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class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Spectrum of collective cell migration</span> </div> </a> <ul id="toc-Spectrum_of_collective_cell_migration-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a 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dir="ltr"><p><b>Collective cell migration</b> describes the movements of group of cells and the <a href="/wiki/Emergence" title="Emergence">emergence</a> of collective behavior from cell-environment interactions and cell-cell communication. Collective cell migration is an essential process in the lives of <a href="/wiki/Multicellular_organisms" class="mw-redirect" title="Multicellular organisms">multicellular organisms</a>, e.g. <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a>, <a href="/wiki/Wound_healing" title="Wound healing">wound healing</a> and cancer spreading (<a href="/wiki/Metastasis" title="Metastasis">metastasis</a>).<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Cells can migrate as a cohesive group (e.g. <a href="/wiki/Epithelial_cells" class="mw-redirect" title="Epithelial cells">epithelial cells</a>) or have transient cell-cell adhesion sites (e.g. <a href="/wiki/Mesenchyme" title="Mesenchyme">mesenchymal</a> cells).<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> They can also migrate in different modes like sheets, strands, tubes, and clusters.<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> While <a href="/wiki/Cell_migration" title="Cell migration">single-cell migration</a> has been extensively studied, collective cell migration is a relatively new field with applications in preventing birth defects or dysfunction of embryos. 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rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><table class="sidebar sidebar-collapse nomobile nowraplinks"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#82C3D8; padding:0.2em; font-size:150%; font-weight:bold;">Microbial and microbot movement</th></tr><tr><td class="sidebar-image"><span typeof="mw:File"><a href="/wiki/File:Flagellum_from_a_gram-negative_bacterium_(unlabelled).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Flagellum_from_a_gram-negative_bacterium_%28unlabelled%29.png/160px-Flagellum_from_a_gram-negative_bacterium_%28unlabelled%29.png" decoding="async" width="160" height="160" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Flagellum_from_a_gram-negative_bacterium_%28unlabelled%29.png/240px-Flagellum_from_a_gram-negative_bacterium_%28unlabelled%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Flagellum_from_a_gram-negative_bacterium_%28unlabelled%29.png/320px-Flagellum_from_a_gram-negative_bacterium_%28unlabelled%29.png 2x" data-file-width="358" data-file-height="357" /></a></span></td></tr><tr><td class="sidebar-content"> <b><a href="/wiki/Microswimmer" title="Microswimmer">Microswimmers</a></b></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)">Taxa</div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Bacterial_motility" title="Bacterial motility">Bacterial motility</a> <br /> <ul><li><a href="/wiki/Run-and-tumble_motion" title="Run-and-tumble motion">run-and-tumble</a></li> <li><a href="/wiki/Twitching_motility" title="Twitching motility">twitching</a></li> <li><a href="/wiki/Gliding_motility" title="Gliding motility">gliding</a></li></ul></li></ul> </div> <div class="hlist"> <ul><li><a href="/wiki/Protist_locomotion" title="Protist locomotion">Protist locomotion</a> <ul><li><a href="/wiki/Amoeboid_movement" title="Amoeboid movement">amoeboids</a></li></ul></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)"><a href="/wiki/Taxis" title="Taxis">Taxis</a></div><div class="sidebar-list-content mw-collapsible-content"><div class="plainlist"> <ul><li><a href="/wiki/Aerotaxis" title="Aerotaxis">Aerotaxis</a> (oxygen)</li> <li><a href="/wiki/Anemotaxis" class="mw-redirect" title="Anemotaxis">Anemotaxis</a> (wind)</li> <li><a href="/wiki/Chemotaxis" title="Chemotaxis">Chemotaxis</a> (chemicals)</li> <li><a href="/wiki/Electrotaxis" title="Electrotaxis">Electrotaxis</a>(electric current)</li> <li><a href="/wiki/Gravitaxis" title="Gravitaxis">Gravitaxis</a> (gravity)</li> <li><a href="/wiki/Magnetotaxis" title="Magnetotaxis">Magnetotaxis</a> (magnetic field)</li> <li><a href="/wiki/Phototaxis" title="Phototaxis">Phototaxis</a> (light)</li> <li><a href="/wiki/Rheotaxis" title="Rheotaxis">Rheotaxis</a> (fluid flow)</li> <li><a href="/wiki/Thermotaxis" title="Thermotaxis">Thermotaxis</a> (temperature)</li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)"><a href="/wiki/Kinesis_(biology)" title="Kinesis (biology)">Kinesis</a></div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Kinesis_(biology)" title="Kinesis (biology)">Kinesis</a> <ul><li><a href="/wiki/Chemokinesis" title="Chemokinesis">chemokinesis</a></li> <li><a href="/wiki/Photokinesis" title="Photokinesis">photokinesis</a></li></ul></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)">Microbots and particles</div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Microbotics" title="Microbotics">Microbotics</a></li> <li><a href="/wiki/Nanorobotics" title="Nanorobotics">Nanorobotics</a></li> <li><a href="/wiki/Nanomotors" class="mw-redirect" title="Nanomotors">Nanomotors</a></li> <li><a href="/wiki/DNA_machine" title="DNA machine">DNA machine</a></li> <li><a href="/wiki/Microparticle" title="Microparticle">Microparticle</a> <ul><li><a href="/wiki/Nanoparticle" title="Nanoparticle">Nanoparticle</a></li> <li><a href="/wiki/Janus_particles" title="Janus particles">Janus particles</a></li> <li><a href="/wiki/Self-propelled_particles" title="Self-propelled particles">Self-propelled particles</a></li></ul></li> <li><a href="/wiki/Swarm_robotics" title="Swarm robotics">Swarm robotics</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)">Biohybrids</div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Biohybrid_microswimmers" class="mw-redirect" title="Biohybrid microswimmers">Biohybrid microswimmers</a></li></ul> </div> <div class="hlist"> <ul><li><ul><li><a href="/wiki/Biohybrid_microswimmer#Bacterial_biohybrids" title="Biohybrid microswimmer">bacterial biohybrids</a></li> <li><a href="/wiki/Protist_locomotion#Biohybrid_microswimmers" title="Protist locomotion">protist biohybrids</a></li> <li><a href="/wiki/Robotic_sperm" title="Robotic sperm">robotic sperm</a></li></ul></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)">Collective motion</div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Active_matter" title="Active matter">Active matter</a></li> <li><a href="/wiki/Bacteria_collective_motion" title="Bacteria collective motion">Bacteria collective motion</a></li> <li><a class="mw-selflink selflink">Collective cell migration</a></li> <li><a href="/wiki/Quorum_sensing" title="Quorum sensing">Quorum sensing</a></li> <li><a href="/wiki/Swarming_motility" title="Swarming motility">Swarming motility</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <b><a href="/wiki/Molecular_motor" title="Molecular motor">Molecular motors</a></b></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)">Biological motors</div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Flagellum" title="Flagellum">Flagellum</a> <ul><li><a href="/wiki/Archaellum" title="Archaellum">archaellum</a></li> <li><a href="/wiki/Cilium" title="Cilium">cilium</a></li> <li><a href="/wiki/Axoneme" title="Axoneme">axoneme</a></li> <li><a href="/wiki/Flagellar_motor_switch" class="mw-redirect" title="Flagellar motor switch">motor switch</a></li> <li><a href="/wiki/Intraflagellar_transport" title="Intraflagellar transport">intraflagellar</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">evolution</a></li></ul></li></ul> </div><div class="hlist"> <ul><li><a href="/wiki/Motor_protein" title="Motor protein">Motor proteins</a> <ul><li><a href="/wiki/Myosin" title="Myosin">myosin</a></li> <li><a href="/wiki/Kinesin" title="Kinesin">kinesin</a></li> <li><a href="/wiki/Dynein" title="Dynein">dynein</a></li></ul></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)"><a href="/wiki/Synthetic_molecular_motor" title="Synthetic molecular motor">Synthetic motors</a></div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Synthetic_molecular_motor" title="Synthetic molecular motor">Synthetic molecular motor</a></li> <li><a href="/wiki/Molecular_modelling" title="Molecular modelling">Molecular modelling</a></li> <li><a href="/wiki/Molecular_propeller" title="Molecular propeller">Molecular propeller</a></li> <li><a href="/wiki/Molecular_sensor" title="Molecular sensor">molecular sensor</a></li> <li><a href="/wiki/Molecular_shuttle" title="Molecular shuttle">molecular shuttle</a></li> <li><a href="/wiki/Molecular_tweezers" title="Molecular tweezers">Molecular tweezers</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;font-size:100%;font-weight:bold;;color: var(--color-base)">Related</div><div class="sidebar-list-content mw-collapsible-content"><div class="hlist"> <ul><li><a href="/wiki/Brownian_motor" title="Brownian motor">Brownian motor</a></li> <li><a href="/wiki/Biochip" title="Biochip">Biochip</a><br /></li> <li><a href="/wiki/Endocytosis" title="Endocytosis">Endocytosis</a></li> <li><a href="/wiki/Axophilic_migration" class="mw-redirect" title="Axophilic migration">Axophilic migration</a></li></ul> </div><div class="hlist"> <ul><li><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</a> <ul><li><a href="/wiki/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">prokaryotic</a><br /></li> <li><a href="/wiki/Cytoskeleton#Eukaryotic_cytoskeleton" title="Cytoskeleton">eukaryotic</a></li> <li><a href="/wiki/Cytoplasmic_streaming" title="Cytoplasmic streaming">cytoplasmic streaming</a></li></ul></li></ul> </div><div class="hlist"> <ul><li><a href="/wiki/Gray_goo" title="Gray goo">Gray goo</a></li> <li><a href="/wiki/Mucilage" title="Mucilage">Mucilage</a></li> <li><a href="/wiki/Molecular_biophysics" title="Molecular biophysics">Molecular biophysics</a></li> <li><a href="/wiki/Molecular_machine" title="Molecular machine">Molecular machine</a></li> <li><a href="/wiki/Nanoengineering" title="Nanoengineering">Nanoengineering</a></li> <li><a href="/wiki/Non-motile_bacteria" title="Non-motile bacteria">Non-motile bacteria</a></li> <li><a href="/wiki/Virophysics" title="Virophysics">Virophysics</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-below hlist" style="background-color: #82C3D8; border-color: #A2B8BF"> <ul><li><span class="nowrap"><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Microswimmers" title="Category:Microswimmers">Category</a></span></li></ul></td></tr><tr><td class="sidebar-navbar"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Microbial_and_microbot_movement" title="Template:Microbial and microbot movement"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/w/index.php?title=Template_talk:Microbial_and_microbot_movement&action=edit&redlink=1" class="new" title="Template talk:Microbial and microbot movement (page does not exist)"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Microbial_and_microbot_movement" title="Special:EditPage/Template:Microbial and microbot movement"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p>The environment of the migrating cell can affect its speed, persistence and direction of migration by stimulating it. The <a href="/wiki/Extracellular_matrix" title="Extracellular matrix">extracellular matrix</a> (ECM) provides not only the structural and biochemical support, but also plays a major role in regulating cell behavior. Different ECM proteins (such as <a href="/wiki/Collagen" title="Collagen">collagen</a>, <a href="/wiki/Elastin" title="Elastin">elastin</a>, <a href="/wiki/Fibronectin" title="Fibronectin">fibronectin</a>, <a href="/wiki/Laminin" title="Laminin">laminin</a>, and others) allow cells to adhere and migrate, while forming <a href="/wiki/Focal_adhesions" class="mw-redirect" title="Focal adhesions">focal adhesions</a> in the front and disassembling them in the back. Using these adhesion sites, cells also sense the mechanical properties of the ECM. Cells can be guided by a gradient of those proteins (<a href="/wiki/Haptotaxis" title="Haptotaxis">haptotaxis</a>) or a gradient of soluble substrates in the liquid phase surrounding the cell (<a href="/wiki/Chemotaxis" title="Chemotaxis">chemotaxis</a>). Cells sense the substrate through their <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptors</a> and migrate toward the concentration (or the opposite direction). Another form of stimulation can be rigidity gradients of the ECM (<a href="/wiki/Durotaxis" title="Durotaxis">durotaxis</a>).<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2017)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Confinement">Confinement</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Collective_cell_migration&action=edit&section=2" title="Edit section: Confinement"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Collective cell migration is enhanced by geometrical confinement of an <a href="/wiki/Extracellular_matrix" title="Extracellular matrix">extracellular matrix</a> <a href="/wiki/Molecule" title="Molecule">molecule</a> (e.g. the <a href="/wiki/Proteoglycan" title="Proteoglycan">proteoglycan</a> <a href="/wiki/Versican" title="Versican">versican</a> in <a href="/wiki/Neural_crest" title="Neural crest">neural crest</a> cells), that acts as a barrier, to promote the <a href="/wiki/Emergence" title="Emergence">emergence</a> of organized migration in separated streams. Confinement is also observed <a href="/wiki/In_vivo" title="In vivo">in vivo</a>, where the optimal width is a function of the number of migrating cells in different streams of different <a href="/wiki/Species" title="Species">species</a>.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Cell-cell_communication">Cell-cell communication</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Collective_cell_migration&action=edit&section=3" title="Edit section: Cell-cell communication"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Migrating isolated cell responds to cues in its environment and changes its behavior accordingly. As cell-cell communication does not play a major role in this case, similar trajectories are observed in different isolated cells. However, when the cell migrates as part of the collective, it not only responds to its environment but also interacts with other cells through soluble substrates and physical contact. These cell-cell communication mechanisms are the main reasons for the difference between efficient migration of the collective and <a href="/wiki/Random_walk" title="Random walk">random walk</a> movements of the isolated cell. Cell-cell communication mechanisms are widely studied experimentally (<a href="/wiki/In_vivo" title="In vivo">in vivo</a> and <a href="/wiki/In_vitro" title="In vitro">in vitro</a>),<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> and computationally (<a href="/wiki/In_silico" title="In silico">in silico</a>).<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Co-attraction">Co-attraction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Collective_cell_migration&action=edit&section=4" title="Edit section: Co-attraction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Co-attraction between collectively migrating cells is the process by which cells of the same type secrete <a href="/wiki/Chemotaxis" title="Chemotaxis">chemo-attractant</a> (e.g. <a href="/wiki/C3a_(complement)" title="C3a (complement)">C3a</a> in neural crest cells), that stimulates other cells in the group that have the <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptors</a> to that chemo-attractant. Cells sense the secreted substrate and respond to the stimulation by moving towards each other's and maintain high cell density.<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><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> </p> <div class="mw-heading mw-heading3"><h3 id="Contact_inhibition_of_locomotion">Contact inhibition of locomotion</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Collective_cell_migration&action=edit&section=5" title="Edit section: Contact inhibition of locomotion"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Contact inhibition of locomotion (CIL) is a process in which the cell changes its direction of movement after colliding into another cell. Those cells could be of the same cell type or different types. The contacts (<a href="/wiki/Cell_junction" title="Cell junction">cell-junctions</a>) are created by <a href="/wiki/Transmembrane_protein" title="Transmembrane protein">transmembrane</a> <a href="/wiki/Glycoprotein" title="Glycoprotein">glycoproteins</a> named <a href="/wiki/Cadherin" title="Cadherin">cadherins</a> (<a href="/wiki/E-cadherin" class="mw-redirect" title="E-cadherin">E-cadherin</a>, <a href="/wiki/N-cadherin" class="mw-redirect" title="N-cadherin">N-cadherin</a> or <a href="/wiki/Cadherin_11" class="mw-redirect" title="Cadherin 11">cadherin 11</a>) and other <a href="/wiki/Protein" title="Protein">proteins</a>. After cell-cell contact, the protrusions of cells in the contact direction are inhibited. In the CIL process, cells migrate away from each other by repolarizing in the new direction, so that new protrusions are formed in the front while contractions pull the back from contact.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2017)">citation needed</span></a></i>]</sup> </p> <ul><li><a href="/wiki/Contact_inhibition" title="Contact inhibition">Contact inhibition of proliferation</a> (CIP) is the inhibition of cell division with increasing percent of confluency. CIP and CIL are two different processes, which are sometimes mistakenly interrelated.<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></li></ul> <div class="mw-heading mw-heading2"><h2 id="Examples_of_studied_systems">Examples of studied systems</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Collective_cell_migration&action=edit&section=6" title="Edit section: Examples of studied systems"><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:Models_for_studying_collective_cell_migration.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Models_for_studying_collective_cell_migration.jpg/350px-Models_for_studying_collective_cell_migration.jpg" decoding="async" width="350" height="392" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Models_for_studying_collective_cell_migration.jpg/525px-Models_for_studying_collective_cell_migration.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/e/e9/Models_for_studying_collective_cell_migration.jpg 2x" data-file-width="567" data-file-height="635" /></a><figcaption> <div class="center" style="width:auto; margin-left:auto; margin-right:auto;"><b>Models for studying collective cell migration</b> <sup id="cite_ref-Barriga2019_10-0" class="reference"><a href="#cite_note-Barriga2019-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><br /><small>Red arrows show the direction of migration for each tissue</small></div></figcaption></figure> <p>Collective cell migration is studied over many model species. </p><p><a href="/wiki/Border_cells" class="mw-redirect" title="Border cells">Border cells</a> in flies (<i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i>): the border cells migrate during the differentiation of egg cells to be ready for fertilization.<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 <a href="/wiki/Lateral_line" title="Lateral line">lateral line</a> in <a href="/wiki/Zebrafish" title="Zebrafish">zebrafish</a>: collective cell migration from head to tails is essential to the development of the sensory system of the fish. The sensors of the lateral line measure the flow over the body-surface of the fish.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Wound_healing" title="Wound healing">Wound healing</a>: collective cell migration is an essential part in this healing process, wound area is closed by the migrating cells.<sup id="cite_ref-pmid28110712_13-0" class="reference"><a href="#cite_note-pmid28110712-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Wound healing is commonly studied in vitro using cell lines such as <a href="/wiki/Madin-Darby_Canine_Kidney_Cells" class="mw-redirect" title="Madin-Darby Canine Kidney Cells">Madin-Darby Canine Kidney cells</a>. </p><p><a href="/wiki/Neural_crest" title="Neural crest">Neural crest</a> cells in <a href="/wiki/Mice" class="mw-redirect" title="Mice">mice</a>,<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Leghorn_chicken" title="Leghorn chicken">Leghorn chicks</a>,<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Amphibian" title="Amphibian">amphibians</a> (<i><a href="/wiki/Xenopus_laevis" class="mw-redirect" title="Xenopus laevis">Xenopus laevis</a></i>),<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Fish" title="Fish">fish</a><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> (<a href="/wiki/Zebrafish" title="Zebrafish">zebrafish</a>): collective migration of neural crest cells occurs during embryo development of vertebrates. They migrate long distances from the head (<a href="/wiki/Neural_tube" title="Neural tube">neural tube</a>) to give rise to different tissues.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p>Spreading of <a href="/wiki/Cancer" title="Cancer">cancer</a> (<a href="/wiki/Metastasis" title="Metastasis">metastasis</a>): common complication of cancer involve formation of new tumors (secondary tumors), as a result of migration of cancer cells from the primary tumor. Similar to collective cell migration in development and wound healing, cancer cells also undergo <a href="/wiki/Epithelial_to_mesenchymal_transition" class="mw-redirect" title="Epithelial to mesenchymal transition">epithelial to mesenchymal transition</a> (EMT), that reduces cell-cell adhesions and allows cancer spreading.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>The diagram on the right shows: </p> <ul><li>A: <a href="/wiki/Border_cells_(Drosophila)" title="Border cells (Drosophila)">Border cell</a> migration in a <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i> embryo. (a) shows border cells migrating in a confined space surrounded by gigantic nurse cells.<sup id="cite_ref-Barriga2019_10-1" class="reference"><a href="#cite_note-Barriga2019-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></li> <li>B: The initial position of <a href="/wiki/Zebrafish" title="Zebrafish">zebrafish</a> posterior <a href="/wiki/Lateral_line" title="Lateral line">lateral line</a> <a href="/wiki/Primordia" class="mw-redirect" title="Primordia">primordia</a> (pLLP) cells. (b) is a sagittal section showing how these cells migrate in a confined space between the somatic mesoderm and epidermis.<sup id="cite_ref-Barriga2019_10-2" class="reference"><a href="#cite_note-Barriga2019-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></li> <li>C: The cephalic <a href="/wiki/Neural_crest" title="Neural crest">neural crest</a> of the clawed frog <i><a href="/wiki/Xenopus" title="Xenopus">Xenopus</a></i> migrating in well-defined streams from dorsal to ventral and anterior. (c) is a transverse section across the head of a <i>Xenopus</i> embryo showing the high degree of confinement experienced by the neural crest while migrating sandwiched between the epidermis and underlying head mesoderm.<sup id="cite_ref-Barriga2019_10-3" class="reference"><a href="#cite_note-Barriga2019-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading2"><h2 id="Mathematical_models">Mathematical models</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Collective_cell_migration&action=edit&section=7" title="Edit section: Mathematical models"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are several mathematical models that describe collective cell motion. Typically, a <a href="/wiki/Classical_mechanics" title="Classical mechanics">Newtonian equation of motion</a> for a system of cells is solved.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> Several <a href="/wiki/Force" title="Force">forces</a> act on each individual cell, examples are <a href="/wiki/Friction" title="Friction">friction</a> (between environment and other cells), chemotaxis and <a href="/wiki/Self-propelled_particles" title="Self-propelled particles">self-propulsion</a>. The latter implies that cells are <a href="/wiki/Active_matter" title="Active matter">active</a> matter far from <a href="/wiki/Thermal_equilibrium" title="Thermal equilibrium">thermal equilibrium</a> that are able to generate force due to myosin-actin contractile motion. An overview over physical description of collective cell migration<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> explains that the following types of models can be used: </p> <ul><li><a href="/wiki/Lattice_model_(physics)" title="Lattice model (physics)">Lattice models</a> (e.g. <a href="/wiki/BIO-LGCA" title="BIO-LGCA">BIO-LGCA</a> models)</li> <li>Models similar to <a href="/wiki/Dissipative_particle_dynamics" title="Dissipative particle dynamics">Dissipative particle dynamics</a> that solve Newton's equation of motion with dissipative and random forces</li> <li>Models where cells depict <a href="/wiki/Voronoi_diagram" title="Voronoi diagram">Voronoi regions</a> and an effective potential (based on Voronoi graphs) for the tissue is used</li> <li>Continuum models, e.g. with the use of a phase field</li> <li>Kinetic theories similar to the <a href="/wiki/Boltzmann_equation" title="Boltzmann equation">Boltzmann equation</a><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup></li></ul> <p>These mathematical models give some insight in complex phenomena like cancer, wound healing<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> and <a href="/wiki/Ectoplasm_(paranormal)" title="Ectoplasm (paranormal)">ectoplasms</a>. </p> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:324px;max-width:324px"><div class="trow"><div class="tsingle" style="width:322px;max-width:322px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Chemotaxis_of_a_cell_up_a_fixed_gradient.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Chemotaxis_of_a_cell_up_a_fixed_gradient.jpg/320px-Chemotaxis_of_a_cell_up_a_fixed_gradient.jpg" decoding="async" width="320" height="177" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Chemotaxis_of_a_cell_up_a_fixed_gradient.jpg/480px-Chemotaxis_of_a_cell_up_a_fixed_gradient.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Chemotaxis_of_a_cell_up_a_fixed_gradient.jpg/640px-Chemotaxis_of_a_cell_up_a_fixed_gradient.jpg 2x" data-file-width="1280" data-file-height="709" /></a></span></div><div class="thumbcaption"><a href="/wiki/Chemotaxis" title="Chemotaxis">Chemotaxis</a> of a cell up a fixed (left, blue) and cell-induced, or self-generated, gradient (right, green) of chemoattractant. Lines show the concentration (c) of chemoattractant along space (x), in which cells (ellipses) migrate. Darker shapes illustrate successive time-points.<sup id="cite_ref-Schumacher2016_25-0" class="reference"><a href="#cite_note-Schumacher2016-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></div></div></div><div class="trow"><div class="tsingle" style="width:322px;max-width:322px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Spectrum_of_cell%E2%80%93cell_interactions_from_repulsive_to_volume_exclusion.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Spectrum_of_cell%E2%80%93cell_interactions_from_repulsive_to_volume_exclusion.jpg/320px-Spectrum_of_cell%E2%80%93cell_interactions_from_repulsive_to_volume_exclusion.jpg" decoding="async" width="320" height="261" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Spectrum_of_cell%E2%80%93cell_interactions_from_repulsive_to_volume_exclusion.jpg/480px-Spectrum_of_cell%E2%80%93cell_interactions_from_repulsive_to_volume_exclusion.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Spectrum_of_cell%E2%80%93cell_interactions_from_repulsive_to_volume_exclusion.jpg/640px-Spectrum_of_cell%E2%80%93cell_interactions_from_repulsive_to_volume_exclusion.jpg 2x" data-file-width="1280" data-file-height="1045" /></a></span></div><div class="thumbcaption">Spectrum of cell–cell interactions, from repulsive interactions (left, blue) to volume exclusion (right, green). With repulsive interactions, cells move away from the point of contact with another cell (second cell shown as stationary for simplicity). With volume exclusion, cells block each other's movement, but can move to any space not occupied by another cell.<sup id="cite_ref-Schumacher2016_25-1" class="reference"><a href="#cite_note-Schumacher2016-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></div></div></div></div></div> <div class="mw-heading mw-heading3"><h3 id="Spectrum_of_collective_cell_migration">Spectrum of collective cell migration</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Collective_cell_migration&action=edit&section=8" title="Edit section: Spectrum of collective cell migration"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the diagram immediately below, different morphologies of collective cell migration are characterized by their cohesiveness during migration (inversely related to density), as well as the number of nearest neighbours with which a cell interacts while moving (i.e. the topological arrangement of individual cells in the population). Cells (ellipses) can migrate in linear chains (top left), with persistent contact to cells either side of them, or along trails formed by preceding cells (bottom left). In migrating sheets, cells may maintain most of their nearest neighbours over time (top right), whereas in streaming migration cell–cell contacts occur at longer range and with potentially frequent neighbour rearrangement (bottom right). These concepts easily extend to three-dimensional migration, in which case the place of migrating sheets can be taken by moving clusters or spheroids.<sup id="cite_ref-Schumacher2016_25-2" class="reference"><a href="#cite_note-Schumacher2016-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Spectrum_of_collective_cell_migration.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/Spectrum_of_collective_cell_migration.jpg/400px-Spectrum_of_collective_cell_migration.jpg" decoding="async" width="400" height="338" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/Spectrum_of_collective_cell_migration.jpg/600px-Spectrum_of_collective_cell_migration.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/60/Spectrum_of_collective_cell_migration.jpg/800px-Spectrum_of_collective_cell_migration.jpg 2x" data-file-width="1280" data-file-height="1083" /></a><figcaption> <div class="center" style="width:auto; margin-left:auto; margin-right:auto;"><b>Spectrum of collective cell migration</b> <sup id="cite_ref-Schumacher2016_25-3" class="reference"><a href="#cite_note-Schumacher2016-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></div></figcaption></figure> <div style="clear:both;" class=""></div> <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=Collective_cell_migration&action=edit&section=9" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Bacteria_collective_motion" title="Bacteria collective motion">Bacteria collective motion</a></li> <li><a href="/wiki/Boids" title="Boids">Boids</a></li> <li><a href="/wiki/Collective_animal_behavior" title="Collective animal behavior">Collective animal behavior</a></li> <li><a href="/wiki/Collective_motion" title="Collective motion">Collective motion</a></li> <li><a href="/wiki/Embryogenesis" class="mw-redirect" title="Embryogenesis">Embryogenesis</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=Collective_cell_migration&action=edit&section=10" 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-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</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="CITEREFFriedlHegerfeldtTusch2004" class="citation journal cs1">Friedl, P; 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