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{{Short description|Process in cell biology}}
{{Short description|Process in cell biology}}


'''Cell extrusion''', discovered in 2001,<ref>{{cite journal |last1=Rosenblatt |first1=Jody |last2=Raff |first2=Martin C. |last3=Cramer |first3=Louise P. |title=An epithelial cell destined for apoptosis signals its neighbors to extrude it by an actin- and myosin-dependent mechanism |journal=Current Biology |date=November 2001 |volume=11 |issue=23 |pages=1847–1857 |doi=10.1016/s0960-9822(01)00587-5|pmid=11728307 |s2cid=5858676 }}</ref> is the process used by [[epithelial cell]]s to remove unwanted or dying cells and at the same time maintaining the integrity of the barrier.<ref name="pathways">{{cite journal |last1=Gudipaty |first1=Swapna Aravind |last2=Rosenblatt |first2=Jody |title=Epithelial cell extrusion: Pathways and pathologies |journal=Seminars in Cell & Developmental Biology |date=July 2017 |volume=67 |pages=132–140 |doi=10.1016/j.semcdb.2016.05.010|pmc=5116298 }}</ref> If cells were to die without extrusion, gaps would be created, compromising the epithelia's function. Cell extrusion occurs primarily in the [[endothelia]] and epithelia where the cells are linked to one another by tight cell-to-cell junctions.<ref>{{cite journal |last1=Ohsawa |first1=Shizue |last2=Vaughen |first2=John |last3=Igaki |first3=Tatsushi |title=Cell Extrusion: A Stress-Responsive Force for Good or Evil in Epithelial Homeostasis |journal=Developmental Cell |date=February 2018 |volume=44 |issue=3 |pages=284–296 |doi=10.1016/j.devcel.2018.01.009}}</ref>
'''Cell extrusion''', discovered in 2001,<ref name=":0">{{cite journal |last1=Rosenblatt |first1=Jody |last2=Raff |first2=Martin C. |last3=Cramer |first3=Louise P. |title=An epithelial cell destined for apoptosis signals its neighbors to extrude it by an actin- and myosin-dependent mechanism |journal=Current Biology |date=November 2001 |volume=11 |issue=23 |pages=1847–1857 |doi=10.1016/s0960-9822(01)00587-5|pmid=11728307 |s2cid=5858676 }}</ref> is a process conserved in [[epithelial cell|epithelial]] from humans to sea sponge<ref>{{Cite journal |last=Dwivedi |first=Vivek K. |last2=Pardo-Pastor |first2=Carlos |last3=Droste |first3=Rita |last4=Kong |first4=Ji Na |last5=Tucker |first5=Nolan |last6=Denning |first6=Daniel P. |last7=Rosenblatt |first7=Jody |last8=Horvitz |first8=H. Robert |date=2021-05 |title=Replication stress promotes cell elimination by extrusion |url=https://pubmed.ncbi.nlm.nih.gov/33953402/ |journal=Nature |volume=593 |issue=7860 |pages=591–596 |doi=10.1038/s41586-021-03526-y |issn=1476-4687 |pmc=8403516 |pmid=33953402}}</ref> to seamlessly remove unwanted or dying cells while maintaining the integrity of the epithelial barrier.<ref name="pathways">{{cite journal |last1=Gudipaty |first1=Swapna Aravind |last2=Rosenblatt |first2=Jody |title=Epithelial cell extrusion: Pathways and pathologies |journal=Seminars in Cell & Developmental Biology |date=July 2017 |volume=67 |pages=132–140 |doi=10.1016/j.semcdb.2016.05.010|pmc=5116298 }}</ref> If cells were to die without extrusion, gaps would be created, compromising the epithelia's function. While cell targeted to die by apoptotic stimuli extrude to prevent gaps from forming<ref name=":0" />, most cells die as a result of extruding live cells<ref name=":1">{{Cite journal |last=Eisenhoffer |first=George T. |last2=Loftus |first2=Patrick D. |last3=Yoshigi |first3=Masaaki |last4=Otsuna |first4=Hideo |last5=Chien |first5=Chi-Bin |last6=Morcos |first6=Paul A. |last7=Rosenblatt |first7=Jody |date=2012-04-15 |title=Crowding induces live cell extrusion to maintain homeostatic cell numbers in epithelia |url=https://pubmed.ncbi.nlm.nih.gov/22504183/ |journal=Nature |volume=484 |issue=7395 |pages=546–549 |doi=10.1038/nature10999 |issn=1476-4687 |pmc=4593481 |pmid=22504183}}</ref>. To maintain epithelial cell number homeostasis, live cells extrude when they become too crowded.


==Triggers==
==Triggers==
Various factors such as [[apoptosis]], overcrowding, [[pathogens]] and [[oncogenic]] markers are triggers of cell extrusion.<ref>{{cite journal |last1=Fadul |first1=John |last2=Zulueta-Coarasa |first2=Teresa |last3=Slattum |first3=Gloria M. |last4=Redd |first4=Nadja M. |last5=Jin |first5=Mauricio Franco |last6=Redd |first6=Michael J. |last7=Daetwyler |first7=Stephan |last8=Hedeen |first8=Danielle |last9=Huisken |first9=Jan |last10=Rosenblatt |first10=Jody |title=KRas-transformed epithelia cells invade and partially dedifferentiate by basal cell extrusion |journal=Nature Communications |date=10 December 2021 |volume=12 |issue=1 |pages=7180 |doi=10.1038/s41467-021-27513-z|s2cid=245021043 }}</ref> When a cell has been triggered to extrude, it can be seamlessly ejected into the apical environment or the basal environment. In the airway epithelia and intestinal endothelia, apical cell extrusions are important because the cell is extruded into the external environment.<ref name="pathways"/> However, this is not so with basal extrusions where the cell is extruded into the tissue. Basal extrusions are associated with the onset of [[metastasis]].<ref>{{cite journal |last1=Fadul |first1=John |last2=Rosenblatt |first2=Jody |title=The forces and fates of extruding cells |journal=Current Opinion in Cell Biology |date=1 October 2018 |volume=54 |pages=66–71 |doi=10.1016/j.ceb.2018.04.007|pmid=29727745 |pmc=6212327 }}</ref>
Various factors such as [[apoptosis]]<ref name=":0" />, overcrowding<ref name=":1" />, [[pathogens]]<ref>{{Cite journal |last=Gudipaty |first=Swapna Aravind |last2=Rosenblatt |first2=Jody |date=2017-07-01 |title=Epithelial cell extrusion: Pathways and pathologies |url=https://www.sciencedirect.com/science/article/pii/S1084952116301367 |journal=Seminars in Cell & Developmental Biology |series=Extracellular Vesicles |language=en |volume=67 |pages=132–140 |doi=10.1016/j.semcdb.2016.05.010 |issn=1084-9521 |pmc=PMC5116298 |pmid=27212253}}</ref> and replicative stress<ref>{{Cite journal |last=Dwivedi |first=Vivek K. |last2=Pardo-Pastor |first2=Carlos |last3=Droste |first3=Rita |last4=Kong |first4=Ji Na |last5=Tucker |first5=Nolan |last6=Denning |first6=Daniel P. |last7=Rosenblatt |first7=Jody |last8=Horvitz |first8=H. Robert |date=2021-05 |title=Replication stress promotes cell elimination by extrusion |url=https://www.nature.com/articles/s41586-021-03526-y |journal=Nature |language=en |volume=593 |issue=7860 |pages=591–596 |doi=10.1038/s41586-021-03526-y |issn=1476-4687}}</ref> can trigger extrusion from epithelia. Additionally, cells transformed with [[oncogenic]] mutations such as [[HRAS]]<ref>{{Cite journal |last=Hogan |first=Catherine |last2=Kajita |first2=Mihoko |last3=Lawrenson |first3=Kate |last4=Fujita |first4=Yasuyuki |date=2011-04 |title=Interactions between normal and transformed epithelial cells: Their contributions to tumourigenesis |url=https://linkinghub.elsevier.com/retrieve/pii/S1357272510004310 |journal=The International Journal of Biochemistry & Cell Biology |language=en |volume=43 |issue=4 |pages=496–503 |doi=10.1016/j.biocel.2010.12.019}}</ref> and [[Src family kinase|Src]] can be ejected from epithelia by a similar extrusion process called Epithelial Defense Against Cancer (EDAC)<ref>{{Cite journal |last=Tanimura |first=Nobuyuki |last2=Fujita |first2=Yasuyuki |date=2020-06 |title=Epithelial defense against cancer (EDAC) |url=https://linkinghub.elsevier.com/retrieve/pii/S1044579X19300331 |journal=Seminars in Cancer Biology |language=en |volume=63 |pages=44–48 |doi=10.1016/j.semcancer.2019.05.011}}</ref>.


==Functions==
==Functions ==
Cell extrusion enables the removal of less fit and excess cells from the epithelia and endothelia. The removal of overcrowded cells enables the epithelia to maintain a [[Homeostasis|homeostatic]] cell population.<ref name="pathways"/>
Cell extrusion enables the removal of less fit and excess cells from the epithelia and endothelia. The removal of overcrowded cells enables the epithelia to maintain a [[Homeostasis|homeostatic]] cell population.<ref name="pathways"/>

While most cells extrude apically out of the epithelia organs encase, some oncogenic drivers of aggressive tumors hijack this process to drive extrusion the opposite direction--basally, back into the [[Stromal cell|stroma]]<ref>{{Cite journal |last=Slattum |first=Gloria |last2=Gu |first2=Yapeng |last3=Sabbadini |first3=Roger |last4=Rosenblatt |first4=Jody |date=2014-01 |title=Autophagy in Oncogenic K-Ras Promotes Basal Extrusion of Epithelial Cells by Degrading S1P |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982213014486 |journal=Current Biology |language=en |volume=24 |issue=1 |pages=19–28 |doi=10.1016/j.cub.2013.11.029 |pmc=PMC3932369 |pmid=24361067}}</ref>.<ref>{{Cite journal |last=Marshall |first=Thomas W. |last2=Lloyd |first2=Isaac E. |last3=Delalande |first3=Jean Marie |last4=Näthke |first4=Inke |last5=Rosenblatt |first5=Jody |date=2011-11 |editor-last=Yap |editor-first=Alpha |title=The tumor suppressor adenomatous polyposis coli controls the direction in which a cell extrudes from an epithelium |url=https://www.molbiolcell.org/doi/10.1091/mbc.e11-05-0469 |journal=Molecular Biology of the Cell |language=en |volume=22 |issue=21 |pages=3962–3970 |doi=10.1091/mbc.e11-05-0469 |issn=1059-1524 |pmc=PMC3204059 |pmid=21900494}}</ref> Cells transformed with KRas, a driver of pancreatic, and some types of lung and colon cancer, causes cells to invaded by aberrant Basal Cell Extrusion (BCE), causing them to lose their surface-associated epithelial determinants and to initiate metastasis<ref>{{Cite journal |last=Fadul |first=John |last2=Zulueta-Coarasa |first2=Teresa |last3=Slattum |first3=Gloria M. |last4=Redd |first4=Nadja M. |last5=Jin |first5=Mauricio Franco |last6=Redd |first6=Michael J. |last7=Daetwyler |first7=Stephan |last8=Hedeen |first8=Danielle |last9=Huisken |first9=Jan |last10=Rosenblatt |first10=Jody |date=2021-12 |title=KRas-transformed epithelia cells invade and partially dedifferentiate by basal cell extrusion |url=https://www.nature.com/articles/s41467-021-27513-z |journal=Nature Communications |language=en |volume=12 |issue=1 |pages=7180 |doi=10.1038/s41467-021-27513-z |issn=2041-1723 |pmc=PMC8664939 |pmid=34893591}}</ref>. Thus, oncogenic transformation can cause cells to mechanically de-differentiate and invade by hijacking a process that normally drives cell death.


== References ==
== References ==

Revision as of 09:33, 14 August 2022

Cell extrusion, discovered in 2001,[1] is a process conserved in epithelial from humans to sea sponge[2] to seamlessly remove unwanted or dying cells while maintaining the integrity of the epithelial barrier.[3] If cells were to die without extrusion, gaps would be created, compromising the epithelia's function. While cell targeted to die by apoptotic stimuli extrude to prevent gaps from forming[1], most cells die as a result of extruding live cells[4]. To maintain epithelial cell number homeostasis, live cells extrude when they become too crowded.

Triggers

Various factors such as apoptosis[1], overcrowding[4], pathogens[5] and replicative stress[6] can trigger extrusion from epithelia. Additionally, cells transformed with oncogenic mutations such as HRAS[7] and Src can be ejected from epithelia by a similar extrusion process called Epithelial Defense Against Cancer (EDAC)[8].

Functions

Cell extrusion enables the removal of less fit and excess cells from the epithelia and endothelia. The removal of overcrowded cells enables the epithelia to maintain a homeostatic cell population.[3]

While most cells extrude apically out of the epithelia organs encase, some oncogenic drivers of aggressive tumors hijack this process to drive extrusion the opposite direction--basally, back into the stroma[9].[10] Cells transformed with KRas, a driver of pancreatic, and some types of lung and colon cancer, causes cells to invaded by aberrant Basal Cell Extrusion (BCE), causing them to lose their surface-associated epithelial determinants and to initiate metastasis[11]. Thus, oncogenic transformation can cause cells to mechanically de-differentiate and invade by hijacking a process that normally drives cell death.

References

  1. ^ a b c Rosenblatt, Jody; Raff, Martin C.; Cramer, Louise P. (November 2001). "An epithelial cell destined for apoptosis signals its neighbors to extrude it by an actin- and myosin-dependent mechanism". Current Biology. 11 (23): 1847–1857. doi:10.1016/s0960-9822(01)00587-5. PMID 11728307. S2CID 5858676.
  2. ^ Dwivedi, Vivek K.; Pardo-Pastor, Carlos; Droste, Rita; Kong, Ji Na; Tucker, Nolan; Denning, Daniel P.; Rosenblatt, Jody; Horvitz, H. Robert (2021-05). "Replication stress promotes cell elimination by extrusion". Nature. 593 (7860): 591–596. doi:10.1038/s41586-021-03526-y. ISSN 1476-4687. PMC 8403516. PMID 33953402. {{cite journal}}: Check date values in: |date= (help)
  3. ^ a b Gudipaty, Swapna Aravind; Rosenblatt, Jody (July 2017). "Epithelial cell extrusion: Pathways and pathologies". Seminars in Cell & Developmental Biology. 67: 132–140. doi:10.1016/j.semcdb.2016.05.010. PMC 5116298.
  4. ^ a b Eisenhoffer, George T.; Loftus, Patrick D.; Yoshigi, Masaaki; Otsuna, Hideo; Chien, Chi-Bin; Morcos, Paul A.; Rosenblatt, Jody (2012-04-15). "Crowding induces live cell extrusion to maintain homeostatic cell numbers in epithelia". Nature. 484 (7395): 546–549. doi:10.1038/nature10999. ISSN 1476-4687. PMC 4593481. PMID 22504183.
  5. ^ Gudipaty, Swapna Aravind; Rosenblatt, Jody (2017-07-01). "Epithelial cell extrusion: Pathways and pathologies". Seminars in Cell & Developmental Biology. Extracellular Vesicles. 67: 132–140. doi:10.1016/j.semcdb.2016.05.010. ISSN 1084-9521. PMC 5116298. PMID 27212253.{{cite journal}}: CS1 maint: PMC format (link)
  6. ^ Dwivedi, Vivek K.; Pardo-Pastor, Carlos; Droste, Rita; Kong, Ji Na; Tucker, Nolan; Denning, Daniel P.; Rosenblatt, Jody; Horvitz, H. Robert (2021-05). "Replication stress promotes cell elimination by extrusion". Nature. 593 (7860): 591–596. doi:10.1038/s41586-021-03526-y. ISSN 1476-4687. {{cite journal}}: Check date values in: |date= (help)
  7. ^ Hogan, Catherine; Kajita, Mihoko; Lawrenson, Kate; Fujita, Yasuyuki (2011-04). "Interactions between normal and transformed epithelial cells: Their contributions to tumourigenesis". The International Journal of Biochemistry & Cell Biology. 43 (4): 496–503. doi:10.1016/j.biocel.2010.12.019. {{cite journal}}: Check date values in: |date= (help)
  8. ^ Tanimura, Nobuyuki; Fujita, Yasuyuki (2020-06). "Epithelial defense against cancer (EDAC)". Seminars in Cancer Biology. 63: 44–48. doi:10.1016/j.semcancer.2019.05.011. {{cite journal}}: Check date values in: |date= (help)
  9. ^ Slattum, Gloria; Gu, Yapeng; Sabbadini, Roger; Rosenblatt, Jody (2014-01). "Autophagy in Oncogenic K-Ras Promotes Basal Extrusion of Epithelial Cells by Degrading S1P". Current Biology. 24 (1): 19–28. doi:10.1016/j.cub.2013.11.029. PMC 3932369. PMID 24361067. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
  10. ^ Marshall, Thomas W.; Lloyd, Isaac E.; Delalande, Jean Marie; Näthke, Inke; Rosenblatt, Jody (2011-11). Yap, Alpha (ed.). "The tumor suppressor adenomatous polyposis coli controls the direction in which a cell extrudes from an epithelium". Molecular Biology of the Cell. 22 (21): 3962–3970. doi:10.1091/mbc.e11-05-0469. ISSN 1059-1524. PMC 3204059. PMID 21900494. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
  11. ^ Fadul, John; Zulueta-Coarasa, Teresa; Slattum, Gloria M.; Redd, Nadja M.; Jin, Mauricio Franco; Redd, Michael J.; Daetwyler, Stephan; Hedeen, Danielle; Huisken, Jan; Rosenblatt, Jody (2021-12). "KRas-transformed epithelia cells invade and partially dedifferentiate by basal cell extrusion". Nature Communications. 12 (1): 7180. doi:10.1038/s41467-021-27513-z. ISSN 2041-1723. PMC 8664939. PMID 34893591. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
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