CINXE.COM
ÉCOLOGIE - Encyclopædia Universalis
<!DOCTYPE html> <html lang="fr-FR" xml:lang="fr" dir="ltr"> <head> <meta charset="utf-8"> <!-- This website is powered by TYPO3 - inspiring people to share! TYPO3 is a free open source Content Management Framework initially created by Kasper Skaarhoj and licensed under GNU/GPL. TYPO3 is copyright 1998-2024 of Kasper Skaarhoj. Extensions are copyright of their respective owners. Information and contribution at https://typo3.org/ --> <title>ÉCOLOGIE - Encyclopædia Universalis</title> <meta http-equiv="x-ua-compatible" content="IE=edge,chrome=1" /> <meta name="generator" content="TYPO3 CMS" /> <meta name="description" content="Le terme écologie (du grec oikos , demeure, et logos , science) a été proposé par Ernst Haeckel en 1866 pour désigner la science qui étudie les rapports..." /> <meta name="robots" content="index, all, follow" /> <meta name="viewport" content="viewport-fit=cover, width=device-width, initial-scale=1.0, minimum-scale=1.0" /> <meta name="author" content="Encyclopædia Universalis" /> <meta property="og:title" content="ÉCOLOGIE" /> <meta property="og:description" content="Le terme écologie (du grec oikos , demeure, et logos , science) a été proposé par Ernst Haeckel en 1866 pour désigner la science qui étudie les rapports entre les organismes et le milieu où ils vivent. Cette définition reste encore valable, mais elle demande à être approfondie et..." /> <meta property="og:url" content="https://www.universalis.fr/encyclopedie/ecologie/" /> <meta property="og:locale" content="fr_FR" /> <meta property="og:site_name" content="Encyclopædia Universalis" /> <meta property="og:type" content="website" /> <meta property="og:image" content="https://www.universalis.fr/_web/static/icons/favicon.png" /> <meta property="og:image:width" content="256" /> <meta property="og:image:height" content="252" /> <meta name="twitter:title" content="ÉCOLOGIE" /> <meta name="twitter:description" content="Le terme écologie (du grec oikos , demeure, et logos , science) a été proposé par Ernst Haeckel en 1866 pour désigner la science qui étudie les rapports entre les organismes et le milieu où ils vivent. Cette définition reste encore valable, mais elle demande à être approfondie et..." /> <meta name="twitter:card" content="summary" /> <meta name="twitter:site" content="@universalisfr" /> <meta name="location" content="France" /> <meta name="rating" content="General" /> <meta name="distribution" content="Global" /> <meta name="resource-type" content="document" /> <meta name="copyright" content="© Copyright 2024 - Universalis" /> <meta name="apple-mobile-web-app-title" content="Universalis" /> <meta name="apple-mobile-web-app-capable" content="yes" /> <meta name="apple-mobile-web-app-status-bar-style" content="default" /> <meta name="format-detection" content="telephone=no,date=no,address=no,email=no,url=no" /> <link rel="stylesheet" href="/_shared/pub/pub.css?1730108805" media="all"> <link rel="stylesheet" href="/_web/assets/css/home.css?1730119190" media="all"> <link rel="stylesheet" href="/_web/assets/css/classification.css?1730119190" media="all"> <link rel="canonical" href="https://www.universalis.fr/encyclopedie/ecologie/" /> <link rel="next" href="https://www.universalis.fr/encyclopedie/ecologie/2-fonctionnement-des-populations/" /> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"\u00c9COLOGIE","headline":"\u00c9COLOGIE","description":"Le terme \u00e9cologie (du grec oikos , demeure, et logos , science) a \u00e9t\u00e9 propos\u00e9 par Ernst Haeckel en 1866 pour d\u00e9signer la science qui \u00e9tudie les rapports entre les organismes et le milieu o\u00f9 ils vivent. Cette d\u00e9finition reste encore valable, mais elle demande \u00e0 \u00eatre approfondie et pr\u00e9cis\u00e9e, car elle est trop g\u00e9n\u00e9rale. Pour la situer par rapport aux autres sciences biologiques, il est commode de consid\u00e9rer les divers niveaux d'organisation de la mati\u00e8re vivante.. Le syst\u00e8me le plus simple ayant toutes les caract\u00e9ristiques fondamentales des \u00eatres vivants est la cellule. Des cellules, associ\u00e9es en tissus et en organes, sont int\u00e9gr\u00e9es en organismes pluricellulaires, animaux ou v\u00e9g\u00e9taux....","url":"https:\/\/www.universalis.fr\/encyclopedie\/ecologie\/","wordCount":"20635","image":{"@type":"ImageObject","url":"https:\/\/www.universalis.fr\/_web\/static\/icons\/favicon.png"},"isPartOf":{"@type":"WebPage","url":"https:\/\/www.universalis.fr\/encyclopedie\/ecologie\/","name":"\u00c9COLOGIE"},"author":[{"@context":"https:\/\/schema.org","@type":"Person","name":"Patrick BLANDIN","familyName":"BLANDIN","givenName":"Patrick","description":"Professeur au Mus\u00e9um national d'histoire naturelle, directeur de la Grande Galerie de l'\u00e9volution.","url":"https:\/\/www.universalis.fr\/auteurs\/patrick-blandin\/"},{"@context":"https:\/\/schema.org","@type":"Person","name":"Denis COUVET","familyName":"COUVET","givenName":"Denis","description":"Professeur au Mus\u00e9um national d'histoire naturelle, Paris.","url":"https:\/\/www.universalis.fr\/auteurs\/denis-couvet\/"},{"@context":"https:\/\/schema.org","@type":"Person","name":"Maxime LAMOTTE","familyName":"LAMOTTE","givenName":"Maxime","description":"Professeur honoraire \u00e0 l'universit\u00e9 de Paris-VI-Pierre-et-Marie-Curie (facult\u00e9 des sciences), ancien directeur du laboratoire de zoologie de l'\u00c9cole normale sup\u00e9rieure.","url":"https:\/\/www.universalis.fr\/auteurs\/maxime-lamotte\/"},{"@context":"https:\/\/schema.org","@type":"Person","name":"Cesare F. SACCHI","familyName":"SACCHI","givenName":"Cesare F.","description":"Professeur d'\u00e9cologie \u00e0 l'universit\u00e9 de Paris-VI, directeur de l'Institut d'\u00e9cologie et d'\u00e9thologie de Pavie.","url":"https:\/\/www.universalis.fr\/auteurs\/cesare-f.-sacchi\/"}],"publisher":{"@type":"Organization","name":"Encyclop\u00e6dia Universalis","alternateName":"Universalis","logo":{"@type":"ImageObject","url":"https:\/\/www.universalis.fr\/_web\/assets\/img\/banniere-fr.png"},"sameAs":["https:\/\/www.facebook.com\/EncyclopedieUniversalis\/","https:\/\/www.linkedin.com\/company\/encyclopaedia-universalis","https:\/\/www.twitter.com\/universalisfr","https:\/\/www.youtube.com\/channel\/UC0srTZiJaRaDw3fAvtCo4Ow"],"slogan":"Passeur de savoir"}}</script> <script type="application/ld+json">[{"@context":"https:\/\/schema.org","@type":"BreadcrumbList","itemListElement":[{"@type":"ListItem","position":1,"name":"Classification","item":"https:\/\/www.universalis.fr\/classification\/"},{"@type":"ListItem","position":2,"name":"Sciences de la vie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/"},{"@type":"ListItem","position":3,"name":"\u00c9cologie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/"}],"numberOfItems":3,"itemListOrder":"Ascending","name":"Classification : \u00c9cologie"},{"@context":"https:\/\/schema.org","@type":"BreadcrumbList","itemListElement":[{"@type":"ListItem","position":1,"name":"Classification","item":"https:\/\/www.universalis.fr\/classification\/"},{"@type":"ListItem","position":2,"name":"Sciences de la vie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/"},{"@type":"ListItem","position":3,"name":"\u00c9cologie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/"},{"@type":"ListItem","position":4,"name":"\u00c9cophysiologie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/ecophysiologie\/"}],"numberOfItems":4,"itemListOrder":"Ascending","name":"Classification : \u00c9cophysiologie"},{"@context":"https:\/\/schema.org","@type":"BreadcrumbList","itemListElement":[{"@type":"ListItem","position":1,"name":"Classification","item":"https:\/\/www.universalis.fr\/classification\/"},{"@type":"ListItem","position":2,"name":"Sciences de la vie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/"},{"@type":"ListItem","position":3,"name":"\u00c9cologie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/"},{"@type":"ListItem","position":4,"name":"Biologie et dynamique des populations","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/biologie-et-dynamique-des-populations\/"}],"numberOfItems":4,"itemListOrder":"Ascending","name":"Classification : Biologie et dynamique des populations"},{"@context":"https:\/\/schema.org","@type":"BreadcrumbList","itemListElement":[{"@type":"ListItem","position":1,"name":"Classification","item":"https:\/\/www.universalis.fr\/classification\/"},{"@type":"ListItem","position":2,"name":"Sciences de la vie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/"},{"@type":"ListItem","position":3,"name":"\u00c9cologie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/"},{"@type":"ListItem","position":4,"name":"Structure et fonctionnement des \u00e9cosyst\u00e8mes","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/structure-et-fonctionnement-des-ecosystemes\/"}],"numberOfItems":4,"itemListOrder":"Ascending","name":"Classification : Structure et fonctionnement des \u00e9cosyst\u00e8mes"},{"@context":"https:\/\/schema.org","@type":"BreadcrumbList","itemListElement":[{"@type":"ListItem","position":1,"name":"Classification","item":"https:\/\/www.universalis.fr\/classification\/"},{"@type":"ListItem","position":2,"name":"Sciences de la vie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/"},{"@type":"ListItem","position":3,"name":"Sciences de la vie: instruments et techniques exp\u00e9rimentales","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/sciences-de-la-vie-instruments-et-techniques-experimentales\/"}],"numberOfItems":3,"itemListOrder":"Ascending","name":"Classification : Sciences de la vie: instruments et techniques exp\u00e9rimentales"},{"@context":"https:\/\/schema.org","@type":"BreadcrumbList","itemListElement":[{"@type":"ListItem","position":1,"name":"Classification","item":"https:\/\/www.universalis.fr\/classification\/"},{"@type":"ListItem","position":2,"name":"Sciences de la vie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/"},{"@type":"ListItem","position":3,"name":"\u00c9cologie","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/"},{"@type":"ListItem","position":4,"name":"\u00c9cologie appliqu\u00e9e","item":"https:\/\/www.universalis.fr\/classification\/sciences-de-la-vie\/ecologie\/ecologie-appliquee\/"}],"numberOfItems":4,"itemListOrder":"Ascending","name":"Classification : \u00c9cologie appliqu\u00e9e"}]</script> <meta name="apple-mobile-web-app-capable" content="yes"/> <meta name="mobile-web-app-capable" content="yes"/> <meta name="apple-mobile-web-app-status-bar-style" content="black"> <meta name="apple-mobile-web-app-title" content="Universalis"> <meta name="theme-color" content="#526D78"> <link rel="preconnect" href="https://statistiques.universalis.fr" /> <link rel="preconnect" href="https://cmp.inmobi.com" /> <link rel="search" type="application/opensearchdescription+xml" title="Universalis FR" href="/_web/static/search-fr.xml" /> <link rel="manifest" href="/_web/static/manifest.json"> <link rel="icon" type="image/png" href="/_web/static/icons/favicon.png"> <link rel="apple-touch-icon" href="/_web/static/icons/fr-apple-touch-icon.png" /> <link rel="apple-touch-icon" sizes="57x57" href="/_web/static/icons/fr-apple-touch-icon-57x57.png" /> <link rel="apple-touch-icon" sizes="72x72" href="/_web/static/icons/fr-apple-touch-icon-72x72.png" /> <link rel="apple-touch-icon" sizes="76x76" href="/_web/static/icons/fr-apple-touch-icon-76x76.png" /> <link rel="apple-touch-icon" sizes="114x114" href="/_web/static/icons/fr-apple-touch-icon-114x114.png" /> <link rel="apple-touch-icon" sizes="120x120" href="/_web/static/icons/fr-apple-touch-icon-120x120.png" /> <link rel="apple-touch-icon" sizes="144x144" href="/_web/static/icons/fr-apple-touch-icon-144x144.png" /> <link rel="apple-touch-icon" sizes="152x152" href="/_web/static/icons/fr-apple-touch-icon-152x152.png" /> <link rel="apple-touch-icon" sizes="180x180" href="/_web/static/icons/fr-apple-touch-icon-180x180.png" /> <link rel="apple-touch-startup-image" media="screen and (device-width: 834px) and (device-height: 1194px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/11__iPad_Pro__10.5__iPad_Pro_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 810px) and (device-height: 1080px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/10.2__iPad_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 744px) and (device-height: 1133px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/8.3__iPad_Mini_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 810px) and (device-height: 1080px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/10.2__iPad_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 820px) and (device-height: 1180px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/10.9__iPad_Air_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 744px) and (device-height: 1133px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/8.3__iPad_Mini_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 1024px) and (device-height: 1366px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/12.9__iPad_Pro_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 428px) and (device-height: 926px) and (-webkit-device-pixel-ratio: 3) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_14_Plus__iPhone_13_Pro_Max__iPhone_12_Pro_Max_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 414px) and (device-height: 896px) and (-webkit-device-pixel-ratio: 3) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_11_Pro_Max__iPhone_XS_Max_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 1024px) and (device-height: 1366px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/12.9__iPad_Pro_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 768px) and (device-height: 1024px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/9.7__iPad_Pro__7.9__iPad_mini__9.7__iPad_Air__9.7__iPad_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 375px) and (device-height: 812px) and (-webkit-device-pixel-ratio: 3) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_13_mini__iPhone_12_mini__iPhone_11_Pro__iPhone_XS__iPhone_X_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 820px) and (device-height: 1180px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/10.9__iPad_Air_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 393px) and (device-height: 852px) and (-webkit-device-pixel-ratio: 3) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_14_Pro_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 414px) and (device-height: 896px) and (-webkit-device-pixel-ratio: 3) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_11_Pro_Max__iPhone_XS_Max_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 414px) and (device-height: 736px) and (-webkit-device-pixel-ratio: 3) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_8_Plus__iPhone_7_Plus__iPhone_6s_Plus__iPhone_6_Plus_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 320px) and (device-height: 568px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/4__iPhone_SE__iPod_touch_5th_generation_and_later_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 393px) and (device-height: 852px) and (-webkit-device-pixel-ratio: 3) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_14_Pro_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 414px) and (device-height: 896px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_11__iPhone_XR_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 375px) and (device-height: 667px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_8__iPhone_7__iPhone_6s__iPhone_6__4.7__iPhone_SE_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 320px) and (device-height: 568px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/4__iPhone_SE__iPod_touch_5th_generation_and_later_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 390px) and (device-height: 844px) and (-webkit-device-pixel-ratio: 3) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_14__iPhone_13_Pro__iPhone_13__iPhone_12_Pro__iPhone_12_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 375px) and (device-height: 812px) and (-webkit-device-pixel-ratio: 3) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_13_mini__iPhone_12_mini__iPhone_11_Pro__iPhone_XS__iPhone_X_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 834px) and (device-height: 1112px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/10.5__iPad_Air_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 390px) and (device-height: 844px) and (-webkit-device-pixel-ratio: 3) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_14__iPhone_13_Pro__iPhone_13__iPhone_12_Pro__iPhone_12_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 768px) and (device-height: 1024px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/9.7__iPad_Pro__7.9__iPad_mini__9.7__iPad_Air__9.7__iPad_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 430px) and (device-height: 932px) and (-webkit-device-pixel-ratio: 3) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_14_Pro_Max_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 428px) and (device-height: 926px) and (-webkit-device-pixel-ratio: 3) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_14_Plus__iPhone_13_Pro_Max__iPhone_12_Pro_Max_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 414px) and (device-height: 896px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_11__iPhone_XR_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 375px) and (device-height: 667px) and (-webkit-device-pixel-ratio: 2) and (orientation: portrait)" href="/_web/static/img/splash/iPhone_8__iPhone_7__iPhone_6s__iPhone_6__4.7__iPhone_SE_portrait.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 834px) and (device-height: 1112px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/10.5__iPad_Air_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 834px) and (device-height: 1194px) and (-webkit-device-pixel-ratio: 2) and (orientation: landscape)" href="/_web/static/img/splash/11__iPad_Pro__10.5__iPad_Pro_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 414px) and (device-height: 736px) and (-webkit-device-pixel-ratio: 3) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_8_Plus__iPhone_7_Plus__iPhone_6s_Plus__iPhone_6_Plus_landscape.png"> <link rel="apple-touch-startup-image" media="screen and (device-width: 430px) and (device-height: 932px) and (-webkit-device-pixel-ratio: 3) and (orientation: landscape)" href="/_web/static/img/splash/iPhone_14_Pro_Max_landscape.png"> <script>!function(a,b,c){if(c in b&&b[c]){var d,e=a.location,f=/^(a|html)$/i;a.addEventListener("click",function(a){for(d=a.target;!f.test(d.nodeName);)d=d.parentNode;"href"in d&&(d.href.indexOf("http")||~d.href.indexOf(e.host))&&(a.preventDefault(),e.href=d.href)},!1),a.querySelector("html").classList.add("standalone")}}(document,window.navigator,"standalone")</script> <script> var _paq = window._paq = window._paq || []; /* tracker methods like "setCustomDimension" should be called before "trackPageView" */ _paq.push(['trackPageView']); _paq.push(['enableLinkTracking']); (function() { var u="//statistiques.universalis.fr/"; _paq.push(['setTrackerUrl', u+'matomo.php']); _paq.push(['setSiteId', '1']); var d=document, g=d.createElement('script'), s=d.getElementsByTagName('script')[0]; g.async=true; g.src=u+'matomo.js'; s.parentNode.insertBefore(g,s); })(); </script> <style>.swiper-container[data-loaded=false] .swiper-wrapper .swiper-slide{display:none}.swiper-container[data-loaded=false] .swiper-wrapper .swiper-slide:first-child{display:block}img{max-width:100%;height:auto;max-height:250px}img.d-formule{max-height:none}strong{font-weight:bold !important}.dark-mode body{background:#29363a !important}.loader{display:none;height:100vh}.loader:after{display:block;margin:auto;content:"";background:url(/_web/assets/img/banniere-fr.png) no-repeat center;background-size:contain;width:50%;height:50%;position:absolute;top:50%;left:50%;transform:translate(-50%, -50%);opacity:.25}#page_3 .pub-container,#page_16 .pub-container{display:none !important}#skip{padding:5px 15px;position:absolute;background:rgba(0,0,0,.85);color:#fff;left:10%;transform:translateY(-100%);transition:transform .3s;border-bottom-left-radius:6px;border-bottom-right-radius:6px;z-index:100;text-decoration:none}#skip:focus{transform:translateY(0%);box-shadow:0 0 10px #333}.login-form-wrapper-mobile{display:none}.site-fr .article-quote{display:none !important}@media(max-width: 768px){.login-form-wrapper-mobile{display:block;text-align:center;margin-top:-1em;margin-bottom:1em}}body.loading{overflow:hidden}body.loading>*{display:none}body.loading .loader{display:block}.dropdown-menu,.d-none,#podcast audio{display:none}.list-inline-item{display:inline-block}.offcanvas{visibility:hidden}.main-navbar-wrapper .btn-universalis{background-color:#e8eced;color:#000;font-weight:normal}.offcanvas{display:none !important;height:0 !important}.offcanvas[data-loaded=true]{display:inherit !important;height:inherit !important}.clearfix{clear:both}.fr .is-edu{display:none !important}#podcast{min-height:90px}#article-medias-strip-junior{min-height:255px}#callSommaire{position:fixed;bottom:10px;left:10px;z-index:2}#qc-cmp2-ui{padding:10px !important}.qc-cmp2-summary-buttons{justify-content:space-around !important}.qc-cmp2-summary-buttons button[mode=secondary],.qc-cmp2-footer button[mode=secondary]{border:none;box-shadow:none}.eCBsvP.eCBsvP{height:auto !important}.qc-cmp-cleanslate{background-color:rgba(33,41,52,.95) !important}.fill-universalis{fill:#1c3479}.fill-universalis-up{fill:#526d78}@media(min-width: 768px){.qc-cmp2-publisher-logo-container{text-align:center}.qc-cmp2-publisher-logo-container figure img{display:block !important}}#page_14 .filter-article-container{display:none}#newsletter_subscription .text-light{color:#fff !important}#bandeau-footer{display:none}</style> <link rel="stylesheet" id="typo-css" href="/_web/assets/css/typo.css?t=20240418-1" /><script>let cssLoaded=!1,bodyLoaded=!1;const loadedTO=setTimeout((()=>{cssLoaded=!0,bodyLoaded=!0,loaded()}),5e3),loaded=function(){cssLoaded&&bodyLoaded&&document.body.classList.contains("loading")&&(document.body.className=document.body.className.replace("loading","").trim(),clearTimeout(loadedTO))};</script> <link rel="preload" href="/_web/assets/css/lazy.css?t=20240418-1" as="style" media="all" crossorigin onload="this.rel=`stylesheet`; cssLoaded = true; loaded()" /><style>.breadcrumb{--bs-breadcrumb-padding-x: 0;--bs-breadcrumb-padding-y: 0;--bs-breadcrumb-margin-bottom: 1rem;--bs-breadcrumb-bg: ;--bs-breadcrumb-border-radius: ;--bs-breadcrumb-divider-color: #6c757d;--bs-breadcrumb-item-padding-x: 0.5rem;--bs-breadcrumb-item-active-color: #6c757d;display:flex;flex-wrap:wrap;padding:var(--bs-breadcrumb-padding-y) var(--bs-breadcrumb-padding-x);margin-bottom:var(--bs-breadcrumb-margin-bottom);font-size:var(--bs-breadcrumb-font-size);list-style:none;background-color:var(--bs-breadcrumb-bg);border-radius:var(--bs-breadcrumb-border-radius)}.breadcrumb-item+.breadcrumb-item{padding-left:var(--bs-breadcrumb-item-padding-x)}.breadcrumb-item+.breadcrumb-item::before{float:left;padding-right:var(--bs-breadcrumb-item-padding-x);color:var(--bs-breadcrumb-divider-color);content:var(--bs-breadcrumb-divider, "/") /* rtl: var(--bs-breadcrumb-divider, "/") */}.breadcrumb-item.active{color:var(--bs-breadcrumb-item-active-color)}.breadcrumb{padding:.5rem 1rem !important;background-color:#f5f5f5 !important;border-radius:.375rem !important}.breadcrumb .breadcrumb-item i.label-sub{font-size:.85em;font-weight:normal}.breadcrumb .breadcrumb-item a{text-decoration:none}.breadcrumb .breadcrumb-item a:hover{text-decoration:underline}.breadcrumb .breadcrumb-item.active{font-weight:bold}.breadcrumb .breadcrumb-item+.breadcrumb-item::before{background-image:url('data:image/svg+xml;charset=UTF-8, <svg aria-hidden="true" focusable="false" data-prefix="far" data-icon="chevron-double-right" role="img" xmlns="http://www.w3.org/2000/svg" viewBox="0 0 448 512" class="svg-inline--fa fa-chevron-double-right fa-w-14 fa-7x"><path style="fill: rgb(97, 104, 118);" d="M57.7 38.1l209.4 209.4c4.7 4.7 4.7 12.3 0 17L57.7 473.9c-4.7 4.7-12.3 4.7-17 0l-19.8-19.8c-4.7-4.7-4.7-12.3 0-17L201.6 256 20.9 74.9c-4.7-4.7-4.7-12.3 0-17l19.8-19.8c4.7-4.7 12.3-4.7 17 0zm143 0l-19.8 19.8c-4.7 4.7-4.7 12.3 0 17L361.6 256 180.9 437.1c-4.7 4.7-4.7 12.3 0 17l19.8 19.8c4.7 4.7 12.3 4.7 17 0l209.4-209.4c4.7-4.7 4.7-12.3 0-17L217.7 38.1c-4.7-4.7-12.3-4.7-17 0z" class=""></path></svg>');background-repeat:no-repeat;background-position:center;content:"";width:.5rem;height:100%;margin-right:.5rem}.list-group{--bs-list-group-color: #212529;--bs-list-group-bg: #fff;--bs-list-group-border-color: rgba(0, 0, 0, 0.125);--bs-list-group-border-width: 1px;--bs-list-group-border-radius: 0.375rem;--bs-list-group-item-padding-x: 1rem;--bs-list-group-item-padding-y: 0.5rem;--bs-list-group-action-color: #495057;--bs-list-group-action-hover-color: #495057;--bs-list-group-action-hover-bg: #f8f9fa;--bs-list-group-action-active-color: #212529;--bs-list-group-action-active-bg: #e9ecef;--bs-list-group-disabled-color: #6c757d;--bs-list-group-disabled-bg: #fff;--bs-list-group-active-color: #fff;--bs-list-group-active-bg: #526d78;--bs-list-group-active-border-color: #526d78;display:flex;flex-direction:column;padding-left:0;margin-bottom:0;border-radius:var(--bs-list-group-border-radius)}.list-group-numbered{list-style-type:none;counter-reset:section}.list-group-numbered>.list-group-item::before{content:counters(section, ".") ". ";counter-increment:section}.list-group-item-action{width:100%;color:var(--bs-list-group-action-color);text-align:inherit}.list-group-item-action:hover,.list-group-item-action:focus{z-index:1;color:var(--bs-list-group-action-hover-color);text-decoration:none;background-color:var(--bs-list-group-action-hover-bg)}.list-group-item-action:active{color:var(--bs-list-group-action-active-color);background-color:var(--bs-list-group-action-active-bg)}.list-group-item{position:relative;display:block;padding:var(--bs-list-group-item-padding-y) var(--bs-list-group-item-padding-x);color:var(--bs-list-group-color);text-decoration:none;background-color:var(--bs-list-group-bg);border:var(--bs-list-group-border-width) solid var(--bs-list-group-border-color)}.list-group-item:first-child{border-top-left-radius:inherit;border-top-right-radius:inherit}.list-group-item:last-child{border-bottom-right-radius:inherit;border-bottom-left-radius:inherit}.list-group-item.disabled,.list-group-item:disabled{color:var(--bs-list-group-disabled-color);pointer-events:none;background-color:var(--bs-list-group-disabled-bg)}.list-group-item.active{z-index:2;color:var(--bs-list-group-active-color);background-color:var(--bs-list-group-active-bg);border-color:var(--bs-list-group-active-border-color)}.list-group-item+.list-group-item{border-top-width:0}.list-group-item+.list-group-item.active{margin-top:calc(-1 * var(--bs-list-group-border-width));border-top-width:var(--bs-list-group-border-width)}.list-group-horizontal{flex-direction:row}.list-group-horizontal>.list-group-item:first-child:not(:last-child){border-bottom-left-radius:var(--bs-list-group-border-radius);border-top-right-radius:0}.list-group-horizontal>.list-group-item:last-child:not(:first-child){border-top-right-radius:var(--bs-list-group-border-radius);border-bottom-left-radius:0}.list-group-horizontal>.list-group-item.active{margin-top:0}.list-group-horizontal>.list-group-item+.list-group-item{border-top-width:var(--bs-list-group-border-width);border-left-width:0}.list-group-horizontal>.list-group-item+.list-group-item.active{margin-left:calc(-1 * var(--bs-list-group-border-width));border-left-width:var(--bs-list-group-border-width)}@media(min-width: 576px){.list-group-horizontal-sm{flex-direction:row}.list-group-horizontal-sm>.list-group-item:first-child:not(:last-child){border-bottom-left-radius:var(--bs-list-group-border-radius);border-top-right-radius:0}.list-group-horizontal-sm>.list-group-item:last-child:not(:first-child){border-top-right-radius:var(--bs-list-group-border-radius);border-bottom-left-radius:0}.list-group-horizontal-sm>.list-group-item.active{margin-top:0}.list-group-horizontal-sm>.list-group-item+.list-group-item{border-top-width:var(--bs-list-group-border-width);border-left-width:0}.list-group-horizontal-sm>.list-group-item+.list-group-item.active{margin-left:calc(-1 * var(--bs-list-group-border-width));border-left-width:var(--bs-list-group-border-width)}}@media(min-width: 768px){.list-group-horizontal-md{flex-direction:row}.list-group-horizontal-md>.list-group-item:first-child:not(:last-child){border-bottom-left-radius:var(--bs-list-group-border-radius);border-top-right-radius:0}.list-group-horizontal-md>.list-group-item:last-child:not(:first-child){border-top-right-radius:var(--bs-list-group-border-radius);border-bottom-left-radius:0}.list-group-horizontal-md>.list-group-item.active{margin-top:0}.list-group-horizontal-md>.list-group-item+.list-group-item{border-top-width:var(--bs-list-group-border-width);border-left-width:0}.list-group-horizontal-md>.list-group-item+.list-group-item.active{margin-left:calc(-1 * var(--bs-list-group-border-width));border-left-width:var(--bs-list-group-border-width)}}@media(min-width: 992px){.list-group-horizontal-lg{flex-direction:row}.list-group-horizontal-lg>.list-group-item:first-child:not(:last-child){border-bottom-left-radius:var(--bs-list-group-border-radius);border-top-right-radius:0}.list-group-horizontal-lg>.list-group-item:last-child:not(:first-child){border-top-right-radius:var(--bs-list-group-border-radius);border-bottom-left-radius:0}.list-group-horizontal-lg>.list-group-item.active{margin-top:0}.list-group-horizontal-lg>.list-group-item+.list-group-item{border-top-width:var(--bs-list-group-border-width);border-left-width:0}.list-group-horizontal-lg>.list-group-item+.list-group-item.active{margin-left:calc(-1 * var(--bs-list-group-border-width));border-left-width:var(--bs-list-group-border-width)}}@media(min-width: 1200px){.list-group-horizontal-xl{flex-direction:row}.list-group-horizontal-xl>.list-group-item:first-child:not(:last-child){border-bottom-left-radius:var(--bs-list-group-border-radius);border-top-right-radius:0}.list-group-horizontal-xl>.list-group-item:last-child:not(:first-child){border-top-right-radius:var(--bs-list-group-border-radius);border-bottom-left-radius:0}.list-group-horizontal-xl>.list-group-item.active{margin-top:0}.list-group-horizontal-xl>.list-group-item+.list-group-item{border-top-width:var(--bs-list-group-border-width);border-left-width:0}.list-group-horizontal-xl>.list-group-item+.list-group-item.active{margin-left:calc(-1 * var(--bs-list-group-border-width));border-left-width:var(--bs-list-group-border-width)}}@media(min-width: 1400px){.list-group-horizontal-xxl{flex-direction:row}.list-group-horizontal-xxl>.list-group-item:first-child:not(:last-child){border-bottom-left-radius:var(--bs-list-group-border-radius);border-top-right-radius:0}.list-group-horizontal-xxl>.list-group-item:last-child:not(:first-child){border-top-right-radius:var(--bs-list-group-border-radius);border-bottom-left-radius:0}.list-group-horizontal-xxl>.list-group-item.active{margin-top:0}.list-group-horizontal-xxl>.list-group-item+.list-group-item{border-top-width:var(--bs-list-group-border-width);border-left-width:0}.list-group-horizontal-xxl>.list-group-item+.list-group-item.active{margin-left:calc(-1 * var(--bs-list-group-border-width));border-left-width:var(--bs-list-group-border-width)}}.list-group-flush{border-radius:0}.list-group-flush>.list-group-item{border-width:0 0 var(--bs-list-group-border-width)}.list-group-flush>.list-group-item:last-child{border-bottom-width:0}.list-group-item-primary{color:#314148;background-color:#dce2e4}.list-group-item-primary.list-group-item-action:hover,.list-group-item-primary.list-group-item-action:focus{color:#314148;background-color:#c6cbcd}.list-group-item-primary.list-group-item-action.active{color:#fff;background-color:#314148;border-color:#314148}.list-group-item-secondary{color:#192023;background-color:#d4d7d8}.list-group-item-secondary.list-group-item-action:hover,.list-group-item-secondary.list-group-item-action:focus{color:#192023;background-color:#bfc2c2}.list-group-item-secondary.list-group-item-action.active{color:#fff;background-color:#192023;border-color:#192023}.list-group-item-success{color:#0f5132;background-color:#d1e7dd}.list-group-item-success.list-group-item-action:hover,.list-group-item-success.list-group-item-action:focus{color:#0f5132;background-color:#bcd0c7}.list-group-item-success.list-group-item-action.active{color:#fff;background-color:#0f5132;border-color:#0f5132}.list-group-item-info{color:#055160;background-color:#cff4fc}.list-group-item-info.list-group-item-action:hover,.list-group-item-info.list-group-item-action:focus{color:#055160;background-color:#badce3}.list-group-item-info.list-group-item-action.active{color:#fff;background-color:#055160;border-color:#055160}.list-group-item-warning{color:#664d03;background-color:#fff3cd}.list-group-item-warning.list-group-item-action:hover,.list-group-item-warning.list-group-item-action:focus{color:#664d03;background-color:#e6dbb9}.list-group-item-warning.list-group-item-action.active{color:#fff;background-color:#664d03;border-color:#664d03}.list-group-item-danger{color:#851a31;background-color:#f8d5dc}.list-group-item-danger.list-group-item-action:hover,.list-group-item-danger.list-group-item-action:focus{color:#851a31;background-color:#dfc0c6}.list-group-item-danger.list-group-item-action.active{color:#fff;background-color:#851a31;border-color:#851a31}.list-group-item-light{color:#5d5e5f;background-color:#fafbfb}.list-group-item-light.list-group-item-action:hover,.list-group-item-light.list-group-item-action:focus{color:#5d5e5f;background-color:#e1e2e2}.list-group-item-light.list-group-item-action.active{color:#fff;background-color:#5d5e5f;border-color:#5d5e5f}.list-group-item-dark{color:#192023;background-color:#d4d7d8}.list-group-item-dark.list-group-item-action:hover,.list-group-item-dark.list-group-item-action:focus{color:#192023;background-color:#bfc2c2}.list-group-item-dark.list-group-item-action.active{color:#fff;background-color:#192023;border-color:#192023}.badge{--bs-badge-padding-x: 0.65em;--bs-badge-padding-y: 0.35em;--bs-badge-font-size:0.75em;--bs-badge-font-weight: 700;--bs-badge-color: #fff;--bs-badge-border-radius: 0.375rem;display:inline-block;padding:var(--bs-badge-padding-y) var(--bs-badge-padding-x);font-size:var(--bs-badge-font-size);font-weight:var(--bs-badge-font-weight);line-height:1;color:var(--bs-badge-color);text-align:center;white-space:nowrap;vertical-align:baseline;border-radius:var(--bs-badge-border-radius)}.badge:empty{display:none}.btn .badge{position:relative;top:-1px}.btn-group,.btn-group-vertical{position:relative;display:inline-flex;vertical-align:middle}.btn-group>.btn,.btn-group-vertical>.btn{position:relative;flex:1 1 auto}.btn-group>.btn-check:checked+.btn,.btn-group>.btn-check:focus+.btn,.btn-group>.btn:hover,.btn-group>.btn:focus,.btn-group>.btn:active,.btn-group>.btn.active,.btn-group-vertical>.btn-check:checked+.btn,.btn-group-vertical>.btn-check:focus+.btn,.btn-group-vertical>.btn:hover,.btn-group-vertical>.btn:focus,.btn-group-vertical>.btn:active,.btn-group-vertical>.btn.active{z-index:1}.btn-toolbar{display:flex;flex-wrap:wrap;justify-content:flex-start}.btn-toolbar .input-group{width:auto}.btn-group{border-radius:.375rem}.btn-group>:not(.btn-check:first-child)+.btn,.btn-group>.btn-group:not(:first-child){margin-left:-1px}.btn-group>.btn:not(:last-child):not(.dropdown-toggle),.btn-group>.btn.dropdown-toggle-split:first-child,.btn-group>.btn-group:not(:last-child)>.btn{border-top-right-radius:0;border-bottom-right-radius:0}.btn-group>.btn:nth-child(n+3),.btn-group>:not(.btn-check)+.btn,.btn-group>.btn-group:not(:first-child)>.btn{border-top-left-radius:0;border-bottom-left-radius:0}.dropdown-toggle-split{padding-right:.5625rem;padding-left:.5625rem}.dropdown-toggle-split::after,.dropup .dropdown-toggle-split::after,.dropend .dropdown-toggle-split::after{margin-left:0}.dropstart .dropdown-toggle-split::before{margin-right:0}.btn-sm+.dropdown-toggle-split,.btn-group-sm>.btn+.dropdown-toggle-split{padding-right:.375rem;padding-left:.375rem}.btn-lg+.dropdown-toggle-split,.btn-group-lg>.btn+.dropdown-toggle-split{padding-right:.75rem;padding-left:.75rem}.btn-group-vertical{flex-direction:column;align-items:flex-start;justify-content:center}.btn-group-vertical>.btn,.btn-group-vertical>.btn-group{width:100%}.btn-group-vertical>.btn:not(:first-child),.btn-group-vertical>.btn-group:not(:first-child){margin-top:-1px}.btn-group-vertical>.btn:not(:last-child):not(.dropdown-toggle),.btn-group-vertical>.btn-group:not(:last-child)>.btn{border-bottom-right-radius:0;border-bottom-left-radius:0}.btn-group-vertical>.btn~.btn,.btn-group-vertical>.btn-group:not(:first-child)>.btn{border-top-left-radius:0;border-top-right-radius:0}.pagination{--bs-pagination-padding-x: 0.75rem;--bs-pagination-padding-y: 0.375rem;--bs-pagination-font-size:0.95rem;--bs-pagination-color: var(--bs-link-color);--bs-pagination-bg: #fff;--bs-pagination-border-width: 1px;--bs-pagination-border-color: #dee2e6;--bs-pagination-border-radius: 0.375rem;--bs-pagination-hover-color: var(--bs-link-hover-color);--bs-pagination-hover-bg: #e9ecef;--bs-pagination-hover-border-color: #dee2e6;--bs-pagination-focus-color: var(--bs-link-hover-color);--bs-pagination-focus-bg: #e9ecef;--bs-pagination-focus-box-shadow: 0 0 0 0.25rem rgba(82, 109, 120, 0.25);--bs-pagination-active-color: #fff;--bs-pagination-active-bg: #526d78;--bs-pagination-active-border-color: #526d78;--bs-pagination-disabled-color: #6c757d;--bs-pagination-disabled-bg: #fff;--bs-pagination-disabled-border-color: #dee2e6;display:flex;padding-left:0;list-style:none}.page-link{position:relative;display:block;padding:var(--bs-pagination-padding-y) var(--bs-pagination-padding-x);font-size:var(--bs-pagination-font-size);color:var(--bs-pagination-color);text-decoration:none;background-color:var(--bs-pagination-bg);border:var(--bs-pagination-border-width) solid var(--bs-pagination-border-color);transition:color .15s ease-in-out,background-color .15s ease-in-out,border-color .15s ease-in-out,box-shadow .15s ease-in-out}@media(prefers-reduced-motion: reduce){.page-link{transition:none}}.page-link:hover{z-index:2;color:var(--bs-pagination-hover-color);background-color:var(--bs-pagination-hover-bg);border-color:var(--bs-pagination-hover-border-color)}.page-link:focus{z-index:3;color:var(--bs-pagination-focus-color);background-color:var(--bs-pagination-focus-bg);outline:0;box-shadow:var(--bs-pagination-focus-box-shadow)}.page-link.active,.active>.page-link{z-index:3;color:var(--bs-pagination-active-color);background-color:var(--bs-pagination-active-bg);border-color:var(--bs-pagination-active-border-color)}.page-link.disabled,.disabled>.page-link{color:var(--bs-pagination-disabled-color);pointer-events:none;background-color:var(--bs-pagination-disabled-bg);border-color:var(--bs-pagination-disabled-border-color)}.page-item:not(:first-child) .page-link{margin-left:-1px}.page-item:first-child .page-link{border-top-left-radius:var(--bs-pagination-border-radius);border-bottom-left-radius:var(--bs-pagination-border-radius)}.page-item:last-child .page-link{border-top-right-radius:var(--bs-pagination-border-radius);border-bottom-right-radius:var(--bs-pagination-border-radius)}.pagination-lg{--bs-pagination-padding-x: 1.5rem;--bs-pagination-padding-y: 0.75rem;--bs-pagination-font-size:1.1875rem;--bs-pagination-border-radius: 0.5rem}.pagination-sm{--bs-pagination-padding-x: 0.5rem;--bs-pagination-padding-y: 0.25rem;--bs-pagination-font-size:0.83125rem;--bs-pagination-border-radius: 0.25rem}.list-group a.list-group-item{border-color:#f8fafc}.list-group a.list-group-item.active{background-color:#526d78}.list-group a.list-group-item h3{margin:0;line-height:1.5;font-size:.95rem;font-family:system-ui,-apple-system,"Segoe UI",Roboto,"Helvetica Neue","Noto Sans","Liberation Sans",Arial,sans-serif,"Apple Color Emoji","Segoe UI Emoji","Segoe UI Symbol","Noto Color Emoji";padding-bottom:0}.list-group a.list-group-item h3:hover{color:#fff}.list-group a.list-group-item h3 .badge{line-height:1.25}.list-group a.list-group-item:hover{background-color:#526d78;color:#fff}.list-group a.list-group-item:hover h3{color:inherit}i.anchor{display:block;position:relative;visibility:hidden;top:calc(-25vh + 15px)}.article__props{font-size:.8rem;color:#000;font-style:italic;padding:.2rem 0;margin:0 auto;display:block}.content h6,.h6,h5,.h5,h4,.h4,h3,.h3,h2,.h2,h1,.h1{margin-bottom:0;padding-bottom:.5rem}titre_main+titre_subs{margin-left:.25em}pc,.pc{text-transform:uppercase;font-size:.8em}h2 .pc,h3 .pc,h4 .pc,h5 .pc,h6 .pc{font-size:inherit !important}.sup-sub{position:relative;margin-right:1rem}.sup-sub sup,.sup-sub sub,.sup-sub inf{position:absolute;display:block;left:0}.sup-sub sup{top:.25rem}.sup-sub sub,.sup-sub inf{bottom:.25rem}p:empty{display:none !important}.content.whole .read-more{display:none}blockquote{position:relative;text-align:center;padding:1rem 1.4rem;width:80%;margin:1rem auto 2rem;font-family:Times;font-style:italic;box-shadow:0 .5rem 1rem rgba(0,0,0,.15);border-radius:.375rem}blockquote:before{position:absolute;color:#526d78;font-size:3rem;content:"«";top:-22px;margin-right:-28px;right:100%}blockquote:after{position:absolute;color:#526d78;font-size:3rem;content:"»";margin-left:-28px;left:100%;top:auto;bottom:-12px}.offcanvas,.offcanvas-xxl,.offcanvas-xl,.offcanvas-lg,.offcanvas-md,.offcanvas-sm{--bs-offcanvas-zindex: 1045;--bs-offcanvas-width: 400px;--bs-offcanvas-height: 30vh;--bs-offcanvas-padding-x: 1rem;--bs-offcanvas-padding-y: 1rem;--bs-offcanvas-color: ;--bs-offcanvas-bg: #fff;--bs-offcanvas-border-width: 1px;--bs-offcanvas-border-color: var(--bs-border-color-translucent);--bs-offcanvas-box-shadow: 0 0.125rem 0.25rem rgba(0, 0, 0, 0.075)}@media(max-width: 575.98px){.offcanvas-sm{position:fixed;bottom:0;z-index:var(--bs-offcanvas-zindex);display:flex;flex-direction:column;max-width:100%;color:var(--bs-offcanvas-color);visibility:hidden;background-color:var(--bs-offcanvas-bg);background-clip:padding-box;outline:0;transition:transform .3s ease-in-out}}@media(max-width: 575.98px)and (prefers-reduced-motion: reduce){.offcanvas-sm{transition:none}}@media(max-width: 575.98px){.offcanvas-sm.offcanvas-start{top:0;left:0;width:var(--bs-offcanvas-width);border-right:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(-100%)}}@media(max-width: 575.98px){.offcanvas-sm.offcanvas-end{top:0;right:0;width:var(--bs-offcanvas-width);border-left:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(100%)}}@media(max-width: 575.98px){.offcanvas-sm.offcanvas-top{top:0;right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-bottom:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(-100%)}}@media(max-width: 575.98px){.offcanvas-sm.offcanvas-bottom{right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-top:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(100%)}}@media(max-width: 575.98px){.offcanvas-sm.showing,.offcanvas-sm.show:not(.hiding){transform:none}}@media(max-width: 575.98px){.offcanvas-sm.showing,.offcanvas-sm.hiding,.offcanvas-sm.show{visibility:visible}}@media(min-width: 576px){.offcanvas-sm{--bs-offcanvas-height: auto;--bs-offcanvas-border-width: 0;background-color:transparent !important}.offcanvas-sm .offcanvas-header{display:none}.offcanvas-sm .offcanvas-body{display:flex;flex-grow:0;padding:0;overflow-y:visible;background-color:transparent !important}}@media(max-width: 767.98px){.offcanvas-md{position:fixed;bottom:0;z-index:var(--bs-offcanvas-zindex);display:flex;flex-direction:column;max-width:100%;color:var(--bs-offcanvas-color);visibility:hidden;background-color:var(--bs-offcanvas-bg);background-clip:padding-box;outline:0;transition:transform .3s ease-in-out}}@media(max-width: 767.98px)and (prefers-reduced-motion: reduce){.offcanvas-md{transition:none}}@media(max-width: 767.98px){.offcanvas-md.offcanvas-start{top:0;left:0;width:var(--bs-offcanvas-width);border-right:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(-100%)}}@media(max-width: 767.98px){.offcanvas-md.offcanvas-end{top:0;right:0;width:var(--bs-offcanvas-width);border-left:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(100%)}}@media(max-width: 767.98px){.offcanvas-md.offcanvas-top{top:0;right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-bottom:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(-100%)}}@media(max-width: 767.98px){.offcanvas-md.offcanvas-bottom{right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-top:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(100%)}}@media(max-width: 767.98px){.offcanvas-md.showing,.offcanvas-md.show:not(.hiding){transform:none}}@media(max-width: 767.98px){.offcanvas-md.showing,.offcanvas-md.hiding,.offcanvas-md.show{visibility:visible}}@media(min-width: 768px){.offcanvas-md{--bs-offcanvas-height: auto;--bs-offcanvas-border-width: 0;background-color:transparent !important}.offcanvas-md .offcanvas-header{display:none}.offcanvas-md .offcanvas-body{display:flex;flex-grow:0;padding:0;overflow-y:visible;background-color:transparent !important}}@media(max-width: 991.98px){.offcanvas-lg{position:fixed;bottom:0;z-index:var(--bs-offcanvas-zindex);display:flex;flex-direction:column;max-width:100%;color:var(--bs-offcanvas-color);visibility:hidden;background-color:var(--bs-offcanvas-bg);background-clip:padding-box;outline:0;transition:transform .3s ease-in-out}}@media(max-width: 991.98px)and (prefers-reduced-motion: reduce){.offcanvas-lg{transition:none}}@media(max-width: 991.98px){.offcanvas-lg.offcanvas-start{top:0;left:0;width:var(--bs-offcanvas-width);border-right:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(-100%)}}@media(max-width: 991.98px){.offcanvas-lg.offcanvas-end{top:0;right:0;width:var(--bs-offcanvas-width);border-left:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(100%)}}@media(max-width: 991.98px){.offcanvas-lg.offcanvas-top{top:0;right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-bottom:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(-100%)}}@media(max-width: 991.98px){.offcanvas-lg.offcanvas-bottom{right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-top:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(100%)}}@media(max-width: 991.98px){.offcanvas-lg.showing,.offcanvas-lg.show:not(.hiding){transform:none}}@media(max-width: 991.98px){.offcanvas-lg.showing,.offcanvas-lg.hiding,.offcanvas-lg.show{visibility:visible}}@media(min-width: 992px){.offcanvas-lg{--bs-offcanvas-height: auto;--bs-offcanvas-border-width: 0;background-color:transparent !important}.offcanvas-lg .offcanvas-header{display:none}.offcanvas-lg .offcanvas-body{display:flex;flex-grow:0;padding:0;overflow-y:visible;background-color:transparent !important}}@media(max-width: 1199.98px){.offcanvas-xl{position:fixed;bottom:0;z-index:var(--bs-offcanvas-zindex);display:flex;flex-direction:column;max-width:100%;color:var(--bs-offcanvas-color);visibility:hidden;background-color:var(--bs-offcanvas-bg);background-clip:padding-box;outline:0;transition:transform .3s ease-in-out}}@media(max-width: 1199.98px)and (prefers-reduced-motion: reduce){.offcanvas-xl{transition:none}}@media(max-width: 1199.98px){.offcanvas-xl.offcanvas-start{top:0;left:0;width:var(--bs-offcanvas-width);border-right:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(-100%)}}@media(max-width: 1199.98px){.offcanvas-xl.offcanvas-end{top:0;right:0;width:var(--bs-offcanvas-width);border-left:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(100%)}}@media(max-width: 1199.98px){.offcanvas-xl.offcanvas-top{top:0;right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-bottom:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(-100%)}}@media(max-width: 1199.98px){.offcanvas-xl.offcanvas-bottom{right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-top:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(100%)}}@media(max-width: 1199.98px){.offcanvas-xl.showing,.offcanvas-xl.show:not(.hiding){transform:none}}@media(max-width: 1199.98px){.offcanvas-xl.showing,.offcanvas-xl.hiding,.offcanvas-xl.show{visibility:visible}}@media(min-width: 1200px){.offcanvas-xl{--bs-offcanvas-height: auto;--bs-offcanvas-border-width: 0;background-color:transparent !important}.offcanvas-xl .offcanvas-header{display:none}.offcanvas-xl .offcanvas-body{display:flex;flex-grow:0;padding:0;overflow-y:visible;background-color:transparent !important}}@media(max-width: 1399.98px){.offcanvas-xxl{position:fixed;bottom:0;z-index:var(--bs-offcanvas-zindex);display:flex;flex-direction:column;max-width:100%;color:var(--bs-offcanvas-color);visibility:hidden;background-color:var(--bs-offcanvas-bg);background-clip:padding-box;outline:0;transition:transform .3s ease-in-out}}@media(max-width: 1399.98px)and (prefers-reduced-motion: reduce){.offcanvas-xxl{transition:none}}@media(max-width: 1399.98px){.offcanvas-xxl.offcanvas-start{top:0;left:0;width:var(--bs-offcanvas-width);border-right:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(-100%)}}@media(max-width: 1399.98px){.offcanvas-xxl.offcanvas-end{top:0;right:0;width:var(--bs-offcanvas-width);border-left:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(100%)}}@media(max-width: 1399.98px){.offcanvas-xxl.offcanvas-top{top:0;right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-bottom:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(-100%)}}@media(max-width: 1399.98px){.offcanvas-xxl.offcanvas-bottom{right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-top:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(100%)}}@media(max-width: 1399.98px){.offcanvas-xxl.showing,.offcanvas-xxl.show:not(.hiding){transform:none}}@media(max-width: 1399.98px){.offcanvas-xxl.showing,.offcanvas-xxl.hiding,.offcanvas-xxl.show{visibility:visible}}@media(min-width: 1400px){.offcanvas-xxl{--bs-offcanvas-height: auto;--bs-offcanvas-border-width: 0;background-color:transparent !important}.offcanvas-xxl .offcanvas-header{display:none}.offcanvas-xxl .offcanvas-body{display:flex;flex-grow:0;padding:0;overflow-y:visible;background-color:transparent !important}}.offcanvas{position:fixed;bottom:0;z-index:var(--bs-offcanvas-zindex);display:flex;flex-direction:column;max-width:100%;color:var(--bs-offcanvas-color);visibility:hidden;background-color:var(--bs-offcanvas-bg);background-clip:padding-box;outline:0;transition:transform .3s ease-in-out}@media(prefers-reduced-motion: reduce){.offcanvas{transition:none}}.offcanvas.offcanvas-start{top:0;left:0;width:var(--bs-offcanvas-width);border-right:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(-100%)}.offcanvas.offcanvas-end{top:0;right:0;width:var(--bs-offcanvas-width);border-left:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateX(100%)}.offcanvas.offcanvas-top{top:0;right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-bottom:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(-100%)}.offcanvas.offcanvas-bottom{right:0;left:0;height:var(--bs-offcanvas-height);max-height:100%;border-top:var(--bs-offcanvas-border-width) solid var(--bs-offcanvas-border-color);transform:translateY(100%)}.offcanvas.showing,.offcanvas.show:not(.hiding){transform:none}.offcanvas.showing,.offcanvas.hiding,.offcanvas.show{visibility:visible}.offcanvas-backdrop{position:fixed;top:0;left:0;z-index:1040;width:100vw;height:100vh;background-color:#000}.offcanvas-backdrop.fade{opacity:0}.offcanvas-backdrop.show{opacity:.5}.offcanvas-header{display:flex;align-items:center;justify-content:space-between;padding:var(--bs-offcanvas-padding-y) var(--bs-offcanvas-padding-x)}.offcanvas-header .btn-close{padding:calc(var(--bs-offcanvas-padding-y) * .5) calc(var(--bs-offcanvas-padding-x) * .5);margin-top:calc(-.5 * var(--bs-offcanvas-padding-y));margin-right:calc(-.5 * var(--bs-offcanvas-padding-x));margin-bottom:calc(-.5 * var(--bs-offcanvas-padding-y))}.offcanvas-title{margin-bottom:0;line-height:1.5}.offcanvas-body{flex-grow:1;padding:var(--bs-offcanvas-padding-y) var(--bs-offcanvas-padding-x);overflow-y:auto}.article__single__toc .float-end{float:right !important}.article__single__toc .list-group-item{border:none;border-radius:0}.article__single__toc .list-group-item i{transition:transform .2s;position:absolute;left:.25rem;margin-top:.25rem}.article__single__toc .list-group-item.expanded i{transform:rotateZ(90deg)}.article__single__toc>.list-group{background:transparent}.article__single__toc>.list-group>.list-group{margin-left:.5em}.article__single__toc>.list-group>.list-group>.list-group{margin-left:.75em;border-left:1px solid #ccc}.article__single__toc>.list-group>.list-group>.list-group>.list-group-item{padding:.5em}#callSommaire{position:fixed;bottom:10px;left:10px;z-index:2}.free-toc-container{position:fixed;bottom:3em;left:.5em;max-width:300px;z-index:10}@media(max-width: 768px){.free-toc-container{bottom:5em}}#free-toc{max-height:calc(100vh - 275px);overflow:auto}.free-toc-container .list-group a{text-decoration:none}.free-toc-container .list-group li:not(.active):hover{background-color:#f5f5f5;color:#fff}.free-toc-container .list-group .active a{color:#fff;pointer-events:none;font-weight:bold}.pagination{--bs-pagination-padding-x: 0.75rem;--bs-pagination-padding-y: 0.375rem;--bs-pagination-font-size:0.95rem;--bs-pagination-color: var(--bs-link-color);--bs-pagination-bg: #fff;--bs-pagination-border-width: 1px;--bs-pagination-border-color: #dee2e6;--bs-pagination-border-radius: 0.375rem;--bs-pagination-hover-color: var(--bs-link-hover-color);--bs-pagination-hover-bg: #e9ecef;--bs-pagination-hover-border-color: #dee2e6;--bs-pagination-focus-color: var(--bs-link-hover-color);--bs-pagination-focus-bg: #e9ecef;--bs-pagination-focus-box-shadow: 0 0 0 0.25rem rgba(82, 109, 120, 0.25);--bs-pagination-active-color: #fff;--bs-pagination-active-bg: #526d78;--bs-pagination-active-border-color: #526d78;--bs-pagination-disabled-color: #6c757d;--bs-pagination-disabled-bg: #fff;--bs-pagination-disabled-border-color: #dee2e6;display:flex;padding-left:0;list-style:none}.page-link{position:relative;display:block;padding:var(--bs-pagination-padding-y) var(--bs-pagination-padding-x);font-size:var(--bs-pagination-font-size);color:var(--bs-pagination-color);text-decoration:none;background-color:var(--bs-pagination-bg);border:var(--bs-pagination-border-width) solid var(--bs-pagination-border-color);transition:color .15s ease-in-out,background-color .15s ease-in-out,border-color .15s ease-in-out,box-shadow .15s ease-in-out}@media(prefers-reduced-motion: reduce){.page-link{transition:none}}.page-link:hover{z-index:2;color:var(--bs-pagination-hover-color);background-color:var(--bs-pagination-hover-bg);border-color:var(--bs-pagination-hover-border-color)}.page-link:focus{z-index:3;color:var(--bs-pagination-focus-color);background-color:var(--bs-pagination-focus-bg);outline:0;box-shadow:var(--bs-pagination-focus-box-shadow)}.page-link.active,.active>.page-link{z-index:3;color:var(--bs-pagination-active-color);background-color:var(--bs-pagination-active-bg);border-color:var(--bs-pagination-active-border-color)}.page-link.disabled,.disabled>.page-link{color:var(--bs-pagination-disabled-color);pointer-events:none;background-color:var(--bs-pagination-disabled-bg);border-color:var(--bs-pagination-disabled-border-color)}.page-item:not(:first-child) .page-link{margin-left:-1px}.page-item:first-child .page-link{border-top-left-radius:var(--bs-pagination-border-radius);border-bottom-left-radius:var(--bs-pagination-border-radius)}.page-item:last-child .page-link{border-top-right-radius:var(--bs-pagination-border-radius);border-bottom-right-radius:var(--bs-pagination-border-radius)}.pagination-lg{--bs-pagination-padding-x: 1.5rem;--bs-pagination-padding-y: 0.75rem;--bs-pagination-font-size:1.1875rem;--bs-pagination-border-radius: 0.5rem}.pagination-sm{--bs-pagination-padding-x: 0.5rem;--bs-pagination-padding-y: 0.25rem;--bs-pagination-font-size:0.83125rem;--bs-pagination-border-radius: 0.25rem}.table{--bs-table-color: var(--bs-body-color);--bs-table-bg: transparent;--bs-table-border-color: var(--bs-border-color);--bs-table-accent-bg: transparent;--bs-table-striped-color: var(--bs-body-color);--bs-table-striped-bg: rgba(0, 0, 0, 0.05);--bs-table-active-color: var(--bs-body-color);--bs-table-active-bg: rgba(0, 0, 0, 0.1);--bs-table-hover-color: var(--bs-body-color);--bs-table-hover-bg: rgba(0, 0, 0, 0.075);width:100%;margin-bottom:1rem;color:var(--bs-table-color);vertical-align:top;border-color:var(--bs-table-border-color)}.table>:not(caption)>*>*{padding:.5rem .5rem;background-color:var(--bs-table-bg);border-bottom-width:1px;box-shadow:inset 0 0 0 9999px var(--bs-table-accent-bg)}.table>tbody{vertical-align:inherit}.table>thead{vertical-align:bottom}.table-group-divider{border-top:2px solid currentcolor}.caption-top{caption-side:top}.table-sm>:not(caption)>*>*{padding:.25rem .25rem}.table-bordered>:not(caption)>*{border-width:1px 0}.table-bordered>:not(caption)>*>*{border-width:0 1px}.table-borderless>:not(caption)>*>*{border-bottom-width:0}.table-borderless>:not(:first-child){border-top-width:0}.table-striped>tbody>tr:nth-of-type(odd)>*{--bs-table-accent-bg: var(--bs-table-striped-bg);color:var(--bs-table-striped-color)}.table-striped-columns>:not(caption)>tr>:nth-child(even){--bs-table-accent-bg: var(--bs-table-striped-bg);color:var(--bs-table-striped-color)}.table-active{--bs-table-accent-bg: var(--bs-table-active-bg);color:var(--bs-table-active-color)}.table-hover>tbody>tr:hover>*{--bs-table-accent-bg: var(--bs-table-hover-bg);color:var(--bs-table-hover-color)}.table-primary{--bs-table-color: #000;--bs-table-bg: #dce2e4;--bs-table-border-color: #c6cbcd;--bs-table-striped-bg: #d1d7d9;--bs-table-striped-color: #000;--bs-table-active-bg: #c6cbcd;--bs-table-active-color: #000;--bs-table-hover-bg: #ccd1d3;--bs-table-hover-color: #000;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-secondary{--bs-table-color: #000;--bs-table-bg: #e2e3e5;--bs-table-border-color: #cbccce;--bs-table-striped-bg: #d7d8da;--bs-table-striped-color: #000;--bs-table-active-bg: #cbccce;--bs-table-active-color: #000;--bs-table-hover-bg: #d1d2d4;--bs-table-hover-color: #000;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-success{--bs-table-color: #000;--bs-table-bg: #d1e7dd;--bs-table-border-color: #bcd0c7;--bs-table-striped-bg: #c7dbd2;--bs-table-striped-color: #000;--bs-table-active-bg: #bcd0c7;--bs-table-active-color: #000;--bs-table-hover-bg: #c1d6cc;--bs-table-hover-color: #000;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-info{--bs-table-color: #000;--bs-table-bg: #cff4fc;--bs-table-border-color: #badce3;--bs-table-striped-bg: #c5e8ef;--bs-table-striped-color: #000;--bs-table-active-bg: #badce3;--bs-table-active-color: #000;--bs-table-hover-bg: #bfe2e9;--bs-table-hover-color: #000;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-warning{--bs-table-color: #000;--bs-table-bg: #fff3cd;--bs-table-border-color: #e6dbb9;--bs-table-striped-bg: #f2e7c3;--bs-table-striped-color: #000;--bs-table-active-bg: #e6dbb9;--bs-table-active-color: #000;--bs-table-hover-bg: #ece1be;--bs-table-hover-color: #000;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-danger{--bs-table-color: #000;--bs-table-bg: #f8d5dc;--bs-table-border-color: #dfc0c6;--bs-table-striped-bg: #eccad1;--bs-table-striped-color: #000;--bs-table-active-bg: #dfc0c6;--bs-table-active-color: #000;--bs-table-hover-bg: #e5c5cc;--bs-table-hover-color: #000;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-light{--bs-table-color: #000;--bs-table-bg: #e8eced;--bs-table-border-color: #d1d4d5;--bs-table-striped-bg: #dce0e1;--bs-table-striped-color: #000;--bs-table-active-bg: #d1d4d5;--bs-table-active-color: #000;--bs-table-hover-bg: #d7dadb;--bs-table-hover-color: #000;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-dark{--bs-table-color: #fff;--bs-table-bg: #29363a;--bs-table-border-color: #3e4a4e;--bs-table-striped-bg: #344044;--bs-table-striped-color: #fff;--bs-table-active-bg: #3e4a4e;--bs-table-active-color: #fff;--bs-table-hover-bg: #394549;--bs-table-hover-color: #fff;color:var(--bs-table-color);border-color:var(--bs-table-border-color)}.table-responsive{overflow-x:auto;-webkit-overflow-scrolling:touch}@media(max-width: 575.98px){.table-responsive-sm{overflow-x:auto;-webkit-overflow-scrolling:touch}}@media(max-width: 767.98px){.table-responsive-md{overflow-x:auto;-webkit-overflow-scrolling:touch}}@media(max-width: 991.98px){.table-responsive-lg{overflow-x:auto;-webkit-overflow-scrolling:touch}}@media(max-width: 1199.98px){.table-responsive-xl{overflow-x:auto;-webkit-overflow-scrolling:touch}}@media(max-width: 1399.98px){.table-responsive-xxl{overflow-x:auto;-webkit-overflow-scrolling:touch}}span.rg{font-weight:bold}.lined-bg{background:repeating-linear-gradient(45deg, #606dbc, #606dbc 10px, #465298 10px, #465298 20px)}@media(min-width: 768px){.geography-list-group{grid-template-columns:repeat(2, 1fr);display:grid;grid-gap:.5rem}.geography-list-group .list-group-item+.list-group-item{border-top-width:1px}}@media(min-width: 1200px){.geography-list-group{grid-template-columns:repeat(3, 1fr);display:grid;grid-gap:.5rem}.geography-list-group .list-group-item+.list-group-item{border-top-width:1px}}table .note{font-size:.8rem;line-height:1.1;position:relative;padding-left:.8rem;display:block;margin-bottom:.5rem;font-style:italic}table .note b{position:absolute;font-style:normal;left:0;top:0}h1 em.label-sub{font-style:normal}.event-date{font-size:1rem;font-style:italic;color:#526d78}.event-label{font-weight:bold;margin-bottom:0}.event-label .event-date{font-size:.8em}.badge{background-color:#e8eced !important;border-radius:0;color:#526d78}.badge.text-bg-primary{color:#fff !important;background-color:#0d6efd !important}.badge.text-bg-success{color:#fff !important;background-color:#198754 !important}.badge .bad a{text-decoration:underline}.event-single .badge{font-size:.85rem;font-weight:normal;margin-top:-0.25rem;padding-left:1rem !important;padding-right:1rem !important}.figure{display:flex;flex-direction:column;justify-content:center}@media(max-width: 768px){.figure{background:#f5f5f5;box-shadow:0 .125rem .25rem rgba(0,0,0,.075);margin-bottom:1em !important;padding-top:.5em !important}.figure .popover{background:transparent}.figure .popover .popover-header{color:#222}}img.media_resized{max-width:100%;height:auto;width:auto;margin:0 auto;max-height:200px}.protected{position:relative}.protected:after{position:absolute;content:"";width:100%;height:100%;top:0;left:0;background:transparent}@media(min-width: 992px){.inline-media-group{display:flex;justify-content:space-around}}@media screen and (max-width: 480px){.nav.nav-tabs{display:block}.nav.nav-tabs .nav-item{display:block !important}.nav.nav-tabs .nav-link{border:none;width:100%;margin:0}.nav.nav-tabs .nav-link.active{background-color:#e8eced;margin:0}}.index-list{border:1px solid #dfdfdf;background-color:#fafafa;border-radius:.375em;box-shadow:0 .125rem .25rem rgba(0,0,0,.075) !important;margin-bottom:1rem !important;margin-right:.25rem !important;margin-left:.25rem !important;padding:1rem}.index-list a{padding:.125em .25em;border-radius:.375em;display:inline-block}.index-list a.active{background:#526d78;color:#fff;text-decoration:none}.index-list a.text-muted{color:#a4a9ae !important}.card{--bs-card-spacer-y: 1rem;--bs-card-spacer-x: 1rem;--bs-card-title-spacer-y: 0.5rem;--bs-card-border-width: 1px;--bs-card-border-color: var(--bs-border-color-translucent);--bs-card-border-radius: 0.375rem;--bs-card-box-shadow: ;--bs-card-inner-border-radius: calc(0.375rem - 1px);--bs-card-cap-padding-y: 0.5rem;--bs-card-cap-padding-x: 1rem;--bs-card-cap-bg: rgba(0, 0, 0, 0.03);--bs-card-cap-color: ;--bs-card-height: ;--bs-card-color: ;--bs-card-bg: #fff;--bs-card-img-overlay-padding: 1rem;--bs-card-group-margin: 0.75rem;position:relative;display:flex;flex-direction:column;min-width:0;height:var(--bs-card-height);word-wrap:break-word;background-color:var(--bs-card-bg);background-clip:border-box;border:var(--bs-card-border-width) solid var(--bs-card-border-color);border-radius:var(--bs-card-border-radius)}.card>hr{margin-right:0;margin-left:0}.card>.list-group{border-top:inherit;border-bottom:inherit}.card>.list-group:first-child{border-top-width:0;border-top-left-radius:var(--bs-card-inner-border-radius);border-top-right-radius:var(--bs-card-inner-border-radius)}.card>.list-group:last-child{border-bottom-width:0;border-bottom-right-radius:var(--bs-card-inner-border-radius);border-bottom-left-radius:var(--bs-card-inner-border-radius)}.card>.card-header+.list-group,.card>.list-group+.card-footer{border-top:0}.card-body{flex:1 1 auto;padding:var(--bs-card-spacer-y) var(--bs-card-spacer-x);color:var(--bs-card-color)}.card-title{margin-bottom:var(--bs-card-title-spacer-y)}.card-subtitle{margin-top:calc(-.5 * var(--bs-card-title-spacer-y));margin-bottom:0}.card-text:last-child{margin-bottom:0}.card-link+.card-link{margin-left:var(--bs-card-spacer-x)}.card-header{padding:var(--bs-card-cap-padding-y) var(--bs-card-cap-padding-x);margin-bottom:0;color:var(--bs-card-cap-color);background-color:var(--bs-card-cap-bg);border-bottom:var(--bs-card-border-width) solid var(--bs-card-border-color)}.card-header:first-child{border-radius:var(--bs-card-inner-border-radius) var(--bs-card-inner-border-radius) 0 0}.card-footer{padding:var(--bs-card-cap-padding-y) var(--bs-card-cap-padding-x);color:var(--bs-card-cap-color);background-color:var(--bs-card-cap-bg);border-top:var(--bs-card-border-width) solid var(--bs-card-border-color)}.card-footer:last-child{border-radius:0 0 var(--bs-card-inner-border-radius) var(--bs-card-inner-border-radius)}.card-header-tabs{margin-right:calc(-.5 * var(--bs-card-cap-padding-x));margin-bottom:calc(-1 * var(--bs-card-cap-padding-y));margin-left:calc(-.5 * var(--bs-card-cap-padding-x));border-bottom:0}.card-header-tabs .nav-link.active{background-color:var(--bs-card-bg);border-bottom-color:var(--bs-card-bg)}.card-header-pills{margin-right:calc(-.5 * var(--bs-card-cap-padding-x));margin-left:calc(-.5 * var(--bs-card-cap-padding-x))}.card-img-overlay{position:absolute;top:0;right:0;bottom:0;left:0;padding:var(--bs-card-img-overlay-padding);border-radius:var(--bs-card-inner-border-radius)}.card-img,.card-img-top,.card-img-bottom{width:100%}.card-img,.card-img-top{border-top-left-radius:var(--bs-card-inner-border-radius);border-top-right-radius:var(--bs-card-inner-border-radius)}.card-img,.card-img-bottom{border-bottom-right-radius:var(--bs-card-inner-border-radius);border-bottom-left-radius:var(--bs-card-inner-border-radius)}.card-group>.card{margin-bottom:var(--bs-card-group-margin)}@media(min-width: 576px){.card-group{display:flex;flex-flow:row wrap}.card-group>.card{flex:1 0 0%;margin-bottom:0}.card-group>.card+.card{margin-left:0;border-left:0}.card-group>.card:not(:last-child){border-top-right-radius:0;border-bottom-right-radius:0}.card-group>.card:not(:last-child) .card-img-top,.card-group>.card:not(:last-child) .card-header{border-top-right-radius:0}.card-group>.card:not(:last-child) .card-img-bottom,.card-group>.card:not(:last-child) .card-footer{border-bottom-right-radius:0}.card-group>.card:not(:first-child){border-top-left-radius:0;border-bottom-left-radius:0}.card-group>.card:not(:first-child) .card-img-top,.card-group>.card:not(:first-child) .card-header{border-top-left-radius:0}.card-group>.card:not(:first-child) .card-img-bottom,.card-group>.card:not(:first-child) .card-footer{border-bottom-left-radius:0}}.alert{--bs-alert-bg: transparent;--bs-alert-padding-x: 1rem;--bs-alert-padding-y: 1rem;--bs-alert-margin-bottom: 1rem;--bs-alert-color: inherit;--bs-alert-border-color: transparent;--bs-alert-border: 1px solid var(--bs-alert-border-color);--bs-alert-border-radius: 0.375rem;position:relative;padding:var(--bs-alert-padding-y) var(--bs-alert-padding-x);margin-bottom:var(--bs-alert-margin-bottom);color:var(--bs-alert-color);background-color:var(--bs-alert-bg);border:var(--bs-alert-border);border-radius:var(--bs-alert-border-radius)}.alert-heading{color:inherit}.alert-link{font-weight:700}.alert-dismissible{padding-right:3rem}.alert-dismissible .btn-close{position:absolute;top:0;right:0;z-index:2;padding:1.25rem 1rem}.alert-primary{--bs-alert-color: #314148;--bs-alert-bg: #dce2e4;--bs-alert-border-color: #cbd3d7}.alert-primary .alert-link{color:#27343a}.alert-secondary{--bs-alert-color: #192023;--bs-alert-bg: #d4d7d8;--bs-alert-border-color: #bfc3c4}.alert-secondary .alert-link{color:#141a1c}.alert-success{--bs-alert-color: #0f5132;--bs-alert-bg: #d1e7dd;--bs-alert-border-color: #badbcc}.alert-success .alert-link{color:#0c4128}.alert-info{--bs-alert-color: #055160;--bs-alert-bg: #cff4fc;--bs-alert-border-color: #b6effb}.alert-info .alert-link{color:#04414d}.alert-warning{--bs-alert-color: #664d03;--bs-alert-bg: #fff3cd;--bs-alert-border-color: #ffecb5}.alert-warning .alert-link{color:#523e02}.alert-danger{--bs-alert-color: #851a31;--bs-alert-bg: #f8d5dc;--bs-alert-border-color: #f5bfcb}.alert-danger .alert-link{color:#6a1527}.alert-light{--bs-alert-color: #5d5e5f;--bs-alert-bg: #fafbfb;--bs-alert-border-color: #f8f9fa}.alert-light .alert-link{color:#4a4b4c}.alert-dark{--bs-alert-color: #192023;--bs-alert-bg: #d4d7d8;--bs-alert-border-color: #bfc3c4}.alert-dark .alert-link{color:#141a1c}.media-landing .media-reference-copy{display:block;position:relative !important;top:auto;left:auto;width:auto;height:auto}.media-landing .media-reference-copy:after{content:"Afficher le média";background:rgba(255,255,255,.1);position:absolute;top:0;left:0;width:100%;height:100%;display:flex;align-items:center;justify-content:center;color:#222;font-size:2em;font-weight:bold;text-shadow:0 0 5px #fff;backdrop-filter:blur(5px)}.media-landing .media-reference-copy img{width:auto !important;height:auto !important}</style><script type="application/ld+json"> { "@context": "https://schema.org", "@type": "WebSite", "name": "Encyclopædia Universalis", "alternateName": "Universalis", "url": "https://www.universalis.fr", "sameAs": [ "https://www.facebook.com/EncyclopedieUniversalis/", "https://www.linkedin.com/company/encyclopaedia-universalis", "https://www.twitter.com/universalisfr", "https://www.youtube.com/channel/UC0srTZiJaRaDw3fAvtCo4Ow" ] } </script> </head> <body id="page_3" class="edu SENIOR loading" > <div class="loader"></div> <a href="#main-content" id="skip"> <i aria-hidden="true" class="fa fa-fw fa-arrow-down"></i> Accéder au contenu </a> <header class="d-none d-md-block d-print-none"> <div class="container row-between"> <a class="header-logo mr-3" title="Retour à la page d'accueil" href="/"> <svg viewBox="0 0 370 76.44" height="70" width="277" id="logo-main"> <text class="fill-universalis text-darkblue" transform="translate(74.25 37.1)" font-family="Arial" font-size="43.37"> <tspan x="0" y="0">UNIVERSALIS</tspan> </text> <text class="text-darkblue subtitle" transform="translate(76.75 66.74)" font-family="Arial" font-size="35.8"> <tspan class="fill-universalis-up" x="0" y="0" font-size=".85em">.fr</tspan> </text> <g> <path class="fill-universalis-up" d="M57,6.36H4.25C1.9,6.36,0,8.27,0,10.62V62.51c0,2.35,1.9,4.25,4.25,4.25H57c2.35,0,4.25-1.9,4.25-4.25V10.62c0-2.35-1.9-4.25-4.25-4.25ZM29.63,31.87h7.33v8.06c-.77-1.77-1.61-2.99-2.96-4.38-1.41-1.44-2.42-2.35-4.36-3.29v-.39Zm2.37,29.2c-.85,.68-1.75,1.28-2.71,1.75-.11,.05-.22,.1-.33,.15H13.39c-1.67-.77-3.19-1.8-4.53-3.14-3.13-3.13-4.69-7.06-4.69-11.77s1.56-8.84,4.67-12.04c3.12-3.19,7.02-4.79,11.72-4.79,2.71,0,5.14,.49,7.3,1.43,1.81,.79,3.44,1.9,4.87,3.36,1.35,1.38,2.4,2.97,3.17,4.74,1,2.31,1.52,4.94,1.52,7.9l-.03,1.51H11.86c.37,2.25,1.36,4.04,2.95,5.37,1.6,1.33,3.64,1.99,6.13,1.99,2.79,0,4.96-.67,6.99-2.49l6.74,3.26c-.84,1.12-1.61,1.9-2.67,2.75Zm24.74-15.61c0,5.53-.44,9.31-1.31,11.35-1.07,2.49-2.64,4.39-4.72,5.72-.24,.15-.49,.3-.75,.44h-14.33c-.44-.23-.87-.49-1.26-.77-.26-.18-.43-.29-.66-.49,.29-.3,2.13-2.54,3.11-3.92,0,0-7.13-3.32-7.3-3.39-.22-1.04-.28-1.34-.44-2.83h7.98c.1,1.11,.27,1.26,.5,1.93,.41,1.15,1.07,2.05,1.97,2.69,.9,.64,2.02,.96,3.34,.96s2.45-.32,3.37-.95c.92-.63,1.61-1.55,2.06-2.78,.33-.91,.5-2.86,.5-5.85v-15.51h7.95v13.42Z"/> <path class="fill-universalis-up" d="M29.44,43.92c-.32-1.09-.87-2.04-1.58-2.88-.44-.52-.93-1-1.52-1.43-1.54-1.11-3.34-1.66-5.38-1.66-2.22,0-4.16,.62-5.83,1.86-1.05,.78-2.02,2.14-2.92,4.11H29.44Z"/> </g> </svg> </a> <div class="p-1 d-flex justify-content-end flex-column" style="height: 80px"> <div class="user-welcome"></div> <div class="login-form-wrapper" style="align-items: flex-end; display: flex; flex-direction: column" data-identification_uri="/identification/"> <div class="login-form"> <div class="btn btn-sm btn-primary keep-label-space-top"> <i aria-hidden="true" class="fa fa-user"></i> Mon Compte </div> </div> </div> <div id="toast-manager"></div> </div> </div> </header> <div class="d-print-none sticky-top"> <div role="navigation" aria-label="Navigation mobile" class="main-navbar-wrapper shadow"> <a class="navbar-brand-md d-none d-md-block" href="/"> <img alt="Universalis" src=" /typo3temp/assets/_processed_/e/b/csm_eu-blanc-2x_3bdd01b6ad.webp " width="32" height="32" alt=""> </a> <div class="container"> <nav class="navbar navbar-expand-lg navbar-dark"> <a class="navbar-brand d-md-none" href="/"> <img alt="Universalis" src=" /typo3temp/assets/_processed_/e/b/csm_eu-blanc-2x_3bdd01b6ad.webp " width="36" height="36" alt=""> </a> <div id="navbar-collapse-grid" class="navbar-collapse collapse pt-2 pt-md-0"> <ul class="navbar-nav me-auto"> <li><a href="/" title="Niveaux" class="nav-link">Accueil</a></li> <li><a href="/atlas/" title="Atlas" class="nav-link">Atlas</a></li><li><a href="/datapays/" title="DataPays" class="nav-link">DataPays</a></li><li><a href="/classification/" title="Classification" class="nav-link">Classification</a></li><li><a href="/dictionnaire/" title="Dictionnaire français Universalis : retrouvez toutes les définitions, étymologies, expressions, synonymes, exemples, et flexions de plus de 120 000 mots de la langue française." class="nav-link">Dictionnaire</a></li><li><a href="https://boutique.universalis.fr" title="Boutique" class="nav-link">Boutique</a></li> </ul> </div> <form method="get" name="searchQuery" class="flex-grow-1 my-1" id="search-form" autocomplete="off" action="/recherche/redirect/"> <div> <input type="hidden" name="tx_solrdata_pi1[__referrer][@extension]" value="SolrData" /> <input type="hidden" name="tx_solrdata_pi1[__referrer][@controller]" value="SearchForm" /> <input type="hidden" name="tx_solrdata_pi1[__referrer][@action]" value="showFr" /> <input type="hidden" name="tx_solrdata_pi1[__referrer][arguments]" value="YTowOnt9789d077876a7a0adeb4a9a4c4853f315af9bb201" /> <input type="hidden" name="tx_solrdata_pi1[__referrer][@request]" value="{"@extension":"SolrData","@controller":"SearchForm","@action":"showFr"}db953a457671272659a1e7710096726899d321a7" /> <input type="hidden" name="tx_solrdata_pi1[__trustedProperties]" value="{"data-action":1,"searchQuery":{"resultType":1,"searchedString":1}}36e15bfcc110b23d6ba91a6810381c8ec6b87146" /> </div> <input type="hidden" name="tx_solrdata_pi1[data-action]" value="https://www.universalis.fr/recherche/___searchedString___/article/1/" /> <input type="hidden" name="tx_solrdata_pi1[searchQuery][resultType]" value="ARTICLE" /> <div class="input-wrapper input-group"> <input placeholder="Rechercher sur Universalis" class="form-control" id="searchedString" autocomplete="off" type="text" name="tx_solrdata_pi1[searchQuery][searchedString]" value="" /> <button type="submit" class="btn btn-sm btn-light"> <span class="d-none d-md-block">Recherche</span> <span class="visually-hidden">Recherche</span> <span class="d-md-none"> <i aria-hidden="true" class="fa fa-fw fa-magnifying-glass"></i></span> </button> </div> </form> <button title="Afficher / masquer le menu du site" class="navbar-toggler ms-1" type="button" data-bs-toggle="collapse" data-bs-target="#navbar-collapse-grid"> <span class="navbar-toggler-icon"></span> </button> </nav> </div> </div> </div> <main class="site-fr container main rounded-bottom shadow mb-3"> <div class="user-welcome"></div> <div class="login-form-wrapper-mobile"> <div class="login-form"> <div class="btn btn-sm btn-primary keep-label-space-top"> <i aria-hidden="true" class="fa fa-user"></i> Mon Compte </div> </div> </div> <div id="ad_topslot" class="pub-container text-center"> <div class="pub-placeholder"></div> </div> <div class="d-lg-none text-center mb-3"> <a href="/abonnement" title="Je m'abonne" data-matomo-category="CTA" data-matomo-name="Clic" data-matomo-value="Top (mobile)"> <img src="/typo3temp/assets/_processed_/1/6/csm_banner-cta_4de68c3a6b.webp" width="728" height="90" alt="Abonnez-vous à Universalis pour 1 euro" class="img-max shadow-sm"/> </a> </div> <div class="row"> <div id="main-content" class="col-lg-8 col-xl-9 pb-4"> <div class="free-toc-container"> <ul id="free-toc" class="collapse list-group shadow-sm mb-1" style="overflow: hidden"> <li class="list-group-item active"> <a class="d-block" title="ÉCOLOGIE, L'individu dans son environnement" href="/encyclopedie/ecologie/"> 1. L'individu dans son environnement </a> </li> <li class="list-group-item "> <a class="d-block" title="ÉCOLOGIE, Fonctionnement des populations" href="/encyclopedie/ecologie/2-fonctionnement-des-populations/"> 2. Fonctionnement des populations </a> </li> <li class="list-group-item "> <a class="d-block" title="ÉCOLOGIE, Organisation fonctionnelle des biocénoses" href="/encyclopedie/ecologie/3-organisation-fonctionnelle-des-biocenoses/"> 3. Organisation fonctionnelle des biocénoses </a> </li> <li class="list-group-item "> <a class="d-block" title="ÉCOLOGIE, Stabilité, adaptabilité et évolution des biocénoses" href="/encyclopedie/ecologie/4-stabilite-adaptabilite-et-evolution-des-biocenoses/"> 4. Stabilité, adaptabilité et évolution des biocénoses </a> </li> <li class="list-group-item "> <a class="d-block" title="ÉCOLOGIE, Méthodologie écologique" href="/encyclopedie/ecologie/5-methodologie-ecologique/"> 5. Méthodologie écologique </a> </li> <li class="list-group-item "> <a class="d-block" title="ÉCOLOGIE, Applications de l'écologie" href="/encyclopedie/ecologie/6-applications-de-l-ecologie/"> 6. Applications de l'écologie </a> </li> <li class="list-group-item "> <a class="d-block" title="ÉCOLOGIE, Bibliographie" href="/encyclopedie/ecologie/7-bibliographie/"> 7. Bibliographie </a> </li> </ul> <button class="btn btn-sm btn-universalis" data-bs-toggle="collapse" data-bs-target="#free-toc" aria-expanded="false" title="Afficher / masquer le sommaire"> <i aria-hidden="true" class="fa fa-fw fa-eye"></i> Sommaire </button> </div> <h1 class="mb-1"> <strong>ÉCOLOGIE</strong> </h1> <ul class="pagination justify-content-center public-links "><li class="page-item active"><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 1" class="page-link" title="Abonnez-vous" data-loaded="true">1</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 2" class="page-link" title="Abonnez-vous" data-loaded="true">2</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 3" class="page-link" title="Abonnez-vous" data-loaded="true">3</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 4" class="page-link" title="Abonnez-vous" data-loaded="true">4</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 5" class="page-link" title="Abonnez-vous" data-loaded="true">5</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 6" class="page-link" title="Abonnez-vous" data-loaded="true">6</a></li></ul><div class="content paged type-article" ><div class="block_1" data-ucode="yC071134_0"><i class="anchor" id="yC071134_0"></i><div class="clearfix"></div><div class="media-container float-md-start me-2 col-md-3 text-center"><figure class="figure text-center w-100 mb-0 p-1" style="position: relative"><a data-ucode="PH995132" data-showMediaPlayer="1" class="protected media-reference-copy " title="Média : Ernst Haeckel" href="/media/ernst-haeckel-ph995132/"><img id="PH995132" loading="lazy" src="/typo3temp/assets/_processed_/a/f/csm_ph995132_29b8b812f6.webp" class="figure-img rounded mx-auto media_resized" alt="Ernst Haeckel - crédits : Hulton-Deutsch Collection/ Corbis/ Getty Images" data-description="En diffusant et en popularisant la théorie darwinienne et en énonçant la « loi de la récapitulation » (selon laquelle le développement embryonnaire serait une répétition de l'histoire évolutive), le zoologiste Ernst Haeckel (1834-1919) a joué un rôle important dans l'histoire des sciences de…" data-copyright="Hulton-Deutsch Collection/ Corbis/ Getty Images" width="864" height="1024" /></a><figcaption class="figure-caption"><p class="m-0 mt-1 fw-bold text-gray-700 text-truncate"> Ernst Haeckel </p><p class="m-0 copyright text-sm text-gray-700 text-truncate"> Hulton-Deutsch Collection/ Corbis/ Getty Images </p></figcaption></figure></div><p>Le terme écologie (du grec <em>oikos</em>, demeure, et <em>logos</em>, science) a été proposé par Ernst Haeckel en 1866 pour désigner la science qui étudie les rapports entre les organismes et le milieu où ils vivent. Cette définition reste encore valable, mais elle demande à être approfondie et précisée, car elle est trop générale. Pour la situer par rapport aux autres sciences biologiques, il est commode de considérer les divers niveaux d'organisation de la matière vivante.</p><p>Le système le plus simple ayant toutes les caractéristiques fondamentales des êtres vivants est la cellule. Des cellules, associées en tissus et en organes, sont intégrées en organismes pluricellulaires, animaux ou végétaux.</p><p>À un niveau d'intégration supérieur, les individus de certaines espèces peuvent constituer des <em>colonies</em>, comme chez les cœlentérés, ou des <em>sociétés</em>, comme chez les termites, les fourmis et les abeilles. À un niveau encore plus élevé se place, pour tous les êtres vivants, la <em>population</em>, ensemble des individus d'une même espèce liés entre eux par des liens parentaux, c'est-à-dire génétiques ; selon que la reproduction est sexuée ou non, ces liens sont évidemment différents.</p><p>Les populations des diverses espèces (bactériennes, végétales, animales...) groupées en un lieu déterminé forment un niveau d'organisation plus complexe encore, la communauté biologique ou <em>biocénose</em> ; le milieu physique et chimique où celle-ci est installée est souvent appelé <em>biotope</em> et leur ensemble est un <em>écosystème</em>. Le niveau ultime d'organisation du <a data-bs-toggle="popover" data-bs-content="Le terme de monde est utilisé dans des contextes très divers pour indiquer, d'une manière ou de l'autre, une totalité d'appartenance. Ainsi on parle du « monde grec » ou du « monde de la Renaissance », pour désigner des ensembles culturels spécifiques, situés dans notre passé historique et auxquels nous avons accès à travers un certain nombre d'œuvres qui sont venues jusqu'à ..." title="Article : MONDE" href="/encyclopedie/monde/">monde</a> vivant est constitué par l'ensemble des <a data-bs-toggle="popover" data-bs-content="Le terme « écosystème » désigne un groupe d'êtres vivants – parfois désigné sous le nom de biocénose (animaux, végétaux et microorganismes) – et de composantes physiques et chimiques – parfois regroupées sous le terme de biotope . Il correspond à une vision intégrative de la nature, centrée sur le concept d’interaction : toute composante de l’écosystème, vi..." title="Article : ÉCOSYSTÈMES" href="/encyclopedie/ecosystemes/">écosystèmes</a> de la planète, c'est-à-dire la <em>biosphère</em>.</p><p>La structure et le fonctionnement des cellules constituent l'objet de la cytologie et de la physiologie cellulaire ; la structure et le fonctionnement des êtres pluricellulaires concernent l'histologie, l'anatomie et la physiologie des organismes. L'écologie a donc pour objet essentiel l'étude des niveaux supérieurs d'organisation de la matière vivante, de la population monospécifique à l'écosystème et à la biosphère : comme pour les niveaux inférieurs, mais cette fois à l'échelle de systèmes particulièrement complexes, il s'agit de décrire des structures, de comprendre des fonctionnements et de reconstituer des évolutions. On ne peut toutefois saisir les phénomènes jouant à un certain niveau d'intégration sans connaître ceux qui interviennent aux niveaux inférieurs : de même que l'organisation des écosystèmes ne peut être comprise qu'en tenant compte du fonctionnement des populations, celui-ci ne peut l'être sans référence aux relations que chaque <a data-bs-toggle="popover" data-bs-content="La notion d'individualité ne se laisse définir ni par les procédés étymologiques habituels aux philosophes depuis que Platon en a établi la coutume dans le Cratyle , ni par une axiomatique rationnelle. Lorsqu'on s'y attache, c'est pour trouver des différences, lui opposer d'autres notions ; tout se passe comme si elle relevait de plusieurs champs sémantiques, sans qu'au premier abord i..." title="Article : INDIVIDU" href="/encyclopedie/individu/">individu</a> entretient avec le milieu.</p><p>Ces relations entre les individus d'une même espèce et le milieu où ils vivent constituent le domaine de l'écologie des populations (autrefois appelée autoécologie). Elles concernent à la fois les individus, pour eux-mêmes, et les populations qu'ils forment. Dans cette perspective, le milieu est conçu comme un ensemble de facteurs dont on distingue deux types : les facteurs <em>abiotiques</em>, liés au milieu physique et chimique, et les facteurs <em>biotiques</em>, liés aux êtres vivants présents dans l'écosystème étudié.</p><p>L'écologie des communautés (autrefois appelée synécologie ou <em>biocénotique</em>) envisage essentiellement la structure et le fonctionnement des écosystèmes, mais son champ s'étend également aux complexes d'écosystèmes, associés par exemple dans un même bassin versant, et finalement à la <a data-bs-toggle="popover" data-bs-content="L'idée de biosphère, ensemble des êtres vivants qui peuplent notre planète, remonte à J.-B. Lamarck. C'est ensuite le géologue autrichien Suess qui, en 1875, a mis en parallèle le terme de biosphère avec ceux d'hydrosphère, d'atmosphère et de lithosphère . Il étendait en fait le concept à tout ce qui constituait ou avait constitué le monde vivant, en y incluant donc aussi, par ..." title="Article : BIOSPHÈRE" href="/encyclopedie/biosphere/">biosphère</a>.</p><p>Discipline biologique, l'écologie ne se contente pas de décrire des structures et d'analyser des fonctionnements : elle tente de les interpréter dans une optique évolutionniste et doit considérer les systèmes qu'elle étudie – populations, écosystèmes – à différentes échelles de temps.<a href="javascript:" title="Lire la suite" class="read-more ms-1">[...]</a></p></div><div class="clearfix"></div></div><ul class="pagination justify-content-center public-links "><li class="page-item active"><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 1" class="page-link" title="Abonnez-vous" data-loaded="true">1</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 2" class="page-link" title="Abonnez-vous" data-loaded="true">2</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 3" class="page-link" title="Abonnez-vous" data-loaded="true">3</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 4" class="page-link" title="Abonnez-vous" data-loaded="true">4</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 5" class="page-link" title="Abonnez-vous" data-loaded="true">5</a></li><li class="page-item "><a href="/abonnement" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="page 6" class="page-link" title="Abonnez-vous" data-loaded="true">6</a></li></ul><div class="folio-block"><p class="folio-block-title text-center text-uc"><strong>La suite de cet article est accessible aux abonnés</strong></p><ul class="list-unstyled"><li><i class="fa fa-fw fa-check-circle"></i> Des contenus variés, complets et fiables</li><li><i class="fa fa-fw fa-check-circle"></i> Accessible sur tous les écrans</li><li><i class="fa fa-fw fa-check-circle"></i> Pas de publicité</li></ul><p class="text-center"><a href="/abonnement/" class="btn btn-cta text-uc" id="folio-button" data-matomo-category="Article" data-matomo-name="Folio" data-matomo-value="Block bottom">Découvrez nos offres </a></p><p class="text-center text-sm"> Déjà abonné ? <a href="/identification/" id="folio-login">Se connecter</a></p></div><div id="ad_mpuslot" class="pub-container text-center"><div class="pub-placeholder"></div></div><section class="mt-3 mb-5"><i class="anchor" id="C071134_AUTHORS"></i><p class="title-minor">Écrit par</p><ul class="list-group"><li class="list-group-item" data-dc-author="BLANDIN, Patrick"><a title="Auteur : Patrick BLANDIN" rel="author" href="/auteurs/patrick-blandin/"><strong>Patrick BLANDIN</strong></a> : professeur au Muséum national d'histoire naturelle, directeur de la Grande Galerie de l'évolution </li><li class="list-group-item" data-dc-author="COUVET, Denis"><a title="Auteur : Denis COUVET" rel="author" href="/auteurs/denis-couvet/"><strong>Denis COUVET</strong></a> : professeur au Muséum national d'histoire naturelle, Paris </li><li class="list-group-item" data-dc-author="LAMOTTE, Maxime"><a title="Auteur : Maxime LAMOTTE" rel="author" href="/auteurs/maxime-lamotte/"><strong>Maxime LAMOTTE</strong></a> : professeur honoraire à l'université de Paris-VI-Pierre-et-Marie-Curie (faculté des sciences), ancien directeur du laboratoire de zoologie de l'École normale supérieure </li><li class="list-group-item" data-dc-author="SACCHI, Cesare F."><a title="Auteur : Cesare F. SACCHI" rel="author" href="/auteurs/cesare-f.-sacchi/"><strong>Cesare F. SACCHI</strong></a> : professeur d'écologie à l'université de Paris-VI, directeur de l'Institut d'écologie et d'éthologie de Pavie </li></ul></section><section class="mb-5"><div class="text-center mb-3 d-print-none"><a class="d-inline-flex flex-row justify-content-center align-items-center btn btn-light btn-large text-start" title="Carte mentale : ÉCOLOGIE" href="/carte-mentale/ecologie/"><i class="fa fa-sitemap fa-rotate-270 me-1 fs-1 text-universalis"></i><p class="m-0"><strong class="fs-4">Carte mentale</strong><br /> Élargissez votre recherche dans Universalis </p></a></div></section> <section class="mb-5 article-classification" data-print-label="la classification" data-print-target=".article-classification"> <i class="anchor" id="yC071134_ROOT_LINES"></i> <p class="title-minor">Classification</p> <nav aria-label="Classification 1"> <ol class="breadcrumb"> <li class="breadcrumb-item"> <a title="Classification : Sciences de la vie" href="/classification/sciences-de-la-vie/"> Sciences de la vie </a> </li> <li class="breadcrumb-item active" data-dc-subject="Écologie"> <a title="Classification : Écologie" href="/classification/sciences-de-la-vie/ecologie/"> Écologie </a> </li> </ol> </nav> <nav aria-label="Classification 2"> <ol class="breadcrumb"> <li class="breadcrumb-item"> <a title="Classification : Sciences de la vie" href="/classification/sciences-de-la-vie/"> Sciences de la vie </a> </li> <li class="breadcrumb-item"> <a title="Classification : Écologie" href="/classification/sciences-de-la-vie/ecologie/"> Écologie </a> </li> <li class="breadcrumb-item active" data-dc-subject="Écophysiologie"> <a title="Classification : Écophysiologie" href="/classification/sciences-de-la-vie/ecologie/ecophysiologie/"> Écophysiologie </a> </li> </ol> </nav> <nav aria-label="Classification 3"> <ol class="breadcrumb"> <li class="breadcrumb-item"> <a title="Classification : Sciences de la vie" href="/classification/sciences-de-la-vie/"> Sciences de la vie </a> </li> <li class="breadcrumb-item"> <a title="Classification : Écologie" href="/classification/sciences-de-la-vie/ecologie/"> Écologie </a> </li> <li class="breadcrumb-item active" data-dc-subject="Biologie et dynamique des populations"> <a title="Classification : Biologie et dynamique des populations" href="/classification/sciences-de-la-vie/ecologie/biologie-et-dynamique-des-populations/"> Biologie et dynamique des populations </a> </li> </ol> </nav> <nav aria-label="Classification 4"> <ol class="breadcrumb"> <li class="breadcrumb-item"> <a title="Classification : Sciences de la vie" href="/classification/sciences-de-la-vie/"> Sciences de la vie </a> </li> <li class="breadcrumb-item"> <a title="Classification : Écologie" href="/classification/sciences-de-la-vie/ecologie/"> Écologie </a> </li> <li class="breadcrumb-item active" data-dc-subject="Structure et fonctionnement des écosystèmes"> <a title="Classification : Structure et fonctionnement des écosystèmes" href="/classification/sciences-de-la-vie/ecologie/structure-et-fonctionnement-des-ecosystemes/"> Structure et fonctionnement des écosystèmes </a> </li> </ol> </nav> <nav aria-label="Classification 5"> <ol class="breadcrumb"> <li class="breadcrumb-item"> <a title="Classification : Sciences de la vie" href="/classification/sciences-de-la-vie/"> Sciences de la vie </a> </li> <li class="breadcrumb-item active" data-dc-subject="Sciences de la vie: instruments et techniques expérimentales"> <a title="Classification : Sciences de la vie: instruments et techniques expérimentales" href="/classification/sciences-de-la-vie/sciences-de-la-vie-instruments-et-techniques-experimentales/"> Sciences de la vie: instruments et techniques expérimentales </a> </li> </ol> </nav> <nav aria-label="Classification 6"> <ol class="breadcrumb"> <li class="breadcrumb-item"> <a title="Classification : Sciences de la vie" href="/classification/sciences-de-la-vie/"> Sciences de la vie </a> </li> <li class="breadcrumb-item"> <a title="Classification : Écologie" href="/classification/sciences-de-la-vie/ecologie/"> Écologie </a> </li> <li class="breadcrumb-item active" data-dc-subject="Écologie appliquée"> <a title="Classification : Écologie appliquée" href="/classification/sciences-de-la-vie/ecologie/ecologie-appliquee/"> Écologie appliquée </a> </li> </ol> </nav> </section> <section class="mb-5 d-print-none"> <i class="anchor" id="yC071134_MEDIAS"></i> <p class="title-minor">Médias</p> <div class="row"> <div class="col-12 col-md-4 text-center p-1"> <figure class="figure text-center w-100 mb-0 p-1" style="position: relative"> <a data-ucode="PH995132" data-showMediaPlayer="1" class="protected media-reference-copy " title="Média : Ernst Haeckel" href="/media/ernst-haeckel-ph995132/"> <img id="PH995132" loading="lazy" src="/typo3temp/assets/_processed_/a/f/csm_ph995132_29b8b812f6.webp" class="figure-img rounded mx-auto media_resized" alt="Ernst Haeckel - crédits : Hulton-Deutsch Collection/ Corbis/ Getty Images" data-description="En diffusant et en popularisant la théorie darwinienne et en énonçant la « loi de la récapitulation » (selon laquelle le développement embryonnaire serait une répétition de l'histoire évolutive), le zoologiste Ernst Haeckel (1834-1919) a joué un rôle important dans l'histoire des sciences de…" data-copyright="Hulton-Deutsch Collection/ Corbis/ Getty Images" width="864" height="1024" /> </a> <figcaption class="figure-caption"> <p class="m-0 mt-1 fw-bold text-gray-700 text-truncate"> Ernst Haeckel </p> <p class="m-0 copyright text-sm text-gray-700 text-truncate"> Hulton-Deutsch Collection/ Corbis/ Getty Images </p> </figcaption> </figure> </div> <div class="col-12 col-md-4 text-center p-1"> <figure class="figure text-center w-100 mb-0 p-1" style="position: relative"> <a data-ucode="V070862A" data-showMediaPlayer="1" class="protected media-reference-copy " title="Média : Flux d'énergie chez les animaux et les végétaux" href="/media/flux-denergie-chez-les-animaux-et-les-vegetaux-v070862a/"> <img id="V070862A" loading="lazy" src="/typo3temp/assets/_processed_/f/5/csm_v070862a_28f2508da1.webp" class="figure-img rounded mx-auto media_resized" alt="Flux d'énergie chez les animaux et les végétaux - crédits : Encyclopædia Universalis France" data-description="Répartition des flux d'énergie : a) chez les végétaux et b) chez les animaux." data-copyright="Encyclopædia Universalis France" width="2000" height="1160" /> </a> <figcaption class="figure-caption"> <p class="m-0 mt-1 fw-bold text-gray-700 text-truncate"> Flux d'énergie chez les animaux et les végétaux </p> <p class="m-0 copyright text-sm text-gray-700 text-truncate"> Encyclopædia Universalis France </p> </figcaption> </figure> </div> <div class="col-12 col-md-4 text-center p-1"> <figure class="figure text-center w-100 mb-0 p-1" style="position: relative"> <a data-ucode="V070862B" data-showMediaPlayer="1" class="protected media-reference-copy " title="Média : Métabolisme d'animaux homéothermes" href="/media/metabolisme-danimaux-homeothermes-v070862b/"> <img id="V070862B" loading="lazy" src="/typo3temp/assets/_processed_/f/4/csm_v070862b_8a9227154c.webp" class="figure-img rounded mx-auto media_resized" alt="Métabolisme d'animaux homéothermes - crédits : Encyclopædia Universalis France" data-description="Variation du métabolisme de base A d'animaux homéothermes en fonction de leur poids W." data-copyright="Encyclopædia Universalis France" width="1220" height="2000" /> </a> <figcaption class="figure-caption"> <p class="m-0 mt-1 fw-bold text-gray-700 text-truncate"> Métabolisme d'animaux homéothermes </p> <p class="m-0 copyright text-sm text-gray-700 text-truncate"> Encyclopædia Universalis France </p> </figcaption> </figure> </div> </div> <div class="text-center"> <a class="btn btn-sm btn-primary" rel="related" href="/medias/ecologie/"> Afficher les 15 médias de l'article <span class="rg">ÉCOLOGIE</span> <i aria-hidden="true" class="fa fa-fw fa-chevron-right"></i> </a> </div> </section> <section class="mb-5 d-print-none learn-more"> <i class="anchor" id="yC071134_REFERENCES"></i> <p class="title-minor">Autres références</p> <ul class="list-group"> <li class="list-group-item"> <li class="list-group-item"> <h3 class="fs-5 m-0 mt-1"> <a href="/encyclopedie/ecologie-et-societe/"> <span class="rg">ÉCOLOGIE ET SOCIÉTÉ</span> </a> </h3> <ul class="list-inline list-inline-bullet mb-1 text-muted"> <li class="list-inline-item text-sm">Écrit par <span rel="author">Robert BARBAULT</span> </li> <li class="list-inline-item text-sm">7 804 mots</li> <li class="list-inline-item text-sm">5 médias </li> </ul> <div class="d-none d-md-inline-block float-start me-3"> <a class="protected" title="Article : &Eacute;COLOGIE ET SOCI&Eacute;T&Eacute;" href="/encyclopedie/ecologie-et-societe/"> <img class="rounded shadow" loading="lazy" alt="Déforestation à Guiroutou, Côte d'Ivoire - crédits : P. Poilecot/ CIRAD" width="125" height="125" src="/typo3temp/assets/_processed_/e/8/csm_pa081494_3542d81eb3.webp"/> </a> </div> <div class="snippet-content mt-3"> <p>L'<a data-bs-toggle="popover" data-bs-content="Le terme écologie (du grec oikos , demeure, et logos , science) a été proposé par Ernst Haeckel en 1866 pour désigner la science qui étudie les rapports entre les organismes et le milieu où ils vivent. Cette définition reste encore valable, mais elle demande à être approfondie et précisée, car elle est trop générale. Pour la situer par rapport aux autres sciences biologiques,..." title="Article : ÉCOLOGIE" href="/encyclopedie/ecologie/">écologie</a>, comme science et vision du <a data-bs-toggle="popover" data-bs-content="Le terme de monde est utilisé dans des contextes très divers pour indiquer, d'une manière ou de l'autre, une totalité d'appartenance. Ainsi on parle du « monde grec » ou du « monde de la Renaissance », pour désigner des ensembles culturels spécifiques, situés dans notre passé historique et auxquels nous avons accès à travers un certain nombre d'œuvres qui sont venues jusqu'à ..." title="Article : MONDE" href="/encyclopedie/monde/">monde</a>, a un destin singulier. Fondée dans la seconde moitié du <span class="pc">xix</span><sup>e</sup> siècle, dans les remous d'une <a data-bs-toggle="popover" data-bs-content="L'opposition entre « pays industrialisés » et « Tiers Monde », caractéristique majeure du monde contemporain, est une conséquence de la révolution industrielle dont la première phase se déroule en Angleterre dans le dernier tiers du xviii e siècle.. Vers 1760, l'Angleterre est encore une société pré-industrielle. Elle doit à sa position dominante dans le grand commerce mari..." title="Article : RÉVOLUTION INDUSTRIELLE" href="/encyclopedie/revolution-industrielle/">révolution industrielle</a> qui va peu à peu mais inexorablement façonner notre monde, elle constitue une réponse de la communauté scientifique, alors largement...</p> </div> </li> </li> <li class="list-group-item"> <h3 class="fs-5 m-0 mt-1"> <a href="/encyclopedie/agence-francaise-pour-la-biodiversite/#c7588"> <span class="rg">AGENCE FRANÇAISE POUR LA BIODIVERSITÉ</span> </a> </h3> <ul class="list-inline list-inline-bullet mb-1 text-muted"> <li class="list-inline-item text-sm">Écrit par <span rel="author">Denis COUVET</span> </li> <li class="list-inline-item text-sm">1 558 mots</li> <li class="list-inline-item text-sm">1 média </li> </ul> <div class="d-none d-md-inline-block float-start me-3"> <a class="protected" title="Article : AGENCE FRAN&Ccedil;AISE POUR LA BIODIVERSIT&Eacute;" href="/encyclopedie/agence-francaise-pour-la-biodiversite/"> <img class="rounded shadow" loading="lazy" alt="Pollinisateur en action - crédits : L. Casagrande-Fioretti" width="125" height="125" src="/typo3temp/assets/_processed_/0/3/csm_ph151399_646fa90b99.webp"/> </a> </div> <div class="snippet-content mt-3"> Une autre mission de l’Agence française pour la biodiversité est d’assurer un appui technique, notamment dans le cadre de la compensation écologique, stade ultime de la séquence « éviter, réduire, compenser » (E.R.C.) imposée lors de la construction d’infrastructures : on évite de détruire la biodiversité... </div> </li> <li class="list-group-item"> <h3 class="fs-5 m-0 mt-1"> <a href="/encyclopedie/ammonification-ammonisation/#c7588"> <span class="rg">AMMONIFICATION </span><span class="rm">ou </span><span class="rg">AMMONISATION</span> </a> </h3> <ul class="list-inline list-inline-bullet mb-1 text-muted"> <li class="list-inline-item text-sm">Écrit par <span rel="author">Didier LAVERGNE</span> </li> <li class="list-inline-item text-sm">1 920 mots</li> <li class="list-inline-item text-sm">1 média </li> </ul> <div class="d-none d-md-inline-block float-start me-3"> <a class="protected" title="Article : AMMONIFICATION ou AMMONISATION" href="/encyclopedie/ammonification-ammonisation/"> <img class="rounded shadow" loading="lazy" alt="Cycle de l'azote - crédits : Planeta Actimedia S.A.© Encyclopædia Universalis France pour la version française." width="125" height="125" src="/typo3temp/assets/_processed_/8/6/csm_vi990044_803d59a4c1.webp"/> </a> </div> <div class="snippet-content mt-3"> On considère qu'en culture, les conditions favorables aux ammonificateurs des sols sont une température de 30 <sup>0</sup>C, et une hydratation à 60 p. 100 de la capacité de rétention de la terre, ce qui garantit une bonne aération. Ces conditions idéales sont loin d'être réunies dans les milieux naturels... </div> </li> <li class="list-group-item"> <h3 class="fs-5 m-0 mt-1"> <a href="/encyclopedie/anthropologie/#c7588"> <span class="rg">ANTHROPOLOGIE</span> </a> </h3> <ul class="list-inline list-inline-bullet mb-1 text-muted"> <li class="list-inline-item text-sm">Écrit par <span rel="author">Élisabeth COPET-ROUGIER</span> et <span rel="author">Christian GHASARIAN</span> </li> <li class="list-inline-item text-sm">16 158 mots</li> <li class="list-inline-item text-sm">1 média </li> </ul> <div class="d-none d-md-inline-block float-start me-3"> <a class="protected" title="Article : ANTHROPOLOGIE" href="/encyclopedie/anthropologie/"> <img class="rounded shadow" loading="lazy" alt="Claude Lévi-Strauss - crédits : Marc Gantier/ Gamma-Rapho/ Getty Images" width="125" height="125" src="/typo3temp/assets/_processed_/f/8/csm_pa082775_36e7ed6627.webp"/> </a> </div> <div class="snippet-content mt-3"> ...des bases matérielles des sociétés humaines en relation avec leur adaptation au milieu. La société passe ainsi au rang d'un sous-système au sein d'un ensemble plus vaste qui inclut la nature animale et végétale, l'écosystème. Avant de comprendre, il est nécessaire d'identifier, de compter, de mesurer... </div> </li> <li class="list-group-item"> <h3 class="fs-5 m-0 mt-1"> <a href="/encyclopedie/anthropologie-des-zoonoses/#c7588"> <span class="rg">ANTHROPOLOGIE DES ZOONOSES</span> </a> </h3> <ul class="list-inline list-inline-bullet mb-1 text-muted"> <li class="list-inline-item text-sm">Écrit par <span rel="author">Frédéric KECK</span> et <span rel="author">Christos LYNTERIS</span> </li> <li class="list-inline-item text-sm">3 954 mots</li> <li class="list-inline-item text-sm">4 médias </li> </ul> <div class="d-none d-md-inline-block float-start me-3"> <a class="protected" title="Article : ANTHROPOLOGIE DES ZOONOSES" href="/encyclopedie/anthropologie-des-zoonoses/"> <img class="rounded shadow" loading="lazy" alt="Chasseurs de rats dans le port de Liverpool - crédits : Wellcome Collection ; CC BY 4.0" width="125" height="125" src="/typo3temp/assets/_processed_/a/1/csm_ph200531_6e84d3f110.webp"/> </a> </div> <div class="snippet-content mt-3"> Alors que le concept de zoonose s’était d’abord développé dans une écologie où l’homme ne joue qu’un rôle accidentel, l’anthropologie est davantage interpellée lorsque la zoonose est caractérisée comme une émergence catastrophique. La fin des années 1970 voit émerger de nouveaux pathogènes impliquant... </div> </li> <li class="list-unstyled text-center mt-3"> <a class="btn btn-sm btn-primary" title="Afficher la liste complète des références" rel="related" href="/index/ecologie/"> Afficher les 72 références <i aria-hidden="true" class="fa fa-fw fa-chevron-right"></i> </a> </li> </ul> </section> </div> <div class="col-lg-4 col-xl-3 px-0 d-none d-lg-block side-bar position-relative"> <div id="ad_rightslot" class="pub-container text-center"> <div class="pub-placeholder"></div> </div> <div id="ad_rightslot2" class="pub-container text-center"> <div class="pub-placeholder"></div> </div> <a class="wrapper cta" href="/abonnement" title="Abonnez-vous à Universalis pour 1 euro" data-matomo-category="CTA" data-matomo-name="Clic" data-matomo-value="Col right"> <img src="/typo3temp/assets/_processed_/b/6/csm_abo-3_f303c484f7.webp" class="full-height" loading="lazy" width="290" height="476" alt="Abonnez-vous à Universalis pour 1 euro"> </a> <div id="newsletter_subscription" class="wrapper p-3"> <p class="fs-3 fw-bold text-center text-light mb-1">Rejoignez-nous</p> <p class="text-center"> Inscrivez-vous à notre <strong>newsletter</strong></p> <div class="input-group mb-3"> <input type="text" class="form-control" placeholder="votre email"> <div class="input-group-append"> <button id="newsletter_subscription_button" class="btn btn-primary text-uppercase" type="button"> <strong>S'inscrire</strong></button> </div> </div> </div> </div> </div> </main> <div id="bandeau-footer"> <div class="container"> <p><a href="/abonnement" title="Abonnez-vous à Universalis pour 1 euro">Accédez à l'intégralité d'Universalis.fr <strong>sans publicité</strong></a></p> </div> </div> <footer class="container footer text-center d-print-none"> <ul class="list-inline list-inline-bullet mb-2"> <li class="list-inline-item"> <a href="http://www.encyclopaedia-universalis.fr/" title="Encyclopædia Universalis" rel="noopener" target="_blank">Encyclopædia Universalis </a> </li> <li class="list-inline-item"> <a href="https://www.encyclopaedia-universalis.fr/nous-contacter/infos/" title="Contactez-nous" rel="noopener nofollow" target="_blank">Contact</a></li> <li class="list-inline-item"> <a title="Mentions légales" rel="nofollow" href="/mentions-legales/">Mentions légales et politique de confidentialité</a> </li> <li class="list-inline-item"> <a href="https://abonnement.universalis.fr/typo3conf/ext/mailer/Resources/Public/Pdf/cgv.pdf" rel="noopener nofollow" target="_blank" title="Conditions générales de vente">CGV</a> </li> <li class="list-inline-item d-none d-md-inline-block"> <a href="javascript:" id="rgpd" rel="nofollow" title="Consentement RGPD">RGPD</a> </li> </ul> <p class="mb-2">Consulter le <a title="Dictionnaire de l'Encyclopædia Universalis" href="/dictionnaire/"> dictionnaire de l'Encyclopædia Universalis </a> </p> <p class="mb-3">© 2024 Encyclopædia Universalis France. Tous droits de propriété industrielle et intellectuelle réservés.</p> </footer> <a href="javascript:" class="btn btn-primary" id="back-to-top" title="Retour en haut de page"><i aria-hidden="true" class="far fa-arrow-up"></i></a> <script> // noinspection ES6ConvertVarToLetConst,JSUnusedGlobalSymbols var subscription_uri = 'https://www.universalis.fr/abonnement/' bodyLoaded = true; loaded(); </script> <script>(()=>{var e,a,r,_,d,c={},b={};function __webpack_require__(e){var a=b[e];if(void 0!==a)return a.exports;var r=b[e]={id:e,loaded:!1,exports:{}};return c[e].call(r.exports,r,r.exports,__webpack_require__),r.loaded=!0,r.exports}__webpack_require__.m=c,__webpack_require__.n=e=>{var a=e&&e.__esModule?()=>e.default:()=>e;return __webpack_require__.d(a,),a},__webpack_require__.d=(e,a)=>{for(var r in a)__webpack_require__.o(a,r)&&!__webpack_require__.o(e,r)&&Object.defineProperty(e,r,{enumerable:!0,get:a[r]})},__webpack_require__.f={},__webpack_require__.e=e=>Promise.all(Object.keys(__webpack_require__.f).reduce(((a,r)=>(__webpack_require__.f[r](e,a),a)),[])),__webpack_require__.u=e=>"js/"+e+"."+{13:"80d7e21db2f95ae99720",26:"378d7d0789f5af40346c",32:"f20d73ca82d98e7527c8",69:"7313a4f3fed161acce4b",89:"c626d929711e7d9749bb",596:"f42a3891c8a58bdfb96a",603:"ea3de1982fc803344bdd",689:"31c396836cb7e99e178d",709:"d4e0658f463af05ae223",897:"3e91b02003555b671e4b",1095:"9e59e92f30df1ec926e4",1203:"76e8c707d971e06822da",1250:"9865f41df828b2e8b447",1292:"93adae3e6784d8a8a976",1349:"9efae443d9392248187c",1377:"446728afc62ef1c19520",1394:"86027a2d101b1eb260c8",1419:"1387d471be2d6ae0a0aa",1641:"20b900e0adda4c80db1d",1651:"c86d270cf7725cab1d79",1956:"74c8ff5a5ca0f4903469",2034:"10aef63b282d61d6c639",2035:"e1aebed4bfd6bd09ee7e",2353:"44bbdb8df13e8a622208",2372:"a22ed3ddae437e986477",2724:"0bf0368274ee9e0747d2",2832:"eb28f9840b1271c7d62a",2936:"62bb94fb29069bd56587",2966:"2c1ff93e3007f7225cb8",2979:"d966b75bfce67342eeac",3107:"4ae39080543abe3da100",3138:"75edb076a18972355143",3195:"8dc5dbdee19baf1191a1",3292:"4204bb21e63dbda3313b",3493:"b843364884901abc3eb0",3559:"0407e4aa7480eddd0ef1",3581:"e65b43625d381405a12c",3716:"d6ab7b12b699eee673e0",3862:"4213b001dbf03cea161e",3963:"49b1c889b28a0c5714d1",4123:"874e7dece76e9760a495",4152:"70172b2b6a2202bc19e4",4254:"05f327b93d437c2f3601",4299:"37495161e93e1b14a62a",4325:"88bc29304269c158b066",4407:"767000e4cd0f3e15307e",4512:"f82c4b423f5c0dbd30dd",4531:"7457626c1fb01d704537",4644:"2721494c4c12cc6d3bc2",4785:"a097bbcd58df09778180",4840:"1b05bd12ede150f03e77",4947:"676f3abafc2b02c5fa31",5015:"211414a16467b37cae34",5058:"c0ca6a30fe8b53f6d2b2",5106:"908216bcff18d3fd7aad",5133:"74d1d5d467c9be5ef47a",5267:"e5a307f4cc5995492285",5274:"0be3aa33d3ffc719dcff",5505:"313286a530596f7a6b25",5662:"9b43b1237225d7ff7542",5667:"851d77804ca881f31ac9",5797:"861f7e8a005e9a4063c6",5865:"f1a798618f3b6f9903b0",5977:"bf7db06c1133a66449f3",5987:"028c01be18ffb17f5482",6017:"f4e0251f6746e3f339f1",6322:"30a126263c6970e6793a",6363:"9d5f249f8ba4467549c4",6574:"6f9df14941f855202a9d",6903:"d6dfc26ae3076c85ed1f",7024:"f436d8b414f7a558c554",7084:"4cdc22dfd7d247240602",7090:"291628b7c2c07c2d6f94",7131:"c27aefd28ade057122d1",7204:"3e5234153f1f490db272",7262:"0380fb22702b7b1fbf61",7269:"a46d5b5fe65c5fd574e8",7329:"430ef38cd030411e470c",7335:"920664487a3d2ae93e13",7345:"66b366ea63a5ec112a1d",7357:"557d366dc6db747c3062",7363:"f3c9aaf269ff7f29b793",7369:"38be51a057fc00f0f7ef",7539:"45b957bf1324ccf78ffe",7697:"35e517bb715e4b5c7b10",7794:"4e5dcc0a46043951fd82",8010:"1d4a0c12caaaca4542da",8113:"1a49932d3f9f58a0e19b",8151:"286ead9d06e13c599aa4",8211:"983a3e4ce7f9d841c8fb",8224:"972a60c780f0448a5f47",8236:"2aead7cb9f823d2b0bd9",8285:"01146be20c37f70177c6",8519:"69cf4d6707f67ab66004",8603:"5723927781ff4ed61db8",8661:"7093c6c8677b30f38493",8852:"d237c16bb6b8e4258674",9019:"633dbe3b28ca96a0abf5",9049:"d9917b1342dda4789bae",9198:"097d936d40dd0f3aa9da",9229:"5d03eb6d1503a7e3708e",9411:"fa23eea5dd4e55cc2da6",9416:"6ee4ffb0e861caab07f2",9529:"f93f4d7733e910b1cad8",9612:"48e1d8ac85448df568dc",9767:"51e75328b2c6a3d3fe16"}[e]+".bundle.js",__webpack_require__.miniCssF=e=>4826===e?"css/index.css":"css/"+e+".css",__webpack_require__.g=function(){if("object"==typeof globalThis)return globalThis;try{return this||new Function("return this")()}catch(e){if("object"==typeof window)return window}}(),__webpack_require__.o=(e,a)=>Object.prototype.hasOwnProperty.call(e,a),e={},a="@universalis/web_fr:",__webpack_require__.l=(r,_,d,c)=>{if(e[r])e[r].push(_);else{var b,f;if(void 0!==d)for(var t=document.getElementsByTagName("script"),i=0;i<t.length;i++){var n=t[i];if(n.getAttribute("src")==r||n.getAttribute("data-webpack")==a+d){b=n;break}}b||(f=!0,(b=document.createElement("script")).charset="utf-8",b.timeout=120,__webpack_require__.nc&&b.setAttribute("nonce",__webpack_require__.nc),b.setAttribute("data-webpack",a+d),b.src=r),e[r]=[_];var onScriptComplete=(a,_)=>{b.onerror=b.onload=null,clearTimeout(u);var d=e[r];if(delete e[r],b.parentNode&&b.parentNode.removeChild(b),d&&d.forEach((e=>e(_))),a)return a(_)},u=setTimeout(onScriptComplete.bind(null,void 0,{type:"timeout",target:b}),12e4);b.onerror=onScriptComplete.bind(null,b.onerror),b.onload=onScriptComplete.bind(null,b.onload),f&&document.head.appendChild(b)}},__webpack_require__.r=e=>{"undefined"!=typeof Symbol&&Symbol.toStringTag&&Object.defineProperty(e,Symbol.toStringTag,{value:"Module"}),Object.defineProperty(e,"__esModule",{value:!0})},__webpack_require__.nmd=e=>(e.paths=[],e.children||(e.children=[]),e),__webpack_require__.p="/_web/assets/",r=(e,a,r)=>{var _=document.createElement("link");return _.rel="stylesheet",_.type="text/css",_.onload=a,_.onerror=function(a){var d=a&&a.target&&a.target.src||e,c=new Error("Loading CSS chunk "+chunkId+" failed.\n("+d+")");c.code="CSS_CHUNK_LOAD_FAILED",c.request=d,_.parentNode.removeChild(_),r(c)},_.href=e,document.getElementsByTagName("head")[0].appendChild(_),_},_=e=>new Promise(((a,_)=>{var d=__webpack_require__.miniCssF(e),c=__webpack_require__.p+d;if(((e,a)=>{for(var r=document.getElementsByTagName("link"),_=0;_<r.length;_++){var d=(b=r[_]).getAttribute("data-href")||b.getAttribute("href");if("stylesheet"===b.rel&&(d===e||d===a))return b}var c=document.getElementsByTagName("style");for(_=0;_<c.length;_++){var b;if((d=(b=c[_]).getAttribute("data-href"))===e||d===a)return b}})(d,c))return a();r(c,a,_)})),d={4826:0},__webpack_require__.f.miniCss=(e,a)=>{d[e]?a.push(d[e]):0!==d[e]&&{1203:1,2724:1,2979:1,4644:1,5667:1,6363:1}[e]&&a.push(d[e]=_(e).then((()=>{d[e]=0}),(a=>{throw delete d[e],a})))},(()=>{var e={4826:0};__webpack_require__.f.j=(a,r)=>{var _=__webpack_require__.o(e,a)?e[a]:void 0;if(0!==_)if(_)r.push(_[2]);else{var d=new Promise(((r,d)=>_=e[a]=[r,d]));r.push(_[2]=d);var c=__webpack_require__.p+__webpack_require__.u(a),b=new Error;__webpack_require__.l(c,(r=>{if(__webpack_require__.o(e,a)&&(0!==(_=e[a])&&(e[a]=void 0),_)){var d=r&&("load"===r.type?"missing":r.type),c=r&&r.target&&r.target.src;b.message="Loading chunk "+a+" failed.\n("+d+": "+c+")",b.name="ChunkLoadError",b.type=d,b.request=c,_[1](b)}}),"chunk-"+a,a)}};var webpackJsonpCallback=(a,r)=>{var _,d,[c,b,f]=r,t=0;if(c.some((a=>0!==e[a]))){for(_ in b)__webpack_require__.o(b,_)&&(__webpack_require__.m[_]=b[_]);if(f)f(__webpack_require__)}for(a&&a(r);t<c.length;t++)d=c[t],__webpack_require__.o(e,d)&&e[d]&&e[d][0](),e[d]=0},a=self.webpackChunk_universalis_web_fr=self.webpackChunk_universalis_web_fr||[];a.forEach(webpackJsonpCallback.bind(null,0)),a.push=webpackJsonpCallback.bind(null,a.push.bind(a))})(),__webpack_require__.nc=void 0,document.addEventListener("DOMContentLoaded",(function(){__webpack_require__.e(3138).then(__webpack_require__.bind(__webpack_require__,73138)).then((function(){Promise.all([__webpack_require__.e(9529),__webpack_require__.e(2372)]).then(__webpack_require__.bind(__webpack_require__,82372)).then((function(e){e.default()}))}))}))})();</script> <script> let eu_article_ucode = "C071134"</script> </body> </html>