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'''Collective cell migration''' describes the movements of group of cells and the [[emergence]] of collective behavior from cell-environment interactions and cell-cell communication. Collective cell migration is an essential process in the lives of [[multicellular organisms]], e.g. [[embryonic development]], [[wound healing]] and cancer spreading ([[metastasis]]).<ref>{{cite journal|last1=Friedl|first1=P|last2=Hegerfeldt|first2=Y|last3=Tusch|first3=M|title=Collective cell migration in morphogenesis and cancer|journal=[[The International Journal of Developmental Biology]]|date=2004|volume=48|issue=5–6|pages=441–9|doi=10.1387/ijdb.041821pf|pmid=15349818|doi-access=free}}</ref> Cells can migrate as a cohesive group (e.g. [[epithelial cells]]) or have transient cell-cell adhesion sites (e.g. [[mesenchyme|mesenchymal]] cells).<ref>{{cite journal|last1=Weijer|first1=CJ|title=Collective cell migration in development|journal=[[Journal of Cell Science]]|date=15 September 2009|volume=122|issue=Pt 18|pages=3215–23|doi=10.1242/jcs.036517|pmid=19726631|doi-access=free}}{{open access}}</ref> They can also migrate in different modes like sheets, strands, tubes, and clusters.<ref>{{cite journal|last1=Friedl|first1=P|title=Prespecification and plasticity: shifting mechanisms of cell migration|journal=[[Current Opinion in Cell Biology]]|date=February 2004|volume=16|issue=1|pages=14–23|doi=10.1016/j.ceb.2003.11.001|pmid=15037300}}</ref> While [[Cell migration|single-cell migration]] has been extensively studied, collective cell migration is a relatively new field with applications in preventing birth defects or dysfunction of embryos. It may improve cancer treatment by enabling doctors to prevent tumors from spreading and forming new tumors.

== Cell-environment interactions ==
{{microbial and microbot movement|collective}}

The environment of the migrating cell can affect its speed, persistence and direction of migration by stimulating it. The [[extracellular matrix]] (ECM) provides not only the structural and biochemical support, but also plays a major role in regulating cell behavior. Different ECM proteins (such as [[collagen]], [[elastin]], [[fibronectin]], [[laminin]], and others) allow cells to adhere and migrate, while forming [[focal adhesions]] 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 ([[haptotaxis]]) or a gradient of soluble substrates in the liquid phase surrounding the cell ([[chemotaxis]]). Cells sense the substrate through their [[receptor (biochemistry)|receptor]]s and migrate toward the concentration (or the opposite direction). Another form of stimulation can be rigidity gradients of the ECM ([[durotaxis]]).{{citation needed|date=May 2017}}

=== Confinement ===
Collective cell migration is enhanced by geometrical confinement of an [[extracellular matrix]] [[molecule]] (e.g. the [[proteoglycan]] [[versican]] in [[neural crest]] cells), that acts as a barrier, to promote the [[emergence]] of organized migration in separated streams. Confinement is also observed [[in vivo]], where the optimal width is a function of the number of migrating cells in different streams of different [[species]].<ref>{{cite journal|last1=Szabó|first1=András|last2=Melchionda|first2=Manuela|last3=Nastasi|first3=Giancarlo|last4=Woods|first4=Mae L.|last5=Campo|first5=Salvatore|last6=Perris|first6=Roberto|last7=Mayor|first7=Roberto|title=In vivo confinement promotes collective migration of neural crest cells|journal=[[Journal of Cell Biology]]|date=6 June 2016|volume=213|issue=5|pages=543–555|doi=10.1083/jcb.201602083|pmid=27241911|pmc=4896058}}</ref>

== Cell-cell communication ==
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 [[random walk]] movements of the isolated cell. Cell-cell communication mechanisms are widely studied experimentally ([[in vivo]] and [[in vitro]]),<ref>{{cite journal|last1=Mayor|first1=R|last2=Etienne-Manneville|first2=S|title=The front and rear of collective cell migration|journal=[[Nature Reviews Molecular Cell Biology|Nature Reviews. Molecular Cell Biology]]|date=February 2016|volume=17|issue=2|pages=97–109|doi=10.1038/nrm.2015.14|pmid=26726037|s2cid=27261044|url=http://discovery.ucl.ac.uk/1483697/1/Mayor_acepted.pdf}}</ref> and computationally ([[in silico]]).<ref>{{cite journal|last1=Szabó|first1=A|last2=Mayor|first2=R|title=Modelling collective cell migration of neural crest|journal=[[Current Opinion in Cell Biology]]|date=October 2016|volume=42|pages=22–28|doi=10.1016/j.ceb.2016.03.023|pmid=27085004|pmc=5017515}}</ref>

=== Co-attraction ===
Co-attraction between collectively migrating cells is the process by which cells of the same type secrete [[chemotaxis|chemo-attractant]] (e.g. [[C3a (complement)|C3a]] in neural crest cells), that stimulates other cells in the group that have the [[receptor (biochemistry)|receptor]]s 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.<ref>{{cite journal|last1=Woods|first1=ML|last2=Carmona-Fontaine|first2=C|last3=Barnes|first3=CP|last4=Couzin|first4=ID|last5=Mayor|first5=R|last6=Page|first6=KM|title=Directional collective cell migration emerges as a property of cell interactions|journal=[[PLoS ONE]]|date=2014|volume=9|issue=9|pages=e104969|doi=10.1371/journal.pone.0104969|pmid=25181349|pmc=4152153|bibcode=2014PLoSO...9j4969W|doi-access=free}}</ref><ref>{{cite journal|last1=Carmona-Fontaine|first1=C|last2=Theveneau|first2=E|last3=Tzekou|first3=A|last4=Tada|first4=M|last5=Woods|first5=M|last6=Page|first6=KM|last7=Parsons|first7=M|last8=Lambris|first8=JD|last9=Mayor|first9=R|title=Complement fragment C3a controls mutual cell attraction during collective cell migration|journal=[[Developmental Cell]]|date=13 December 2011|volume=21|issue=6|pages=1026–37|doi=10.1016/j.devcel.2011.10.012|pmid=22118769|pmc=3272547}}</ref>

=== Contact inhibition of locomotion ===
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 ([[Cell junction|cell-junctions]]) are created by [[Transmembrane protein|transmembrane]] [[glycoprotein]]s named [[cadherin]]s ([[E-cadherin]], [[N-cadherin]] or [[cadherin 11]]) and other [[protein]]s. 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.{{citation needed|date=May 2017}}
* [[Contact inhibition|Contact inhibition of proliferation]] (CIP) is the inhibition of cell division with increasing percent of confluency. CIP and CIL are two different processes, which are sometimes mistakenly interrelated.<ref>{{cite journal|last1=Stoker|first1=MG|last2=Rubin|first2=H|title=Density dependent inhibition of cell growth in culture|journal=Nature|date=8 July 1967|volume=215|issue=5097|pages=171–2|pmid=6049107|doi=10.1038/215171a0|bibcode=1967Natur.215..171S|s2cid=4150783}}</ref>

== Examples of studied systems ==
[[File:Models for studying collective cell migration.jpg|thumb|upright=1.6| {{center|'''Models for studying collective cell migration'''{{hsp}}<ref name=Barriga2019>{{cite journal |doi = 10.1016/j.semcdb.2018.05.027|title = Adjustable viscoelasticity allows for efficient collective cell migration|year = 2019|last1 = Barriga|first1 = Elias H.|last2 = Mayor|first2 = Roberto|journal = Seminars in Cell & Developmental Biology|volume = 93|pages = 55–68|pmid = 29859995|pmc = 6854469}} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</ref><br /><small>Red arrows show the direction of migration for each tissue</small>}}]]

Collective cell migration is studied over many model species.

[[Border cells]] in flies (''[[Drosophila melanogaster]]''): the border cells migrate during the differentiation of egg cells to be ready for fertilization.<ref>{{cite journal|last1=Bianco|first1=A|last2=Poukkula|first2=M|last3=Cliffe|first3=A|last4=Mathieu|first4=J|last5=Luque|first5=CM|last6=Fulga|first6=TA|last7=Rørth|first7=P|title=Two distinct modes of guidance signalling during collective migration of border cells|journal=Nature|date=19 July 2007|volume=448|issue=7151|pages=362–5|doi=10.1038/nature05965|pmid=17637670|bibcode=2007Natur.448..362B|s2cid=4369682}}</ref>

The [[lateral line]] in [[zebrafish]]: 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.<ref>{{cite journal|last1=Dalle Nogare|first1=D|last2=Somers|first2=K|last3=Rao|first3=S|last4=Matsuda|first4=M|last5=Reichman-Fried|first5=M|last6=Raz|first6=E|last7=Chitnis|first7=AB|title=Leading and trailing cells cooperate in collective migration of the zebrafish posterior lateral line primordium|journal=Development|date=August 2014|volume=141|issue=16|pages=3188–96|doi=10.1242/dev.106690|pmid=25063456|pmc=4197546}}</ref>

[[Wound healing]]: collective cell migration is an essential part in this healing process, wound area is closed by the migrating cells.<ref name="pmid28110712">{{cite journal | vauthors = Grada A | title = Analysis of Collective Cell Migration Using the Wound Healing Assay | journal = J Invest Dermatol | volume = 137 | issue = 2 | pages = e11–e16 | date = February 2017 | pmid = 28110712 | doi=10.1016/j.jid.2016.11.020 | doi-access = free }}</ref><ref>{{cite journal|last1=Trepat|first1=Xavier|last2=Wasserman|first2=Michael R.|last3=Angelini|first3=Thomas E.|last4=Millet|first4=Emil|last5=Weitz|first5=David A.|last6=Butler|first6=James P.|last7=Fredberg|first7=Jeffrey J.|title=Physical forces during collective cell migration|journal=[[Nature Physics]]|date=3 May 2009|volume=5|issue=6|pages=426–430|doi=10.1038/nphys1269|bibcode=2009NatPh...5..426T|doi-access=free}}</ref> Wound healing is commonly studied in vitro using cell lines such as [[Madin-Darby Canine Kidney Cells|Madin-Darby Canine Kidney cells]].

[[Neural crest]] cells in [[mice]],<ref>{{cite journal|last1=Trainor|first1=PA|title=Specification of neural crest cell formation and migration in mouse embryos|journal=Seminars in Cell & Developmental Biology|date=December 2005|volume=16|issue=6|pages=683–93|doi=10.1016/j.semcdb.2005.06.007|pmid=16043371}}</ref> [[Leghorn chicken|Leghorn chicks]],<ref>{{cite journal|last1=Johnston|first1=MC|title=A radioautographic study of the migration and fate of cranial neural crest cells in the chick embryo|journal=[[The Anatomical Record]]|date=October 1966|volume=156|issue=2|pages=143–55|doi=10.1002/ar.1091560204|pmid=5969670|s2cid=11251193}}</ref> [[amphibian]]s (''[[Xenopus laevis]]''),<ref>{{cite journal|last1=Sadaghiani|first1=B|last2=Thiébaud|first2=CH|title=Neural crest development in the ''Xenopus laevis'' embryo, studied by interspecific transplantation and scanning electron microscopy|journal=Developmental Biology|date=November 1987|volume=124|issue=1|pages=91–110|doi=10.1016/0012-1606(87)90463-5|pmid=3666314}}</ref> and [[fish]]<ref>{{cite journal|last1=Smith|first1=M.|last2=Hickman|first2=A.|last3=Amanze|first3=D.|last4=Lumsden|first4=A.|last5=Thorogood|first5=P.|title=Trunk neural crest origin of caudal fin mesenchyme in the zebrafish ''Brachydanio rerio''|journal=Proceedings of the Royal Society B: Biological Sciences|date=23 May 1994|volume=256|issue=1346|pages=137–145|doi=10.1098/rspb.1994.0061|jstor=50346|bibcode=1994RSPSB.256..137S|s2cid=86494317}}</ref> ([[zebrafish]]): collective migration of neural crest cells occurs during embryo development of vertebrates. They migrate long distances from the head ([[neural tube]]) to give rise to different tissues.<ref>Le Douarin, Nicole, and Chaya Kalcheim. The neural crest. No. 36. Cambridge University Press, 1999.</ref>

Spreading of [[cancer]] ([[metastasis]]): 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 [[epithelial to mesenchymal transition]] (EMT), that reduces cell-cell adhesions and allows cancer spreading.<ref>{{cite journal|last1=Thiery|first1=JP|last2=Acloque|first2=H|last3=Huang|first3=RY|last4=Nieto|first4=MA|title=Epithelial-mesenchymal transitions in development and disease|journal=[[Cell (journal)|Cell]]|date=25 November 2009|volume=139|issue=5|pages=871–90|doi=10.1016/j.cell.2009.11.007|pmid=19945376|doi-access=free}}{{open access}}</ref>

The diagram on the right shows:
* A: [[Border cells (Drosophila)|Border cell]] migration in a ''[[Drosophila]]'' embryo. (a) shows border cells migrating in a confined space surrounded by gigantic nurse cells.<ref name=Barriga2019 />
* B: The initial position of [[zebrafish]] posterior [[lateral line]] [[primordia]]  ({{not a typo|pLLP}}) cells. (b) is a sagittal section showing how these cells migrate in a confined space between the somatic mesoderm and epidermis.<ref name=Barriga2019 />
* C: The cephalic [[neural crest]] of the clawed frog ''[[Xenopus]]'' migrating in well-defined streams from dorsal to ventral and anterior. (c) is a transverse section across the head of a ''Xenopus'' embryo showing the high degree of confinement experienced by the neural crest while migrating sandwiched between the epidermis and underlying head mesoderm.<ref name=Barriga2019 />

== Mathematical models ==
There are several mathematical models that describe collective cell motion. Typically, a [[Classical mechanics|Newtonian equation of motion]] for a system of cells is solved.<ref>{{Cite journal|last1=Akiyama|first1=Masakazu|last2=Sushida|first2=Takamichi|last3=Ishida|first3=Sumire|last4=Haga|first4=Hisashi|date=2017|title=Mathematical model of collective cell migrations based on cell polarity|journal=Development, Growth & Differentiation|language=en|volume=59|issue=5|pages=471–490|doi=10.1111/dgd.12381|pmid=28714585|issn=1440-169X|doi-access=free}}</ref> Several [[force]]s act on each individual cell, examples are [[friction]] (between environment and other cells), chemotaxis and [[Self-propelled particles|self-propulsion]]. The latter implies that cells are [[Active matter|active]] matter far from [[thermal equilibrium]] that are able to generate force due to myosin-actin contractile motion. An overview over physical description of collective cell migration<ref>{{Cite journal|last1=Alert|first1=Ricard|last2=Trepat|first2=Xavier|date=2020-03-10|title=Physical Models of Collective Cell Migration|url=https://www.annualreviews.org/doi/10.1146/annurev-conmatphys-031218-013516|journal=Annual Review of Condensed Matter Physics|volume=11|issue=1|pages=77–101|doi=10.1146/annurev-conmatphys-031218-013516|issn=1947-5454|arxiv=1905.07675|bibcode=2020ARCMP..11...77A |s2cid=159041328}}</ref> explains that the following types of models can be used:
* [[Lattice model (physics)|Lattice models]] (e.g. [[BIO-LGCA]] models)
* Models similar to [[Dissipative particle dynamics]] that solve Newton's equation of motion with dissipative and random forces
* Models where cells depict [[Voronoi diagram|Voronoi regions]] and an effective potential (based on Voronoi graphs) for the tissue is used
* Continuum models, e.g. with the use of a phase field
* Kinetic theories similar to the [[Boltzmann equation]]<ref>{{Cite journal |last1=Chauviere |first1=A. |last2=Hillen |first2=T. |last3=Preziosi |first3=L. |date=2007 |title=Modeling cell movement in anisotropic and heterogeneous network tissues |url=http://dx.doi.org/10.3934/nhm.2007.2.333 |journal=Networks & Heterogeneous Media |volume=2 |issue=2 |pages=333–357 |doi=10.3934/nhm.2007.2.333 |issn=1556-181X}}</ref>

These mathematical models give some insight in complex phenomena like cancer, wound healing<ref>{{Cite journal|last1=Arciero|first1=Julia C.|last2=Mi|first2=Qi|last3=Branca|first3=Maria F.|last4=Hackam|first4=David J.|last5=Swigon|first5=David|date=2011-02-02|title=Continuum Model of Collective Cell Migration in Wound Healing and Colony Expansion|url= |journal=Biophysical Journal|language=English|volume=100|issue=3|pages=535–543|doi=10.1016/j.bpj.2010.11.083|issn=0006-3495|pmid=21281567|pmc=3030184|bibcode=2011BpJ...100..535A }}</ref> and [[Ectoplasm (paranormal)|ectoplasms]].

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 | caption1  = [[Chemotaxis]] 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.<ref name=Schumacher2016 />
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 | caption2  = 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.<ref name=Schumacher2016 />
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===Spectrum of collective cell migration===
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.<ref name=Schumacher2016 />

[[File:Spectrum of collective cell migration.jpg|thumb|upright=1.8|left| {{center|'''Spectrum of collective cell migration'''{{hsp}}<ref name=Schumacher2016>{{cite journal |doi = 10.1098/rsob.160056|title = Multidisciplinary approaches to understanding collective cell migration in developmental biology|year = 2016|last1 = Schumacher|first1 = Linus J.|last2 = Kulesa|first2 = Paul M.|last3 = McLennan|first3 = Rebecca|last4 = Baker|first4 = Ruth E.|last5 = Maini|first5 = Philip K.|journal = Open Biology|volume = 6|issue = 6|page = 160056|pmid = 27278647|pmc = 4929938}} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</ref>}}]]

{{clear}}

== See also ==
* [[Bacteria collective motion]]
* [[Boids]]
* [[Collective animal behavior]]
* [[Collective motion]]
* [[Embryogenesis]]

== References ==
{{reflist|30em}}

== External links ==
* [http://www.cs.tau.ac.il/thesis/thesis/Zaritsky.Assaf-PhD-Thesis.pdf Computational Methods to Quantify and Model Collective Cell Migration]
* [http://www.cell.com/trends/cell-biology/pdf/S0962-8924(15)00165-8.pdf Taking Aim at Moving Targets in Computational Cell Migration]

[[Category:Cells]]
[[Category:Cancer]]

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