1px;
            background: #fff
        }

        .search-input-layout {
            margin-right: -12rem
        }

        .search-input-field {
            margin-right: 12rem
        }

        .search-input-suffix {
            width: 7rem;
            min-width: 70px
        }

        .search-input-suffix-inner {
            padding: .5rem 1px .5rem 5px;
            border-left: 1px solid #069
        }

        .search-input-suffix select {
            width: 100%
        }

        .extend-suffix .search-input-layout {
            margin-right: -17rem
        }

        .extend-suffix .search-input-field {
            margin-right: 17rem
        }

        .extend-suffix .search-input-suffix {
            width: 12rem;
            min-width: 120px
        }

        .search-input-box {
            margin-bottom: .4rem;
        }

        @media only screen and (max-width: 40em) {
            .search-input-layout, .extend-suffix .search-input-layout {
                margin-right: 0
            }

            .search-input-field, .extend-suffix .search-input-field {
                margin-right: 5rem
            }

            .search-input-suffix, .extend-suffix .search-input-suffix {
                position: absolute;
                right: 0;
                top: 5rem
            }

            .search-input-suffix-inner {
                border: 0
            }

            .search-input-field input, .search-input-field input[type=search] {
                font-size: 2rem;
                line-height: 1.75
            }
        }

        .icon {
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        }

        .icon-center {
            background-position: 50% 50%;
        }

        .ie9 .search-btn {
            background-size: 16px auto
        }

        .search-btn {
            width: 4rem;
            min-width: 40px;
            height: 4rem;
            min-height: 40px;
            font-size: 1.2rem;
            background-image: url(data:image/svg+xml;base64,PHN2ZyB4bWxucz0iaHR0cDovL3d3dy53My5vcmcvMjAwMC9zdmciIHdpZHRoPSIxNiIgaGVpZ2h0PSIxNiIgdmlld0JveD0iMCAwIDE2IDE2Ij48cGF0aCBmaWxsPSIjZmZmIiBkPSJNMTAuNiAxLjNjMi4zIDAgNC4yIDEuOSA0LjIgNC4ycy0xLjkgNC4yLTQuMiA0LjItNC4yLTItNC4yLTQuMyAxLjktNC4xIDQuMi00LjFtMC0xLjNjLTMgMC01LjQgMi40LTUuNCA1LjRzMi40IDUuNCA1LjQgNS40IDUuNC0yLjQgNS40LTUuNC0yLjQtNS40LTUuNC01LjR6bS0zLjUgMTEuMWwtNC45IDQuOS0yLjItMi4yIDQuOS00LjggMi4yIDIuMXoiLz48L3N2Zz4=), none;
            background-size: 40% auto;
            background-color: #069;
            border: 1px solid #069;
            color: transparent;
        }

        .search-btn:hover {
            background-image: url(data:image/svg+xml;base64,PHN2ZyB4bWxucz0iaHR0cDovL3d3dy53My5vcmcvMjAwMC9zdmciIHdpZHRoPSIxNiIgaGVpZ2h0PSIxNiIgdmlld0JveD0iMCAwIDE2IDE2Ij48cGF0aCBmaWxsPSIjMDY5IiBkPSJNMTAuNiAxLjNjMi4zIDAgNC4yIDEuOSA0LjIgNC4ycy0xLjkgNC4yLTQuMiA0LjItNC4yLTItNC4yLTQuMyAxLjktNC4xIDQuMi00LjFtMC0xLjNjLTMgMC01LjQgMi40LTUuNCA1LjRzMi40IDUuNCA1LjQgNS40IDUuNC0yLjQgNS40LTUuNC0yLjQtNS40LTUuNC01LjR6bS0zLjUgMTEuMWwtNC45IDQuOS0yLjItMi4yIDQuOS00LjggMi4yIDIuMXoiLz48L3N2Zz4=), none;
            background-color: #fff
        }

        @media screen and (-ms-high-contrast: active),(-ms-high-contrast: none) {
            .search-btn {
                background-size: 16px auto
            }
        }

        .search-input-field input::-webkit-input-placeholder {
            color: #999
        }

        .search-input-field input:-moz-placeholder {
            color: #999;
            opacity: 1
        }

        .search-input-field input::-moz-placeholder {
            color: #999;
            opacity: 1
        }

        .search-input-field input:-ms-input-placeholder {
            color: #999
        }

        html {
            height: 100%;
            overflow-y: scroll;
            font-size: 62.5%
        }

        body {
            min-height: 100%;
            font-family: Source Sans Pro, arial, helvetica, sans-serif;
            line-height: 1.76;
            color: #222;
            background: #eee;
            font-size: 1.7em;
            letter-spacing: .01em
        }

        figure {
            margin: 0
        }

        body, div, dl, dt, dd, pre, code, form, fieldset, legend, input, button, textarea, p, blockquote, th, td {
            margin: 0;
            padding: 0
        }

        h1, h2, h3, h4, h5, h6, small {
            margin: 0;
            padding: 0
        }

        abbr[title] {
            text-decoration: none
        }

        .content {
            max-width: 1000px;
            margin: 0 auto
        }

        .main-column {
            width: 60.2%;
            margin-right: 8.6%
        }

        .z-index-100 {
            z-index: 100
        }

        .z-index-50 {
            z-index: 50
        }

        .z-index-1 {
            z-index: 1
        }

        @media only screen and (min-width: 75.06em) {
            .full-size-width-160 {
                width: 160px
            }
        }

        @media only screen and (max-width: 30em) {
            .mq480-max-width-100 {
                max-width: 100px
            }
        }

        .separator {
            margin-top: 0;
            position: relative;
            border: none;
            border-top: 1px solid #999
        }

        .separator-light {
            border-color: #eee
        }

        .composite-layer {
            -webkit-transform: translateZ(0);
            -moz-transform: translateZ(0);
            transform: translateZ(0)
        }

        @media only screen and (max-width: 75em) {
            .mq1200-kill-flex {
                display: block
            }
        }

        @media only screen and (max-width: 54.688em) {
            .mq875-kill-flex {
                display: block
            }
        }

        @media only screen and (max-width: 40em) {
            .mq640-kill-flex {
                display: block
            }
        }

        @media only screen and (max-width: 30em) {
            .mq480-kill-flex {
                display: block
            }
        }

        .grid-inline, .grid {
            margin-right: 3.2%
        }

        .last {
            margin-right: 0
        }

        .grid-12 {
            width: 100%
        }

        p, ul, ol {
            margin-top: 0;
            margin-bottom: 28px
        }

        .suppress-bottom-margin > * {
            margin-bottom: 0
        }

        .add-top-margin > * + * {
            margin-top: 28px
        }

        .add-top-margin-small > * + * {
            margin-top: 14px
        }

        .h4 {
            color: #666;
            font-family: Source Sans Pro, arial, helvetica, sans-serif;
            font-weight: 700;
            text-transform: uppercase;
            -webkit-font-smoothing: antialiased;
            -moz-osx-font-smoothing: grayscale
        }

        h5, .h5 {
            font-size: 1.2rem
        }

        h1, .h1, h2, .h2, h3, .h3, h5, .h5 {
            font-family: Lora, Palatino, Times, Times New Roman, serif;
            font-weight: 400
        }

        .serif {
            font-family: Lora, Palatino, Times, Times New Roman, serif
        }

        .wf-loading h1, .wf-loading .h1, .wf-loading h2, .wf-loading .h2, .wf-loading h3, .wf-loading .h3, .wf-loading h5, .wf-loading .h5, .wf-loading .serif {
            font-family: Palatino, Times, Times New Roman, serif
        }

        .wf-active .serif, .wf-active h1, .wf-active .h1, .wf-active h2, .wf-active .h2, .wf-active h3, .wf-active .h3, .wf-active h5, .wf-active .h5 {
            font-family: Lora, Palatino, Times, Times New Roman, serif
        }

        .contrast-text, .contrast-text a {
            color: #fff
        }

        .text-gray-light {
            color: #666
        }

        .equalize-line-height {
            line-height: 1em
        }

        .extra-tight-line-height {
            line-height: 1.3
        }

        .tighten-line-height {
            line-height: 1.4
        }

        .doctype-line-height {
            line-height: 24px
        }

        .text11 {
            font-size: 1.1rem
        }

        fieldset {
            border: 0
        }

        input + label {
            padding-left: .5em
        }

        .background-white {
            background-color: #fff
        }

        .background-gray-dark {
            background-color: #333
        }

        [data-hellip] {
            max-height: 12.5rem;
            overflow: hidden
        }

        @media only screen and (max-width: 54.688em) {
            [data-hellip] {
                max-height: 15rem;
                overflow: hidden
            }
        }

        .ma0 {
            margin: 0
        }

        .mt20 {
            margin-top: 20px
        }

        .mt70 {
            margin-top: 70px
        }

        .mb20 {
            margin-bottom: 20px
        }

        .mb30 {
            margin-bottom: 30px
        }

        .pa20 {
            padding: 20px
        }

        .pt15 {
            padding-top: 15px
        }

        .pt30 {
            padding-top: 30px
        }

        .pb30 {
            padding-bottom: 30px;
        }

        .pb40 {
            padding-bottom: 40px
        }

        @media only screen and (max-width: 75em) {
            .mq1200-padded {
                padding-left: 20px;
                padding-right: 20px
            }
        }

        @media only screen and (max-width: 75em) {
            .mq1200-pl0 {
                padding-left: 0
            }

            .mq1200-pl20 {
                padding-left: 20px
            }
        }

        @media only screen and (max-width: 54.688em) {
            .mq875-ml0 {
                margin-left: 0
            }

            .mq875-mb20 {
                margin-bottom: 20px
            }

            .mq875-mt30 {
                margin-top: 30px
            }

            .mq875-pa0 {
                padding: 0
            }

            .mq875-pl0 {
                padding-left: 0
            }

            .mq875-pl20 {
                padding-left: 20px
            }

            .mq875-pr20 {
                padding-right: 20px
            }
        }

        @media only screen and (max-width: 40em) {
            .mq640-pa4 {
                padding: 4px
            }

            .mq640-pt10 {
                padding-top: 10px
            }

            .mq640-pb10 {
                padding-bottom: 10px
            }

            .mq640-pt20 {
                padding-top: 20px
            }

            .mq640-pl20 {
                padding-left: 20px
            }

            .mq640-pl0 {
                padding-left: 0
            }

            .mq640-pr5 {
                padding-right: 5px
            }

            .mq640-pr20 {
                padding-right: 20px
            }

            .mq640-pb30 {
                padding-bottom: 30px
            }

            .mq640-mt0 {
                margin-top: 0
            }

            .mq640-ml0 {
                margin-left: 0
            }

            .mq640-mb0 {
                margin-bottom: 0
            }

            .mq640-mb5 {
                margin-bottom: 5px
            }

            .mq640-mt20 {
                margin-top: 20px
            }

            .mq640-mb20 {
                margin-bottom: 20px
            }

            .mq640-mb30 {
                margin-bottom: 30px
            }
        }

        @media only screen and (max-width: 30em) {
            .mq480-mr0 {
                margin-right: 0
            }

            .mq480-mb10 {
                margin-bottom: 10px
            }

            .mq480-pt10 {
                padding-top: 10px
            }
        }

        .table {
            display: table
        }

        .table-row {
            display: table-row
        }

        .table-cell {
            display: table-cell
        }

        @media only screen and (max-width: 75em) {
            .mq1200-block {
                display: block
            }
        }

        @media only screen and (max-width: 54.688em) {
            .mq875-block {
                display: block
            }
        }

        @media only screen and (max-width: 40em) {
            .mq640-block {
                display: block
            }
        }

        @media only screen and (max-width: 30em) {
            .mq480-block {
                display: block
            }
        }

        .js .js-author-etal {
            display: none;
            visibility: hidden
        }

        .js .js-authors-expanded .js-author-etal {
            display: inline;
            visibility: visible
        }

        @media only screen and (max-width: 75em) {
            .mq1200-hide, .js .js-mq1200-hide {
                display: none;
                visibility: hidden
            }

            .mq1200-show, .js .js-mq1200-show {
                display: block;
                visibility: visible
            }

            .mq1200-show-inline, .js .js-mq1200-show-inline {
                display: inline;
                visibility: visible
            }

            .mq1200-visually-show {
                clip: auto;
                height: auto;
                position: static !important;
                width: auto
            }
        }

        @media only screen and (max-width: 54.688em) {
            .mq875-hide, .js .js-mq875-hide {
                display: none;
                visibility: hidden
            }

            .mq875-show, .js .js-mq875-show {
                display: block;
                visibility: visible
            }

            .mq875-show-inline, .js .js-mq875-show-inline {
                display: inline;
                visibility: visible
            }

            .mq875-visually-show {
                clip: auto;
                height: auto;
                position: static !important;
                width: auto
            }
        }

        @media only screen and (max-width: 40em) {
            .mq640-hide, .js .js-mq640-hide {
                display: none;
                visibility: hidden
            }

            .mq640-show, .js .js-mq640-show {
                display: block;
                visibility: visible
            }

            .mq640-show-inline, .js .js-mq640-show-inline {
                display: inline;
                visibility: visible
            }

            .mq640-visually-show {
                clip: auto;
                height: auto;
                position: static !important;
                width: auto
            }
        }

        @media only screen and (max-width: 30em) {
            .mq480-hide, .js .js-mq480-hide {
                display: none;
                visibility: hidden
            }

            .mq480-show, .js .js-mq480-show {
                display: block;
                visibility: visible
            }

            .mq480-show-inline, .js .js-mq480-show-inline {
                display: inline;
                visibility: visible
            }

            .mq480-visually-show {
                clip: auto;
                height: auto;
                position: static !important;
                width: auto
            }

            .js .js-smaller-author-etal {
                display: none;
                visibility: hidden
            }

            .js .js-authors-expanded .js-smaller-author-etal {
                display: inline;
                visibility: visible
            }
        }

        .container-type-hero:first-child .hero {
            top: -100px;
            margin-bottom: -100px;
            position: relative
        }

        .container-type-hero + .container-type-article-grid, .container-type-hero + .container-type-mixed-grid {
            margin-top: -30px
        }

        .container-type-hero + .container-type-article-grid .article-grid-heading, .container-type-hero + .container-type-mixed-grid .mixed-grid-heading {
            width: 1px;
            height: 1px;
            position: absolute !important;
            clip: rect(1px, 1px, 1px, 1px)
        }

        .advert img, .advert object, .advert embed {
            max-width: 100%
        }

        .billboard-container img, .billboard-container object, .billboard-container embed, .billboard-inner iframe {
            max-height: 250px
        }

        .leaderboard-container img, .leaderboard-container object, .leaderboard-container embed, .leaderboard-container iframe {
            max-height: 90px
        }

        .medium-rectangle object, .medium-rectangle embed, .medium-rectangle iframe {
            max-height: 250px
        }

        .skyscraper img, .skyscraper object, .skyscraper embed, .skyscraper iframe, .skyscraper a {
            display: block;
            margin: 0 auto
        }

        @media only screen and (max-width: 45.75em) {
            .leaderboard-container object, .leaderboard-container embed {
                display: none
            }

            .billboard-container object, .leaderboard-container embed {
                display: none
            }
        }

        a {
            text-decoration: none;
            vertical-align: baseline;
            color: #069
        }

        a:hover, a:focus {
            text-decoration: underline;
            -webkit-text-decoration-skip: skip;
            text-decoration-skip: ink
        }

        .block-link:hover > span, .block-link:focus > span {
            text-decoration: underline;
            -webkit-text-decoration-skip: skip;
            text-decoration-skip: ink
        }

        .background-brand-secondary .tools-popup a {
            color: #069
        }

        .tab-like button {
            color: #069;
            border: solid #069;
            border-width: 1px 1px 1px 0;
            background-color: #dae5ea;
            position: relative;
            width: 100%
        }

        .tab-like:first-child button {
            border-left: 1px solid #069
        }

        .tab-like button:focus {
            text-decoration: none;
            outline: thin dotted
        }

        .tab-like button.selected {
            color: #222;
            background-color: #fff;
            border-bottom: 1px solid #fff;
            font-weight: 700;
            -webkit-font-smoothing: antialiased;
            -moz-osx-font-smoothing: grayscale
        }

        .tab-like:first-child button:before, .tab-like button.selected:after {
            content: '';
            width: 1px;
            height: 1px;
            background-color: #069;
            position: absolute;
            bottom: -1px;
            right: -1px
        }

        .tab-like:first-child button:before {
            left: -1px
        }

        nav ul, nav ol {
            list-style: none;
            list-style-image: none
        }

        .ie11-list-hack {
            list-style-image: url(data:image/gif;base64,R0lGODlhAQABAIAAAAAAAP///yH5BAEAAAAALAAAAAABAAEAAAIBRAA7)
        }

        .horizontal-list > li {
            float: left
        }

        .inline-list > li, .inline-list > dt, .inline-list > dd {
            display: inline
        }

        .piped-list > li {
            padding: 0 8px;
            border-left: 1px solid #dadada
        }

        .piped-list > li:first-child {
            border: 0;
            padding-left: 0
        }

        .indented-list p, .indented-list ul, .indented-list h1, .indented-list h3 {
            margin-left: 50px
        }

        table {
            width: 100%;
            margin-bottom: 0;
            border-collapse: collapse;
            border-spacing: 0
        }

        .table-group > table {
            border-bottom: 1px solid #dadada
        }

        .table-group > .last-table {
            border-bottom: 0
        }

        .table-group > table + table {
            margin-top: 24px;
            border-top: 1px solid #dadada
        }

        th, td {
            padding: 5px 10px;
            border-right: 1px solid #dadada;
            text-align: left
        }

        .match-spacing {
            padding: 5px 10px
        }

        th {
            vertical-align: bottom
        }

        th[valign=top], .row-scope {
            vertical-align: top
        }

        th[valign=middle] {
            vertical-align: middle
        }

        td {
            vertical-align: top
        }

        .table-highlight tr:hover, .table-highlight tr:hover th, .table-highlight tr:hover td {
            background-color: #ff9
        }

        .last-cell {
            border-right: 0
        }

        thead tr {
            border-bottom: 1px solid #dadada
        }

        tbody tr {
            border-top: 1px solid #dadada
        }

        tbody tr:first-child {
            border-top: 0
        }

        th, tr {
            background: #eee
        }

        .odd {
            background: #fff
        }

        tfoot tr {
            border-top: 2px solid #dadada
        }

        tfoot td {
            font-weight: 700;
            -webkit-font-smoothing: antialiased;
            -moz-osx-font-smoothing: grayscale
        }

        .js .tab-single + .tab-content > div {
            border: 0;
            padding: 0
        }

        .image-constraint-stretch img {
            max-width: none;
            max-height: none;
            width: 25%
        }

        @media only screen and (min-width: 75.06em) {
            .full-size-last {
                margin-right: 0
            }

            .full-size-clear {
                clear: both
            }

            .full-size-kill-left {
                margin-left: 0
            }
        }

        @media only screen and (max-width: 75em) and (min-width: 54.74em) {
            .just-mq1200-last {
                margin-right: 0
            }

            .just-mq1200-clear {
                clear: both
            }

            .just-mq1200-kill-left {
                margin-left: 0
            }
        }

        @media only screen and (max-width: 54.688em) and (min-width: 40.06em) {
            .just-mq875-last {
                margin-right: 0
            }

            .just-mq875-clear {
                clear: both
            }

            .just-mq875-kill-left {
                margin-left: 0
            }
        }

        @media only screen and (max-width: 40em) and (min-width: 30.06em) {
            .just-mq640-last {
                margin-right: 0
            }

            .just-mq640-clear {
                clear: both
            }

            .just-mq640-kill-left {
                margin-left: 0
            }
        }

        @media only screen and (max-width: 30em) {
            .just-mq480-last {
                margin-right: 0
            }

            .just-mq480-clear {
                clear: both
            }

            .just-mq480-kill-left {
                margin-left: 0
            }
        }

        @media only screen and (max-width: 75em) {
            .mq1200-grid-12 {
                width: 100%
            }

            .mq1200-grid-4 {
                width: 31.2%
            }
        }

        @media only screen and (max-width: 54.688em) {
            .mq875-grid-12 {
                width: 100%
            }

            .mq875-grid-10 {
                width: 82.8%
            }

            .mq875-grid-8 {
                width: 65.6%
            }

            .mq875-grid-7 {
                width: 57%
            }

            .mq875-grid-6 {
                width: 48.4%
            }

            .mq875-grid-4 {
                width: 31.2%
            }

            .mq875-grid-3 {
                width: 22.6%
            }

            .mq875-grid-1 {
                width: 5.4%
            }

            .mq875-kill-left {
                margin-left: 0
            }
        }

        @media only screen and (max-width: 40em) {
            .mq640-last {
                margin-right: 0
            }

            .mq640-clear {
                clear: both
            }

            .mq640-grid-12 {
                width: 100%
            }

            .mq640-grid-9 {
                width: 74.2%
            }

            .mq640-grid-6 {
                width: 48.4%
            }

            .mq640-grid-4 {
                width: 31.2%
            }

            .mq640-kill-left {
                margin-left: 0
            }
        }

        @media only screen and (max-width: 30em) {
            .mq480-grid-12 {
                width: 100%
            }
        }

        body:before {
            content: 'full';
            position: absolute;
            top: -1px;
            left: -1px;
            height: 0;
            width: 0;
            overflow: hidden
        }

        @media only screen and (max-width: 75em) {
            body:before {
                content: 'mq1200'
            }
        }

        @media only screen and (max-width: 54.688em) {
            body:before {
                content: 'mq875'
            }
        }

        @media only screen and (max-width: 40em) {
            body:before {
                content: 'mq640'
            }
        }

        @media only screen and (max-width: 30em) {
            body:before {
                content: 'mq480'
            }
        }

        .skiplink {
            width: 0;
            height: 0;
            display: block;
            overflow: hidden
        }

        .skiplink:focus {
            width: auto;
            height: auto;
            overflow: visible;
            margin: 2px 0 1px;
            padding: 2px
        }

        .breadcrumbs {
            padding: 10px 20px
        }

        .breadcrumbs ol {
            overflow: hidden;
            line-height: 1.1em;
            white-space: nowrap;
            text-overflow: ellipsis
        }

        .small-header-icons {
            padding-left: 7px;
            padding-right: 7px;
            margin-top: 14px;
            width: 60px
        }

        .small-header-icons:hover {
            text-decoration: none
        }

        @media only screen and (max-width: 75em) {
            .breadcrumbs {
                padding: 10px 20px 10px 0
            }
        }

        .sticky-header {
            width: 100%;
            height: 39px;
            position: fixed;
            top: 0
        }

        .sticky-header .menu-button {
            top: 0;
            margin: 7px 0
        }

        .header-logo-container {
            padding: 15px 20px
        }

        .sticky-header .header-logo-container {
            padding: 7px 0 7px 20px;
            height: 25px
        }

        .sticky-header .header-logo {
            height: 25px
        }

        .sticky-header .header-logo img {
            max-height: 25px
        }

        .sticky-header .small-header-icons {
            border-left: 1px solid #8d8d78;
            padding: 7px 16px;
            background-color: #5a5a62;
            margin-top: 0;
            margin-right: 0;
            width: auto
        }

        .sticky-header .small-header-icons:first-of-type {
            border-left: 0
        }

        .sticky-header .small-header-main {
            width: auto;
            margin: 0
        }

        .header-logo-container, .header-logo {
            height: 50px;
            overflow: hidden
        }

        .header-logo img {
            max-height: 50px;
            margin: 0;
            padding: 0
        }

        .header-logo-primary, .sticky-header .header-logo-secondary {
            display: inline;
            visibility: visible
        }

        .header-logo-secondary, .sticky-header .header-logo-primary {
            display: none;
            visibility: hidden
        }

        @media only screen and (max-width: 75em) {
            .sticky-header .header-logo-container, .header-logo-container {
                padding-left: 0
            }

            .banner {
                padding-left: 20px;
                padding-right: 0
            }
        }

        @media only screen and (max-width: 54.688em) {
            .menu-button {
                margin: 1px 20px 1px 0
            }

            .header-logo-container {
                padding: 7px 20px 7px 0;
                height: 25px
            }

            .header-logo {
                height: 25px
            }

            .header-logo img {
                max-height: 25px
            }

            .header-logo-primary {
                display: none;
                visibility: hidden
            }

            .small-header-main {
                width: auto;
                margin: 0
            }

            .menu .menu-cell ul {
                margin: 0
            }

            .menu .menu-cell li {
                display: block;
                padding: 20px;
                float: none;
                border-bottom: 1px solid #dadada
            }

            .menu .background-gradient-brand-gray {
                background-image: none
            }

            .menu h2 {
                margin: 0;
                padding: 20px
            }
        }

        @media only screen and (max-width: 40em) {
            .menu-button, .sticky-header .menu-button {
                margin-right: 4px
            }

            .sticky-header .header-logo-primary, .header-logo-primary {
                display: inline;
                visibility: visible
            }

            .header-logo-secondary, .sticky-header .header-logo-secondary {
                display: none;
                visibility: hidden
            }
        }

        .js .menu {
            position: absolute;
            overflow-y: auto;
            left: 0;
            right: 0;
            top: -500px;
            opacity: 0;
            margin: 0
        }

        .js .sticky-header ~ .menu {
            position: fixed
        }

        .js .menu-open {
            opacity: 1
        }

        .beta-switch {
            display: block;
            background-color: #069;
            color: #fff
        }

        .beta-switch a, .beta-switch a:visited {
            color: #fff;
            text-decoration: underline;
            -webkit-text-decoration-skip: skip;
            text-decoration-skip: ink
        }

        @media only screen and (max-width: 75em) and (min-width: 54.74em) {
            .banner {
                padding-right: 20px
            }
        }

        .stack sup, .stack sub {
            display: table-row;
            line-height: 1
        }

        svg {
            float: left
        }

        path {
            fill: none
        }

        text {
            font-size: 14px;
            fill: #666
        }

        line {
            stroke: #dadada;
            stroke-width: 1;
            stroke-dasharray: 2 2;
            fill: none
        }

        .map-controls a {
            background-color: #fff;
            background-color: rgba(255, 255, 255, .8)
        }

        @media only screen and (max-width: 62.5em) {
            text {
                font-size: 13px
            }
        }

        @media only screen and (max-width: 48em) {
            text {
                font-size: 12px
            }
        }

        .grade-c-show {
            display: none;
            visibility: hidden
        }

        .grade-c-invisible {
            visibility: visible
        }

        .container {
            max-width: none;
            padding: 0;
            padding: unset
        }

        .inline-list.js-list-authors > li {
            padding-right: 0
        }

        @media only screen and (min-width: 350px) {
            [data-back-link] {
                float: left
            }
        }

        .issue-research li:first-child {
            border-top: 0
        }

        @media only screen and (min-width: 40em) {
            .issue-research li:nth-child(-n+2) {
                border-top: 0
            }
        }

        .background-gradient-brand-primary {
            background-color: -webkit-gradient(left top, right top, color-stop(50%, #940720), color-stop(50%, transparent));
            /* for Android 4.3- */
            background: url("data:image/svg+xml;charset=utf-8,%3Csvg%20xmlns=%22http:%2F%2Fwww.w3.org%2F2000%2Fsvg%22%20preserveAspectRatio=%22none%22%20version=%221.0%22%20width=%22100%25%22%20height=%22100%25%22%20xmlns:xlink=%22http:%2F%2Fwww.w3.org%2F1999%2Fxlink%22%3E%3Cdefs%3E%3ClinearGradient%20id=%22myLinearGradient%22%20x1=%220%25%22%20y1=%220%25%22%20x2=%22100%25%22%20y2=%220%25%22%20spreadMethod=%22pad%22%3E%3Cstop%20offset=%220%25%22%20stop-color=%22%23940720%22%20stop-opacity=%221%22%2F%3E%3Cstop%20offset=%2250%25%22%20stop-color=%22%23940720%22%20stop-opacity=%221%22%2F%3E%3Cstop%20offset=%2250%25%22%20stop-color=%22%23940720%22%20stop-opacity=%220%22%2F%3E%3Cstop%20offset=%22100%25%22%20stop-color=%22%23940720%22%20stop-opacity=%220%22%2F%3E%3C%2FlinearGradient%3E%3C%2Fdefs%3E%3Crect%20width=%22100%25%22%20height=%22100%25%22%20style=%22fill:url(%23myLinearGradient);%22%20%2F%3E%3C%2Fsvg%3E") repeat-y;
        }

        .background-brand-primary a {
            color: #fff;
        }

        .background-brand-secondary a {
            color: #fff;
        }

        @media only screen and (max-width: 54.688em) {
            .banner {
                background-color: #940720;
            }

            .nav-border .menu-cell li {
                border-bottom: 1px solid #a12a3f;
            }
        }

        .ie7 .small-header-main,
        .ie8 .small-header-main {
            background-color: #940720;
        }

        .header-primary-color {
            background-color: #940720;
        }
    </style>
    <meta name="robots" content="noarchive">
    <meta name="access" content="Yes">
</head>
<!--[if IE 9]>
<body class="ie9 home-page"><![endif]-->
<!--[if gt IE 9]><!-->

<body class="home-page">
<!--<![endif]-->
<div role="banner" class="position-relative cleared z-index-50" data-test="top-containers">
    <header>
        <div class="hide-print container background-gray-dark">
            <div class="content mq1200-padded">
                <a href="#content" id="skiplink" class="skiplink contrast-text js-no-scroll" data-track="click"
                   data-track-source="skip-link" data-track-dest="anchor:Skip to main content">Skip to main content</a>
            </div>
        </div>
        <div class="text-orange content grade-c-show">
            <p>Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To
                obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility
                mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site
                without styles and JavaScript.</p>
        </div>
        <div class="container js-header-container mb20 position-relative z-index-50" data-ui="header-container"
             data-container-type="header">
            <div class="cleared clear hide-print position-relative background-gradient-brand-primary js-header default-header z-index-100 composite-layer tighten-line-height">
                <div class="banner js-banner content mq1200-padded position-relative">
                    <div class="inner-banner cleared">
                        <div class="main-column small-header-main pin-left">
                            <div class="header-logo-container header-primary-color">
                                <h1 class="visually-hidden" data-test="product-title">Nature</h1>
                                <a href="/" class="header-logo inline-block equalize-line-height"
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