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href="#Formation_of_the_floral_meristem_or_the_inflorescence"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Formation of the floral meristem or the inflorescence</span> </div> </a> <ul id="toc-Formation_of_the_floral_meristem_or_the_inflorescence-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Floral_architecture" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Floral_architecture"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Floral architecture</span> </div> </a> <button aria-controls="toc-Floral_architecture-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Floral architecture subsection</span> </button> <ul id="toc-Floral_architecture-sublist" class="vector-toc-list"> <li id="toc-The_ABC_model" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#The_ABC_model"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>The ABC model</span> </div> </a> <ul id="toc-The_ABC_model-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetic_analysis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_analysis"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Genetic analysis</span> </div> </a> <ul id="toc-Genetic_analysis-sublist" class="vector-toc-list"> <li id="toc-Analysis_of_mutants" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Analysis_of_mutants"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.1</span> <span>Analysis of mutants</span> </div> </a> <ul id="toc-Analysis_of_mutants-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Techniques_for_detecting_differential_expression" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Techniques_for_detecting_differential_expression"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.2</span> <span>Techniques for detecting differential expression</span> </div> </a> <ul id="toc-Techniques_for_detecting_differential_expression-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genes_exhibiting_type-A_function" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genes_exhibiting_type-A_function"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Genes exhibiting type-A function</span> </div> </a> <ul id="toc-Genes_exhibiting_type-A_function-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genes_exhibiting_type-B_function" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genes_exhibiting_type-B_function"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Genes exhibiting type-B function</span> </div> </a> <ul id="toc-Genes_exhibiting_type-B_function-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genes_exhibiting_type-C_function" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genes_exhibiting_type-C_function"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Genes exhibiting type-C function</span> </div> </a> <ul id="toc-Genes_exhibiting_type-C_function-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genes_exhibiting_type-D_and_E_functions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genes_exhibiting_type-D_and_E_functions"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.6</span> <span>Genes exhibiting type-D and E functions</span> </div> </a> <ul id="toc-Genes_exhibiting_type-D_and_E_functions-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" 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class="firstHeading mw-first-heading"><span class="mw-page-title-main">ABC model of flower development</span></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/Desenvolupament_floral" title="Desenvolupament floral – Catalan" lang="ca" hreflang="ca" data-title="Desenvolupament floral" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://es.wikipedia.org/wiki/Desarrollo_floral" title="Desarrollo floral – Spanish" lang="es" hreflang="es" data-title="Desarrollo floral" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Mod%C3%A8le_ABC" title="Modèle ABC – French" lang="fr" hreflang="fr" data-title="Modèle ABC" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Modello_ABC_di_sviluppo_fiorale" title="Modello ABC di sviluppo fiorale – Italian" lang="it" hreflang="it" data-title="Modello ABC di sviluppo fiorale" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/ABC-model_van_bloemontwikkeling" title="ABC-model van bloemontwikkeling – Dutch" lang="nl" hreflang="nl" data-title="ABC-model van bloemontwikkeling" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/ABC%E3%83%A2%E3%83%87%E3%83%AB" title="ABCモデル – Japanese" lang="ja" hreflang="ja" data-title="ABCモデル" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Controle_do_florescimento" title="Controle do florescimento – Portuguese" lang="pt" hreflang="pt" data-title="Controle do florescimento" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/ABC-%D0%BC%D0%BE%D0%B4%D0%B5%D0%BB%D1%8C_%D1%80%D0%B0%D0%B7%D0%B2%D0%B8%D1%82%D0%B8%D1%8F_%D1%86%D0%B2%D0%B5%D1%82%D0%BA%D0%B0" title="ABC-модель развития цветка – Russian" lang="ru" hreflang="ru" data-title="ABC-модель развития цветка" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li 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free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></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">Model for genetics of flower development</div> <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">"ABC model" redirects here. For the model of attitudes used in psychology, see <a href="/wiki/Attitude_(psychology)" title="Attitude (psychology)">Attitude (psychology)</a>.</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:ABC_Model.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/ABC_Model.svg/440px-ABC_Model.svg.png" decoding="async" width="440" height="336" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/ABC_Model.svg/660px-ABC_Model.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6e/ABC_Model.svg/880px-ABC_Model.svg.png 2x" data-file-width="936" data-file-height="715" /></a><figcaption>ABC model of flower development guided by three groups of <a href="/wiki/Homeotic_gene" title="Homeotic gene">homeotic genes</a>.</figcaption></figure> <p>The <b>ABC model of flower development</b> is a <a href="/wiki/Scientific_model" class="mw-redirect" title="Scientific model">scientific model</a> of the process by which <a href="/wiki/Flowering_plant" title="Flowering plant">flowering plants</a> produce a pattern of <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> in <a href="/wiki/Meristem" title="Meristem">meristems</a> that leads to the appearance of an organ oriented towards <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual reproduction</a>, a flower. There are three <a href="/wiki/Physiology" title="Physiology">physiological</a> developments that must occur in order for this to take place: firstly, the plant must pass from sexual immaturity into a sexually mature state (i.e. a transition towards flowering); secondly, the transformation of the <a href="/wiki/Apical_meristem" class="mw-redirect" title="Apical meristem">apical meristem's</a> function from a vegetative meristem into a floral meristem or <a href="/wiki/Inflorescence" title="Inflorescence">inflorescence</a>; and finally the growth of the flower's individual organs. The latter phase has been modelled using the <b>ABC model</b>, which aims to describe the biological basis of the process from the perspective of <a href="/wiki/Molecular_genetics" title="Molecular genetics">molecular</a> and <a href="/wiki/Developmental_genetics" class="mw-redirect" title="Developmental genetics">developmental</a> genetics. </p><p>An external <a href="/wiki/Stimulus_(physiology)" title="Stimulus (physiology)">stimulus</a> is required in order to trigger the <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiation</a> of the meristem into a flower meristem. This stimulus will activate <a href="/wiki/Mitosis" title="Mitosis">mitotic</a> cell division in the apical meristem, particularly on its sides where new <a href="/wiki/Primordium" title="Primordium">primordia</a> are formed. This same stimulus will also cause the meristem to follow a <a href="/wiki/Biological_development" class="mw-redirect" title="Biological development">developmental</a> pattern that will lead to the growth of floral meristems as opposed to vegetative meristems. The main difference between these two types of meristem, apart from the obvious disparity between the objective organ, is the verticillate (or whorled) <a href="/wiki/Phyllotaxis" title="Phyllotaxis">phyllotaxis</a>, that is, the absence of <a href="/wiki/Plant_stem" title="Plant stem">stem</a> elongation among the successive <a href="/wiki/Whorl_(botany)" title="Whorl (botany)">whorls or verticils</a> of the primordium. These verticils follow an acropetal development, giving rise to <a href="/wiki/Sepal" title="Sepal">sepals</a>, <a href="/wiki/Petal" title="Petal">petals</a>, <a href="/wiki/Stamen" title="Stamen">stamens</a> and <a href="/wiki/Carpel" class="mw-redirect" title="Carpel">carpels</a>. Another difference from vegetative axillary meristems is that the floral meristem is "determined", which means that, once differentiated, its cells will no longer <a href="/wiki/Cell_cycle" title="Cell cycle">divide</a>.<sup id="cite_ref-azcón-bieto_1-0" class="reference"><a href="#cite_note-azcón-bieto-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>The identity of the organs present in the four floral verticils is a consequence of the interaction of at least three types of <a href="/wiki/Gene_products" class="mw-redirect" title="Gene products">gene products</a>, each with distinct functions. According to the ABC model, functions A and C are required in order to determine the identity of the verticils of the <a href="/wiki/Perianth" title="Perianth">perianth</a> and the reproductive verticils, respectively. These functions are exclusive and the absence of one of them means that the other will determine the identity of all the floral verticils. The B function allows the differentiation of petals from sepals in the secondary verticil, as well as the differentiation of the stamen from the carpel on the tertiary verticil. </p><p><a href="/wiki/Johann_Wolfgang_von_Goethe" title="Johann Wolfgang von Goethe">Goethe</a>'s <b>foliar theory</b> was formulated in the 18th century and it suggests that the constituent parts of a flower are structurally modified leaves, which are functionally specialized for reproduction or protection. The theory was first published in 1790 in the essay "Metamorphosis of Plants" ("<i>Versuch die Metamorphose der Pflanzen zu erklären</i>").<sup id="cite_ref-D2_2-0" class="reference"><a href="#cite_note-D2-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> where Goethe wrote: </p> <style data-mw-deduplicate="TemplateStyles:r1244412712">.mw-parser-output .templatequote{overflow:hidden;margin:1em 0;padding:0 32px}.mw-parser-output .templatequotecite{line-height:1.5em;text-align:left;margin-top:0}@media(min-width:500px){.mw-parser-output .templatequotecite{padding-left:1.6em}}</style><blockquote class="templatequote"><p>"...we may equally well say that a stamen is a contracted petal, as that a petal is a stamen in a state of expansion; or that a sepal is a contracted stem leaf approaching a certain stage of refinement, as that a stem leaf is a sepal expanded by the influx of cruder saps".<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup></p></blockquote> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Floral_transition">Floral transition</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=1" title="Edit section: Floral transition"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Vegetative_phase_change" title="Vegetative phase change">transition from</a> the <a href="/wiki/Vegetative_phase" class="mw-redirect" title="Vegetative phase">vegetative phase</a> to a <a href="/wiki/Flower" title="Flower">reproductive phase</a> involves a dramatic change in the plant's vital cycle, perhaps the most important one, as the process must be carried out correctly in order to guarantee that the plant produces <a href="/wiki/Kinship" title="Kinship">descendants</a>. This transition is characterised by the induction and development of the meristem of the inflorescence, which will produce a collection of flowers or one flower. This <a href="/wiki/Morphogenesis" title="Morphogenesis">morphogenetic</a> change contains both endogenous and exogenous elements: For example, in order for the change to be initiated the plant must have a certain number of <a href="/wiki/Leaf" title="Leaf">leaves</a> and contain a certain level of total <a href="/wiki/Biomass" title="Biomass">biomass</a>. Certain environmental conditions are also required such as a characteristic <a href="/wiki/Photoperiod" class="mw-redirect" title="Photoperiod">photoperiod</a>. <a href="/wiki/Plant_hormone" title="Plant hormone">Plant hormones</a> play an important part in the process, with the <a href="/wiki/Gibberellin" title="Gibberellin">gibberellins</a> having a particularly important role.<sup id="cite_ref-blázquez_4-0" class="reference"><a href="#cite_note-blázquez-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>There are many <a href="/wiki/Signal_transduction" title="Signal transduction">signals</a> that regulate the <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a> of the process. The following three genes in <i><a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a></i> possess both common and independent functions in floral transition: <i>FLOWERING LOCUS T</i> (<i>FT</i>), <i>LEAFY</i> (<i>LFY</i>), <i>SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1</i> (<i>SOC1</i>, also called <i>AGAMOUS-LIKE20</i>).<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> <i>SOC1</i> is a <a href="/wiki/MADS-box" title="MADS-box">MADS-box</a>-type gene, which integrates responses to photoperiod, <a href="/wiki/Vernalization" title="Vernalization">vernalization</a> and gibberellins.<sup id="cite_ref-blázquez_4-1" class="reference"><a href="#cite_note-blázquez-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Formation_of_the_floral_meristem_or_the_inflorescence">Formation of the floral meristem or the inflorescence</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=2" title="Edit section: Formation of the floral meristem or the inflorescence"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Meristem" title="Meristem">meristem</a> can be defined as the tissue or group of plant tissues that contain undifferentiated <a href="/wiki/Stem_cells" class="mw-redirect" title="Stem cells">stem cells</a>, which are capable of producing any type of cell tissue. Their maintenance and development, both in the vegetative meristem or the meristem of the inflorescence is controlled by genetic <a href="/wiki/Cell_fate_determination" title="Cell fate determination">cell fate determination</a> mechanisms. This means that a number of genes will directly regulate, for example, the maintenance of the stem cell's characteristics (gene <i>WUSCHEL</i> or <i>WUS</i>), and others will act via <a href="/wiki/Negative_feedback" title="Negative feedback">negative feedback</a> mechanisms in order to inhibit a characteristic (gene <i>CLAVATA</i> or <i>CLV</i>). In this way both mechanisms give rise to a <a href="/wiki/Feedback_loop" class="mw-redirect" title="Feedback loop">feedback loop</a>, which along with other elements lend a great deal of robustness to the system.<sup id="cite_ref-Brand_2000_6-0" class="reference"><a href="#cite_note-Brand_2000-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Along with the <i>WUS</i> gene the <i>SHOOTMERISTEMLESS</i> (<i>STM</i>) gene also represses the differentiation of the meristematic dome. This gene acts by inhibiting the possible <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiation</a> of the stem cells but still allows <a href="/wiki/Cell_division" title="Cell division">cell division</a> in the daughter cells, which, had they been allowed to differentiate, would have given rise to distinct organs.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Floral_architecture">Floral architecture</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=3" title="Edit section: Floral architecture"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Mature_flower_diagram.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Mature_flower_diagram.svg/330px-Mature_flower_diagram.svg.png" decoding="async" width="330" height="190" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Mature_flower_diagram.svg/495px-Mature_flower_diagram.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Mature_flower_diagram.svg/660px-Mature_flower_diagram.svg.png 2x" data-file-width="693" data-file-height="400" /></a><figcaption>Anatomy of a flower</figcaption></figure> <p>A flower's anatomy, as defined by the presence of a series of organs (sepals, petals, stamens and carpels) positioned according to a given pattern, facilitate <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual reproduction</a> in <a href="/wiki/Flowering_plant" title="Flowering plant">flowering plants</a>. The flower arises from the activity of three classes of genes, which regulate floral development:<sup id="cite_ref-taiz_8-0" class="reference"><a href="#cite_note-taiz-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <ul><li><b>Meristem identity genes</b>, which code for the <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> required to initiate the induction of the identity genes. They are positive regulators of organ identity during floral development.</li> <li><b>Organ identity genes</b>, which directly control organ identity and also code for transcription factors that control the expression of other genes, whose products are implicated in the formation or function of the distinct organs of the flower.</li> <li><b>Cadastral genes</b>, which act as spatial regulators for the organ identity genes by defining boundaries for their expression. In this way they control the extent to which genes interact thereby regulating whether they act in the same place at the same time.</li></ul> <div class="mw-heading mw-heading3"><h3 id="The_ABC_model">The ABC model</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=4" title="Edit section: The ABC model"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ABC model of flower development was first formulated by George Haughn and Chris Somerville in 1988.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> It was first used as a model to describe the collection of genetic mechanisms that establish floral organ identity in the <a href="/wiki/Rosids" title="Rosids">Rosids</a>, as exemplified by <i>Arabidopsis thaliana</i>, and the <a href="/wiki/Asterids" title="Asterids">Asterids</a>, as demonstrated by <i>Antirrhinum majus</i>. Both species have four verticils (sepals, petals, stamens and carpels), which are defined by the differential expression of a number of <a href="/wiki/Homeotic_gene" title="Homeotic gene">homeotic genes</a> present in each verticil. This means that the sepals are solely characterized by the expression of A genes, while the petals are characterized by the co-expression of A and B genes. The B and C genes establish the identity of the stamens and the carpels only require C genes to be active. Type A and C genes are reciprocally antagonistic.<sup id="cite_ref-Bowman_10-0" class="reference"><a href="#cite_note-Bowman-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>The fact that these homeotic genes determine an organ's identity becomes evident when a gene that represents a particular function, for example the A gene, is not expressed. In <i>Arabidopsis</i> this loss results in a flower which is composed of one verticil of carpels, another containing stamens and another of carpels.<sup id="cite_ref-Bowman_10-1" class="reference"><a href="#cite_note-Bowman-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> This method for studying gene function uses <a href="/wiki/Reverse_genetics" title="Reverse genetics">reverse genetics</a> techniques to produce transgenic plants that contain a mechanism for <a href="/wiki/Gene_silencing" title="Gene silencing">gene silencing</a> through <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a>. In other studies, using <a href="/wiki/Forward_genetics" title="Forward genetics">forward genetics</a> techniques such as <a href="/wiki/Genetic_mapping" class="mw-redirect" title="Genetic mapping">genetic mapping</a>, it is the analysis of the <a href="/wiki/Phenotype" title="Phenotype">phenotypes</a> of flowers with structural anomalies that leads to the <a href="/wiki/Cloning" title="Cloning">cloning</a> of the gene of interest. The flowers may possess a non-functional or <a href="/wiki/Gene_expression" title="Gene expression">over expressed</a> <a href="/wiki/Allele" title="Allele">allele</a> for the gene being studied.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>The existence of two supplementary functions, D and E, have also been proposed in addition to the A, B and C functions already discussed. Function D specifies the identity of the <a href="/wiki/Ovule" title="Ovule">ovule</a>, as a separate reproductive function from the development of the carpels, which occurs after their determination.<sup id="cite_ref-Angenent_12-0" class="reference"><a href="#cite_note-Angenent-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Function E relates to a physiological requirement that is a characteristic of all floral verticils, although, it was initially described as necessary for the development of the three innermost verticils (Function E <i>sensu stricto</i>).<sup id="cite_ref-pelaz_13-0" class="reference"><a href="#cite_note-pelaz-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> However, its broader definition (<i>sensu lato</i>) suggests that it is required in the four verticils.<sup id="cite_ref-ditta_14-0" class="reference"><a href="#cite_note-ditta-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Therefore, when Function D is lost the structure of the ovules becomes similar to that of leaves and when Function E is lost <i>sensu stricto</i>, the floral organs of the three outer most verticils are transformed into sepals,<sup id="cite_ref-pelaz_13-1" class="reference"><a href="#cite_note-pelaz-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> while on losing Function E <i>sensu lato</i>, all the verticils are similar to leaves.<sup id="cite_ref-ditta_14-1" class="reference"><a href="#cite_note-ditta-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Gene_product" title="Gene product">gene products</a> of genes with D and E functions are also MADS-box genes.<sup id="cite_ref-Ma_(2005)_15-0" class="reference"><a href="#cite_note-Ma_(2005)-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genetic_analysis">Genetic analysis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=5" title="Edit section: Genetic analysis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Arabidopsis_thaliana-flower.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Arabidopsis_thaliana-flower.jpg/220px-Arabidopsis_thaliana-flower.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Arabidopsis_thaliana-flower.jpg/330px-Arabidopsis_thaliana-flower.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/91/Arabidopsis_thaliana-flower.jpg/440px-Arabidopsis_thaliana-flower.jpg 2x" data-file-width="600" data-file-height="600" /></a><figcaption>Flower of <i>A. thaliana</i>.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Antirrhinum_majus6.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Antirrhinum_majus6.jpg/220px-Antirrhinum_majus6.jpg" decoding="async" width="220" height="293" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Antirrhinum_majus6.jpg/330px-Antirrhinum_majus6.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Antirrhinum_majus6.jpg/440px-Antirrhinum_majus6.jpg 2x" data-file-width="1512" data-file-height="2016" /></a><figcaption>Flowers of <i>A. majus</i>.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Petunia_x_hybrida_a1.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Petunia_x_hybrida_a1.JPG/220px-Petunia_x_hybrida_a1.JPG" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Petunia_x_hybrida_a1.JPG/330px-Petunia_x_hybrida_a1.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Petunia_x_hybrida_a1.JPG/440px-Petunia_x_hybrida_a1.JPG 2x" data-file-width="1916" data-file-height="1437" /></a><figcaption>Flowers of <i>Petunia hybrid</i>.</figcaption></figure> <p>The methodology for studying flower development involves two steps. Firstly, the identification of the exact genes required for determining the identity of the floral meristem. In <i>A. thaliana</i> these include APETALA1 (<i>AP1</i>) and LEAFY (<i>LFY</i>). Secondly, genetic analysis is carried out on the aberrant <a href="/wiki/Phenotype" title="Phenotype">phenotypes</a> for the relative characteristics of the flowers, which allows the characterization of the <a href="/wiki/Homeotic_genes" class="mw-redirect" title="Homeotic genes">homeotic genes</a> implicated in the process. </p> <div class="mw-heading mw-heading4"><h4 id="Analysis_of_mutants">Analysis of mutants</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=6" title="Edit section: Analysis of mutants"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are a great many <a href="/wiki/Mutation" title="Mutation">mutations</a> that affect floral <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>, although the analysis of these mutants is a recent development. Supporting evidence for the existence of these mutations comes from the fact that a large number affect the identity of floral organs. For example, some organs develop in a location where others should develop. This is called <a href="/wiki/Homeotic_mutation" class="mw-redirect" title="Homeotic mutation">homeotic mutation</a>, which is analogous to <a href="/wiki/Hox_gene#Genes_regulated_by_Hox_proteins" title="Hox gene">HOX gene mutations</a> found in <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i>. In <i>Arabidopsis</i> and <i><a href="/wiki/Antirrhinum" title="Antirrhinum">Antirrhinum</a></i>, the two taxa on which models are based, these mutations always affect adjacent verticils. This allows the characterization of three classes of mutation, according to which verticils are affected: </p> <ul><li>Mutations in type A genes – These mutations affect the calyx and corolla, which are the outermost verticils. In these mutants, such as APETALA2 in <i>A. thaliana</i>, carpels develop instead of sepals and stamen in place of petals. This means that, the verticils of the <a href="/wiki/Perianth" title="Perianth">perianth</a> are transformed into reproductive verticils.</li> <li>Mutations in type B genes – These mutations affect the corolla and the stamen, which are the intermediate verticils. Two mutations have been found in <i>A. thaliana</i>, APETALA3 and PISTILLATA, which cause development of sepals instead of petals and carpels in the place of stamen.</li> <li>Mutations in type C genes – These mutations affect the reproductive verticils, namely the stamen and the carpels. The <i>A. thaliana</i> mutant of this type is called AGAMOUS, it possesses a phenotype containing petals instead of stamen and sepals instead of carpels.</li></ul> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Arabidopsis_mutants.jpg" class="mw-file-description" title="AGAMOUS mutation resulting in a double-flowered Arabidopsis"><img alt="AGAMOUS mutation resulting in a double-flowered Arabidopsis" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/Arabidopsis_mutants.jpg/120px-Arabidopsis_mutants.jpg" decoding="async" width="120" height="90" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/Arabidopsis_mutants.jpg/180px-Arabidopsis_mutants.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0a/Arabidopsis_mutants.jpg/240px-Arabidopsis_mutants.jpg 2x" data-file-width="900" data-file-height="675" /></a></span></div> <div class="gallerytext"> AGAMOUS mutation resulting in a <a href="/wiki/Double-flowered" title="Double-flowered">double-flowered</a> <a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis</a></div> </li> </ul> <div class="mw-heading mw-heading4"><h4 id="Techniques_for_detecting_differential_expression">Techniques for detecting differential expression</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=7" title="Edit section: Techniques for detecting differential expression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Cloning" title="Cloning">Cloning</a> studies have been carried out on <a href="/wiki/DNA" title="DNA">DNA</a> in the genes associated with the affected homeotic functions in the mutants discussed above. These studies used <a href="/wiki/Serial_analysis_of_gene_expression" title="Serial analysis of gene expression">serial analysis of gene expression</a> throughout floral development to show patterns of tissue expression, which, in general, correspond with the predictions of the ABC model. </p><p>The nature of these genes corresponds to that of <a href="/wiki/Transcription_factors" class="mw-redirect" title="Transcription factors">transcription factors</a>, which, as expected, have analogous structures to a group of factors contained in <a href="/wiki/Yeast" title="Yeast">yeasts</a> and <a href="/wiki/Animal_cells" class="mw-redirect" title="Animal cells">animal cells</a>. This group is called MADS, which is an acronym for the different factors contained in the group. These MADS factors have been detected in all the vegetable species studied, although the involvement of other elements involved in the <a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">regulation of gene expression</a> cannot be discounted.<sup id="cite_ref-taiz_8-1" class="reference"><a href="#cite_note-taiz-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genes_exhibiting_type-A_function">Genes exhibiting type-A function</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=8" title="Edit section: Genes exhibiting type-A function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In <i>A. thaliana</i>, function A is mainly represented by two genes <i>APETALA1</i> (<i>AP1)</i> and <i>APETALA2</i> (<i>AP2</i>)<sup id="cite_ref-Bowman_1989_16-0" class="reference"><a href="#cite_note-Bowman_1989-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> <i>AP1</i> is a MADS-box type gene, while <i>AP2</i> belongs to the family of genes that contains AP2, which it gives its name to and which consists of <a href="/wiki/Transcription_factors" class="mw-redirect" title="Transcription factors">transcription factors</a> that are only found in plants.<sup id="cite_ref-Jofuku_1994_17-0" class="reference"><a href="#cite_note-Jofuku_1994-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> AP2 has also been shown to complex with the co-repressor TOPLESS (TPL) in developing floral buds to repress the C-class gene <i>AGAMOUS</i> (<i>AG</i>).<sup id="cite_ref-APETALA2_18-0" class="reference"><a href="#cite_note-APETALA2-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> However, <i>AP2</i> is not expressed in the shoot apical meristem (SAM), which contains the latent stem cell population throughout the adult life of <i>Arabidopsis</i>, and so it is speculated that TPL works with some other A-class gene in the SAM to repress <i>AG</i>.<sup id="cite_ref-APETALA2_18-1" class="reference"><a href="#cite_note-APETALA2-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><i>AP1</i> functions as a type A gene, both in controlling the identity of sepals and petals, and it also acts in the <a href="/wiki/Floral_meristem" class="mw-redirect" title="Floral meristem">floral meristem</a>. <i>AP2</i> not only functions in the first two verticils, but also in the remaining two, in developing ovules and even in leaves. It is also likely that <a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">post-transcriptional regulation</a> exists, which controls its A function, or even that it has other purposes in the determination of organ identity independent of that mentioned here.<sup id="cite_ref-Jofuku_1994_17-1" class="reference"><a href="#cite_note-Jofuku_1994-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>In <i>Antirrhinum</i>, the <a href="/wiki/Homology_(biology)" title="Homology (biology)">orthologous gene</a> to <i>AP1</i> is <i>SQUAMOSA</i> (<i>SQUA</i>), which also has a particular impact on the floral meristem. The homologs for <i>AP2</i> are <i>LIPLESS1</i> (<i>LIP1</i>) and <i>LIPLESS2</i> (<i>LIP2</i>), which have a redundant function and are of special interest in the development of sepals, petals and ovules.<sup id="cite_ref-Keck_2003_19-0" class="reference"><a href="#cite_note-Keck_2003-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p>A total of three genes have been isolated from <i>Petunia hybrida</i> that are similar to <i>AP2</i>: <i>P. hybrida APETALA2A</i> (<i>PhAP2A</i>), <i>PhAP2B</i> and <i>PhAP2C</i>. <i>PhAP2A</i> is, to a large degree, homologous with the <i>AP2</i> gene of <i>Arabidopsis</i>, both in its sequence and in its expression pattern, which suggests that the two genes are orthologs. The proteins <i>PhAP2B</i> and <i>PhAP2C</i>, on the other hand, are slightly different, even though they belong to the family of transcription factors that are similar to <i>AP2</i>. In addition they are expressed in different ways, although they are very similar in comparison with <i>PhAP2A</i>. In fact, the mutants for these genes do not show the usual phenotype, that of the <a href="/wiki/Null_alleles" class="mw-redirect" title="Null alleles">null alleles</a> of A genes.<sup id="cite_ref-Maes_2001_20-0" class="reference"><a href="#cite_note-Maes_2001-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> A true A-function gene has not been found in Petunia; though a part of the A-function (the inhibition of the C in the outer two whorls) has been largely attributed to miRNA169 (colloquially called BLIND)<sup>ref</sup>. </p> <div class="mw-heading mw-heading3"><h3 id="Genes_exhibiting_type-B_function">Genes exhibiting type-B function</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=9" title="Edit section: Genes exhibiting type-B function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In <i>A. thaliana</i> the type-B function mainly arises from two genes, <i>APETALA3</i> (<i>AP3</i>) and <i>PISTILLATA</i> (<i>PI</i>), both of which are MADS-box genes. A mutation of either of these genes causes the homeotic conversion of petals into sepals and of stamens into carpels.<sup id="cite_ref-autogenerated1_21-0" class="reference"><a href="#cite_note-autogenerated1-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> This also occurs in its orthologs in <i>A. majus</i>, which are <span class="anchor" id="DEFICIENS"></span>DEFICIENS (<i>DEF</i>) and <i>GLOBOSA</i> (<i>GLO</i>) respectively.<sup id="cite_ref-Sommer_1990_22-0" class="reference"><a href="#cite_note-Sommer_1990-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> For both species the active form of binding with DNA is that derived from the heterodimer: AP3 and PI, or DEF and GLO, <a href="/wiki/Dimer_(chemistry)" class="mw-redirect" title="Dimer (chemistry)">dimerize</a>. This is the form in which they are able to function.<sup id="cite_ref-Riechmann_1996_23-0" class="reference"><a href="#cite_note-Riechmann_1996-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>The <i>GLO</i>/<i>PI</i> lines that have been duplicated in <i>Petunia</i> contain <i>P. hybrida GLOBOSA1</i> (<i>PhGLO1</i>, also called <i>FBP1</i>) and also <i>PhGLO2</i> (also called <i>PMADS2</i> or <i>FBP3</i>). For the functional elements equivalent to <i>AP3</i>/<i>DEF</i> in <i>Petunia</i> there is both a gene that possesses a relatively similar sequence, called <i>PhDEF</i> and there is also an atypical B function gene called PhTM6. <a href="/wiki/Phylogenetic" class="mw-redirect" title="Phylogenetic">Phylogenetic</a> studies have placed the first three within the «euAP3» lineage, while PhTM6 belongs to that of «paleoAP3».<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> It is worth pointing out that, in terms of evolutionary history, the appearance of the euAP3 line seems to be related with the emergence of <a href="/wiki/Dicotyledon" title="Dicotyledon">dicotyledons</a>, as representatives of euAP3-type B function genes are present in dicotyledons while paleoAP3 genes are present in monocotyledons and basal angiosperms, among others.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>As discussed above, the floral organs of eudicotyledonous angiosperms are arranged in 4 different verticils, containing the sepals, petals, stamen and carpels. The ABC model states that the identity of these organs is determined by the homeotic genes A, A+B, B+C and C, respectively. In contrast with the sepal and petal verticils of the eudicots, the perigone of many plants of the family <a href="/wiki/Liliaceae" title="Liliaceae">Liliaceae</a> have two nearly identical external petaloid verticils (the <a href="/wiki/Tepal" title="Tepal">tepals</a>). In order to explain the floral morphology of the Liliaceae, van Tunen <i>et al.</i> proposed a modified ABC model in 1993. This model suggests that class B genes are not only expressed in verticils 2 and 3, but also in 1. It therefore follows that the organs of verticils 1 and 2 express class A and B genes and this is how they have a petaloid structure. This theoretical model has been experimentally proven through the cloning and characterization of homologs of the <i>Antirrhinum</i> genes <i>GLOBOSA</i> and <i>DEFICIENS</i> in a Liliaceae, the <a href="/wiki/Tulip" title="Tulip">tulip</a> <i>Tulipa gesneriana</i>. These genes are expressed in verticils 1,2 and 3.<sup id="cite_ref-Akira_Kanno_26-0" class="reference"><a href="#cite_note-Akira_Kanno-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> The homologs <i>GLOBOSA</i> and <i>DEFICIENS</i> have also been isolated and characterized in <i>Agapanthus praecox</i> ssp. <i>orientalis</i> (<a href="/wiki/Agapanthaceae" class="mw-redirect" title="Agapanthaceae">Agapanthaceae</a>), which is phylogenetically distant from the model organisms. In this study the genes were called <i>ApGLO</i> and <i>ApDEF</i>, respectively. Both contain <a href="/wiki/Open_reading_frame" title="Open reading frame">open reading frames</a> that code for proteins with 210 to 214 <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a>. Phylogenetic analysis of these sequences indicated that they belong to B gene family of the <a href="/wiki/Monocotyledon" title="Monocotyledon">monocotyledons</a>. <a href="/wiki/In_situ_hybridization" title="In situ hybridization">In situ hybridization</a> studies revealed that both sequences are expressed in verticil 1 as well as in 2 and 3. When taken together, these observations show that the floral development mechanism of <i><a href="/wiki/Agapanthus" title="Agapanthus">Agapanthus</a></i> also follows the modified ABC model.<sup id="cite_ref-Toru_Nakamura_27-0" class="reference"><a href="#cite_note-Toru_Nakamura-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genes_exhibiting_type-C_function">Genes exhibiting type-C function</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=10" title="Edit section: Genes exhibiting type-C function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In <i>A. thaliana</i>, the C function is derived from one MADS-box type gene called <i>AGAMOUS</i> (<i>AG</i>), which intervenes both in the establishment of stamen and carpel identity as well as in the determination of the floral meristem.<sup id="cite_ref-Bowman_1989_16-1" class="reference"><a href="#cite_note-Bowman_1989-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Therefore, the <i>AG</i> mutants are devoid of <a href="/wiki/Androecium" class="mw-redirect" title="Androecium">androecium</a> and <a href="/wiki/Gynoecium" title="Gynoecium">gynoecium</a> and they have petals and sepals in their place. In addition, the growth in the centre of the flower is undifferentiated, therefore the petals and sepals grow in repetitive verticils. </p><p>The <i>PLENA</i> (<i>PLE</i>) gene is present in <i>A. majus</i>, in place of the <i>AG</i> gene, although it is not an ortholog. However, the <i>FARINELLI</i> (<i>FAR</i>) gene is an ortholog, which is specific to the development of the <a href="/wiki/Anther" class="mw-redirect" title="Anther">anthers</a> and the maturation of <a href="/wiki/Pollen" title="Pollen">pollen</a>.<sup id="cite_ref-brendan_1999_28-0" class="reference"><a href="#cite_note-brendan_1999-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>In <i>Petunia</i>, <i>Antirrhinum</i> and in <a href="/wiki/Maize" title="Maize">maize</a> the C function is controlled by a number of genes that act in the same manner. The genes that are closer homologs of <i>AG</i> in <i>Petunia</i> are <i>pMADS3</i> and <i>floral-binding protein 6</i> (<i>FBP6</i>).<sup id="cite_ref-brendan_1999_28-1" class="reference"><a href="#cite_note-brendan_1999-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genes_exhibiting_type-D_and_E_functions">Genes exhibiting type-D and E functions</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=11" title="Edit section: Genes exhibiting type-D and E functions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The D function genes were discovered in 1995. These genes are MADS-box proteins and they have a function that is distinct from those previously described, although they have a certain homology with C function genes. These genes are called <i>FLORAL BINDING PROTEIN7</i> (<i>FBP7</i>) and <i>FLORAL BINDING PROTEIN1L</i> (<i>FBP1l</i>).<sup id="cite_ref-Angenent_12-1" class="reference"><a href="#cite_note-Angenent-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> It was found that, in <i>Petunia</i>, they are involved in the development of the ovule. Equivalent genes were later found in <i>Arabidopsis</i>,<sup id="cite_ref-Favaro_29-0" class="reference"><a href="#cite_note-Favaro-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> where they are also involved in controlling the development of carpels and the ovule and even with structures related to <a href="/wiki/Seed_dispersal" title="Seed dispersal">seed dispersal</a>. </p><p>The appearance of interesting phenotypes in <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a> studies in <i>Petunia</i> and <a href="/wiki/Tomato" title="Tomato">tomato</a> led, in 1994, to the definition of a new type of function in the floral development model. The E function was initially thought to be only involved in the development of the three innermost verticils, however, subsequent work found that its expression was required in all the floral verticils.<sup id="cite_ref-pelaz_13-2" class="reference"><a href="#cite_note-pelaz-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=12" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Flower" title="Flower">Flower</a></li> <li><a href="/wiki/MADS-box" title="MADS-box">MADS-box</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Plant_evolutionary_developmental_biology" title="Plant evolutionary developmental biology">Plant evolutionary developmental biology</a></li> <li><a href="/wiki/Superman_(gene)" title="Superman (gene)">Superman (gene)</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=13" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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-azcón-bieto-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-azcón-bieto_1-0">^</a></b></span> <span class="reference-text"><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="CITEREFAzcón-Bieto2000" class="citation book cs1">Azcón-Bieto; et al. (2000). <i>Fundamentos de fisiología vegetal</i>. McGraw-Hill/Interamericana de España, SAU. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-84-486-0258-1" title="Special:BookSources/978-84-486-0258-1"><bdi>978-84-486-0258-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Fundamentos+de+fisiolog%C3%ADa+vegetal&rft.pub=McGraw-Hill%2FInteramericana+de+Espa%C3%B1a%2C+SAU&rft.date=2000&rft.isbn=978-84-486-0258-1&rft.au=Azc%C3%B3n-Bieto&rfr_id=info%3Asid%2Fen.wikipedia.org%3AABC+model+of+flower+development" class="Z3988"></span><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (April 2013)">page needed</span></a></i>]</sup></span> </li> <li id="cite_note-D2-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-D2_2-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDornelasDornelas2005" class="citation journal cs1">Dornelas, Marcelo Carnier; Dornelas, Odair (2005). <a rel="nofollow" class="external text" href="https://doi.org/10.1590%2FS1677-04202005000400001">"From leaf to flower: Revisiting Goethe's concepts on the ¨metamorphosis¨ of plants"</a>. <i>Brazilian Journal of Plant Physiology</i>. <b>17</b> (4): 335–344. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1590%2FS1677-04202005000400001">10.1590/S1677-04202005000400001</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Brazilian+Journal+of+Plant+Physiology&rft.atitle=From+leaf+to+flower%3A+Revisiting+Goethe%27s+concepts+on+the+%C2%A8metamorphosis%C2%A8+of+plants&rft.volume=17&rft.issue=4&rft.pages=335-344&rft.date=2005&rft_id=info%3Adoi%2F10.1590%2FS1677-04202005000400001&rft.aulast=Dornelas&rft.aufirst=Marcelo+Carnier&rft.au=Dornelas%2C+Odair&rft_id=https%3A%2F%2Fdoi.org%2F10.1590%252FS1677-04202005000400001&rfr_id=info%3Asid%2Fen.wikipedia.org%3AABC+model+of+flower+development" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">Goethe J.W. von (1790) Versuch die Metamorphose der Pflanzen zu erklaren. 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Motte, Patrick; Keck, Emma; Saedler, Heinz; Sommer, Hans; Schwarz-Sommer, Zsuzsanna (1999). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1171478">"PLENA and FARINELLI: Redundancy and regulatory interactions between two Antirrhinum MADS-box factors controlling flower development"</a>. <i>The EMBO Journal</i>. <b>18</b> (14): 4023–34. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Femboj%2F18.14.4023">10.1093/emboj/18.14.4023</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1171478">1171478</a></span>. <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/10406807">10406807</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+EMBO+Journal&rft.atitle=PLENA+and+FARINELLI%3A+Redundancy+and+regulatory+interactions+between+two+Antirrhinum+MADS-box+factors+controlling+flower+development&rft.volume=18&rft.issue=14&rft.pages=4023-34&rft.date=1999&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1171478%23id-name%3DPMC&rft_id=info%3Apmid%2F10406807&rft_id=info%3Adoi%2F10.1093%2Femboj%2F18.14.4023&rft.aulast=Davies&rft.aufirst=Brendan&rft.au=Motte%2C+Patrick&rft.au=Keck%2C+Emma&rft.au=Saedler%2C+Heinz&rft.au=Sommer%2C+Hans&rft.au=Schwarz-Sommer%2C+Zsuzsanna&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1171478&rfr_id=info%3Asid%2Fen.wikipedia.org%3AABC+model+of+flower+development" class="Z3988"></span></span> </li> <li id="cite_note-Favaro-29"><span class="mw-cite-backlink"><b><a href="#cite_ref-Favaro_29-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFavaroPinyopichBattagliaKooiker2003" class="citation journal cs1">Favaro, R; Pinyopich, A; Battaglia, R; Kooiker, M; Borghi, L; Ditta, G; Yanofsky, MF; Kater, MM; Colombo, L (2003). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC280564">"MADS-box protein complexes control carpel and ovule development in Arabidopsis"</a>. <i>The Plant Cell</i>. <b>15</b> (11): 2603–11. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1105%2Ftpc.015123">10.1105/tpc.015123</a>. <a href="/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/3872057">3872057</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC280564">280564</a></span>. <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/14555696">14555696</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Plant+Cell&rft.atitle=MADS-box+protein+complexes+control+carpel+and+ovule+development+in+Arabidopsis&rft.volume=15&rft.issue=11&rft.pages=2603-11&rft.date=2003&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC280564%23id-name%3DPMC&rft_id=info%3Apmid%2F14555696&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F3872057%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.1105%2Ftpc.015123&rft.aulast=Favaro&rft.aufirst=R&rft.au=Pinyopich%2C+A&rft.au=Battaglia%2C+R&rft.au=Kooiker%2C+M&rft.au=Borghi%2C+L&rft.au=Ditta%2C+G&rft.au=Yanofsky%2C+MF&rft.au=Kater%2C+MM&rft.au=Colombo%2C+L&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC280564&rfr_id=info%3Asid%2Fen.wikipedia.org%3AABC+model+of+flower+development" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Sources">Sources</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=14" title="Edit section: Sources"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="General_texts">General texts</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=15" title="Edit section: General texts"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSoltisSoltisLeebens-Mack2006" class="citation book cs1">Soltis, DE; <a href="/wiki/Pamela_S._Soltis" title="Pamela S. Soltis">Soltis, PS</a>; Leebens-Mack, J, eds. (2006). <i>Advances in botanical research: Developmental genetics of the flower</i>. New York, NY: Academic Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-005944-7" title="Special:BookSources/978-0-12-005944-7"><bdi>978-0-12-005944-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Advances+in+botanical+research%3A+Developmental+genetics+of+the+flower&rft.pub=New+York%2C+NY%3A+Academic+Press&rft.date=2006&rft.isbn=978-0-12-005944-7&rfr_id=info%3Asid%2Fen.wikipedia.org%3AABC+model+of+flower+development" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWolpertBeddingtonJessellLawrence2002" class="citation book cs1">Wolpert, Lewis; Beddington, R.; Jessell, T.; Lawrence, P.; Meyerowitz, E.; Smith, W. (2002). <i>Principles of Development</i> (Second ed.). Oxford: Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-879291-8" title="Special:BookSources/978-0-19-879291-8"><bdi>978-0-19-879291-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Principles+of+Development&rft.place=Oxford&rft.edition=Second&rft.pub=Oxford+University+Press&rft.date=2002&rft.isbn=978-0-19-879291-8&rft.aulast=Wolpert&rft.aufirst=Lewis&rft.au=Beddington%2C+R.&rft.au=Jessell%2C+T.&rft.au=Lawrence%2C+P.&rft.au=Meyerowitz%2C+E.&rft.au=Smith%2C+W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AABC+model+of+flower+development" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ABC_model_of_flower_development&action=edit&section=16" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMurai,_K.2013" class="citation journal cs1">Murai, K. (2013). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4844379">"Homeotic Genes and the ABCDE Model for Floral Organ Formation in Wheat"</a>. <i>Plants</i>. <b>2</b> (3): 379–395. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.3390%2Fplants2030379">10.3390/plants2030379</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4844379">4844379</a></span>. <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/27137382">27137382</a>.</cite><span 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botany">History</a></li> <li><a href="/wiki/Outline_of_botany" title="Outline of botany">Outline</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Branches_of_botany" title="Branches of botany">Subdisciplines</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaeobotany" class="mw-redirect" title="Archaeobotany">Archaeobotany</a></li> <li><a href="/wiki/Astrobotany" title="Astrobotany">Astrobotany</a></li> <li><a href="/wiki/Bryology" title="Bryology">Bryology</a></li> <li><a href="/wiki/Dendrology" title="Dendrology">Dendrology</a></li> <li><a href="/wiki/Ethnobotany" title="Ethnobotany">Ethnobotany</a></li> <li><a href="/wiki/Paleobotany" title="Paleobotany">Paleobotany</a></li> <li><a href="/wiki/Phycology" title="Phycology">Phycology</a></li> <li><a href="/wiki/Phytochemistry" title="Phytochemistry">Phytochemistry</a></li> <li><a href="/wiki/Phytogeography" title="Phytogeography">Phytogeography</a> <ul><li><a href="/wiki/Geobotanical_prospecting" title="Geobotanical prospecting">Geobotany</a></li></ul></li> <li><a href="/wiki/Plant_anatomy" title="Plant anatomy">Plant anatomy</a></li> <li><a href="/wiki/Plant_ecology" title="Plant ecology">Plant ecology</a></li> <li><a href="/wiki/Plant_pathology" title="Plant pathology">Plant pathology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant" title="Plant">Plant</a> groups</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Algae" title="Algae">Algae</a></li> <li><a href="/wiki/Archaeplastida" title="Archaeplastida">Archaeplastida</a></li> <li><a href="/wiki/Bryophyte" title="Bryophyte">Bryophyte</a></li> <li><a href="/wiki/Non-vascular_plant" title="Non-vascular plant">Non-vascular plants</a></li> <li><a href="/wiki/Vascular_plant" title="Vascular plant">Vascular plants</a></li> <li><a href="/wiki/Fern" title="Fern">Fern</a></li> <li><a href="/wiki/Lycophyte" title="Lycophyte">Lycophyte</a></li> <li><a href="/wiki/Spermatophyte" class="mw-redirect" title="Spermatophyte">Spermatophytes</a></li> <li><a href="/wiki/Gymnosperm" title="Gymnosperm">Gymnosperm</a></li> <li><a href="/wiki/Flowering_plant" title="Flowering plant">Angiosperm</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_anatomy" title="Plant anatomy">Plant anatomy</a> <div class="hlist"><ul><li><a href="/wiki/Plant_morphology" title="Plant morphology">Plant morphology</a><br /><span class="nobold">(<a href="/wiki/Glossary_of_plant_morphology" title="Glossary of plant morphology">glossary</a>)</span></li></ul></div></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%"><a href="/wiki/Plant_cell" title="Plant cell">Plant cells</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_wall" title="Cell wall">Cell wall</a></li> <li><a href="/wiki/Phragmoplast" title="Phragmoplast">Phragmoplast</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li> <li><a href="/wiki/Plasmodesma" title="Plasmodesma">Plasmodesma</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%"><a href="/wiki/Tissue_(biology)" title="Tissue (biology)">Tissues</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cork_cambium" title="Cork cambium">Cork</a></li> <li><a href="/wiki/Ground_tissue" title="Ground tissue">Ground tissue</a> <ul><li><a href="/wiki/Leaf#Mesophyll" title="Leaf">Mesophyll</a></li></ul></li> <li><a href="/wiki/Meristem" title="Meristem">Meristem</a></li> <li><a href="/wiki/Storage_organ" title="Storage organ">Storage organs</a></li> <li><a href="/wiki/Vascular_tissue" title="Vascular tissue">Vascular tissue</a> <ul><li><a href="/wiki/Vascular_bundle" title="Vascular bundle">Vascular bundle</a></li></ul></li> <li><a href="/wiki/Wood" title="Wood">Wood</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%">Vegetative</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bulb" title="Bulb">Bulb</a></li> <li><a href="/wiki/Root" title="Root">Root</a></li> <li><a href="/wiki/Rhizoid" title="Rhizoid">Rhizoid</a></li> <li><a href="/wiki/Rhizome" title="Rhizome">Rhizome</a></li> <li><a href="/wiki/Shoot_(botany)" title="Shoot (botany)">Shoot</a> <ul><li><a href="/wiki/Bud" title="Bud">Bud</a></li> <li><a href="/wiki/Leaf" title="Leaf">Leaf</a> <ul><li><a href="/wiki/Cataphyll" title="Cataphyll">Cataphyll</a></li> <li><a href="/wiki/Petiole_(botany)" title="Petiole (botany)">Petiole</a></li></ul></li> <li><a href="/wiki/Sessility_(botany)" title="Sessility (botany)">Sessility</a></li> <li><a href="/wiki/Plant_stem" title="Plant stem">Stem</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%"><a href="/wiki/Plant_reproductive_morphology" title="Plant reproductive morphology">Reproductive</a><br />(incl. Flower)</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archegonium" title="Archegonium">Archegonium</a></li> <li><a href="/wiki/Antheridium" title="Antheridium">Antheridium</a></li> <li><a href="/wiki/Stamen" title="Stamen">Androecium</a> <ul><li><a href="/wiki/Pollen" title="Pollen">Pollen</a></li> <li><a href="/wiki/Stamen" title="Stamen">Stamen</a> <ul><li><a href="/wiki/Anther" class="mw-redirect" title="Anther">Anther</a></li> <li><a href="/wiki/Stamen#Morphology_and_terminology" title="Stamen">Filament</a></li></ul></li> <li><a href="/wiki/Staminode" title="Staminode">Staminode</a></li> <li><a href="/wiki/Tapetum_(botany)" title="Tapetum (botany)">Tapetum</a></li></ul></li> <li><a href="/wiki/Flower" title="Flower">Flower</a> <ul><li><a href="/wiki/Aestivation_(botany)" title="Aestivation (botany)">Aestivation</a></li> <li><a class="mw-selflink selflink">Flower development</a></li> <li><a href="/wiki/Floral_diagram" title="Floral diagram">Floral diagram</a></li> <li><a href="/wiki/Floral_formula" title="Floral formula">Floral formula</a></li> <li><a href="/wiki/Floral_symmetry" title="Floral symmetry">Floral symmetry</a></li> <li><a href="/wiki/Whorl_(botany)" title="Whorl (botany)">Whorl</a></li></ul></li> <li><a href="/wiki/Fruit" title="Fruit">Fruit</a> <ul><li><a href="/wiki/Fruit_anatomy" class="mw-redirect" title="Fruit anatomy">Anatomy</a></li> <li><a href="/wiki/Berry_(botany)" title="Berry (botany)">Berry</a></li> <li><a href="/wiki/Capsule_(fruit)" title="Capsule (fruit)">Capsule</a></li> <li><a href="/wiki/Nut_(fruit)" title="Nut (fruit)">Nut</a></li> <li><a href="/wiki/Pyrena" title="Pyrena">Pyrena</a></li> <li><a href="/wiki/Seed" title="Seed">Seed</a> <ul><li><a href="/wiki/Seed_dispersal" title="Seed dispersal">Dispersal</a></li> <li><a href="/wiki/Endosperm" title="Endosperm">Endosperm</a></li></ul></li></ul></li> <li><a href="/wiki/Gametophyte" title="Gametophyte">Gametophyte</a></li> <li><a href="/wiki/Column_(botany)" title="Column (botany)">Gynandrium</a></li> <li><a href="/wiki/Gynoecium" title="Gynoecium">Gynoecium</a> <ul><li><a href="/wiki/Carpel" class="mw-redirect" title="Carpel">Carpel</a> <ul><li><a href="/wiki/Ovary_(botany)" title="Ovary (botany)">Ovary</a> <ul><li><a href="/wiki/Locule" title="Locule">Locule</a></li> <li><a href="/wiki/Ovule" title="Ovule">Ovule</a></li></ul></li> <li><a href="/wiki/Stigma_(botany)" title="Stigma (botany)">Stigma</a></li> <li><a href="/wiki/Style_(botany)" title="Style (botany)">Style</a></li></ul></li></ul></li> <li><a href="/wiki/Hypanthium" title="Hypanthium">Hypanthium (Floral cup)</a></li> <li><a href="/wiki/Inflorescence" title="Inflorescence">Inflorescence</a> <ul><li><a href="/wiki/Bract" title="Bract">Bract</a></li> <li><a href="/wiki/Pedicel_(botany)" title="Pedicel (botany)">Pedicellate</a></li> <li><a href="/wiki/Raceme" title="Raceme">Raceme</a></li> <li><a href="/wiki/Umbel" title="Umbel">Umbel</a></li></ul></li> <li><a href="/wiki/Perianth" title="Perianth">Perianth</a> <ul><li><a href="/wiki/Tepal" title="Tepal">Tepal</a></li> <li><a href="/wiki/Petal" title="Petal">Petal</a></li> <li><a href="/wiki/Sepal" title="Sepal">Sepal</a></li></ul></li> <li><a href="/wiki/Embryo#Plant_embryos" title="Embryo">Plant embryo</a></li> <li><a href="/wiki/Receptacle_(botany)" title="Receptacle (botany)">Receptacle</a></li> <li><a href="/wiki/Sporophyll" title="Sporophyll">Sporophyll</a></li> <li><a href="/wiki/Sporophyte" title="Sporophyte">Sporophyte</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%">Surface structures</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Plant_cuticle" title="Plant cuticle">Cuticle</a></li> <li><a href="/wiki/Epicuticular_wax" title="Epicuticular wax">Epicuticular wax</a></li> <li><a href="/wiki/Epidermis_(botany)" title="Epidermis (botany)">Epidermis</a></li> <li><a href="/wiki/Nectar" title="Nectar">Nectar</a></li> <li><a href="/wiki/Stoma" title="Stoma">Stoma</a></li> <li><a href="/wiki/Thorns,_spines,_and_prickles" title="Thorns, spines, and prickles">Thorns, spines, and prickles</a></li> <li><a href="/wiki/Trichome" title="Trichome">Trichome</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_physiology" title="Plant physiology">Plant physiology</a><br />Materials</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Aleurone" title="Aleurone">Aleurone</a></li> <li><a href="/wiki/Apical_dominance" title="Apical dominance">Apical dominance</a></li> <li><a href="/wiki/Bulk_movement" title="Bulk movement">Bulk flow</a></li> <li><a href="/wiki/Cellulose" title="Cellulose">Cellulose</a></li> <li><a href="/wiki/Plant_nutrition" title="Plant nutrition">Nutrition</a></li> <li><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a> <ul><li><a href="/wiki/Chlorophyll" title="Chlorophyll">Chlorophyll</a></li></ul></li> <li><a href="/wiki/Phytomelanin" title="Phytomelanin">Phytomelanin</a></li> <li><a href="/wiki/Plant_hormone" title="Plant hormone">Plant hormones</a></li> <li><a href="/wiki/Respiratory_system#Plants" title="Respiratory system">Respiration</a> <ul><li><a href="/wiki/Gas_exchange#Plants" title="Gas exchange">Gas Exchange</a></li> <li><a href="/wiki/Cellular_respiration" title="Cellular respiration">Cellular respiration</a></li></ul></li> <li><a href="/wiki/Sap" title="Sap">Sap</a></li> <li><a href="/wiki/Starch" title="Starch">Starch</a></li> <li><a href="/wiki/Sugar" title="Sugar">Sugar</a></li> <li><a href="/wiki/Transpiration" title="Transpiration">Transpiration</a></li> <li><a href="/wiki/Turgor_pressure" title="Turgor pressure">Turgor pressure</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%">Plant growth<br />and habit</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Habit_(biology)#Structure" title="Habit (biology)">Habit</a> <ul><li><a href="/wiki/Cushion_plant" title="Cushion plant">Cushion plants</a></li> <li><a href="/wiki/Rosette_(botany)" title="Rosette (botany)">Rosettes</a></li> <li><a href="/wiki/Shrub" title="Shrub">Shrubs</a> <ul><li><a href="/wiki/Prostrate_shrub" title="Prostrate shrub">Prostrate shrubs</a></li> <li><a href="/wiki/Subshrub" title="Subshrub">Subshrubs</a></li></ul></li> <li><a href="/wiki/Succulent_plant" title="Succulent plant">Succulent plants</a></li> <li><a href="/wiki/Tree" title="Tree">Trees</a></li> <li><a href="/wiki/Vine" title="Vine">Vines</a> <ul><li><a href="/wiki/Liana" title="Liana">Lianas</a></li></ul></li></ul></li> <li><a href="/wiki/Herbaceous_plant" title="Herbaceous plant">Herbaceous plants</a></li> <li><a href="/wiki/Secondary_growth" title="Secondary growth">Secondary growth</a></li> <li><a href="/wiki/Woody_plant" title="Woody plant">Woody plants</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_reproduction" title="Plant reproduction">Reproduction</a> <div class="hlist"><ul><li><a href="/wiki/Plant_evolution" title="Plant evolution">Evolution</a></li><li><a href="/wiki/Plant_ecology" title="Plant ecology">Ecology</a></li></ul></div></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alternation_of_generations" title="Alternation of generations">Alternation of generations</a></li> <li><a href="/wiki/Double_fertilization" title="Double fertilization">Double fertilization</a></li> <li><a href="/wiki/Plant_evolutionary_developmental_biology" title="Plant evolutionary developmental biology">Evolutionary development</a></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Evolutionary history</a> <ul><li><a href="/wiki/Timeline_of_plant_evolution" title="Timeline of plant evolution">timeline</a></li></ul></li> <li><a href="/wiki/Flora" title="Flora">Flora</a></li> <li><a href="/wiki/Germination" title="Germination">Germination</a></li> <li><a href="/wiki/Pollination" title="Pollination">Pollination</a> <ul><li><a href="/wiki/Artificial_pollination" class="mw-redirect" title="Artificial pollination">Artificial</a></li> <li><a href="/wiki/Pollinator" title="Pollinator">Pollinators</a></li> <li><a href="/wiki/Pollen_tube" title="Pollen tube">Pollen tube</a></li> <li><a href="/wiki/Self-pollination" title="Self-pollination">Self</a></li></ul></li> <li><a href="/wiki/Sporangium" title="Sporangium">Sporangium</a> <ul><li><a href="/wiki/Microsporangia" class="mw-redirect" title="Microsporangia">Microsporangia</a> <ul><li><a href="/wiki/Microspore" title="Microspore">Microspore</a></li></ul></li> <li><a href="/wiki/Sporangium" title="Sporangium">Megasporangium</a> <ul><li><a href="/wiki/Megaspore" title="Megaspore">Megaspore</a></li></ul></li> <li><a href="/wiki/Spore" title="Spore">Spore</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_taxonomy" title="Plant taxonomy">Plant taxonomy</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Biological classification</a></li> <li><a href="/wiki/Botanical_nomenclature" title="Botanical nomenclature">Botanical nomenclature</a> <ul><li><a href="/wiki/Botanical_name" title="Botanical name">Botanical name</a></li> <li><a href="/wiki/Correct_name" title="Correct name">Correct name</a></li> <li><a href="/wiki/Author_citation_(botany)" title="Author citation (botany)">Author citation</a></li> <li><a href="/wiki/International_Code_of_Nomenclature_for_algae,_fungi,_and_plants" title="International Code of Nomenclature for algae, fungi, and plants">International Code of Nomenclature (ICN)</a></li> <li><a href="/wiki/International_Code_of_Nomenclature_for_Cultivated_Plants" title="International Code of Nomenclature for Cultivated Plants">ICN for Cultivated Plants (ICNCP)</a></li></ul></li> <li><a href="/wiki/Cultivated_plant_taxonomy" title="Cultivated plant taxonomy">Cultivated plant taxonomy</a> <ul><li><a href="/wiki/Citrus_taxonomy" title="Citrus taxonomy">Citrus taxonomy</a></li> <li><a href="/wiki/Cultigen" title="Cultigen">Cultigen</a> <ul><li><a href="/wiki/Cultivar" title="Cultivar">Cultivar</a></li> <li><a href="/wiki/Cultivar_group" title="Cultivar group">Group</a></li> <li><a href="/wiki/Grex_(horticulture)" title="Grex (horticulture)">Grex</a></li></ul></li></ul></li> <li><a href="/wiki/History_of_plant_systematics" title="History of plant systematics">History of plant systematics</a></li> <li><a href="/wiki/Herbarium" title="Herbarium">Herbarium</a></li> <li><a href="/wiki/International_Association_for_Plant_Taxonomy" title="International Association for Plant Taxonomy">International Association for Plant Taxonomy</a> (IAPT)</li> <li><a href="/wiki/List_of_systems_of_plant_taxonomy" title="List of systems of plant taxonomy">Plant taxonomy systems</a></li> <li><a href="/wiki/Taxonomic_rank" title="Taxonomic rank">Taxonomic rank</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%">Practice</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Agronomy" title="Agronomy">Agronomy</a></li> <li><a href="/wiki/Floriculture" title="Floriculture">Floriculture</a></li> <li><a href="/wiki/Forestry" title="Forestry">Forestry</a></li> <li><a href="/wiki/Horticulture" title="Horticulture">Horticulture</a></li> <li><a href="/wiki/Phytochemical" title="Phytochemical">Phytochemical</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><div class="hlist"><ul><li>Lists</li><li>Related topics</li></ul></div></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Glossary_of_botanical_terms" title="Glossary of botanical terms">Botanical terms</a></li> <li><a href="/wiki/List_of_botanists" title="List of botanists">Botanists</a> <ul><li><a href="/wiki/List_of_botanists_by_author_abbreviation_(W%E2%80%93Z)" title="List of botanists by author abbreviation (W–Z)">by author abbreviation</a></li></ul></li> <li><a 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