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Fc\u03b3 receptor (Fc\u03b3R)-mediated phagocytosis is initiated when these receptors are activated after binding immunoglobulin G (IgG). Receptor activation initiates a signaling cascade that leads to the formation of the phagocytic cup and culminates with ingestion of the foreign particle. In the experimental system termed \u201cfrustrated phagocytosis\u201d, cells attempt to internalize micropatterned disks of IgG. Cells that engage in frustrated phagocytosis form \u201crosettes\u201d of actin-enriched structures called podosomes around the IgG disk. The mechanism that generates the rosette pattern is unknown. We present data that supports the involvement of Cdc42, a member of the Rho family of GTPases, in pattern formation. Cdc42 acts downstream of receptor activation, upstream of actin polymerization, and is known to play a role in polarity establishment. Reaction-diffusion models for GTPase spatiotemporal dynamics exist. We demonstrate how the addition of negative feedback and minor changes to these models can generate the experimentally observed rosette pattern of podosomes. We show that this pattern formation can occur through two general mechanisms. In the first mechanism, an intermediate species forms a ring of high activity around the IgG disk, which then promotes rosette organization. The second mechanism does not require initial ring formation but relies on spatial gradients of intermediate chemical species that are selectively activated over the IgG patch. Finally, we analyze the models to suggest experiments to test their validity.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1010092","type":"journal-article","created":{"date-parts":[[2022,10,3]],"date-time":"2022-10-03T13:51:50Z","timestamp":1664805110000},"page":"e1010092","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":12,"title":["Spatial models of pattern formation during phagocytosis"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6240-9950","authenticated-orcid":true,"given":"John Cody","family":"Herron","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8882-6859","authenticated-orcid":true,"given":"Shiqiong","family":"Hu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5979-3082","authenticated-orcid":true,"given":"Bei","family":"Liu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9015-5246","authenticated-orcid":true,"given":"Takashi","family":"Watanabe","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1970-7562","authenticated-orcid":true,"given":"Klaus M.","family":"Hahn","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4601-2117","authenticated-orcid":true,"given":"Timothy C.","family":"Elston","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2022,10,3]]},"reference":[{"key":"pcbi.1010092.ref001","doi-asserted-by":"crossref","first-page":"100354","DOI":"10.1016\/j.coisb.2021.100354","article-title":"Shapes of cell signaling","volume":"27","author":"R Linding","year":"2021","journal-title":"Curr Opin Syst Biol"},{"issue":"6916","key":"pcbi.1010092.ref002","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1038\/nature01148","article-title":"Rho GTPases in cell biology","volume":"420","author":"S Etienne-Manneville","year":"2002","journal-title":"Nature"},{"issue":"8","key":"pcbi.1010092.ref003","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1038\/nrm.2016.67","article-title":"Regulating Rho GTPases and their regulators","volume":"17","author":"RG Hodge","year":"2016","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"2","key":"pcbi.1010092.ref004","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1083\/jcb.201612069","article-title":"Rho GTPase signaling complexes in cell migration and invasion","volume":"217","author":"CD Lawson","year":"2018","journal-title":"J Cell Biol"},{"issue":"10","key":"pcbi.1010092.ref005","doi-asserted-by":"crossref","first-page":"1955","DOI":"10.1016\/j.cellsig.2013.04.009","article-title":"Signaling networks of Rho GTPases in cell motility","volume":"25","author":"S Hanna","year":"2013","journal-title":"Cell Signal"},{"issue":"2","key":"pcbi.1010092.ref006","doi-asserted-by":"crossref","first-page":"170","DOI":"10.4161\/cam.5.2.14403","article-title":"Dynamics of the Rho-family small GTPases in actin regulation and motility","volume":"5","author":"D Spiering","year":"2011","journal-title":"Cell Adhes Migr"},{"issue":"9","key":"pcbi.1010092.ref007","doi-asserted-by":"crossref","first-page":"3684","DOI":"10.1529\/biophysj.107.120824","article-title":"Wave-Pinning and Cell Polarity from a Bistable Reaction-Diffusion System","volume":"94","author":"Y Mori","year":"2008","journal-title":"Biophys J"},{"issue":"4","key":"pcbi.1010092.ref008","doi-asserted-by":"crossref","first-page":"e1006095","DOI":"10.1371\/journal.pcbi.1006095","article-title":"Principles that govern competition or co-existence in Rho-GTPase driven polarization","volume":"14","author":"JG Chiou","year":"2018","journal-title":"PLOS Comput Biol."},{"key":"pcbi.1010092.ref009","doi-asserted-by":"crossref","first-page":"e58768","DOI":"10.7554\/eLife.58768","article-title":"How cells determine the number of polarity sites.","volume":"10","author":"Chiou J geng","year":"2021","journal-title":"eLife"},{"issue":"3","key":"pcbi.1010092.ref010","doi-asserted-by":"crossref","first-page":"e0213188","DOI":"10.1371\/journal.pone.0213188","article-title":"Small GTPase patterning: How to stabilise cluster coexistence","volume":"14","author":"B Jacobs","year":"2019","journal-title":"PLOS ONE"},{"issue":"9","key":"pcbi.1010092.ref011","doi-asserted-by":"crossref","first-page":"2011","DOI":"10.3390\/cells9092011","article-title":"Compete or Coexist? 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