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class="vector-toc-text"> 1.1 Records of herbivores </div> </a> <ul id="toc-Records_of_herbivores-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Co-evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Co-evolution"> <div class="vector-toc-text"> 1.2 Co-evolution </div> </a> <ul id="toc-Co-evolution-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Types" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Types"> <div class="vector-toc-text"> 2 Types </div> </a> <button aria-controls="toc-Types-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Types subsection </button> <ul id="toc-Types-sublist" class="vector-toc-list"> <li id="toc-Chemical_defenses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Chemical_defenses"> <div class="vector-toc-text"> 2.1 Chemical defenses </div> </a> <ul id="toc-Chemical_defenses-sublist" class="vector-toc-list"> <li id="toc-Antiherbivory_compounds" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Antiherbivory_compounds"> <div class="vector-toc-text"> 2.1.1 Antiherbivory compounds </div> </a> <ul id="toc-Antiherbivory_compounds-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Enzymes" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Enzymes"> <div class="vector-toc-text"> 2.1.2 Enzymes </div> </a> <ul id="toc-Enzymes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mechanical_defenses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mechanical_defenses"> <div class="vector-toc-text"> 2.2 Mechanical defenses </div> </a> <ul id="toc-Mechanical_defenses-sublist" class="vector-toc-list"> <li id="toc-Spines_and_thorns" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Spines_and_thorns"> <div class="vector-toc-text"> 2.2.1 Spines and thorns </div> </a> <ul id="toc-Spines_and_thorns-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Thigmonastic_movements" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Thigmonastic_movements"> <div class="vector-toc-text"> 2.2.2 Thigmonastic movements </div> </a> <ul id="toc-Thigmonastic_movements-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Carnivorous_plants" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Carnivorous_plants"> <div class="vector-toc-text"> 2.3 Carnivorous plants </div> </a> <ul id="toc-Carnivorous_plants-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mimicry_and_camouflage" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mimicry_and_camouflage"> <div class="vector-toc-text"> 2.4 Mimicry and camouflage </div> </a> <ul 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</div> </a> <ul id="toc-Pharmaceutical-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> 5 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 6 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> 7 Further reading </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 8 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id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading">Plant defense against herbivory</h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. 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href="https://ca.wikipedia.org/wiki/Mecanismes_de_defensa_de_les_plantes_contra_els_herb%C3%ADvors" title="Mecanismes de defensa de les plantes contra els herbívors – Catalan" lang="ca" hreflang="ca" data-title="Mecanismes de defensa de les plantes contra els herbívors" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Pflanzliche_Abwehr_von_Herbivoren" title="Pflanzliche Abwehr von Herbivoren – German" lang="de" hreflang="de" data-title="Pflanzliche Abwehr von Herbivoren" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Defensas_vegetales_contra_la_herbivor%C3%ADa" title="Defensas vegetales contra la herbivoría – Spanish" lang="es" hreflang="es" data-title="Defensas vegetales contra la herbivoría" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AF%D9%81%D8%A7%D8%B9_%DA%AF%DB%8C%D8%A7%D9%87_%D8%AF%D8%B1_%D8%A8%D8%B1%D8%A7%D8%A8%D8%B1_%D8%B9%D9%84%D9%81%E2%80%8C%D8%AE%D9%88%D8%A7%D8%B1" title="دفاع گیاه در برابر علف‌خوار – Persian" lang="fa" hreflang="fa" data-title="دفاع گیاه در برابر علف‌خوار" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/D%C3%A9fense_des_plantes_contre_les_herbivores" title="Défense des plantes contre les herbivores – French" lang="fr" hreflang="fr" data-title="Défense des plantes contre les herbivores" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Pertahanan_tanaman_terhadap_herbivora" title="Pertahanan tanaman terhadap herbivora – Indonesian" lang="id" hreflang="id" data-title="Pertahanan tanaman terhadap herbivora" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Difesa_vegetale_contro_gli_erbivori" title="Difesa vegetale contro gli erbivori – Italian" lang="it" hreflang="it" data-title="Difesa vegetale contro gli erbivori" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%AA%D7%92%D7%95%D7%A0%D7%A0%D7%95%D7%AA_%D7%A6%D7%9E%D7%97%D7%99%D7%9D_%D7%9E%D7%98%D7%A8%D7%99%D7%A4%D7%94" title="התגוננות צמחים מטריפה – Hebrew" lang="he" hreflang="he" data-title="התגוננות צמחים מטריפה" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Afweer_van_planten_tegen_herbivoren" title="Afweer van planten tegen herbivoren – Dutch" lang="nl" hreflang="nl" data-title="Afweer van planten tegen herbivoren" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Obrona_przed_ro%C5%9Blino%C5%BCercami" title="Obrona przed roślinożercami – Polish" lang="pl" hreflang="pl" data-title="Obrona przed roślinożercami" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Defesas_vegetais_contra_a_herbivoria" title="Defesas vegetais contra a herbivoria – Portuguese" lang="pt" hreflang="pt" data-title="Defesas vegetais contra a herbivoria" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Defence_against_herbivory" title="Defence against herbivory – Simple English" lang="en-simple" hreflang="en-simple" data-title="Defence against herbivory" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a 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id="mw-indicator-good-star" class="mw-indicator"><div class="mw-parser-output"><a href="/wiki/Wikipedia:Good_articles*" title="This is a good article. Click here for more information."><img alt="This is a good article. Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle">(Redirected from <a href="/w/index.php?title=Plant_defenses_against_herbivory&redirect=no" class="mw-redirect" title="Plant defenses against herbivory">Plant defenses against herbivory</a>)</div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Evolutionary mechanism</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Foxglove2.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Foxglove2.jpg/300px-Foxglove2.jpg" decoding="async" width="300" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Foxglove2.jpg/450px-Foxglove2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/05/Foxglove2.jpg/600px-Foxglove2.jpg 2x" data-file-width="1280" data-file-height="960" /></a><figcaption><a href="/wiki/Foxglove" class="mw-redirect" title="Foxglove">Foxgloves</a> produce toxic chemicals including <a href="/wiki/Cardiac_glycosides" class="mw-redirect" title="Cardiac glycosides">cardiac</a> and <a href="/wiki/Steroid" title="Steroid">steroidal</a> glycosides, deterring herbivory.</figcaption></figure> Plant defense against herbivory or host-plant resistance is a range of <a href="/wiki/Adaptation" title="Adaptation">adaptations</a> <a href="/wiki/Evolution" title="Evolution">evolved</a> by <a href="/wiki/Plants" class="mw-redirect" title="Plants">plants</a> which improve their <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">survival and reproduction</a> by reducing the impact of <a href="/wiki/Herbivore" title="Herbivore">herbivores</a>. Many plants produce <a href="/wiki/Secondary_metabolites" class="mw-redirect" title="Secondary metabolites">secondary metabolites</a>, known as <a href="/wiki/Heterotelergones" class="mw-redirect" title="Heterotelergones">allelochemicals</a>, that influence the behavior, growth, or survival of herbivores. These chemical defenses can act as repellents or toxins to herbivores or reduce plant digestibility. Another defensive strategy of plants is changing their attractiveness. <a href="/wiki/Plant_perception_(physiology)" title="Plant perception (physiology)">Plants can sense being touched</a>,<a href="#cite_note-1">[1]</a> and they can respond with strategies to defend against herbivores. To prevent overconsumption by large herbivores, plants alter their appearance by changing their size or quality, reducing the rate at which they are consumed.<a href="#cite_note-Kersch-Becker_20171120-2">[2]</a> Other defensive strategies used by plants include escaping or avoiding herbivores at any time in any place – for example, by growing in a location where plants are not easily found or accessed by herbivores or by changing seasonal growth patterns. Another approach diverts herbivores toward eating non-essential parts or enhances the ability of a plant to recover from the damage caused by herbivory. Some plants encourage the presence of <a href="/wiki/Predator" class="mw-redirect" title="Predator">natural enemies</a> of herbivores, which in turn protect the plant. Each type of defense can be either constitutive (always present in the plant) or <a href="/wiki/Inducible_plant_defenses_against_herbivory" title="Inducible plant defenses against herbivory">induced</a> (produced in reaction to damage or stress caused by herbivores). Historically, insects have been the most significant herbivores, and the evolution of land plants is closely associated with the <a href="/wiki/Evolution_of_insects" title="Evolution of insects">evolution of insects</a>. While most plant defenses are directed against insects, other defenses have evolved that are aimed at <a href="/wiki/Vertebrate" title="Vertebrate">vertebrate</a> herbivores, such as <a href="/wiki/Bird" title="Bird">birds</a> and <a href="/wiki/Mammal" title="Mammal">mammals</a>. The study of plant defenses against herbivory is important from an evolutionary viewpoint; for the direct impact that these defenses have on <a href="/wiki/Agriculture" title="Agriculture">agriculture</a>, including human and livestock food sources; as beneficial 'biological control agents' in <a href="/wiki/Biological_pest_control" title="Biological pest control">biological pest control</a> programs; and in the search for <a href="/wiki/Medicinal_plants" title="Medicinal plants">plants of medical importance</a>. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_defensive_traits">Evolution of defensive traits</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=1" title="Edit section: Evolution of defensive traits">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:InsectPlantEvol.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bc/InsectPlantEvol.svg/330px-InsectPlantEvol.svg.png" decoding="async" width="330" height="172" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bc/InsectPlantEvol.svg/495px-InsectPlantEvol.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bc/InsectPlantEvol.svg/660px-InsectPlantEvol.svg.png 2x" data-file-width="553" data-file-height="288" /></a><figcaption>Timeline of plant evolution and the beginnings of different modes of insect herbivory</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Evolutionary history of plants</a></div> The earliest land plants evolved from aquatic plants around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=450">450</a> million years ago (Ma) in the <a href="/wiki/Ordovician" title="Ordovician">Ordovician</a> period. Many plants have adapted to an iodine-deficient terrestrial environment by removing iodine from their metabolism; in fact, iodine is essential only for animal cells.<a href="#cite_note-3">[3]</a> An important antiparasitic action is caused by the blockage in the transport of iodide of animal cells, inhibiting <a href="/wiki/Sodium-iodide_symporter" class="mw-redirect" title="Sodium-iodide symporter">sodium-iodide symporter</a> (NIS). Many plant pesticides are glycosides (such as cardiac <a href="/wiki/Digitoxin" title="Digitoxin">digitoxin</a>) and <a href="/wiki/Cyanogenic_glycosides" class="mw-redirect" title="Cyanogenic glycosides">cyanogenic glycosides</a> that liberate <a href="/wiki/Cyanide" title="Cyanide">cyanide</a>, which, by blocking <a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">cytochrome c oxidase</a> and <a href="/wiki/Sodium-iodide_symporter" class="mw-redirect" title="Sodium-iodide symporter">NIS</a>, is poisonous only for a large part of parasites and herbivores and not for the plant cells, in which it seems useful in the <a href="/wiki/Seed_dormancy" title="Seed dormancy">seed dormancy</a> phase. Iodide is not a pesticide but is oxidized, by vegetable peroxidase to iodine, which is a strong oxidant able to kill bacteria, fungi, and protozoa.<a href="#cite_note-4">[4]</a> The <a href="/wiki/Cretaceous" title="Cretaceous">Cretaceous</a> period saw the appearance of more plant defense mechanisms. The diversification of flowering plants (<a href="/wiki/Angiosperm" class="mw-redirect" title="Angiosperm">angiosperms</a>) at that time is associated with the sudden burst of <a href="/wiki/Speciation" title="Speciation">speciation</a> in insects.<a href="#cite_note-Ehrlich-5">[5]</a> This diversification of insects represented a major selective force in plant evolution and led to the selection of plants that had defensive adaptations. Early insect herbivores were <a href="/wiki/Mandibulate" class="mw-redirect" title="Mandibulate">mandibulate</a> and bit or chewed vegetation, but the evolution of vascular plants lead to the co-evolution of other forms of herbivory, such as sap-sucking, <a href="/wiki/Leaf_miner" title="Leaf miner">leaf mining</a>, <a href="/wiki/Gall" title="Gall">gall</a> forming and nectar-feeding.<a href="#cite_note-Labandeira-6">[6]</a> The relative abundance of different species of plants in ecological communities including forests and grasslands may be determined in part by the level of defensive compounds in the different species.<a href="#cite_note-7">[7]</a> Since the cost of replacing damaged leaves is higher in conditions where resources are scarce, it may be that plants growing in areas where water and nutrients are scarce invest more resources into anti-herbivore defenses, resulting in slower plant growth.<a href="#cite_note-Wittstock-et-al-2002-8">[8]</a> <div class="mw-heading mw-heading3"><h3 id="Records_of_herbivores">Records of herbivores</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=2" title="Edit section: Records of herbivores">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Viburnum_lesquereuxii_fossil.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/Viburnum_lesquereuxii_fossil.jpg/220px-Viburnum_lesquereuxii_fossil.jpg" decoding="async" width="220" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/Viburnum_lesquereuxii_fossil.jpg/330px-Viburnum_lesquereuxii_fossil.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/76/Viburnum_lesquereuxii_fossil.jpg/440px-Viburnum_lesquereuxii_fossil.jpg 2x" data-file-width="2000" data-file-height="1432" /></a><figcaption>Viburnum lesquereuxii leaf with insect damage; Dakota Sandstone (Cretaceous) of Ellsworth County, Kansas. Scale bar is 10 mm.</figcaption></figure> Knowledge of herbivory in geological time comes from three sources: fossilized plants, which may preserve evidence of defense (such as spines) or herbivory-related damage; the observation of plant debris in fossilised <a href="/wiki/Coprolite" title="Coprolite">animal feces</a>; and the structure of herbivore mouthparts.<a href="#cite_note-Labandeira1998-9">[9]</a> Long thought to be a <a href="/wiki/Mesozoic" title="Mesozoic">Mesozoic</a> phenomenon, evidence for herbivory is found almost as soon as fossils can show it. As previously discussed, the first land plants emerged around 450 million years ago; however, herbivory, and therefore the need for plant defenses, undoubtedly evolved among aquatic organisms in ancient lakes and oceans.<a href="#cite_note-10">[10]</a> Within 20 million years of the first fossils of sporangia and stems towards the close of the Silurian, around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=420">420</a> million years ago, there is evidence that plants were being consumed.<a href="#cite_note-Labandeira2007-11">[11]</a> Animals fed on the spores of early Devonian plants, and the <a href="/wiki/Rhynie_chert" title="Rhynie chert">Rhynie chert</a> provides evidence that organisms fed on plants using a "pierce and suck" technique.<a href="#cite_note-Labandeira1998-9">[9]</a> During the ensuing 75 million years, plants evolved a range of more complex organs – from roots to seeds. There was a gap of 50 to 100 million years between each organ's evolution and its being eaten.<a href="#cite_note-Labandeira2007-11">[11]</a> Hole feeding and skeletonization are recorded in the <a href="/wiki/Cisuralian" title="Cisuralian">early Permian</a>, with surface fluid feeding evolving by the end of that period.<a href="#cite_note-Labandeira1998-9">[9]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Plain_tiger_moat.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Plain_tiger_moat.JPG/220px-Plain_tiger_moat.JPG" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Plain_tiger_moat.JPG/330px-Plain_tiger_moat.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Plain_tiger_moat.JPG/440px-Plain_tiger_moat.JPG 2x" data-file-width="640" data-file-height="480" /></a><figcaption>A plain tiger <a href="/wiki/Danaus_chrysippus" title="Danaus chrysippus">Danaus chrysippus</a> caterpillar making a moat to block defensive chemicals of <a href="/wiki/Calotropis" title="Calotropis">Calotropis</a> before feeding</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Co-evolution">Co-evolution</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=3" title="Edit section: Co-evolution">edit</a>]</div> Herbivores are dependent on plants for food and have evolved mechanisms to obtain this food despite the evolution of a diverse arsenal of plant defenses. <a href="/wiki/Herbivore_adaptations_to_plant_defense" title="Herbivore adaptations to plant defense">Herbivore adaptations to plant defense</a> have been likened to offensive traits and consist of adaptations that allow increased feeding and use of a host plant.<a href="#cite_note-Karban-12">[12]</a> Relationships between herbivores and their host plants often result in reciprocal evolutionary change, called <a href="/wiki/Co-evolution" class="mw-redirect" title="Co-evolution">co-evolution</a>. When an herbivore eats a plant, it <a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">selects</a> for plants that can mount a defensive response. In cases where this relationship demonstrates specificity (the evolution of each trait is due to the other) and reciprocity (both traits must evolve), the species are thought to have co-evolved.<a href="#cite_note-Futuyma-13">[13]</a> The "escape and radiation" mechanism for co-evolution presents the idea that adaptations in herbivores and their host plants have been the driving force behind <a href="/wiki/Speciation" title="Speciation">speciation</a><a href="#cite_note-Ehrlich-5">[5]</a><a href="#cite_note-Thompson-14">[14]</a> and have played a role in the radiation of insect species during the age of <a href="/wiki/Angiosperm" class="mw-redirect" title="Angiosperm">angiosperms</a>.<a href="#cite_note-15">[15]</a> Some herbivores have evolved ways to hijack plant defenses to their own benefit by sequestering these chemicals and using them to protect themselves from predators.<a href="#cite_note-Ehrlich-5">[5]</a> Plant defenses against herbivores are generally not complete, so plants tend to evolve some <a href="/wiki/Plant_tolerance_to_herbivory" title="Plant tolerance to herbivory">tolerance to herbivory</a>.<a href="#cite_note-16">[16]</a> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=4" title="Edit section: Types">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Inducible_plant_defenses_against_herbivory" title="Inducible plant defenses against herbivory">Inducible plant defenses against herbivory</a></div> Plant defenses can be classified as constitutive or induced. Constitutive defenses are always present, while induced defenses are produced or mobilized to the site where a plant is injured. There is wide variation in the composition and concentration of constitutive defenses; these range from mechanical defenses to digestibility reducers and toxins. Many external mechanical defenses and quantitative defenses are constitutive, as they require large amounts of resources to produce and are costly to mobilize.<a href="#cite_note-17">[17]</a> A variety of molecular and biochemical approaches are used to determine the mechanisms of constitutive and induced defensive responses.<a href="#cite_note-18">[18]</a><a href="#cite_note-19">[19]</a><a href="#cite_note-20">[20]</a><a href="#cite_note-21">[21]</a> Induced defenses include secondary metabolites and morphological and physiological changes.<a href="#cite_note-Karban97-22">[22]</a> An advantage of inducible, as opposed to constitutive defenses, is that they are only produced when needed, and are therefore potentially less costly, especially when herbivory is variable.<a href="#cite_note-Karban97-22">[22]</a> Modes of induced defence include <a href="/wiki/Systemic_acquired_resistance" title="Systemic acquired resistance">systemic acquired resistance</a><a href="#cite_note-23">[23]</a> and <a href="/wiki/Plant-induced_systemic_resistance" title="Plant-induced systemic resistance">plant-induced systemic resistance</a>.<a href="#cite_note-24">[24]</a> <div class="mw-heading mw-heading3"><h3 id="Chemical_defenses">Chemical defenses</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=5" title="Edit section: Chemical defenses">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Diospyros_kaki_fruit.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/12/Diospyros_kaki_fruit.jpg/170px-Diospyros_kaki_fruit.jpg" decoding="async" width="170" height="234" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/12/Diospyros_kaki_fruit.jpg/255px-Diospyros_kaki_fruit.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/12/Diospyros_kaki_fruit.jpg/340px-Diospyros_kaki_fruit.jpg 2x" data-file-width="800" data-file-height="1103" /></a><figcaption><a href="/wiki/Persimmon" title="Persimmon">Persimmon</a>, genus <a href="/wiki/Diospyros" title="Diospyros">Diospyros</a>, has a high <a href="/wiki/Tannin" title="Tannin">tannin</a> content which gives immature fruit, seen above, an <a href="/wiki/Astringent" title="Astringent">astringent</a> and <a href="/wiki/Bitter_(taste)" class="mw-redirect" title="Bitter (taste)">bitter</a> <a href="/wiki/Flavor_(taste)" class="mw-redirect" title="Flavor (taste)">flavor</a>.</figcaption></figure> The evolution of chemical defenses in plants is linked to the emergence of chemical substances that are not involved in the essential photosynthetic and metabolic activities. These substances, <a href="/wiki/Secondary_metabolite" title="Secondary metabolite">secondary metabolites</a>, are organic compounds that are not directly involved in the normal growth, development or reproduction of organisms,<a href="#cite_note-Fraenkel-25">[25]</a> and often produced as by-products during the synthesis of primary metabolic products.<a href="#cite_note-Whittaker1970-26">[26]</a> Examples of these byproducts include phenolics, flavonoids, and tannins.<a href="#cite_note-27">[27]</a> Although these secondary metabolites have been thought to play a major role in defenses against herbivores,<a href="#cite_note-Ehrlich-5">[5]</a><a href="#cite_note-Fraenkel-25">[25]</a><a href="#cite_note-28">[28]</a> a meta-analysis of recent relevant studies has suggested that they have either a more minimal (when compared to other non-secondary metabolites, such as primary chemistry and physiology) or more complex involvement in defense.<a href="#cite_note-29">[29]</a> Plants can communicate through the air. Pheromone release and other scents can be detected by leaves and regulate plant immune response. In other words, plants produce volatile organic compounds (VOC) to warn other plants of danger and change their behavioral state to better respond to threats and survival.<a href="#cite_note-:3-30">[30]</a> These warning signals produced by infected neighboring trees allow the undamaged trees to provocatively activate the necessary defense mechanisms. Within the plant itself, it transmits warning, nonvolatile signals as well as airborne signals to surrounding undamaged trees to strengthen their defense/immune system. For instance, poplar and sugar maple trees demonstrated that they received tannins from nearby damaged trees.<a href="#cite_note-:3-30">[30]</a> In sagebrush, damaged plants send out airborne compounds, such as methyl jasmonate, to undamaged plants to increase proteinase inhibitor production and resistance to herbivory.<a href="#cite_note-:3-30">[30]</a> The release of unique VOCs and extrafloral nectar (EFN) allow plants to protect themselves against herbivores by attracting animals from the third trophic level. For example, caterpillar-damaged plants guide parasitic wasps to prey on victims through the release of chemical signals.<a href="#cite_note-:1-31">[31]</a>The sources of these compounds are most likely from glands in the leaves which are ruptured upon the chewing of an herbivore.<a href="#cite_note-:1-31">[31]</a> The injury by herbivores induces the release of linolenic acid and other enzymatic reactions in an octadecanoid cascade, leading to the synthesis of jasmonic acid, a hormone which plays a central role in regulating immune responses. Jasmonic acid induces the release of VOCs and EFN which attract parasitic wasps and predatory mites to detect and feed on herbivores.<a href="#cite_note-32">[32]</a> These volatile organic compounds can also be released to other nearby plants to be prepared for the potential threats. The volatile compounds emitted by plants are easily detected by third trophic level organisms as these signals are unique to herbivore damage.<a href="#cite_note-:1-31">[31]</a> An experiment conducted to measure the VOCs from growing plants shows that signals are released instantaneously upon the herbivory damage and slowly dropped after the damage stopped. It was also observed that plants release the strongest signals during the time of day which animals tend to forage.<a href="#cite_note-:1-31">[31]</a> Since trees are sessile, they have established unique internal defense systems. For instance, when some trees experience herbivory, they release compounds that make their vegetation less palatable. The herbivores saliva left on the leaves of the tree sends a chemical signal to the tree's cells. The tree cells respond by increasing the concentration of salicylic acid (hormone) production.<a href="#cite_note-:2-33">[33]</a> Salicylic acid is a phytohormone that is one of the essential hormones for regulating plants' immune systems.<a href="#cite_note-34">[34]</a> This hormone then signals to increase the production of tree chemicals called tannins within its leaves.<a href="#cite_note-:2-33">[33]</a> <div class="mw-heading mw-heading4"><h4 id="Antiherbivory_compounds">Antiherbivory compounds</h4>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=6" title="Edit section: Antiherbivory compounds">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Secondary_metabolite" title="Secondary metabolite">Secondary metabolite</a>, <a href="/wiki/Antifeedant" title="Antifeedant">Antifeedant</a>, and <a href="/wiki/Phytoalexin" title="Phytoalexin">Phytoalexin</a></div> Plants have evolved many secondary metabolites involved in plant defense, which are collectively known as antiherbivory compounds and can be classified into three sub-groups: <a href="/wiki/Nitrogen" title="Nitrogen">nitrogen</a> compounds (including alkaloids, cyanogenic glycosides, glucosinolates and benzoxazinoids), terpenoids, and phenolics.<a href="#cite_note-35">[35]</a> <a href="/wiki/Alkaloid" title="Alkaloid">Alkaloids</a> are derived from various <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a>. Over 3000 alkaloids are known, including <a href="/wiki/Nicotine" title="Nicotine">nicotine</a>, <a href="/wiki/Caffeine" title="Caffeine">caffeine</a>, <a href="/wiki/Morphine" title="Morphine">morphine</a>, <a href="/wiki/Cocaine" title="Cocaine">cocaine</a>, <a href="/wiki/Colchicine" title="Colchicine">colchicine</a>, <a href="/wiki/Ergoline" title="Ergoline">ergolines</a>, <a href="/wiki/Strychnine" title="Strychnine">strychnine</a>, and <a href="/wiki/Quinine" title="Quinine">quinine</a>.<a href="#cite_note-36">[36]</a> Alkaloids have <a href="/wiki/Pharmacological" class="mw-redirect" title="Pharmacological">pharmacological</a> effects on humans and other animals. Some alkaloids can inhibit or activate <a href="/wiki/Enzyme" title="Enzyme">enzymes</a>, or alter <a href="/wiki/Carbohydrate" title="Carbohydrate">carbohydrate</a> and fat storage by inhibiting the formation <a href="/wiki/Phosphodiester" class="mw-redirect" title="Phosphodiester">phosphodiester</a> bonds involved in their breakdown.<a href="#cite_note-Roberts-37">[37]</a> Certain alkaloids bind to <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acids</a> and can inhibit synthesis of proteins and affect <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> mechanisms. Alkaloids can also affect <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> and <a href="/wiki/Cytoskeletal" class="mw-redirect" title="Cytoskeletal">cytoskeletal</a> structure causing the cells to weaken, collapse, or leak, and can affect <a href="/wiki/Nerve" title="Nerve">nerve</a> transmission.<a href="#cite_note-38">[38]</a> Although alkaloids act on a diversity of metabolic systems in humans and other animals, they almost uniformly invoke an aversively <a href="/wiki/Bitter_(taste)#Bitterness" class="mw-redirect" title="Bitter (taste)">bitter taste</a>.<a href="#cite_note-Rhoades1979-39">[39]</a> <a href="/wiki/Cyanogenic_glycoside" class="mw-redirect" title="Cyanogenic glycoside">Cyanogenic glycosides</a> are stored in inactive forms in plant <a href="/wiki/Vacuole" title="Vacuole">vacuoles</a>. They become toxic when herbivores eat the plant and break cell membranes allowing the glycosides to come into contact with <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> releasing <a href="/wiki/Hydrogen_cyanide" title="Hydrogen cyanide">hydrogen cyanide</a> which blocks cellular respiration.<a href="#cite_note-40">[40]</a> <a href="/wiki/Glucosinolates" class="mw-redirect" title="Glucosinolates">Glucosinolates</a> are activated in much the same way as cyanogenic glucosides, and the products can cause <a href="/wiki/Gastroenteritis" title="Gastroenteritis">gastroenteritis</a>, salivation, diarrhea, and irritation of the mouth.<a href="#cite_note-Rhoades1979-39">[39]</a> <a href="/w/index.php?title=Benzoxazinoids&action=edit&redlink=1" class="new" title="Benzoxazinoids (page does not exist)">Benzoxazinoids</a>, such as <a href="/wiki/DIMBOA" title="DIMBOA">DIMBOA</a>, are secondary defence metabolites characteristic of certain grasses (<a href="/wiki/Poaceae" title="Poaceae">Poaceae</a>). Like cyanogenic glycosides, they are stored as inactive glucosides in the plant vacuole.<a href="#cite_note-41">[41]</a> Upon tissue disruption they get into contact with β-glucosidases from the chloroplasts, which enzymatically release the toxic aglucones. Whereas some benzoxazinoids are constitutively present, others are only synthesized following herbivore infestation, and thus, considered <a href="/wiki/Inducible_plant_defenses_against_herbivory" title="Inducible plant defenses against herbivory">inducible plant defenses against herbivory</a>.<a href="#cite_note-42">[42]</a> The <a href="/wiki/Terpenoids" class="mw-redirect" title="Terpenoids">terpenoids</a>, sometimes referred to as isoprenoids, are organic chemicals similar to <a href="/wiki/Terpene" title="Terpene">terpenes</a>, derived from five-carbon <a href="/wiki/Isoprene" title="Isoprene">isoprene</a> units. There are over 10,000 known types of terpenoids.<a href="#cite_note-43">[43]</a> Most are multicyclic structures which differ from one another in both functional groups, and in basic carbon skeletons.<a href="#cite_note-Terp-44">[44]</a> Monoterpenoids, containing 2 isoprene units, are <a href="/wiki/Volatility_(chemistry)" title="Volatility (chemistry)">volatile</a> <a href="/wiki/Essential_oil" title="Essential oil">essential oils</a> such as <a href="/wiki/Citronella_oil" title="Citronella oil">citronella</a>, <a href="/wiki/Limonene" title="Limonene">limonene</a>, <a href="/wiki/Menthol" title="Menthol">menthol</a>, <a href="/wiki/Camphor" title="Camphor">camphor</a>, and <a href="/wiki/Pinene" title="Pinene">pinene</a>. Diterpenoids, 4 isoprene units, are widely distributed in <a href="/wiki/Latex" title="Latex">latex</a> and <a href="/wiki/Resins" class="mw-redirect" title="Resins">resins</a>, and can be quite toxic. Diterpenes are responsible for making <a href="/wiki/Rhododendron" title="Rhododendron">Rhododendron</a> leaves poisonous. Plant <a href="/wiki/Steroid" title="Steroid">steroids</a> and <a href="/wiki/Sterol" title="Sterol">sterols</a> are also produced from terpenoid precursors, including <a href="/wiki/Vitamin_D" title="Vitamin D">vitamin D</a>, <a href="/wiki/Glycosides" class="mw-redirect" title="Glycosides">glycosides</a> (such as <a href="/wiki/Digitalis" title="Digitalis">digitalis</a>) and <a href="/wiki/Saponin" title="Saponin">saponins</a> (which lyse <a href="/wiki/Red_blood_cell" title="Red blood cell">red blood cells</a> of herbivores).<a href="#cite_note-45">[45]</a> Phenolics, sometimes called phenols, consist of an <a href="/wiki/Aromatic" class="mw-redirect" title="Aromatic">aromatic</a> 6-carbon ring bonded to a <a href="/wiki/Hydroxyl" class="mw-redirect" title="Hydroxyl">hydroxy</a> group. Some phenols have <a href="/wiki/Antiseptic" title="Antiseptic">antiseptic</a> properties, while others disrupt <a href="/wiki/Endocrine" class="mw-redirect" title="Endocrine">endocrine</a> activity. Phenolics range from simple <a href="/wiki/Tannin" title="Tannin">tannins</a> to the more complex <a href="/wiki/Flavonoid" title="Flavonoid">flavonoids</a> that give plants much of their red, blue, yellow, and white pigments. Complex phenolics called <a href="/wiki/Polyphenol" title="Polyphenol">polyphenols</a> are capable of producing many different types of effects on humans, including <a href="/wiki/Polyphenol_antioxidant" class="mw-redirect" title="Polyphenol antioxidant">antioxidant</a> properties. Some examples of phenolics used for defense in plants are: <a href="/wiki/Lignin" title="Lignin">lignin</a>, <a href="/wiki/Silymarin" class="mw-redirect" title="Silymarin">silymarin</a> and <a href="/wiki/Cannabinoid" title="Cannabinoid">cannabinoids</a>.<a href="#cite_note-Phenols-46">[46]</a> <a href="/wiki/Proanthocyanidin" title="Proanthocyanidin">Condensed tannins</a>, polymers composed of 2 to 50 (or more) flavonoid molecules, inhibit herbivore digestion by binding to consumed plant proteins and making them more difficult for animals to digest, and by interfering with protein absorption and <a href="/wiki/Digestive_enzyme" title="Digestive enzyme">digestive enzymes</a>.<a href="#cite_note-47">[47]</a> In addition, some plants use <a href="/wiki/Fatty_acid" title="Fatty acid">fatty acid</a> derivatives, <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> and even <a href="/wiki/Peptide" title="Peptide">peptides</a><a href="#cite_note-48">[48]</a> as defenses. The <a href="/wiki/Cholinergic" title="Cholinergic">cholinergic toxin</a>, <a href="/wiki/Cicutoxin" title="Cicutoxin">cicutoxin</a> of <a href="/wiki/Water_hemlock" class="mw-redirect" title="Water hemlock">water hemlock</a>, is a <a href="/wiki/Polyyne" title="Polyyne">polyyne</a> derived from the fatty acid metabolism.<a href="#cite_note-49">[49]</a> <a href="/wiki/Oxalyldiaminopropionic_acid" title="Oxalyldiaminopropionic acid">Oxalyldiaminopropionic acid</a> is a neurotoxic amino acid produced as a defensive metabolite in the grass pea (<a href="/wiki/Lathyrus_sativus" title="Lathyrus sativus">Lathyrus sativus</a>).<a href="#cite_note-50">[50]</a> The synthesis of <a href="/wiki/Fluoroacetic_acid" title="Fluoroacetic acid">fluoroacetate</a> in several plants is an example of the use of small molecules to disrupt the metabolism of herbivores, in this case the <a href="/wiki/Citric_acid_cycle" title="Citric acid cycle">citric acid cycle</a>.<a href="#cite_note-51">[51]</a> Plants interact by producing <a href="/wiki/Allelochemicals" class="mw-redirect" title="Allelochemicals">allelochemicals</a> which interfere with the growth of other plants (allelopathy). These have a role in plant defense and may be used to suppress competitors such as weeds of crops. A result may be larger plants better able to survive damage by herbivores.<a href="#cite_note-Latif_Chiapusio_2017-52">[52]</a> <div class="mw-heading mw-heading4"><h4 id="Enzymes">Enzymes</h4>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=7" title="Edit section: Enzymes">edit</a>]</div> Premier examples are substances activated by the <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> <a href="/wiki/Myrosinase" title="Myrosinase">myrosinase</a>. This enzyme converts <a href="/wiki/Glucosinolate" title="Glucosinolate">glucosinolates</a> to various compounds that are toxic to <a href="/wiki/Herbivorous" class="mw-redirect" title="Herbivorous">herbivorous</a> insects. One product of this enzyme is <a href="/wiki/Allyl_isothiocyanate" title="Allyl isothiocyanate">allyl isothiocyanate</a>, the pungent ingredient in <a href="/wiki/Horseradish_sauce" class="mw-redirect" title="Horseradish sauce">horseradish sauces</a>. <figure class="mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:Myrosinase_general_mechanism.png" class="mw-file-description"><img alt="mechanism of glucosinolate hydrolysis by myrosinase" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Myrosinase_general_mechanism.png/760px-Myrosinase_general_mechanism.png" decoding="async" width="760" height="159" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Myrosinase_general_mechanism.png/1140px-Myrosinase_general_mechanism.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/38/Myrosinase_general_mechanism.png/1520px-Myrosinase_general_mechanism.png 2x" data-file-width="2791" data-file-height="584" /></a><figcaption>Biosynthesis of antifeedants by the action of myrosinase.</figcaption></figure> The myrosinase is released only upon crushing the flesh of horseradish. Since allyl isothiocyanate is harmful to the plant as well as the insect, it is stored in the harmless form of the glucosinolate, separate from the myrosinase enzyme.<a href="#cite_note-Cole-53">[53]</a> <div class="mw-heading mw-heading3"><h3 id="Mechanical_defenses">Mechanical defenses</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=8" title="Edit section: Mechanical defenses">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Raspberry_cane_-_closeup_in_winter_-_P.2005.03.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ed/Raspberry_cane_-_closeup_in_winter_-_P.2005.03.jpg/220px-Raspberry_cane_-_closeup_in_winter_-_P.2005.03.jpg" decoding="async" width="220" height="177" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ed/Raspberry_cane_-_closeup_in_winter_-_P.2005.03.jpg/330px-Raspberry_cane_-_closeup_in_winter_-_P.2005.03.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/e/ed/Raspberry_cane_-_closeup_in_winter_-_P.2005.03.jpg 2x" data-file-width="437" data-file-height="352" /></a><figcaption>The prickles on the stem of this <a href="/wiki/Raspberry" title="Raspberry">raspberry</a> plant serve as a mechanical defense against herbivory.</figcaption></figure> See the review of mechanical defenses by Lucas et al., 2000, which remains relevant and well regarded in the subject as of 2018<a class="external text" href="https://en.wikipedia.org/w/index.php?title=Plant_defense_against_herbivory&action=edit">[update]</a>.<a href="#cite_note-Lucas-2000-54">[54]</a> Many plants have external structural defenses that discourage herbivory. Structural defenses can be described as morphological or physical traits that give the plant a fitness advantage by deterring herbivores from feeding.<a href="#cite_note-:0-55">[55]</a> Depending on the herbivore's physical characteristics (i.e. size and defensive armor), plant structural defenses on stems and leaves can deter, injure, or kill the grazer. Some defensive compounds are produced internally but are released onto the plant's surface; for example, <a href="/wiki/Resins" class="mw-redirect" title="Resins">resins</a>, <a href="/wiki/Lignin" title="Lignin">lignins</a>, <a href="/wiki/Silica" class="mw-redirect" title="Silica">silica</a>, and wax cover the <a href="/wiki/Epidermis_(botany)" title="Epidermis (botany)">epidermis</a> of <a href="/wiki/Terrestrial_plant" title="Terrestrial plant">terrestrial plants</a> and alter the texture of the plant tissue. The leaves of <a href="/wiki/Holly" title="Holly">holly</a> plants, for instance, are very smooth and slippery making feeding difficult. Some plants produce <a href="/wiki/Gummosis" title="Gummosis">gummosis</a> or sap that traps insects.<a href="#cite_note-56">[56]</a> <div class="mw-heading mw-heading4"><h4 id="Spines_and_thorns">Spines and thorns</h4>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=9" title="Edit section: Spines and thorns">edit</a>]</div> A plant's leaves and stem may be covered with sharp prickles, spines, thorns or <a href="/wiki/Trichomes" class="mw-redirect" title="Trichomes">trichomes</a>- hairs on the leaf often with barbs, sometimes containing <a href="/wiki/Irritant_(biology)" class="mw-redirect" title="Irritant (biology)">irritants</a> or poisons. Plant structural features like spines, thorns and <a href="/wiki/Awn_(botany)" title="Awn (botany)"> awns</a> reduce feeding by large ungulate herbivores (e.g. <a href="/wiki/Kudu" title="Kudu">kudu</a>, <a href="/wiki/Impala" title="Impala">impala</a>, and <a href="/wiki/Capra_(genus)" title="Capra (genus)">goats</a>) by restricting the herbivores' feeding rate, or by wearing down the molars.<a href="#cite_note-57">[57]</a> Trichomes are frequently associated with lower rates of plant tissue digestion by insect herbivores.<a href="#cite_note-:0-55">[55]</a> <a href="/wiki/Raphides" class="mw-redirect" title="Raphides">Raphides</a> are sharp needles of <a href="/wiki/Calcium_oxalate" title="Calcium oxalate">calcium oxalate</a> or <a href="/wiki/Calcium_carbonate" title="Calcium carbonate">calcium carbonate</a> in plant tissues, making ingestion painful, damaging a herbivore's mouth and gullet and causing more efficient delivery of the plant's toxins. The structure of a plant, its branching and leaf arrangement may also be evolved to reduce herbivore impact. The shrubs of New Zealand have evolved special wide branching adaptations believed to be a response to browsing birds such as the <a href="/wiki/Moa" title="Moa">moas</a>.<a href="#cite_note-58">[58]</a> Similarly, African <a href="/wiki/Acacia" title="Acacia">Acacia</a> trees have long spines low in the canopy, but very short spines high in the canopy, which is comparatively safe from herbivores such as giraffes.<a href="#cite_note-59">[59]</a><a href="#cite_note-60">[60]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cocos_nucifera_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-187.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Cocos_nucifera_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-187.jpg/170px-Cocos_nucifera_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-187.jpg" decoding="async" width="170" height="254" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Cocos_nucifera_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-187.jpg/255px-Cocos_nucifera_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-187.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/32/Cocos_nucifera_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-187.jpg/340px-Cocos_nucifera_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-187.jpg 2x" data-file-width="1578" data-file-height="2362" /></a><figcaption><a href="/wiki/Coconut_palm" class="mw-redirect" title="Coconut palm">Coconut palms</a> protect their fruit by surrounding it with multiple layers of armor.</figcaption></figure> Trees such as palms protect their fruit by multiple layers of armor, needing efficient tools to break through to the seed contents. Some plants, notably the <a href="/wiki/Grass" class="mw-redirect" title="Grass">grasses</a>, use indigestible <a href="/wiki/Silica" class="mw-redirect" title="Silica">silica</a> (and many plants use other relatively indigestible materials such as <a href="/wiki/Lignin" title="Lignin">lignin</a>) to defend themselves against vertebrate and invertebrate herbivores.<a href="#cite_note-61">[61]</a> Plants take up <a href="/wiki/Silicon" title="Silicon">silicon</a> from the soil and deposit it in their tissues in the form of solid silica <a href="/wiki/Phytolith" title="Phytolith">phytoliths</a>. These mechanically reduce the digestibility of plant tissue, causing rapid wear to vertebrate teeth and to insect mandibles,<a href="#cite_note-62">[62]</a> and are effective against herbivores above and below ground.<a href="#cite_note-63">[63]</a> The mechanism may offer future sustainable pest-control strategies.<a href="#cite_note-64">[64]</a> <div class="mw-heading mw-heading4"><h4 id="Thigmonastic_movements">Thigmonastic movements</h4>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=10" title="Edit section: Thigmonastic movements">edit</a>]</div> <a href="/wiki/Thigmonasty" title="Thigmonasty">Thigmonastic movements</a>, those that occur in response to touch, are used as a defense in some plants. The leaves of the <a href="/wiki/Sensitive_plant" class="mw-redirect" title="Sensitive plant">sensitive plant</a>, Mimosa pudica, close up rapidly in response to direct touch, vibration, or even electrical and thermal stimuli. The <a href="/wiki/Proximate_cause" title="Proximate cause">proximate cause</a> of this mechanical response is an abrupt change in the <a href="/wiki/Turgor" class="mw-redirect" title="Turgor">turgor</a> pressure in the <a href="/wiki/Pulvini" class="mw-redirect" title="Pulvini">pulvini</a> at the base of leaves resulting from <a href="/wiki/Osmosis" title="Osmosis">osmotic</a> phenomena. This is then spread via both electrical and chemical means through the plant; only a single leaflet need be disturbed. This response lowers the surface area available to herbivores, which are presented with the underside of each leaflet, and results in a wilted appearance. It may also physically dislodge small herbivores, such as insects.<a href="#cite_note-Raven_2005-65">[65]</a> <div class="mw-heading mw-heading3"><h3 id="Carnivorous_plants">Carnivorous plants</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=11" title="Edit section: Carnivorous plants">edit</a>]</div> <a href="/wiki/Carnivorous_plant" title="Carnivorous plant">Carnivory in plants</a> has evolved at least six times independently, some examples include the <a href="/wiki/Venus_flytrap" title="Venus flytrap">Venus flytrap</a>, <a href="/wiki/Pitcher_plant" title="Pitcher plant">pitcher plant</a>, and <a href="/wiki/Butterwort" class="mw-redirect" title="Butterwort">butterwort</a>.<a href="#cite_note-66">[66]</a> Many of these plants have evolved in poor nutrient soil. To get sufficient nutrients in these conditions they must use an alternative method. They use insects and small birds as a way to gain the minerals they need through carnivory. Carnivorous plants do not use carnivory as self-defense, but to get the nutrients they need.<a href="#cite_note-67">[67]</a> <div class="mw-heading mw-heading3"><h3 id="Mimicry_and_camouflage">Mimicry and camouflage</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=12" title="Edit section: Mimicry and camouflage">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Mimicry_in_plants" title="Mimicry in plants">Mimicry in plants</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Lithops_salicola_GotBot_2015_001.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Lithops_salicola_GotBot_2015_001.jpg/220px-Lithops_salicola_GotBot_2015_001.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Lithops_salicola_GotBot_2015_001.jpg/330px-Lithops_salicola_GotBot_2015_001.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Lithops_salicola_GotBot_2015_001.jpg/440px-Lithops_salicola_GotBot_2015_001.jpg 2x" data-file-width="5496" data-file-height="3670" /></a><figcaption>The cryptic <a href="/wiki/Lithops_salicola" title="Lithops salicola">Lithops salicola</a>, a pebble plant, <a href="/wiki/Camouflage" title="Camouflage">camouflaged</a> as small stones, reducing the chance that it will be seen by herbivores</figcaption></figure> Some plants make use of various forms of <a href="/wiki/Mimicry_in_plants" title="Mimicry in plants">mimicry</a> to reduce herbivory. One mechanism is to mimic the presence of insect eggs on their leaves, dissuading insect species from laying their eggs there. Because female butterflies are less likely to lay their eggs on plants that already have butterfly eggs, some species of <a href="/wiki/Neotropical" class="mw-redirect" title="Neotropical">neotropical</a> <a href="/wiki/Vine" title="Vine">vines</a> of the <a href="/wiki/Genus" title="Genus">genus</a> <a href="/wiki/Passiflora" title="Passiflora">Passiflora</a> (passion flowers) make use of <a href="/wiki/Gilbertian_mimicry" title="Gilbertian mimicry">Gilbertian mimicry</a>: they possess physical structures resembling the yellow eggs of <a href="/wiki/Heliconius" title="Heliconius">Heliconius</a> <a href="/wiki/Butterflies" class="mw-redirect" title="Butterflies">butterflies</a> on their leaves, which discourage <a href="/wiki/Oviposition" class="mw-redirect" title="Oviposition">oviposition</a> by butterflies.<a href="#cite_note-68">[68]</a> Other plants make use of <a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian mimicry</a>, with structures that imitate thorns or other objects to dissuade herbivores directly.<a href="#cite_note-69">[69]</a> A further approach is <a href="/wiki/Camouflage" title="Camouflage">camouflage</a>; the vine <a href="/wiki/Boquila_trifoliolata" class="mw-redirect" title="Boquila trifoliolata">Boquila trifoliolata</a> mimics the leaves of its host plant,<a href="#cite_note-70">[70]</a> while the pebble plant <a href="/wiki/Lithops" title="Lithops">Lithops</a> makes itself hard to spot among the stones of the Southern African environment.<a href="#cite_note-71">[71]</a> <div class="mw-heading mw-heading3"><h3 id="Indirect_defenses">Indirect defenses</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=13" title="Edit section: Indirect defenses">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Acacia-collinsii.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Acacia-collinsii.jpg/170px-Acacia-collinsii.jpg" decoding="async" width="170" height="255" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Acacia-collinsii.jpg/255px-Acacia-collinsii.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/00/Acacia-collinsii.jpg/340px-Acacia-collinsii.jpg 2x" data-file-width="533" data-file-height="800" /></a><figcaption>The large and directly defensive thorn-like stipules of <a href="/wiki/Vachellia_collinsii" title="Vachellia collinsii">Vachellia collinsii</a> are also hollow and offer shelter for ants, which indirectly protect the plant against herbivores.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></div> Another category of plant defenses are those features that indirectly protect the plant by enhancing the probability of attracting the <a href="/wiki/Natural_enemy" class="mw-redirect" title="Natural enemy">natural enemies</a> of herbivores. Such an arrangement is known as <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutualism</a>, in this case of the "<a href="/wiki/The_enemy_of_my_enemy_is_my_friend" title="The enemy of my enemy is my friend">enemy of my enemy</a>" variety. One such feature are <a href="/wiki/Semiochemical" title="Semiochemical">semiochemicals</a>, given off by plants. Semiochemicals are a group of <a href="/wiki/Volatile_organic_compounds" class="mw-redirect" title="Volatile organic compounds">volatile organic compounds</a> involved in interactions between organisms. One group of semiochemicals are <a href="/wiki/Heterotelergones" class="mw-redirect" title="Heterotelergones">allelochemicals</a>; consisting of <a href="/wiki/Allomone" title="Allomone">allomones</a>, which play a defensive role in <a href="/wiki/Interspecies_communication" title="Interspecies communication">interspecies communication</a>, and <a href="/wiki/Kairomone" title="Kairomone">kairomones</a>, which are used by members of higher <a href="/wiki/Trophic_level" title="Trophic level">trophic levels</a> to locate food sources. When a plant is attacked it releases allelochemics containing an abnormal ratio of these <style data-mw-deduplicate="TemplateStyles:r1238216509">.mw-parser-output .vanchor>:target~.vanchor-text{background-color:#b1d2ff}@media screen{html.skin-theme-clientpref-night .mw-parser-output .vanchor>:target~.vanchor-text{background-color:#0f4dc9}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .vanchor>:target~.vanchor-text{background-color:#0f4dc9}}</style>herbivore-induced plant volatiles (HIPVs).<a href="#cite_note-Dicke-72">[72]</a><a href="#cite_note-73">[73]</a> Predators sense these volatiles as food cues, attracting them to the damaged plant, and to feeding herbivores. The subsequent reduction in the number of herbivores confers a <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> benefit to the plant and demonstrates the indirect defensive capabilities of semiochemicals.<a href="#cite_note-Schuman2012-74">[74]</a> Induced volatiles also have drawbacks, however; some studies have suggested that these volatiles attract herbivores.<a href="#cite_note-Dicke-72">[72]</a> <a href="/wiki/Crop_domestication" class="mw-redirect" title="Crop domestication">Crop domestication</a> has increased <a href="/wiki/Crop_yield" title="Crop yield">yield</a> sometimes at the expense of HIPV production. Orre Gordon et al 2013 tests several methods of artificially restoring the plant-predator partnership, by combining <a href="/wiki/Companion_planting" title="Companion planting">companion planting</a> and synthetic predator <a href="/wiki/Attractant" title="Attractant">attractants</a>. They describe several strategies which work and several which do not.<a href="#cite_note-Tooker-et-al-2020-75">[75]</a> Plants sometimes provide housing and food items for natural enemies of herbivores, known as "biotic" defense mechanisms, to maintain their presence. For example, trees from the genus <a href="/wiki/Macaranga" title="Macaranga">Macaranga</a> have adapted their thin stem walls to create ideal housing for ants (genus <a href="/wiki/Crematogaster" title="Crematogaster">Crematogaster</a>), which, in turn, protects the plant from herbivores.<a href="#cite_note-Heil-76">[76]</a> In addition to providing housing, the plant also provides the ant with its exclusive food source; from the food bodies produced by the plant. Similarly, several <a href="/wiki/Acacia" title="Acacia">Acacia</a> tree species have developed stipular spines (direct defenses) that are swollen at the base, forming a hollow structure that provides housing for protective ants. These Acacia trees also produce <a href="/wiki/Nectar" title="Nectar">nectar</a> in <a href="/wiki/Extrafloral_nectaries" class="mw-redirect" title="Extrafloral nectaries">extrafloral nectaries</a> on their leaves as food for the ants.<a href="#cite_note-77">[77]</a> <a href="/wiki/Plant_use_of_endophytic_fungi_in_defense" title="Plant use of endophytic fungi in defense">Plant use of endophytic fungi in defense</a> is common. Most plants have <a href="/wiki/Endophyte" title="Endophyte">endophytes</a>, microbial organisms that live within them. While some cause disease, others protect plants from herbivores and <a href="/wiki/Pathogen" title="Pathogen">pathogenic</a> microbes.<a href="#cite_note-78">[78]</a> Endophytes can help the plant by producing toxins harmful to other organisms that would attack the plant, such as alkaloid producing <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> which are common in grasses such as <a href="/wiki/Tall_fescue" class="mw-redirect" title="Tall fescue">tall fescue</a> (Festuca arundinacea), which is infected by Neotyphodium coenophialum.<a href="#cite_note-Raven_2005-65">[65]</a><a href="#cite_note-79">[79]</a> Trees of the same species form alliances with other tree species to improve their survival rate. They communicate and have dependent relationships through connections below the soil called underground mycorrhiza networks, which allows them to share water/nutrients and various signals for predatory attacks while also protecting its immune system.<a href="#cite_note-:4-80">[80]</a> Within a forest of trees, the ones getting attacked send communication distress signals that alerts neighboring trees to alter their behavior (defense).<a href="#cite_note-:4-80">[80]</a> The tree and fungi relationship is a symbiotic relationship. <a href="#cite_note-:4-80">[80]</a> Fungi, intertwined with the trees' roots, support communication between trees to locate nutrients. In return, the fungi receive some of the sugar that trees photosynthesize. Trees send out several forms of communication including chemical, hormonal, and slow pulsing electric signals. Farmers investigated the electrical signals between trees, using a voltage-based signal system, similar to an animal's nervous system, where a tree faces distress and releases a warning signal to surrounding trees.<a href="#cite_note-:4-80">[80]</a> <div class="mw-heading mw-heading3"><h3 id="Leaf_shedding_and_color">Leaf shedding and color</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=14" title="Edit section: Leaf shedding and color">edit</a>]</div> There have been suggestions that <a href="/wiki/Abscission" title="Abscission">leaf shedding</a> may be a response that provides protection against diseases and certain kinds of pests such as <a href="/wiki/Leaf_miner" title="Leaf miner">leaf miners</a> and <a href="/wiki/Gall" title="Gall">gall</a> forming insects.<a href="#cite_note-81">[81]</a> Other responses such as the change of leaf <a href="/wiki/Biological_pigment" title="Biological pigment">colors</a> prior to fall have also been suggested as adaptations that may help undermine the camouflage of herbivores.<a href="#cite_note-82">[82]</a> <a href="/wiki/Autumn_leaf_color" title="Autumn leaf color">Autumn leaf color</a> has also been suggested to act as an <a href="/wiki/Handicap_principle" title="Handicap principle">honest warning signal</a> of defensive commitment towards insect pests that migrate to the trees in autumn.<a href="#cite_note-83">[83]</a><a href="#cite_note-84">[84]</a> <div class="mw-heading mw-heading2"><h2 id="Costs_and_benefits">Costs and benefits</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=15" title="Edit section: Costs and benefits">edit</a>]</div> Defensive structures and chemicals are costly as they require resources that could otherwise be used by plants to maximize growth and reproduction. In some situations, plant growth slows down when most of the nutrients are being used for the generation of toxins or regeneration of plant parts.<a href="#cite_note-85">[85]</a> Many models have been proposed to explore how and why some plants make this investment in defenses against herbivores.<a href="#cite_note-Wittstock-et-al-2002-8">[8]</a> <div class="mw-heading mw-heading3"><h3 id="Optimal_defense_hypothesis">Optimal defense hypothesis</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=16" title="Edit section: Optimal defense hypothesis">edit</a>]</div> The optimal defense hypothesis attempts to explain how the kinds of defenses a particular plant might use reflect the threats each individual plant faces.<a href="#cite_note-Stamp-86">[86]</a> This model considers three main factors, namely: risk of attack, value of the plant part, and the cost of defense.<a href="#cite_note-Rhoades-87">[87]</a><a href="#cite_note-88">[88]</a> The first factor determining optimal defense is risk: how likely is it that a plant or certain plant parts will be attacked? This is also related to the plant apparency hypothesis, which states that a plant will invest heavily in broadly effective defenses when the plant is easily found by herbivores.<a href="#cite_note-Feeny-89">[89]</a> Examples of apparent plants that produce generalized protections include long-living trees, shrubs, and perennial grasses.<a href="#cite_note-Feeny-89">[89]</a> Unapparent plants, such as short-lived plants of early <a href="/wiki/Ecological_succession" title="Ecological succession">successional</a> stages, on the other hand, preferentially invest in small amounts of qualitative toxins that are effective against all but the most specialized herbivores.<a href="#cite_note-Feeny-89">[89]</a> The second factor is the value of protection: would the plant be less able to survive and reproduce after removal of part of its structure by a herbivore? Not all plant parts are of equal evolutionary value, thus valuable parts contain more defenses. A plant's stage of development at the time of feeding also affects the resulting change in fitness. Experimentally, the fitness value of a plant structure is determined by removing that part of the plant and observing the effect.<a href="#cite_note-McKey1-90">[90]</a> In general, <a href="/wiki/Biological_reproduction" class="mw-redirect" title="Biological reproduction">reproductive</a> parts are not as easily replaced as <a href="/wiki/Vegetation" title="Vegetation">vegetative</a> parts, terminal leaves have greater value than <a href="/wiki/Anatomical_terms_of_location" title="Anatomical terms of location">basal</a> leaves, and the loss of plant parts mid-season has a greater negative effect on fitness than removal at the beginning or end of the season.<a href="#cite_note-91">[91]</a><a href="#cite_note-92">[92]</a> Seeds in particular tend to be very well protected. For example, the seeds of many edible fruits and nuts contain cyanogenic glycosides such as <a href="/wiki/Amygdalin" title="Amygdalin">amygdalin</a>. This results from the need to balance the effort needed to make the fruit attractive to animal dispersers while ensuring that the seeds are not destroyed by the animal.<a href="#cite_note-93">[93]</a><a href="#cite_note-94">[94]</a> The final consideration is cost: how much will a particular defensive strategy cost a plant in energy and materials? This is particularly important, as energy spent on defense cannot be used for other functions, such as reproduction and growth. The optimal defense hypothesis predicts that plants will allocate more energy towards defense when the benefits of protection outweigh the costs, specifically in situations where there is high herbivore pressure.<a href="#cite_note-95">[95]</a><a href="#cite_note-96">[96]</a> <div class="mw-heading mw-heading3"><h3 id="Carbon:nutrient_balance_hypothesis">Carbon:nutrient balance hypothesis</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=17" title="Edit section: Carbon:nutrient balance hypothesis">edit</a>]</div> The carbon:nutrient balance hypothesis, also known as the environmental constraint hypothesis or Carbon Nutrient Balance Model (CNBM), states that the various types of plant defenses are responses to variations in the levels of <a href="/wiki/Nutrient" title="Nutrient">nutrients</a> in the environment. This hypothesis predicts the Carbon/Nitrogen ratio in plants determines which secondary metabolites will be synthesized. For example, plants growing in <a href="/wiki/Nitrogen" title="Nitrogen">nitrogen</a>-poor <a href="/wiki/Soil" title="Soil">soils</a> will use <a href="/wiki/Carbon" title="Carbon">carbon</a>-based defenses (mostly digestibility reducers), while those growing in low-carbon environments (such as shady conditions) are more likely to produce nitrogen-based toxins. The hypothesis further predicts that plants can change their defenses in response to changes in nutrients. For example, if plants are grown in low-nitrogen conditions, then these plants will implement a defensive strategy composed of constitutive carbon-based defenses. If nutrient levels subsequently increase, by for example the addition of <a href="/wiki/Fertilizer" title="Fertilizer">fertilizers</a>, these carbon-based defenses will decrease.<a href="#cite_note-Bryant-97">[97]</a><a href="#cite_note-98">[98]</a> <div class="mw-heading mw-heading3"><h3 id="Growth_rate_hypothesis">Growth rate hypothesis</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=18" title="Edit section: Growth rate hypothesis">edit</a>]</div> The growth rate hypothesis, also known as the resource availability hypothesis, states that defense strategies are determined by the inherent growth rate of the plant, which is in turn determined by the resources available to the plant. A major assumption is that available resources are the <a href="/wiki/Limiting_factor" title="Limiting factor">limiting factor</a> in determining the maximum growth rate of a plant species. This model predicts that the level of defense investment will increase as the potential of growth decreases.<a href="#cite_note-Coley-99">[99]</a> Additionally, plants in resource-poor areas, with inherently slow-growth rates, tend to have long-lived leaves and twigs, and the loss of plant appendages may result in a loss of scarce and valuable nutrients.<a href="#cite_note-100">[100]</a> One test of this model involved a reciprocal transplants of seedlings of 20 species of trees between <a href="/wiki/Clay" title="Clay">clay</a> <a href="/wiki/Soil" title="Soil">soils</a> (nutrient rich) and white <a href="/wiki/Sand" title="Sand">sand</a> (nutrient poor) to determine whether trade-offs between growth rate and defenses restrict species to one habitat. When planted in white sand and protected from herbivores, seedlings originating from clay outgrew those originating from the nutrient-poor sand, but in the presence of herbivores the seedlings originating from white sand performed better, likely due to their higher levels of constitutive carbon-based defenses. These finding suggest that defensive strategies limit the habitats of some plants.<a href="#cite_note-101">[101]</a> <div class="mw-heading mw-heading3"><h3 id="Growth-differentiation_balance_hypothesis">Growth-differentiation balance hypothesis</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=19" title="Edit section: Growth-differentiation balance hypothesis">edit</a>]</div> The growth-differentiation balance hypothesis states that plant defenses are a result of a tradeoff between "growth-related processes" and "differentiation-related processes" in different environments.<a href="#cite_note-Loomis-102">[102]</a> <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">Differentiation</a>-related processes are defined as "processes that enhance the structure or function of existing cells (i.e. maturation and specialization)."<a href="#cite_note-Stamp-86">[86]</a> A plant will produce chemical defenses only when energy is available from <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>, and plants with the highest concentrations of secondary metabolites are the ones with an intermediate level of available resources.<a href="#cite_note-Loomis-102">[102]</a> <div class="mw-heading mw-heading3"><h3 id="Synthesis_tradeoffs">Synthesis tradeoffs</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=20" title="Edit section: Synthesis tradeoffs">edit</a>]</div> The vast majority of plant resistances to herbivores are either unrelated to each other, or are positively correlated. However there are some negative correlations: In <a href="/wiki/Pastinaca_sativa" class="mw-redirect" title="Pastinaca sativa">Pastinaca sativa</a>'s resistances to various biotypes of <a href="/wiki/Depressaria_pastinacella" class="mw-redirect" title="Depressaria pastinacella">Depressaria pastinacella</a>, because the <a href="/wiki/Secondary_metabolites" class="mw-redirect" title="Secondary metabolites">secondary metabolites</a> involved are negatively correlated with each other; and in the resistances of <a href="/wiki/Diplacus_aurantiacus" title="Diplacus aurantiacus">Diplacus aurantiacus</a>.<a href="#cite_note-Strauss-Irwin-2004-103">[103]</a> In <a href="/wiki/Brassica_rapa" title="Brassica rapa">Brassica rapa</a>, resistance to <a href="/wiki/Peronospora_parasitica" class="mw-redirect" title="Peronospora parasitica">Peronospora parasitica</a> and growth rate are negatively correlated.<a href="#cite_note-Strauss-Irwin-2004-103">[103]</a> <div class="mw-heading mw-heading3"><h3 id="Mutualism_and_overcompensation_of_plants">Mutualism and overcompensation of plants</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=21" title="Edit section: Mutualism and overcompensation of plants">edit</a>]</div> Many plants do not have secondary metabolites, chemical processes, or mechanical defenses to help them fend off herbivores.<a href="#cite_note-104">[104]</a> Instead, these plants rely on overcompensation (which is regarded as a form of mutualism) when they are attacked by herbivores. Overcompensation is defined as having higher fitness when attacked by a herbivore. This a mutual relationship; the herbivore is satisfied with a meal, while the plant starts growing the missing part quickly. These plants have a higher chance of reproducing, and their fitness is increased.<a href="#cite_note-105">[105]</a><a href="#cite_note-106">[106]</a> <div class="mw-heading mw-heading2"><h2 id="Importance_to_humans">Importance to humans</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=22" title="Edit section: Importance to humans">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Agriculture">Agriculture</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=23" title="Edit section: Agriculture">edit</a>]</div> <a href="/wiki/Crop" title="Crop">Crop</a> plants can be bred for their ability to resist herbivory, thus protecting themselves from damage with reduced use of <a href="/wiki/Pesticide" title="Pesticide">pesticides</a>.<a href="#cite_note-Mitchell_Brennan_2016-107">[107]</a> In addition, <a href="/wiki/Biological_pest_control" title="Biological pest control">biological pest control</a> sometimes makes use of plant defenses to reduce crop damage by herbivores. Techniques include <a href="/wiki/Polyculture" title="Polyculture">polyculture</a>, the planting together of two or more species such as a primary crop and a secondary plant. This can allow the secondary plant's defensive chemicals to protect the crop planted with it.<a href="#cite_note-Parolin_Bresch_2012-108">[108]</a> The variation of plant susceptibility to pests was probably known even in the early stages of agriculture in humans. In historic times, the observation of such variations in susceptibility have provided solutions for major <a href="/wiki/Socio-economic" class="mw-redirect" title="Socio-economic">socio-economic</a> problems. The <a href="/wiki/Hemiptera" title="Hemiptera">hemipteran</a> pest insect <a href="/wiki/Phylloxera" title="Phylloxera">phylloxera</a> was introduced from North America to France in 1860 and in 25 years it destroyed nearly a third (100,000 km2) of French <a href="/wiki/Vineyard" title="Vineyard">vineyards</a>. <a href="/wiki/Charles_Valentine_Riley" title="Charles Valentine Riley">Charles Valentine Riley</a> noted that the American species <a href="/wiki/Vitis_labrusca" title="Vitis labrusca">Vitis labrusca</a> was resistant to Phylloxera. Riley, with J. E. Planchon, helped save the <a href="/wiki/French_wine" title="French wine">French wine</a> industry by suggesting the <a href="/wiki/Grafting" title="Grafting">grafting</a> of the susceptible but high quality grapes onto Vitis labrusca root stocks.<a href="#cite_note-Rutgers-109">[109]</a> The formal study of plant resistance to herbivory was first covered extensively in 1951 by <a href="/wiki/Reginald_Henry_Painter" title="Reginald Henry Painter">Reginald Henry Painter</a>, who is widely regarded as the founder of this area of research, in his book Plant Resistance to Insects.<a href="#cite_note-Painter-110">[110]</a> While this work pioneered further research in the US, the work of Chesnokov was the basis of further research in the USSR.<a href="#cite_note-111">[111]</a> Fresh growth of grass is sometimes high in <a href="/wiki/Prussic_acid" class="mw-redirect" title="Prussic acid">prussic acid</a> content and can cause poisoning of grazing livestock. The production of <a href="/wiki/Cyanogenic" class="mw-redirect" title="Cyanogenic">cyanogenic</a> chemicals in grasses is primarily a defense against herbivores.<a href="#cite_note-112">[112]</a><a href="#cite_note-113">[113]</a> The human innovation of cooking may have been particularly helpful in overcoming many of the defensive chemicals of plants. Many <a href="/wiki/Enzyme_inhibitor" title="Enzyme inhibitor">enzyme inhibitors</a> in <a href="/wiki/Cereal_grain" class="mw-redirect" title="Cereal grain">cereal grains</a> and <a href="/wiki/Pulse_(legume)" class="mw-redirect" title="Pulse (legume)">pulses</a>, such as <a href="/wiki/Trypsin" title="Trypsin">trypsin</a> inhibitors prevalent in pulse crops, are denatured by cooking, making them digestible.<a href="#cite_note-114">[114]</a><a href="#cite_note-115">[115]</a> It has been known since the late 17th century that plants contain <a href="https://en.wiktionary.org/wiki/noxious" class="extiw" title="wiktionary:noxious">noxious</a> chemicals which are avoided by insects. These chemicals have been used by man as early insecticides; in 1690 nicotine was extracted from tobacco and used as a contact insecticide. In 1773, insect infested plants were treated with nicotine <a href="/wiki/Fumigation" title="Fumigation">fumigation</a> by heating tobacco and blowing the smoke over the plants.<a href="#cite_note-116">[116]</a> The flowers of <a href="/wiki/Chrysanthemum" title="Chrysanthemum">Chrysanthemum</a> species contain <a href="/wiki/Pyrethrin" title="Pyrethrin">pyrethrin</a> which is a potent insecticide. In later years, the applications of plant resistance became an important area of research in <a href="/wiki/Agriculture" title="Agriculture">agriculture</a> and <a href="/wiki/Plant_breeding" title="Plant breeding">plant breeding</a>, particularly because they can serve as a safe and low-cost alternative to the use of <a href="/wiki/Pesticide" title="Pesticide">pesticides</a>.<a href="#cite_note-smith-117">[117]</a> The important role of secondary plant substances in plant defense was described in the late 1950s by <a href="/wiki/Vincent_Dethier" title="Vincent Dethier">Vincent Dethier</a> and G.S. Fraenkel.<a href="#cite_note-Fraenkel-25">[25]</a><a href="#cite_note-118">[118]</a> The use of botanical pesticides is widespread, including azadirachtin from the <a href="/wiki/Neem" class="mw-redirect" title="Neem">neem</a> (Azadirachta indica), d-Limonene from <a href="/wiki/Citrus" title="Citrus">Citrus</a> species, rotenone from <a href="/wiki/Derris" title="Derris">Derris</a>, capsaicin from <a href="/wiki/Chili_pepper" title="Chili pepper">chili pepper</a>, and pyrethrum from <a href="/wiki/Chrysanthemum" title="Chrysanthemum">Chrysanthemum</a>.<a href="#cite_note-119">[119]</a> The selective breeding of crop plants often involves selection against the plant's intrinsic resistance strategies. This makes crop plant varieties particularly susceptible to pests unlike their wild relatives. In breeding for host-plant resistance, it is often the wild relatives that provide the source of resistance <a href="/wiki/Gene" title="Gene">genes</a>. These genes are incorporated using conventional approaches to plant breeding, but have been augmented by <a href="/wiki/Recombinant_DNA" title="Recombinant DNA">recombinant</a> techniques, which allow introduction of genes from completely unrelated organisms. The most famous <a href="/wiki/Transgenic" class="mw-redirect" title="Transgenic">transgenic</a> approach is the introduction of genes from the bacterial species, <a href="/wiki/Bacillus_thuringiensis" title="Bacillus thuringiensis">Bacillus thuringiensis</a>, into plants. The bacterium produces proteins that, when ingested, kill <a href="/wiki/Lepidoptera" title="Lepidoptera">lepidopteran</a> <a href="/wiki/Caterpillar" title="Caterpillar">caterpillars</a>. The gene encoding for these highly toxic proteins, when introduced into the host plant genome, confers resistance against caterpillars, when the same toxic proteins are produced within the plant. This approach is controversial, however, due to the possibility of <a href="/wiki/Ecological" class="mw-redirect" title="Ecological">ecological</a> and <a href="/wiki/Toxicological" class="mw-redirect" title="Toxicological">toxicological</a> side effects.<a href="#cite_note-120">[120]</a> <div class="mw-heading mw-heading3"><h3 id="Pharmaceutical">Pharmaceutical</h3>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=24" title="Edit section: Pharmaceutical">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Mandragora_Tacuinum_Sanitatis.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Mandragora_Tacuinum_Sanitatis.jpg/170px-Mandragora_Tacuinum_Sanitatis.jpg" decoding="async" width="170" height="187" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Mandragora_Tacuinum_Sanitatis.jpg/255px-Mandragora_Tacuinum_Sanitatis.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Mandragora_Tacuinum_Sanitatis.jpg/340px-Mandragora_Tacuinum_Sanitatis.jpg 2x" data-file-width="458" data-file-height="505" /></a><figcaption>Illustration from the 15th-century manuscript <a href="/wiki/Tacuinum_Sanitatis" title="Tacuinum Sanitatis">Tacuinum Sanitatis</a> detailing the beneficial and harmful properties of <a href="/wiki/Mandragora_(genus)" title="Mandragora (genus)">Mandrakes</a></figcaption></figure> Many currently available <a href="/wiki/Pharmaceutical" class="mw-redirect" title="Pharmaceutical">pharmaceuticals</a> are derived from the secondary metabolites plants use to protect themselves from herbivores, including <a href="/wiki/Opium" title="Opium">opium</a>, <a href="/wiki/Aspirin" title="Aspirin">aspirin</a>, <a href="/wiki/Cocaine" title="Cocaine">cocaine</a>, and <a href="/wiki/Atropine" title="Atropine">atropine</a>.<a href="#cite_note-pmid11395950-121">[121]</a> These chemicals have evolved to affect the biochemistry of insects in very specific ways. However, many of these biochemical pathways are conserved in vertebrates, including humans, and the chemicals act on human biochemistry in ways similar to that of insects. It has therefore been suggested that the study of plant-insect interactions may help in <a href="/wiki/Bioprospecting" title="Bioprospecting">bioprospecting</a>.<a href="#cite_note-122">[122]</a> There is evidence that humans began using plant alkaloids in medical preparations as early as 3000 <a href="/wiki/Before_Christ" class="mw-redirect" title="Before Christ">B.C.</a><a href="#cite_note-Roberts-37">[37]</a> Although the active components of most medicinal plants have been isolated only recently (beginning in the early 19th century) these substances have been used as drugs throughout the human history in potions, medicines, teas and as <a href="/wiki/Poison" title="Poison">poisons</a>. For example, to combat herbivory by the larvae of some Lepidoptera species, <a href="/wiki/Cinchona" title="Cinchona">Cinchona</a> trees produce a variety of alkaloids, the most familiar of which is <a href="/wiki/Quinine" title="Quinine">quinine</a>. Quinine is extremely bitter, making the bark of the tree quite unpalatable.<a href="#cite_note-123">[123]</a> Throughout history mandrakes (<a href="/wiki/Mandragora_officinarum" title="Mandragora officinarum">Mandragora officinarum</a>) have been highly sought after for their reputed <a href="/wiki/Aphrodisiac" title="Aphrodisiac">aphrodisiac</a> properties. However, the roots of the mandrake plant also contain large quantities of the alkaloid <a href="/wiki/Hyoscine_hydrobromide" class="mw-redirect" title="Hyoscine hydrobromide">scopolamine</a>, which, at high doses, acts as a <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a> <a href="/wiki/Depressant" title="Depressant">depressant</a>, and makes the plant highly toxic to herbivores. Scopolamine was later found to be medicinally used for pain management prior to and during <a href="/wiki/Childbirth" title="Childbirth">labor</a>; in smaller doses it is used to prevent <a href="/wiki/Motion_sickness" title="Motion sickness">motion sickness</a>.<a href="#cite_note-124">[124]</a> One of the most well-known medicinally valuable <a href="/wiki/Terpene" title="Terpene">terpenes</a> is an <a href="/wiki/Antineoplastic" class="mw-redirect" title="Antineoplastic">anticancer</a> drug, <a href="/wiki/Taxol" class="mw-redirect" title="Taxol">taxol</a>, isolated from the bark of the <a href="/wiki/Pacific_yew" class="mw-redirect" title="Pacific yew">Pacific yew</a>, Taxus brevifolia, in the early 1960s.<a href="#cite_note-125">[125]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=25" title="Edit section: See also">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 22em;"> <ul><li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptation</a></li> <li><a href="/wiki/Biopesticide" title="Biopesticide">Biopesticide</a></li> <li><a href="/wiki/Chemical_ecology" title="Chemical ecology">Chemical ecology</a></li> <li><a href="/wiki/List_of_beneficial_weeds" title="List of beneficial weeds">List of beneficial weeds</a></li> <li><a href="/wiki/List_of_companion_plants" title="List of companion plants">List of companion plants</a></li> <li><a href="/wiki/List_of_pest-repelling_plants" title="List of pest-repelling plants">List of pest-repelling plants</a></li> <li><a href="/wiki/Plant_disease_resistance" title="Plant disease resistance">Plant disease resistance</a></li> <li><a href="/wiki/Plant_tolerance_to_herbivory" title="Plant tolerance to herbivory">Plant tolerance to herbivory</a></li> <li><a href="/wiki/Plant_communication" title="Plant communication">Plant communication</a></li> <li><a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=26" title="Edit section: References">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-1"><a href="#cite_ref-1">^</a> Boyd, Jade (2012). 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Rice University. <a rel="nofollow" class="external free" href="http://news.rice.edu/2012/04/09/a-bit-touchy-plants-insect-defenses-activated-by-touch-2/">http://news.rice.edu/2012/04/09/a-bit-touchy-plants-insect-defenses-activated-by-touch-2/</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20120512012546/http://news.rice.edu/2012/04/09/a-bit-touchy-plants-insect-defenses-activated-by-touch-2/">Archived</a> 2012-05-12 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> </li> <li id="cite_note-Kersch-Becker_20171120-2"><a href="#cite_ref-Kersch-Becker_20171120_2-0">^</a> <style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFKersch-BeckerKesslerThaler2017" class="citation journal cs1">Kersch-Becker, Mônica F.; Kessler, André; Thaler, Jennifer S. 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Willoughby: MeisterPro. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-892829-11-5" title="Special:BookSources/978-1-892829-11-5"><bdi>978-1-892829-11-5</bdi></a>. <q>firat Ware</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Pesticide+Book&rft.place=Willoughby&rft.pub=MeisterPro&rft.date=2004&rft.isbn=978-1-892829-11-5&rft.au=George+W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"> </li> <li id="cite_note-smith-117"><a href="#cite_ref-smith_117-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMichael_Smith2005" class="citation book cs1">Michael Smith, C. (2005). Plant Resistance to Arthropods: Molecular and Conventional Approaches. 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Clemson University Home & Garden Information Center. Retrieved 2007-05-27.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Less+toxic+insecticides&rft.pub=Clemson+University+Home+%26+Garden+Information+Center&rft.aulast=Russ&rft.aufirst=Karen&rft_id=http%3A%2F%2Fwww.clemson.edu%2Fextension%2Fhgic%2Fpests%2Fpdf%2Fhgic2770.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"> </li> <li id="cite_note-120"><a href="#cite_ref-120">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFvan_Emden1999" class="citation journal cs1">van Emden, H.F. (November 1999). "Transgenic Host Plant Resistance to Insects—Some Reservations". 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Chemoecology. 1 (1): 38–40. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1990Checo...1R..38E">1990Checo...1R..38E</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF01240585">10.1007/BF01240585</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:19744770">19744770</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Chemoecology&rft.atitle=Prospecting+for+nature%27s+chemical+riches&rft.volume=1&rft.issue=1&rft.pages=38-40&rft.date=1990-03&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A19744770%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1007%2FBF01240585&rft_id=info%3Abibcode%2F1990Checo...1R..38E&rft.aulast=Eisner&rft.aufirst=Thomas&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"> </li> <li id="cite_note-123"><a href="#cite_ref-123">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlbert_T._Sneden" class="citation web cs1">Albert T. Sneden. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070205203742/http://www.people.vcu.edu/~asneden/The%20Quinine%20Alkaloids.pdf">"The Quinine Alkaloids"</a> (PDF). Medicinal Chemistry and Drug Design. Archived from <a rel="nofollow" class="external text" href="http://www.people.vcu.edu/~asneden/The%20Quinine%20Alkaloids.pdf">the original</a> (PDF) on 2007-02-05. Retrieved 2007-05-23.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Medicinal+Chemistry+and+Drug+Design&rft.atitle=The+Quinine+Alkaloids&rft.au=Albert+T.+Sneden&rft_id=http%3A%2F%2Fwww.people.vcu.edu%2F~asneden%2FThe%2520Quinine%2520Alkaloids.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"> </li> <li id="cite_note-124"><a href="#cite_ref-124">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlbert_T._Sneden" class="citation web cs1">Albert T. Sneden. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070927010636/http://www.people.vcu.edu/~asneden/tropane%20alkaloids.pdf">"The Tropane Alkaloids"</a> (PDF). Medicinal Chemistry and Drug Design. Archived from <a rel="nofollow" class="external text" href="http://www.people.vcu.edu/~asneden/tropane%20alkaloids.pdf">the original</a> (PDF) on 2007-09-27. Retrieved 2007-05-23.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Medicinal+Chemistry+and+Drug+Design&rft.atitle=The+Tropane+Alkaloids&rft.au=Albert+T.+Sneden&rft_id=http%3A%2F%2Fwww.people.vcu.edu%2F~asneden%2Ftropane%2520alkaloids.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"> </li> <li id="cite_note-125"><a href="#cite_ref-125">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlbert_T._Sneden" class="citation web cs1">Albert T. Sneden. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070927010639/http://www.people.vcu.edu/~asneden/Taxol.pdf">"Taxol (Paclitaxe)"</a> (PDF). Medicinal Chemistry and Drug Design. Archived from <a rel="nofollow" class="external text" href="http://www.people.vcu.edu/~asneden/Taxol.pdf">the original</a> (PDF) on 2007-09-27. Retrieved 2007-05-23.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Medicinal+Chemistry+and+Drug+Design&rft.atitle=Taxol+%28Paclitaxe%29&rft.au=Albert+T.+Sneden&rft_id=http%3A%2F%2Fwww.people.vcu.edu%2F~asneden%2FTaxol.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=27" title="Edit section: Further reading">edit</a>]</div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Fritz, Robert S.; Sims, Ellen L., eds. (1992). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=NkF0xuz664gC&q=Herbivores:+Their+interaction+with+secondary+plant">Plant resistance to herbivores and pathogens: ecology, evolution, and genetics</a>. Chicago: University of Chicago Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-26553-7" title="Special:BookSources/978-0-226-26553-7"><bdi>978-0-226-26553-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Plant+resistance+to+herbivores+and+pathogens%3A+ecology%2C+evolution%2C+and+genetics&rft.place=Chicago&rft.pub=University+of+Chicago+Press&rft.date=1992&rft.isbn=978-0-226-26553-7&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DNkF0xuz664gC%26q%3DHerbivores%3A%2BTheir%2Binteraction%2Bwith%2Bsecondary%2Bplant&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Jolivet, Pierre (1998). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=J5sTAB_mymwC&q=Herbivores:+Their+interaction+with+secondary+plant">Interrelationship Between Insects and Plants</a>. Boca Raton: CRC. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-57444-052-2" title="Special:BookSources/978-1-57444-052-2"><bdi>978-1-57444-052-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Interrelationship+Between+Insects+and+Plants&rft.place=Boca+Raton&rft.pub=CRC&rft.date=1998&rft.isbn=978-1-57444-052-2&rft.au=Jolivet%2C+Pierre&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DJ5sTAB_mymwC%26q%3DHerbivores%3A%2BTheir%2Binteraction%2Bwith%2Bsecondary%2Bplant&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Karban, Richard; Baldwin, Ian T. (1997). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=thHTfR3ICZAC&q=Herbivores:+Their+interaction+with+secondary+plant">Induced responses to herbivory</a>. Chicago: University of Chicago Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-42495-8" title="Special:BookSources/978-0-226-42495-8"><bdi>978-0-226-42495-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Induced+responses+to+herbivory&rft.place=Chicago&rft.pub=University+of+Chicago+Press&rft.date=1997&rft.isbn=978-0-226-42495-8&rft.au=Karban%2C+Richard&rft.au=Baldwin%2C+Ian+T.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DthHTfR3ICZAC%26q%3DHerbivores%3A%2BTheir%2Binteraction%2Bwith%2Bsecondary%2Bplant&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Speight, Martin R.; Hunter, Mark D.; Watt, Allan D. (1999). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=JiO_Nnk5Y8AC&q=%22Insect+ecology%22&pg=PP1">Ecology of insects: concepts and applications</a>. Oxford: Blackwell Science. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-86542-745-7" title="Special:BookSources/978-0-86542-745-7"><bdi>978-0-86542-745-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Ecology+of+insects%3A+concepts+and+applications&rft.place=Oxford&rft.pub=Blackwell+Science&rft.date=1999&rft.isbn=978-0-86542-745-7&rft.au=Speight%2C+Martin+R.&rft.au=Hunter%2C+Mark+D.&rft.au=Watt%2C+Allan+D.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DJiO_Nnk5Y8AC%26q%3D%2522Insect%2Becology%2522%26pg%3DPP1&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Thompson, John N. (1994). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=InCAChmWM1QC&q=Herbivores:+Their+interaction+with+secondary+plant">The coevolutionary process</a>. Chicago: University of Chicago Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-79759-5" title="Special:BookSources/978-0-226-79759-5"><bdi>978-0-226-79759-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+coevolutionary+process&rft.place=Chicago&rft.pub=University+of+Chicago+Press&rft.date=1994&rft.isbn=978-0-226-79759-5&rft.au=Thompson%2C+John+N.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DInCAChmWM1QC%26q%3DHerbivores%3A%2BTheir%2Binteraction%2Bwith%2Bsecondary%2Bplant&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Wiens, D. (1978). "Mimicry in Plants". Evolutionary Biology. Vol. 11. pp. 365–403. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-1-4615-6956-5_6">10.1007/978-1-4615-6956-5_6</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4615-6958-9" title="Special:BookSources/978-1-4615-6958-9"><bdi>978-1-4615-6958-9</bdi></a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3282713">3282713</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/22182416">22182416</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Mimicry+in+Plants&rft.btitle=Evolutionary+Biology&rft.pages=365-403&rft.date=1978&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3282713%23id-name%3DPMC&rft_id=info%3Apmid%2F22182416&rft_id=info%3Adoi%2F10.1007%2F978-1-4615-6956-5_6&rft.isbn=978-1-4615-6958-9&rft.aulast=Wiens&rft.aufirst=D.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+defense+against+herbivory" class="Z3988"></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=Plant_defense_against_herbivory&action=edit&section=28" title="Edit section: External links">edit</a>]</div> <ul><li>Bruce A. Kimball <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070205144747/http://www.colostate.edu/Depts/Entomology/courses/en570/papers_1996/kimball.html">Evolutionary Plant Defense Strategies Life Histories and Contributions to Future Generations</a></li> <li><a rel="nofollow" class="external text" href="http://www2.mcdaniel.edu/Biology/botf99/herbnew/aintro.htm">Plant Defense Systems & Medicinal Botany</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070105193927/http://www2.mcdaniel.edu/Biology/botf99/herbnew/aintro.htm">Archived</a> 2007-01-05 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="http://www.johnsankey.ca/senecio.html">Herbivore Defenses of Senecio viscosus L.</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20120107045212/http://richannel.org/christmas-lectures-2009-sue-hartley--the-animals-strike-back">Sue Hartley Royal Institution Christmas Lectures 2009: The Animals Strike Back</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist 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title="Special:EditPage/Template:Modelling ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ecology:_Modelling_ecosystems:_Trophic_components" style="font-size:114%;margin:0 4em"><a href="/wiki/Ecology" title="Ecology">Ecology</a>: <a href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: <a href="/wiki/Trophic_level" title="Trophic level">Trophic</a> components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">General</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiotic_component" title="Abiotic component">Abiotic component</a></li> <li><a href="/wiki/Abiotic_stress" title="Abiotic stress">Abiotic stress</a></li> <li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behaviour</a></li> <li><a href="/wiki/Biogeochemical_cycle" title="Biogeochemical cycle">Biogeochemical cycle</a></li> <li><a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li> <li><a href="/wiki/Biotic_component" class="mw-redirect" title="Biotic component">Biotic component</a></li> <li><a href="/wiki/Biotic_stress" title="Biotic stress">Biotic stress</a></li> <li><a href="/wiki/Carrying_capacity" title="Carrying capacity">Carrying capacity</a></li> <li><a href="/wiki/Competition_(biology)" title="Competition (biology)">Competition</a></li> <li><a href="/wiki/Ecosystem" title="Ecosystem">Ecosystem</a></li> <li><a href="/wiki/Ecosystem_ecology" title="Ecosystem ecology">Ecosystem ecology</a></li> <li><a href="/wiki/Ecosystem_model" title="Ecosystem model">Ecosystem model</a></li> <li><a href="/wiki/Green_world_hypothesis" title="Green world hypothesis">Green world hypothesis</a></li> <li><a href="/wiki/Keystone_species" title="Keystone species">Keystone species</a></li> <li><a href="/wiki/List_of_feeding_behaviours" title="List of feeding behaviours">List of feeding behaviours</a></li> <li><a href="/wiki/Metabolic_theory_of_ecology" title="Metabolic theory of ecology">Metabolic theory of ecology</a></li> <li><a href="/wiki/Productivity_(ecology)" title="Productivity (ecology)">Productivity</a></li> <li><a href="/wiki/Resource_(biology)" title="Resource (biology)">Resource</a></li> <li><a href="/wiki/Restoration_ecology" class="mw-redirect" title="Restoration ecology">Restoration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Autotroph" title="Autotroph">Producers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Autotroph" title="Autotroph">Autotrophs</a></li> <li><a href="/wiki/Chemosynthesis" title="Chemosynthesis">Chemosynthesis</a></li> <li><a href="/wiki/Chemotroph" title="Chemotroph">Chemotrophs</a></li> <li><a href="/wiki/Foundation_species" title="Foundation species">Foundation species</a></li> <li><a href="/wiki/Kinetotroph" title="Kinetotroph">Kinetotrophs</a></li> <li><a href="/wiki/Mixotroph" title="Mixotroph">Mixotrophs</a></li> <li><a href="/wiki/Myco-heterotrophy" title="Myco-heterotrophy">Myco-heterotrophy</a></li> <li><a href="/wiki/Mycotroph" title="Mycotroph">Mycotroph</a></li> <li><a href="/wiki/Organotroph" title="Organotroph">Organotrophs</a></li> <li><a href="/wiki/Photoheterotroph" title="Photoheterotroph">Photoheterotrophs</a></li> <li><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a></li> <li><a href="/wiki/Photosynthetic_efficiency" title="Photosynthetic efficiency">Photosynthetic efficiency</a></li> <li><a href="/wiki/Phototroph" title="Phototroph">Phototrophs</a></li> <li><a href="/wiki/Primary_nutritional_groups" title="Primary nutritional groups">Primary nutritional groups</a></li> <li><a href="/wiki/Primary_production" title="Primary production">Primary production</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Consumer_(food_chain)" title="Consumer (food chain)">Consumers</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Apex_predator" title="Apex predator">Apex predator</a></li> <li><a href="/wiki/Bacterivore" title="Bacterivore">Bacterivore</a></li> <li><a href="/wiki/Carnivore" title="Carnivore">Carnivores</a></li> <li><a href="/wiki/Chemoorganotroph" class="mw-redirect" title="Chemoorganotroph">Chemoorganotroph</a></li> <li><a href="/wiki/Foraging" title="Foraging">Foraging</a></li> <li><a href="/wiki/Generalist_and_specialist_species" title="Generalist and specialist species">Generalist and specialist species</a></li> <li><a href="/wiki/Intraguild_predation" title="Intraguild predation">Intraguild predation</a></li> <li><a href="/wiki/Herbivore" title="Herbivore">Herbivores</a></li> <li><a href="/wiki/Heterotroph" title="Heterotroph">Heterotroph</a></li> <li><a href="/wiki/Heterotrophic_nutrition" title="Heterotrophic nutrition">Heterotrophic nutrition</a></li> <li><a href="/wiki/Insectivore" title="Insectivore">Insectivore</a></li> <li><a href="/wiki/Mesopredator" title="Mesopredator">Mesopredators</a></li> <li><a href="/wiki/Mesopredator_release_hypothesis" title="Mesopredator release hypothesis">Mesopredator release hypothesis</a></li> <li><a href="/wiki/Omnivore" title="Omnivore">Omnivores</a></li> <li><a href="/wiki/Optimal_foraging_theory" title="Optimal foraging theory">Optimal foraging theory</a></li> <li><a href="/wiki/Planktivore" title="Planktivore">Planktivore</a></li> <li><a href="/wiki/Predation" title="Predation">Predation</a></li> <li><a href="/wiki/Prey_switching" title="Prey switching">Prey switching</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Decomposer" title="Decomposer">Decomposers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chemoorganoheterotrophy" class="mw-redirect" title="Chemoorganoheterotrophy">Chemoorganoheterotrophy</a></li> <li><a href="/wiki/Decomposition" title="Decomposition">Decomposition</a></li> <li><a href="/wiki/Detritivore" title="Detritivore">Detritivores</a></li> <li><a href="/wiki/Detritus" title="Detritus">Detritus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Microorganism#Habitats_and_ecology" title="Microorganism">Microorganisms</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteriophage" title="Bacteriophage">Bacteriophage</a></li> <li><a href="/wiki/Lithoautotroph" title="Lithoautotroph">Lithoautotroph</a></li> <li><a href="/wiki/Lithotroph" title="Lithotroph">Lithotrophy</a></li> <li><a href="/wiki/Marine_microorganisms" title="Marine microorganisms">Marine</a></li> <li><a href="/wiki/Microbial_cooperation" title="Microbial cooperation">Microbial cooperation</a></li> <li><a href="/wiki/Microbial_ecology" title="Microbial ecology">Microbial ecology</a></li> <li><a href="/wiki/Microbial_food_web" title="Microbial food web">Microbial food web</a></li> <li><a href="/wiki/Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li> <li><a href="/wiki/Microbial_loop" title="Microbial loop">Microbial loop</a></li> <li><a href="/wiki/Microbial_mat" title="Microbial mat">Microbial mat</a></li> <li><a href="/wiki/Microbial_metabolism" title="Microbial metabolism">Microbial metabolism</a></li> <li><a href="/wiki/Phage_ecology" title="Phage ecology">Phage ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Food_web" title="Food web">Food webs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biomagnification" title="Biomagnification">Biomagnification</a></li> <li><a href="/wiki/Ecological_efficiency" title="Ecological efficiency">Ecological efficiency</a></li> <li><a href="/wiki/Ecological_pyramid" title="Ecological pyramid">Ecological pyramid</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Energy flow</a></li> <li><a href="/wiki/Food_chain" title="Food chain">Food chain</a></li> <li><a href="/wiki/Trophic_level" title="Trophic level">Trophic level</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Example webs</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lake_ecosystem#Trophic_relationships" title="Lake ecosystem">Lakes</a></li> <li><a href="/wiki/River_ecosystem#Trophic_relationships" title="River ecosystem">Rivers</a></li> <li><a href="/wiki/Soil_food_web" title="Soil food web">Soil</a></li> <li><a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></li> <li><a href="/wiki/Marine_food_web" title="Marine food web">Marine food webs</a> <ul><li><a href="/wiki/Cold_seep" title="Cold seep">cold seeps</a></li> <li><a href="/wiki/Hydrothermal_vent#Biological_communities" title="Hydrothermal vent">hydrothermal vents</a></li> <li><a href="/wiki/Intertidal_ecology" title="Intertidal ecology">intertidal</a></li> <li><a href="/wiki/Kelp_forest#Trophic_ecology" title="Kelp forest">kelp forests</a></li> <li><a href="/wiki/North_Pacific_Gyre" title="North Pacific Gyre">North Pacific Gyre</a></li> <li><a href="/wiki/Ecology_of_the_San_Francisco_Estuary#Food_web" title="Ecology of the San Francisco Estuary">San Francisco Estuary</a></li> <li><a href="/wiki/Tide_pool" title="Tide pool">tide pool</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Processes</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ascendency" title="Ascendency">Ascendency</a></li> <li><a href="/wiki/Bioaccumulation" title="Bioaccumulation">Bioaccumulation</a></li> <li><a href="/wiki/Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li> <li><a href="/wiki/Climax_community" title="Climax community">Climax community</a></li> <li><a href="/wiki/Competitive_exclusion_principle" title="Competitive exclusion principle">Competitive exclusion principle</a></li> <li><a href="/wiki/Consumer%E2%80%93resource_interactions" title="Consumer–resource interactions">Consumer–resource interactions</a></li> <li><a href="/wiki/Copiotroph" title="Copiotroph">Copiotrophs</a></li> <li><a href="/wiki/Dominance_(ecology)" title="Dominance (ecology)">Dominance</a></li> <li><a href="/wiki/Ecological_network" title="Ecological network">Ecological network</a></li> <li><a href="/wiki/Ecological_succession" title="Ecological succession">Ecological succession</a></li> <li><a href="/wiki/Energy_quality" title="Energy quality">Energy quality</a></li> <li><a href="/wiki/Energy_systems_language" title="Energy systems language">Energy systems language</a></li> <li><a href="/wiki/F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li> <li><a href="/wiki/Feed_conversion_ratio" title="Feed conversion ratio">Feed conversion ratio</a></li> <li><a href="/wiki/Feeding_frenzy" title="Feeding frenzy">Feeding frenzy</a></li> <li><a href="/wiki/Mesotrophic_soil" title="Mesotrophic soil">Mesotrophic soil</a></li> <li><a href="/wiki/Nutrient_cycle" title="Nutrient cycle">Nutrient cycle</a></li> <li><a href="/wiki/Oligotroph" title="Oligotroph">Oligotroph</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li> <li><a href="/wiki/Trophic_cascade" title="Trophic cascade">Trophic cascade</a></li> <li><a href="/wiki/Trophic_mutualism" title="Trophic mutualism">Trophic mutualism</a></li> <li><a href="/wiki/Trophic_state_index" title="Trophic state index">Trophic state index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Defense, counter</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptations</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Herbivore_adaptations_to_plant_defense" title="Herbivore adaptations to plant defense">Herbivore adaptations to plant defense</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a class="mw-selflink selflink">Plant defense against herbivory</a></li> <li><a href="/wiki/Shoaling_and_schooling#Predator_avoidance" title="Shoaling and schooling">Predator avoidance in schooling fish</a></li></ul> </div></td></tr></tbody></table></div><div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Other_components" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Modelling_ecosystems" title="Template:Modelling ecosystems"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Modelling_ecosystems" title="Template talk:Modelling ecosystems"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Modelling_ecosystems" title="Special:EditPage/Template:Modelling ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ecology:_Modelling_ecosystems:_Other_components" style="font-size:114%;margin:0 4em"><a href="/wiki/Ecology" title="Ecology">Ecology</a>: <a href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: Other components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Population_ecology" title="Population ecology">Population ecology</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abundance_(ecology)" title="Abundance (ecology)">Abundance</a></li> <li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Consumer-resource_model" title="Consumer-resource model">Consumer-resource model</a></li> <li><a href="/wiki/Depensation" title="Depensation">Depensation</a></li> <li><a href="/wiki/Ecological_yield" title="Ecological yield">Ecological yield</a></li> <li><a href="/wiki/Effective_population_size" title="Effective population size">Effective population size</a></li> <li><a href="/wiki/Intraspecific_competition" title="Intraspecific competition">Intraspecific competition</a></li> <li><a href="/wiki/Logistic_function" title="Logistic function">Logistic function</a></li> <li><a href="/wiki/Malthusian_growth_model" title="Malthusian growth model">Malthusian growth model</a></li> <li><a href="/wiki/Maximum_sustainable_yield" title="Maximum sustainable yield">Maximum sustainable yield</a></li> <li><a href="/wiki/Overpopulation" title="Overpopulation">Overpopulation</a></li> <li><a href="/wiki/Overexploitation" title="Overexploitation">Overexploitation</a></li> <li><a href="/wiki/Population_cycle" title="Population cycle">Population cycle</a></li> <li><a href="/wiki/Population_dynamics" title="Population dynamics">Population dynamics</a></li> <li><a href="/wiki/Population_model" title="Population model">Population modeling</a></li> <li><a href="/wiki/Population_size" title="Population size">Population size</a></li> <li><a href="/wiki/Lotka%E2%80%93Volterra_equations" title="Lotka–Volterra equations">Predator–prey (Lotka–Volterra) equations</a></li> <li><a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">Recruitment</a></li> <li><a href="/wiki/Small_population_size" title="Small population size">Small population size</a></li> <li><a href="/wiki/Ecological_stability" title="Ecological stability">Stability</a> <ul><li><a href="/wiki/Ecological_resilience" title="Ecological resilience">Resilience</a></li> <li><a href="/wiki/Resistance_(ecology)" title="Resistance (ecology)">Resistance</a></li></ul></li> <li><a href="/wiki/Random_generalized_Lotka%E2%80%93Volterra_model" title="Random generalized Lotka–Volterra model">Random generalized Lotka–Volterra model</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Density_dependence" title="Density dependence">Density-dependent inhibition</a></li> <li><a href="/wiki/Ecological_effects_of_biodiversity" title="Ecological effects of biodiversity">Ecological effects of biodiversity</a></li> <li><a href="/wiki/Ecological_extinction" title="Ecological extinction">Ecological extinction</a></li> <li><a href="/wiki/Endemism" title="Endemism">Endemic species</a></li> <li><a href="/wiki/Flagship_species" title="Flagship species">Flagship species</a></li> <li><a href="/wiki/Ordination_(statistics)" title="Ordination (statistics)">Gradient analysis</a></li> <li><a href="/wiki/Bioindicator" title="Bioindicator">Indicator species</a></li> <li><a href="/wiki/Introduced_species" title="Introduced species">Introduced species</a></li> <li><a href="/wiki/Invasive_species" title="Invasive species">Invasive species</a> / <a href="/wiki/Native_species" title="Native species">Native species</a></li> <li><a href="/wiki/Latitudinal_gradients_in_species_diversity" title="Latitudinal gradients in species diversity">Latitudinal gradients in species diversity</a></li> <li><a href="/wiki/Minimum_viable_population" title="Minimum viable population">Minimum viable population</a></li> <li><a href="/wiki/Unified_neutral_theory_of_biodiversity" title="Unified neutral theory of biodiversity">Neutral theory</a></li> <li><a href="/wiki/Occupancy%E2%80%93abundance_relationship" title="Occupancy–abundance relationship">Occupancy–abundance relationship</a></li> <li><a href="/wiki/Population_viability_analysis" title="Population viability analysis">Population viability analysis</a></li> <li><a href="/wiki/Priority_effect" title="Priority effect">Priority effect</a></li> <li><a href="/wiki/Rapoport%27s_rule" title="Rapoport's rule">Rapoport's rule</a></li> <li><a href="/wiki/Relative_abundance_distribution" title="Relative abundance distribution">Relative abundance distribution</a></li> <li><a href="/wiki/Relative_species_abundance" title="Relative species abundance">Relative species abundance</a></li> <li><a href="/wiki/Species_diversity" title="Species diversity">Species diversity</a></li> <li><a href="/wiki/Species_homogeneity" title="Species homogeneity">Species homogeneity</a></li> <li><a href="/wiki/Species_richness" title="Species richness">Species richness</a></li> <li><a href="/wiki/Species_distribution" title="Species distribution">Species distribution</a></li> <li><a href="/wiki/Species%E2%80%93area_relationship" title="Species–area relationship">Species–area curve</a></li> <li><a href="/wiki/Umbrella_species" title="Umbrella species">Umbrella species</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species interaction</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antibiosis" title="Antibiosis">Antibiosis</a></li> <li><a href="/wiki/Biological_interaction" title="Biological interaction">Biological interaction</a></li> <li><a href="/wiki/Commensalism" title="Commensalism">Commensalism</a></li> <li><a href="/wiki/Community_(ecology)" title="Community (ecology)">Community ecology</a></li> <li><a href="/wiki/Ecological_facilitation" title="Ecological facilitation">Ecological facilitation</a></li> <li><a href="/wiki/Interspecific_competition" title="Interspecific competition">Interspecific competition</a></li> <li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li> <li><a href="/wiki/Parasitism" title="Parasitism">Parasitism</a></li> <li><a href="/wiki/Storage_effect" title="Storage effect">Storage effect</a></li> <li><a href="/wiki/Symbiosis" title="Symbiosis">Symbiosis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Spatial_ecology" title="Spatial ecology">Spatial ecology</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Cross-boundary_subsidy" title="Cross-boundary subsidy">Cross-boundary subsidy</a></li> <li><a href="/wiki/Cline_(biology)" title="Cline (biology)">Ecocline</a></li> <li><a href="/wiki/Ecotone" title="Ecotone">Ecotone</a></li> <li><a href="/wiki/Ecotype" title="Ecotype">Ecotype</a></li> <li><a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">Disturbance</a></li> <li><a href="/wiki/Edge_effects" title="Edge effects">Edge effects</a></li> <li><a href="/wiki/Foster%27s_rule" title="Foster's rule">Foster's rule</a></li> <li><a href="/wiki/Habitat_fragmentation" title="Habitat fragmentation">Habitat fragmentation</a></li> <li><a href="/wiki/Ideal_free_distribution" title="Ideal free distribution">Ideal free distribution</a></li> <li><a href="/wiki/Intermediate_disturbance_hypothesis" title="Intermediate disturbance hypothesis">Intermediate disturbance hypothesis</a></li> <li><a href="/wiki/Insular_biogeography" title="Insular biogeography">Insular biogeography</a></li> <li><a href="/wiki/Land_change_modeling" title="Land change modeling">Land change modeling</a></li> <li><a href="/wiki/Landscape_ecology" title="Landscape ecology">Landscape ecology</a></li> <li><a href="/wiki/Landscape_epidemiology" title="Landscape epidemiology">Landscape epidemiology</a></li> <li><a href="/wiki/Landscape_limnology" title="Landscape limnology">Landscape limnology</a></li> <li><a href="/wiki/Metapopulation" title="Metapopulation">Metapopulation</a></li> <li><a href="/wiki/Patch_dynamics" title="Patch dynamics">Patch dynamics</a></li> <li><a href="/wiki/R/K_selection_theory" title="R/K selection theory">r/K selection theory</a></li> <li><a href="/wiki/Resource_selection_function" title="Resource selection function">Resource selection function</a></li> <li><a href="/wiki/Source%E2%80%93sink_dynamics" title="Source–sink dynamics">Source–sink dynamics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Ecological_niche" title="Ecological niche">Niche</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ecological_niche" title="Ecological niche">Ecological niche</a></li> <li><a href="/wiki/Ecological_trap" title="Ecological trap">Ecological trap</a></li> <li><a href="/wiki/Ecosystem_engineer" title="Ecosystem engineer">Ecosystem engineer</a></li> <li><a href="/wiki/Species_distribution_modelling" title="Species distribution modelling">Environmental niche modelling</a></li> <li><a href="/wiki/Guild_(ecology)" title="Guild (ecology)">Guild</a></li> <li><a href="/wiki/Habitat" title="Habitat">Habitat</a> <ul><li><a href="/wiki/Marine_habitat" title="Marine habitat">marine habitats</a></li></ul></li> <li><a href="/wiki/Limiting_similarity" title="Limiting similarity">Limiting similarity</a></li> <li><a href="/wiki/Niche_apportionment_models" title="Niche apportionment models">Niche apportionment models</a></li> <li><a href="/wiki/Niche_construction" title="Niche construction">Niche construction</a></li> <li><a href="/wiki/Niche_differentiation" class="mw-redirect" title="Niche differentiation">Niche differentiation</a></li> <li><a href="/wiki/Ontogenetic_niche_shift" title="Ontogenetic niche shift">Ontogenetic niche shift</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Non-trophic_networks" title="Non-trophic networks">Other networks</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Assembly_rules" title="Assembly rules">Assembly rules</a></li> <li><a href="/wiki/Bateman%27s_principle" title="Bateman's principle">Bateman's principle</a></li> <li><a href="/wiki/Bioluminescence" title="Bioluminescence">Bioluminescence</a></li> <li><a href="/wiki/Ecological_collapse" class="mw-redirect" title="Ecological collapse">Ecological collapse</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological debt</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological deficit</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Ecological energetics</a></li> <li><a href="/wiki/Ecological_indicator" title="Ecological indicator">Ecological indicator</a></li> <li><a href="/wiki/Ecological_threshold" title="Ecological threshold">Ecological threshold</a></li> <li><a href="/wiki/Ecosystem_diversity" title="Ecosystem diversity">Ecosystem diversity</a></li> <li><a href="/wiki/Emergence" title="Emergence">Emergence</a></li> <li><a href="/wiki/Extinction_debt" title="Extinction debt">Extinction debt</a></li> <li><a href="/wiki/Kleiber%27s_law" title="Kleiber's law">Kleiber's law</a></li> <li><a href="/wiki/Liebig%27s_law_of_the_minimum" title="Liebig's law of the minimum">Liebig's law of the minimum</a></li> <li><a href="/wiki/Marginal_value_theorem" title="Marginal value theorem">Marginal value theorem</a></li> <li><a href="/wiki/Thorson%27s_rule" title="Thorson's rule">Thorson's rule</a></li> <li><a href="/wiki/Xerosere" title="Xerosere">Xerosere</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allometry" title="Allometry">Allometry</a></li> <li><a href="/wiki/Alternative_stable_state" title="Alternative stable state">Alternative stable state</a></li> <li><a href="/wiki/Balance_of_nature" title="Balance of nature">Balance of nature</a></li> <li><a href="/wiki/Biological_data_visualization" title="Biological data visualization">Biological data visualization</a></li> <li><a href="/wiki/Ecological_economics" title="Ecological economics">Ecological economics</a></li> <li><a href="/wiki/Ecological_footprint" title="Ecological footprint">Ecological footprint</a></li> <li><a href="/wiki/Ecological_forecasting" title="Ecological forecasting">Ecological forecasting</a></li> 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