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<title>Secretion</title>

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<span id="openzim-page-title" class="mw-page-title-main"><span class="mw-page-title-main">Secretion</span></span>

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<p><b>Secretion</b> is the movement of material from one point to another, such as a secreted <a href="Chemical_substance" title="Chemical substance">chemical substance</a> from a <a href="Cell_(biology)" title="Cell (biology)">cell</a> or <a href="Gland" title="Gland">gland</a>. In contrast, <a href="Excretion" title="Excretion">excretion</a> is the removal of certain substances or waste products from a cell or organism. The classical mechanism of cell secretion is via secretory portals at the <a href="Cell_membrane" title="Cell membrane">plasma membrane</a> called <a href="Porosomes" class="mw-redirect" title="Porosomes">porosomes</a>.<sup id="cite_ref-pmid_22659300_1-0" class="reference"><a href="#cite_note-pmid_22659300-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Porosomes are permanent cup-shaped <a href="Lipoprotein" title="Lipoprotein">lipoprotein</a> structures embedded in the cell membrane, where secretory vesicles transiently dock and fuse to release intra-vesicular contents from the cell.

</p><p><a href="Bacterial_secretion_system" title="Bacterial secretion system">Secretion in bacterial species</a> means the transport or translocation of effector molecules. For example: <a href="Proteins" class="mw-redirect" title="Proteins">proteins</a>, <a href="Enzymes" class="mw-redirect" title="Enzymes">enzymes</a> or <a href="Toxin" title="Toxin">toxins</a> (such as <a href="Cholera_toxin" title="Cholera toxin">cholera toxin</a> in <a href="Pathogenic_bacteria" title="Pathogenic bacteria">pathogenic bacteria</a> e.g. <i><a href="Vibrio_cholerae" title="Vibrio cholerae">Vibrio cholerae</a></i>) from across the interior (<a href="Cytoplasm" title="Cytoplasm">cytoplasm</a> or <a href="Cytosol" title="Cytosol">cytosol</a>) of a bacterial cell to its exterior. Secretion is a very important mechanism in bacterial functioning and operation in their natural surrounding environment for adaptation and survival.

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<div class="mw-heading mw-heading2"><h2 id="In_eukaryotic_cells">In eukaryotic cells</h2></div>

<div class="mw-heading mw-heading3"><h3 id="Mechanism">Mechanism</h3></div>

<p><a href="Eukaryote" title="Eukaryote">Eukaryotic</a> <a href="Cell_(biology)" title="Cell (biology)">cells</a>, including <a href="Human_cells" class="mw-redirect" title="Human cells">human cells</a>, have a highly <a href="Evolution" title="Evolution">evolved</a> process of secretion. Proteins <a href="Protein_targeting" title="Protein targeting">targeted</a> for the outside are <a href="Protein_synthesis" class="mw-redirect" title="Protein synthesis">synthesized</a> by <a href="Ribosome" title="Ribosome">ribosomes</a> docked to the rough <a href="Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a> (ER). As they are synthesized, these proteins translocate into the ER <a href="Lumen_(anatomy)" title="Lumen (anatomy)">lumen</a>, where they are <a href="Glycosylation" title="Glycosylation">glycosylated</a> and where molecular <a href="Chaperone_(protein)" title="Chaperone (protein)">chaperones</a> aid <a href="Protein_folding" title="Protein folding">protein folding</a>. <a href="Misfolded_proteins" class="mw-redirect" title="Misfolded proteins">Misfolded proteins</a> are usually identified here and retrotranslocated by <a href="Endoplasmic-reticulum-associated_protein_degradation" title="Endoplasmic-reticulum-associated protein degradation">ER-associated degradation</a> to the <a href="Cytosol" title="Cytosol">cytosol</a>, where they are degraded by a <a href="Proteasome" title="Proteasome">proteasome</a>. The <a href="Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicles</a> containing the properly folded proteins then enter the <a href="Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a>.

</p><p>In the Golgi apparatus, the glycosylation of the proteins is modified and further <a href="Post-translational_modification" title="Post-translational modification">post-translational modifications</a>, including cleavage and functionalization, may occur. The <a href="Proteins" class="mw-redirect" title="Proteins">proteins</a> are then moved into secretory vesicles which travel along the <a href="Cytoskeleton" title="Cytoskeleton">cytoskeleton</a> to the edge of the cell. More modification can occur in the secretory vesicles (for example <a href="Insulin" title="Insulin">insulin</a> is cleaved from <a href="Proinsulin" title="Proinsulin">proinsulin</a> in the secretory vesicles).

</p><p>Eventually, there is <a href="Vesicle_fusion" title="Vesicle fusion">vesicle fusion</a> with the <a href="Cell_membrane" title="Cell membrane">cell membrane</a> at porosomes, by a process called <a href="Exocytosis" title="Exocytosis">exocytosis</a>, dumping its contents out of the cell's environment.<sup id="cite_ref-Anderson_2-0" class="reference"><a href="#cite_note-Anderson-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>

</p><p>Strict <a href="Biochemical" class="mw-redirect" title="Biochemical">biochemical</a> control is maintained over this sequence by usage of a <a href="PH" title="PH">pH</a> gradient: the pH of the cytosol is 7.4, the ER's pH is 7.0, and the cis-golgi has a pH of 6.5. Secretory vesicles have pHs ranging between 5.0 and 6.0; some secretory vesicles evolve into <a href="Lysosome" title="Lysosome">lysosomes</a>, which have a pH of 4.8.

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<div class="mw-heading mw-heading4"><h4 id="Nonclassical_secretion">Nonclassical secretion</h4></div>

<p>There are many proteins like <a href="FGF1" class="mw-redirect" title="FGF1">FGF1</a> (aFGF), <a href="FGF2" class="mw-redirect" title="FGF2">FGF2</a> (bFGF), <a href="Interleukin-1" class="mw-redirect" title="Interleukin-1">interleukin-1</a> (IL1) etc. which do not have a signal sequence. They do not use the classical ER-Golgi pathway. These are secreted through various nonclassical pathways.

</p><p>At least four nonclassical (unconventional) protein secretion pathways have been described.<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> They include:

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<ul><li>direct protein translocation across the plasma membrane likely through <a href="Membrane_transport_protein" title="Membrane transport protein">membrane transport proteins</a></li>

<li><a href="Bleb_(cell_biology)" title="Bleb (cell biology)">blebbing</a></li>

<li>lysosomal secretion</li>

<li>release via exosomes derived from multivesicular bodies</li></ul>

<p>In addition, proteins can be released from cells by mechanical or physiological wounding<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> and through non-lethal, transient <a href="Oncotic_pressure" title="Oncotic pressure">oncotic pores</a> in the plasma membrane induced by washing cells with serum-free media or buffers.<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>

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<div class="mw-heading mw-heading3"><h3 id="In_human_tissues">In human tissues</h3></div>

<p>Many <a href="Human_cell_types" class="mw-redirect" title="Human cell types">human cell types</a> have the ability to be secretory cells. They have a well-developed <a href="Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a>, and <a href="Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a> to fulfill this function. <a href="Tissue_(biology)" title="Tissue (biology)">Tissues</a> that produce secretions include the <a href="Gastrointestinal_tract" title="Gastrointestinal tract">gastrointestinal tract</a>, which secretes <a href="Digestive_enzyme" title="Digestive enzyme">digestive enzymes</a> and <a href="Gastric_acid" title="Gastric acid">gastric acid</a>, the <a href="Lung" title="Lung">lungs</a>, which secrete <a href="Surfactant" title="Surfactant">surfactants</a>, and <a href="Sebaceous_gland" title="Sebaceous gland">sebaceous glands</a>, which secrete <a href="Sebum" class="mw-redirect" title="Sebum">sebum</a> to lubricate the skin and hair. <a href="Meibomian_gland" title="Meibomian gland">Meibomian glands</a> in the <a href="Eyelid" title="Eyelid">eyelid</a> secrete <a href="Meibum" class="mw-redirect" title="Meibum">meibum</a> to lubricate and protect the eye.

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<div class="mw-heading mw-heading2"><h2 id="In_gram-negative_bacteria">In gram-negative bacteria</h2></div>

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</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Bacterial_secretion_system" title="Bacterial secretion system">Bacterial secretion system</a></div>

<p>Secretion is not unique to eukaryotes – it is also present in bacteria and archaea as well. <a href="ATP_binding_cassette" class="mw-redirect" title="ATP binding cassette">ATP binding cassette</a> (ABC) type transporters are common to the three domains of life. Some secreted proteins are translocated across the cytoplasmic membrane by the <a href="SecYEG" class="mw-redirect" title="SecYEG">SecYEG</a> <a href="Translocon" title="Translocon">translocon</a>, one of two translocation systems, which requires the presence of an N-terminal signal peptide on the secreted protein. Others are translocated across the cytoplasmic membrane by the <a href="Twin-arginine_translocation_pathway" title="Twin-arginine translocation pathway">twin-arginine translocation pathway</a> (Tat). <a href="Gram-negative_bacteria" title="Gram-negative bacteria">Gram-negative bacteria</a> have two membranes, thus making secretion topologically more complex. There are at least six specialized secretion systems in Gram-negative bacteria.<sup id="cite_ref-WooldridgeK_6-0" class="reference"><a href="#cite_note-WooldridgeK-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup>

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<div class="mw-heading mw-heading3"><h3 id="Type_I_secretion_system_(T1SS_or_TOSS)">Type I secretion system (T1SS or TOSS)</h3></div>

<p>Type I secretion is a chaperone dependent secretion system employing the Hly and Tol gene clusters. The process begins as a leader sequence on the protein to be secreted is recognized by HlyA and binds HlyB on the membrane. This signal sequence is extremely specific for the ABC transporter. The HlyAB complex stimulates HlyD which begins to uncoil and reaches the outer membrane where TolC recognizes a terminal molecule or signal on HlyD. HlyD recruits TolC to the inner membrane and HlyA is excreted outside of the outer membrane via a long-tunnel protein channel.

</p><p>Type I secretion system transports various molecules, from ions, drugs, to proteins of various sizes (20 – 900 kDa). The molecules secreted vary in size from the small <i><a href="Escherichia_coli" title="Escherichia coli">Escherichia coli</a></i> peptide colicin V, (10 kDa) to the <i><a href="Pseudomonas_fluorescens" title="Pseudomonas fluorescens">Pseudomonas fluorescens</a></i> cell adhesion protein LapA of 520 kDa.<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> The best characterized are the <a href="RTX_toxin" title="RTX toxin">RTX toxins</a> and the lipases. Type I secretion is also involved in export of non-proteinaceous substrates like cyclic β-glucans and polysaccharides.

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<div class="mw-heading mw-heading3"><h3 id="Type_II_secretion_system_(T2SS)">Type II secretion system (T2SS)</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Type_II_secretion_system" title="Type II secretion system">Type II secretion system</a></div>

<p>Proteins secreted through the type II system, or main terminal branch of the general secretory pathway, depend on the Sec or Tat system for initial transport into the <a href="Periplasm" title="Periplasm">periplasm</a>. Once there, they pass through the outer membrane via a multimeric (12–14 subunits) complex of pore forming secretin proteins. In addition to the secretin protein, 10–15 other inner and outer membrane proteins compose the full secretion apparatus, many with as yet unknown function. Gram-negative <a href="Pilus#Type_IV_pili" title="Pilus">type IV pili</a> use a modified version of the type II system for their biogenesis, and in some cases certain proteins are shared between a pilus complex and type II system within a single bacterial species.

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<div class="mw-heading mw-heading3"><h3 id="Type_III_secretion_system_(T3SS_or_TTSS)">Type III secretion system (T3SS or TTSS)</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Type_III_secretion_system" title="Type III secretion system">Type III secretion system</a></div>

<p>It is homologous to the basal body in bacterial flagella. It is like a molecular syringe through which a bacterium (e.g. certain types of <i><a href="Salmonella" title="Salmonella">Salmonella</a></i>, <i><a href="Shigella" title="Shigella">Shigella</a></i>, <i><a href="Yersinia" title="Yersinia">Yersinia</a></i>, <i><a href="Vibrio" title="Vibrio">Vibrio</a></i>) can inject proteins into eukaryotic cells. The low Ca<sup>2+</sup> concentration in the cytosol opens the gate that regulates T3SS. One such mechanism to detect low calcium concentration has been illustrated by the lcrV (Low Calcium Response) antigen utilized by <i><a href="Yersinia_pestis" title="Yersinia pestis">Yersinia pestis</a></i>, which is used to detect low calcium concentrations and elicits T3SS attachment. The Hrp system in plant pathogens inject harpins and pathogen effector proteins through similar mechanisms into plants. This secretion system was first discovered in <i><a href="Yersinia_pestis" title="Yersinia pestis">Yersinia pestis</a></i> and showed that toxins could be injected directly from the bacterial cytoplasm into the cytoplasm of its host's cells rather than simply be secreted into the extracellular medium.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup>

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<div class="mw-heading mw-heading3"><h3 id="Type_IV_secretion_system_(T4SS_or_TFSS)">Type IV secretion system (T4SS or TFSS)</h3></div>

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</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">T4SS</th></tr><tr><td colspan="2" class="infobox-image"><div class="infobox-caption">Type IV secretion system</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd; color:inherit;">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;">Symbol</th><td class="infobox-data" style="background-color: #eee; color:inherit;">T4SS</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="Pfam" title="Pfam">Pfam</a></th><td class="infobox-data pfam" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF07996">PF07996</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="InterPro" title="InterPro">InterPro</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/IPR012991">IPR012991</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="Structural_Classification_of_Proteins" class="mw-redirect" title="Structural Classification of Proteins">SCOP2</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="http://scop2.mrc-lmb.cam.ac.uk/search?t=txt;q=1gl7">1gl7</a> / <a rel="nofollow" class="external text" href="https://scop.berkeley.edu/pdb/code=1gl7">SCOPe</a> / <a rel="nofollow" class="external text" href="http://supfam.org/SUPERFAMILY/cgi-bin/search.cgi?search_field=1gl7">SUPFAM</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="TCDB" class="mw-redirect" title="TCDB">TCDB</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="http://www.tcdb.org/search/result.php?tc=3.A.7">3.A.7</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="Orientations_of_Proteins_in_Membranes_database" title="Orientations of Proteins in Membranes database">OPM superfamily</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://opm.phar.umich.edu/protein_superfamilies/215">215</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="Orientations_of_Proteins_in_Membranes_database" title="Orientations of Proteins in Membranes database">OPM protein</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://opm.phar.umich.edu/proteins?search=3jqo">3jqo</a></td></tr><tr><td colspan="2" class="infobox-full-data" style="background-color: #eee; color:inherit;"><table class="infobox mw-collapsible mw-collapsed" style="float:none; clear:none; margin:0; border-width:0; border-collapse:collapse; text-align:left; width:100%"><tbody><tr><th colspan="2" class="infobox-header" style="background-color: #ddd; color:inherit;">Available protein structures:</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="Pfam" title="Pfam">Pfam</a>

 </th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="http://pfam.xfam.org/family/PF07996?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF07996">ECOD</a>

 </td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="Protein_Data_Bank" title="Protein Data Bank">PDB</a></th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.rcsb.org/search?q=rcsb_polymer_entity_annotation.annotation_id:PF07996%20AND%20rcsb_polymer_entity_annotation.type:Pfam">RCSB PDB</a>; <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/entry/search/index?pfam_accession:PF07996">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&other_db_field=PF07996">PDBj</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="PDBsum" title="PDBsum">PDBsum</a></th><td class="infobox-data" style="background-color: #eee; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/thornton-srv/databases/cgi-bin/pdbsum/GetPfamStr.pl?pfam_id=PF07996">structure summary</a></td></tr></tbody></table></td></tr></tbody></table>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Type_IV_secretion_system" title="Type IV secretion system">Type IV secretion system</a></div>

<p>It is homologous to <a href="Bacterial_conjugation" title="Bacterial conjugation">conjugation</a> machinery of bacteria, the <a href="Conjugative_pili" class="mw-redirect" title="Conjugative pili">conjugative pili</a>. It is capable of transporting both DNA and proteins. It was discovered in <i>Agrobacterium tumefaciens</i>, which uses this system to introduce the T-DNA portion of the Ti plasmid into the plant host, which in turn causes the affected area to develop into a crown gall (tumor). <i><a href="Helicobacter_pylori" title="Helicobacter pylori">Helicobacter pylori</a></i> uses a type IV secretion system to deliver <a href="CagA" class="mw-redirect" title="CagA">CagA</a> into gastric epithelial cells, which is associated with gastric carcinogenesis.<sup id="cite_ref-pmid16367902_9-0" class="reference"><a href="#cite_note-pmid16367902-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> <i><a href="Bordetella_pertussis" title="Bordetella pertussis">Bordetella pertussis</a></i>, the causative agent of whooping cough, secretes the <a href="Pertussis_toxin" title="Pertussis toxin">pertussis toxin</a> partly through the type IV system. <i><a href="Legionella_pneumophila" title="Legionella pneumophila">Legionella pneumophila</a></i>, the causing agent of legionellosis (Legionnaires' disease) utilizes a type IVB secretion system, known as the icm/dot (<b>i</b>ntra<b>c</b>ellular <b>m</b>ultiplication / <b>d</b>efect in <b>o</b>rganelle <b>t</b>rafficking genes) system, to translocate numerous <a href="Bacterial_effector_protein" title="Bacterial effector protein">effector proteins</a> into its eukaryotic host.<sup id="cite_ref-pmid15035043_10-0" class="reference"><a href="#cite_note-pmid15035043-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The prototypic Type IVA secretion system is the VirB complex of <i><a href="Agrobacterium_tumefaciens" title="Agrobacterium tumefaciens">Agrobacterium tumefaciens</a></i>.<sup id="cite_ref-pmid16153176_11-0" class="reference"><a href="#cite_note-pmid16153176-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup>

</p><p>Protein members of this family are components of the type IV secretion system. They mediate <a href="Intracellular" class="mw-redirect" title="Intracellular">intracellular</a> transfer of <a href="Macromolecule" title="Macromolecule">macromolecules</a> via a <a href="Nuclear_receptor#Mechanism_of_action" title="Nuclear receptor">mechanism</a> ancestrally related to that of <a href="Bacterial_conjugation" title="Bacterial conjugation">bacterial conjugation</a> machineries.<sup id="cite_ref-pmid15546668_12-0" class="reference"><a href="#cite_note-pmid15546668-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid14673074_13-0" class="reference"><a href="#cite_note-pmid14673074-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading4"><h4 id="Function">Function</h4></div>

<p>The Type IV secretion system (T4SS) is the general mechanism by which bacterial cells secrete or take up macromolecules. Their precise mechanism remains unknown. T4SS is encoded on <a href="Gram_negative_bacteria" class="mw-redirect" title="Gram negative bacteria">Gram-negative</a> conjugative elements in <a href="Bacteria" title="Bacteria">bacteria</a>. T4SS are cell envelope-spanning complexes, or, in other words, 11–13 core proteins that form a channel through which DNA and proteins can travel from the cytoplasm of the donor cell to the cytoplasm of the recipient cell. T4SS also secrete <a href="Virulence" title="Virulence">virulence</a> factor proteins directly into host cells as well as taking up DNA from the medium during natural <a href="Transformation_(bacteria)" class="mw-redirect" title="Transformation (bacteria)">transformation</a>.<sup id="cite_ref-pmid12855161_14-0" class="reference"><a href="#cite_note-pmid12855161-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading4"><h4 id="Structure">Structure</h4></div>

<p>As shown in the above figure, TraC, in particular consists of a three helix bundle and a loose globular appendage.<sup id="cite_ref-pmid14673074_13-1" class="reference"><a href="#cite_note-pmid14673074-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading4"><h4 id="Interactions">Interactions</h4></div>

<p>T4SS has two effector proteins: firstly, ATS-1, which stands for Anaplasma translocated substrate 1, and secondly <a href="Ankyrin_repeat" title="Ankyrin repeat">AnkA</a>, which stands for ankyrin repeat domain-containing protein A. Additionally, T4SS coupling proteins are VirD4, which bind to VirE2.<sup id="cite_ref-pmid20670295_15-0" class="reference"><a href="#cite_note-pmid20670295-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading3"><h3 id="Type_V_secretion_system_(T5SS)">Type V secretion system (T5SS)</h3></div>

<div role="note" class="hatnote navigation-not-searchable">See also: <a href="Trimeric_autotransporter_adhesin#Type_V_secretion_system_(T5SS)" title="Trimeric autotransporter adhesin">Trimeric autotransporter adhesin § Type V secretion system (T5SS)</a></div>

<p>Also called the autotransporter system,<sup id="cite_ref-Thanassi2005_16-0" class="reference"><a href="#cite_note-Thanassi2005-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> type V secretion involves use of the <i>Sec</i> system for crossing the inner membrane. Proteins which use this pathway have the capability to form a <a href="Beta-barrel" class="mw-redirect" title="Beta-barrel">beta-barrel</a> with their C-terminus which inserts into the outer membrane, allowing the rest of the peptide (the passenger domain) to reach the outside of the cell. Often, autotransporters are cleaved, leaving the beta-barrel domain in the outer membrane and freeing the passenger domain. Some researchers believe remnants of the autotransporters gave rise to the <a href="Porin_(protein)" title="Porin (protein)">porins</a> which form similar beta-barrel structures. A common example of an autotransporter that uses this secretion system is the <a href="Trimeric_Autotransporter_Adhesins_(TAA)" class="mw-redirect" title="Trimeric Autotransporter Adhesins (TAA)">Trimeric Autotransporter Adhesins</a>.<sup id="cite_ref-pmid17482513_17-0" class="reference"><a href="#cite_note-pmid17482513-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading3"><h3 id="Type_VI_secretion_system_(T6SS)">Type VI secretion system (T6SS)</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Type_VI_secretion_system" title="Type VI secretion system">Type VI secretion system</a></div>

<p>Type VI secretion systems were originally identified in 2006 by the group of <a href="John_Mekalanos" title="John Mekalanos">John Mekalanos</a> at the Harvard Medical School (Boston, USA) in two bacterial pathogens, <i><a href="Vibrio_cholerae" title="Vibrio cholerae">Vibrio cholerae</a></i> and <i><a href="Pseudomonas_aeruginosa" title="Pseudomonas aeruginosa">Pseudomonas aeruginosa</a></i>.<sup id="cite_ref-pmid16432199_18-0" class="reference"><a href="#cite_note-pmid16432199-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid16763151_19-0" class="reference"><a href="#cite_note-pmid16763151-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> These were identified when mutations in the Hcp and VrgG genes in <i>Vibrio cholerae</i> led to decreased virulence and pathogenicity. Since then, Type VI secretion systems have been found in a quarter of all proteobacterial genomes, including animal, plant, human pathogens, as well as soil, environmental or marine bacteria.<sup id="cite_ref-pmid18289922_20-0" class="reference"><a href="#cite_note-pmid18289922-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid18617888_21-0" class="reference"><a href="#cite_note-pmid18617888-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> While most of the early studies of Type VI secretion focused on its role in the pathogenesis of higher organisms, more recent studies suggested a broader physiological role in defense against simple eukaryotic predators and its role in inter-bacteria interactions.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Coulthurst_2013_S0923-2508_23-0" class="reference"><a href="#cite_note-Coulthurst_2013_S0923-2508-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> The Type VI secretion system gene clusters contain from 15 to more than 20 genes, two of which, Hcp and VgrG, have been shown to be nearly universally secreted substrates of the system. Structural analysis of these and other proteins in this system bear a striking resemblance to the tail spike of the T4 phage, and the activity of the system is thought to functionally resemble phage infection.<sup id="cite_ref-Silverman_2012_453-472_24-0" class="reference"><a href="#cite_note-Silverman_2012_453-472-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading3"><h3 id="Type_VII_secretion_system_(T7SS)">Type VII secretion system (T7SS)</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Type_VII_secretion_system" title="Type VII secretion system">Type VII secretion system</a></div>

<div class="mw-heading mw-heading3"><h3 id="Type_VIII_secretion_system_(T8SS)">Type VIII secretion system (T8SS)</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Type_VIII_secretion_system" title="Type VIII secretion system">Type VIII secretion system</a></div>

<div class="mw-heading mw-heading3"><h3 id="Type_IX_secretion_system_(T9SS)">Type IX secretion system (T9SS)</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Type_IX_secretion_system" title="Type IX secretion system">Type IX secretion system</a></div>

<div class="mw-heading mw-heading3"><h3 id="Release_of_outer_membrane_vesicles">Release of outer membrane vesicles</h3></div>

<p>In addition to the use of the multiprotein complexes listed above, Gram-negative bacteria possess another method for release of material: the formation of <a href="Bacterial_outer_membrane_vesicles" class="mw-redirect" title="Bacterial outer membrane vesicles">bacterial outer membrane vesicles</a>.<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> Portions of the outer membrane pinch off, forming nano-scale spherical structures made of a lipopolysaccharide-rich lipid bilayer enclosing periplasmic materials, and are deployed for <a href="Membrane_vesicle_trafficking" title="Membrane vesicle trafficking">membrane vesicle trafficking</a> to manipulate environment or invade at <a href="Host%E2%80%93pathogen_interface" class="mw-redirect" title="Host–pathogen interface">host–pathogen interface</a>. Vesicles from a number of bacterial species have been found to contain virulence factors, some have immunomodulatory effects, and some can directly adhere to and intoxicate host cells. release of vesicles has been demonstrated as a general response to stress conditions, the process of loading cargo proteins seems to be selective.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading2"><h2 id="In_gram-positive_bacteria">In gram-positive bacteria</h2></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Bacterial_secretion_system" title="Bacterial secretion system">Bacterial secretion system</a></div>

<p>In some <i>Staphylococcus</i> and <i>Streptococcus</i> species, the accessory secretory system handles the export of highly repetitive adhesion glycoproteins.

</p>

<ul><li><a href="Bacterial_effector_protein" title="Bacterial effector protein">Bacterial effector protein</a></li>

<li><a href="Bacterial_outer_membrane_vesicles" class="mw-redirect" title="Bacterial outer membrane vesicles">Bacterial outer membrane vesicles</a></li>

<li><a href="Host%E2%80%93pathogen_interaction" title="Host–pathogen interaction">Host–pathogen interaction</a></li>

<li><a href="Membrane_vesicle_trafficking" title="Membrane vesicle trafficking">Membrane vesicle trafficking</a></li>

<li><a href="Secretomics" title="Secretomics">Secretomics</a></li>

<li><a href="Secretory_protein" title="Secretory protein">Secretory proteins</a></li>

<li><a href="Secretor_status" title="Secretor status">Secretor status</a></li></ul>

<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>

</p>

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<li id="cite_note-26"><span class="mw-cite-backlink"><b><a href="#cite_ref-26">^</a></b></span> <span class="reference-text"><cite id="CITEREFMcBroomKuehn2007" class="citation journal cs1">McBroom AJ, Kuehn MJ (January 2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1868505">"Release of outer membrane vesicles by Gram-negative bacteria is a novel envelope stress response"</a>. <i>Molecular Microbiology</i>. <b>63</b> (2): <span class="nowrap">545–</span>58. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1365-2958.2006.05522.x">10.1111/j.1365-2958.2006.05522.x</a>. <a href="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/PMC1868505">1868505</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17163978">17163978</a>.</cite></span>

</li>

<li id="cite_note-27"><span class="mw-cite-backlink"><b><a href="#cite_ref-27">^</a></b></span> <span class="reference-text">Z. Esna Ashari, N. Dasgupta, K. Brayton & S. Broschat, “<a rel="nofollow" class="external text" href="https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0197041#sec011">An optimal set of features for predicting type IV secretion system effector proteins for a subset of species based on a multi-level feature selection approach</a>”, PLOS ONE Journal, 2018, 13, e0197041. (doi.org/10.1371/journal.pone.0197041.)</span>

</li>

</ol></div>

<div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2></div>

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<ul><li><cite id="CITEREFAlbertsJohnsonLewisRaff2002" class="citation book cs1">Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P, eds. (2002). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21054/?term=secretion">"Search: Secretion"</a>. <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21054/"><i>Molecular Biology of the Cell</i></a> (4th ed.). New York: Garland Science. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-8153-3218-3</bdi>.</cite></li>

<li><cite id="CITEREFWhite2000" class="citation book cs1">White D (2000). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/physiologybioche00whit"><i>The Physiology and Biochemistry of Prokaryotes</i></a></span> (2nd ed.). Oxford University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-19-512579-5</bdi>.</cite></li>

<li><cite id="CITEREFAvon" class="citation web cs1">Avon D. <a rel="nofollow" class="external text" href="http://cellsalive.com/">"Home page"</a>. <i>Cells alive!</i>.</cite></li></ul>

</div>

<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>

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<div class="side-box-text plainlist">Look up <i><b><a href="https://en.wiktionary.org/wiki/Special:Search/secretion" class="extiw external" title="wiktionary:Special:Search/secretion">secretion</a></b></i> in Wiktionary, the free dictionary.</div></div>

</div>

<ul><li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Secretions">Secretions</a> at the U.S. National Library of Medicine <a href="Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li>

<li>T5SS / Autotransporter illustration at <a rel="nofollow" class="external text" href="https://www.uni-muenster.de/Chemie.pz/forschen/ag/jose/topicsofresearch.html">Uni Münster</a></li></ul>

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</style><div id="Branches_of_biology59" style="font-size:114%;margin:0 4em"><a href="Outline_of_biology#Branches" title="Outline of biology">Branches of biology</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Abiogenesis" title="Abiogenesis">Abiogenesis</a></li>

<li><a href="Aerobiology" title="Aerobiology">Aerobiology</a></li>

<li><a href="Agronomy" title="Agronomy">Agronomy</a></li>

<li><a href="Agrostology" title="Agrostology">Agrostology</a></li>

<li><a href="Anatomy" title="Anatomy">Anatomy</a></li>

<li><a href="Human_evolution" title="Human evolution">Anthropogeny</a></li>

<li><a href="Anthropology" title="Anthropology">Anthropology</a></li>

<li><a href="Astrobiology" title="Astrobiology">Astrobiology</a></li>

<li><a href="Bacteriology" title="Bacteriology">Bacteriology</a></li>

<li><a href="Biochemistry" title="Biochemistry">Biochemistry</a></li>

<li><a href="Biogeography" title="Biogeography">Biogeography</a></li>

<li><a href="Biogeology" title="Biogeology">Biogeology</a></li>

<li><a href="Bioinformatics" title="Bioinformatics">Bioinformatics</a></li>

<li><a href="Biological_engineering" title="Biological engineering">Biological engineering</a></li>

<li><a href="Mathematical_and_theoretical_biology" title="Mathematical and theoretical biology">Biomathematics</a></li>

<li><a href="Biomechanics" title="Biomechanics">Biomechanics</a></li>

<li><a href="Biophysics" title="Biophysics">Biophysics</a></li>

<li><a href="Biosemiotics" title="Biosemiotics">Biosemiotics</a></li>

<li><a href="Biostatistics" title="Biostatistics">Biostatistics</a></li>

<li><a href="Biotechnology" title="Biotechnology">Biotechnology</a></li>

<li><a href="Botany" title="Botany">Botany</a></li>

<li><a href="Cell_biology" title="Cell biology">Cell biology</a></li>

<li><a href="Cellular_microbiology" title="Cellular microbiology">Cellular microbiology</a></li>

<li><a href="Chemical_biology" title="Chemical biology">Chemical biology</a></li>

<li><a href="Chronobiology" title="Chronobiology">Chronobiology</a></li>

<li><a href="Cognitive_biology" title="Cognitive biology">Cognitive biology</a></li>

<li><a href="Computational_biology" title="Computational biology">Computational biology</a></li>

<li><a href="Conservation_biology" title="Conservation biology">Conservation biology</a></li>

<li><a href="Cryobiology" title="Cryobiology">Cryobiology</a></li>

<li><a href="Cytogenetics" title="Cytogenetics">Cytogenetics</a></li>

<li><a href="Dendrology" title="Dendrology">Dendrology</a></li>

<li><a href="Developmental_biology" title="Developmental biology">Developmental biology</a></li>

<li><a href="Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li>

<li><a href="Ecology" title="Ecology">Ecology</a></li>

<li><a href="Embryology" title="Embryology">Embryology</a></li>

<li><a href="Epidemiology" title="Epidemiology">Epidemiology</a></li>

<li><a href="Epigenetics" title="Epigenetics">Epigenetics</a></li>

<li><a href="Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li>

<li><a href="Freshwater_biology" title="Freshwater biology">Freshwater biology</a></li>

<li><a href="Generative_biology" class="mw-redirect" title="Generative biology">Generative biology</a></li>

<li><a href="Genetics" title="Genetics">Genetics</a></li>

<li><a href="Genomics" title="Genomics">Genomics</a></li>

<li><a href="Geobiology" title="Geobiology">Geobiology</a></li>

<li><a href="Gerontology" title="Gerontology">Gerontology</a></li>

<li><a href="Herpetology" title="Herpetology">Herpetology</a></li>

<li><a href="Histology" title="Histology">Histology</a></li>

<li><a href="Human_biology" title="Human biology">Human biology</a></li>

<li><a href="Ichthyology" title="Ichthyology">Ichthyology</a></li>

<li><a href="Immunology" title="Immunology">Immunology</a></li>

<li><a href="Lipidology" title="Lipidology">Lipidology</a></li>

<li><a href="Mammalogy" title="Mammalogy">Mammalogy</a></li>

<li><a href="Marine_biology" title="Marine biology">Marine biology</a></li>

<li><a href="Microbiology" title="Microbiology">Microbiology</a></li>

<li><a href="Molecular_biology" title="Molecular biology">Molecular biology</a></li>

<li><a href="Mycology" title="Mycology">Mycology</a></li>

<li><a href="Neontology" title="Neontology">Neontology</a></li>

<li><a href="Neuroscience" title="Neuroscience">Neuroscience</a></li>

<li><a href="Nutritional_science" title="Nutritional science">Nutrition</a></li>

<li><a href="Ornithology" title="Ornithology">Ornithology</a></li>

<li><a href="Osteology" title="Osteology">Osteology</a></li>

<li><a href="Paleontology" title="Paleontology">Paleontology</a></li>

<li><a href="Parasitology" title="Parasitology">Parasitology</a></li>

<li><a href="Pathology" title="Pathology">Pathology</a></li>

<li><a href="Pharmacology" title="Pharmacology">Pharmacology</a></li>

<li><a href="Photobiology" title="Photobiology">Photobiology</a></li>

<li><a href="Phycology" title="Phycology">Phycology</a></li>

<li><a href="Phylogenetics" title="Phylogenetics">Phylogenetics</a></li>

<li><a href="Physiology" title="Physiology">Physiology</a></li>

<li><a href="Pomology" title="Pomology">Pomology</a></li>

<li><a href="Primatology" title="Primatology">Primatology</a></li>

<li><a href="Proteomics" title="Proteomics">Proteomics</a></li>

<li><a href="Protistology" title="Protistology">Protistology</a></li>

<li><a href="Quantum_biology" title="Quantum biology">Quantum biology</a></li>

<li><a href="Relational_biology" class="mw-redirect" title="Relational biology">Relational biology</a></li>

<li><a href="Reproductive_biology" title="Reproductive biology">Reproductive biology</a></li>

<li><a href="Sociobiology" title="Sociobiology">Sociobiology</a></li>

<li><a href="Structural_biology" title="Structural biology">Structural biology</a></li>

<li><a href="Synthetic_biology" title="Synthetic biology">Synthetic biology</a></li>

<li><a href="Systematics" title="Systematics">Systematics</a></li>

<li><a href="Systems_biology" title="Systems biology">Systems biology</a></li>

<li><a href="Taxonomy_(biology)" title="Taxonomy (biology)">Taxonomy</a></li>

<li><a href="Teratology" title="Teratology">Teratology</a></li>

<li><a href="Toxicology" title="Toxicology">Toxicology</a></li>

<li><a href="Virology" title="Virology">Virology</a></li>

<li><a href="Virophysics" title="Virophysics">Virophysics</a></li>

<li><a href="Welfare_biology" title="Welfare biology">Welfare biology</a></li>

<li><a href="Xenobiology" title="Xenobiology">Xenobiology</a></li>

<li><a href="Zoology" title="Zoology">Zoology</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="History_of_biology" title="History of biology">History of biology</a></li>

<li><a href="Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a></li>

<li><a href="Timeline_of_biology_and_organic_chemistry" title="Timeline of biology and organic chemistry">Timeline of biology and organic chemistry</a></li></ul>

</div></td></tr></tbody></table></div>

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</style></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q84230#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata1970" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q84230#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata1970" style="font-size:114%;margin:0 4em">Authority control databases </div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">International</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Secretion"><a rel="nofollow" class="external text" href="https://id.worldcat.org/fast/1110707">FAST</a></span></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Sekretion"><a rel="nofollow" class="external text" href="https://d-nb.info/gnd/4124179-4">Germany</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Secretion"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85119448">United States</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Sécrétion"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb119332509">France</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Sécrétion"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb119332509">BnF data</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="分泌"><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/00561023">Japan</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://www.nli.org.il/en/authorities/987007529421505171">Israel</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="https://lux.collections.yale.edu/view/concept/cf163a56-6b60-41c6-a4be-0076d10824fb">Yale LUX</a></span></li></ul></div></td></tr></tbody></table></div></div><div><div class="zim-footer">

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