{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T13:17:37Z","timestamp":1790947057235,"version":"4.1.0"},"reference-count":94,"publisher":"Rockefeller University Press","issue":"4","license":[{"start":{"date-parts":[[2026,1,27]],"date-time":"2026-01-27T00:00:00Z","timestamp":1769472000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["rupress.org"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2026,4,6]]},"abstract":"<jats:p>BAR domain\u2013containing proteins are key regulators of endocytosis and actin remodeling. Their function in morphogenesis remains to be investigated. We report that the I-BAR domain\u2013containing protein, missing-in-metastasis (MIM) (also called MTSS1), promotes branched actin network formation and endocytosis to drive rapid, cyclical plasma membrane remodeling during syncytial divisions in Drosophila embryos. Actin-rich villous protrusions in the apical caps in interphase are depleted in metaphase, concurrent with furrow extension between adjacent nuclei. MIM depletion results in a loss of furrow extension and in longer, more abundant apical protrusions containing the formin diaphanous. Branched actin networks promoted by MIM are in balance with bundled actin networks induced by RhoGEF2 and diaphanous. Cyclical recruitment of MIM to the cortex promotes localization of active Rac, the WAVE regulatory complex, and the Arp2\/3 complex to drive endocytic membrane remodeling. These findings identify MIM as an integrator of actin and endocytic dynamics that enables rapid membrane remodeling during Drosophila syncytial division cycles.<\/jats:p>","DOI":"10.1083\/jcb.202502184","type":"journal-article","created":{"date-parts":[[2026,1,8]],"date-time":"2026-01-08T07:21:09Z","timestamp":1767856869000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":1,"title":["MIM triggers formin to Arp2\/3-based actin assembly in membrane remodeling in\n                    <i>Drosophila<\/i>\n                    embryos"],"prefix":"10.1083","volume":"225","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3520-3516","authenticated-orcid":true,"given":"Debasmita","family":"Mitra","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/028qa3n13","id-type":"ROR","asserted-by":"publisher"}],"name":"Indian Institute of Science Education and Research Pune 1 , Pune,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-4199-0527","authenticated-orcid":true,"given":"Georgina K.","family":"Goddard","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/027m9bs27","id-type":"ROR","asserted-by":"publisher"}],"name":"The University of Manchester 2 , Manchester,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-1746-2772","authenticated-orcid":true,"given":"Sanjana","family":"S","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/028qa3n13","id-type":"ROR","asserted-by":"publisher"}],"name":"Indian Institute of Science Education and Research Pune 1 , Pune,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-2269-2396","authenticated-orcid":true,"given":"Aparna","family":"K","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/028qa3n13","id-type":"ROR","asserted-by":"publisher"}],"name":"Indian Institute of Science Education and Research Pune 1 , Pune,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8869-4654","authenticated-orcid":true,"given":"Tom H.","family":"Millard","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/027m9bs27","id-type":"ROR","asserted-by":"publisher"}],"name":"The University of Manchester 2 , Manchester,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4262-0238","authenticated-orcid":true,"given":"Richa","family":"Rikhy","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/028qa3n13","id-type":"ROR","asserted-by":"publisher"}],"name":"Indian Institute of Science Education and Research Pune 1 , Pune,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"291","published-online":{"date-parts":[[2026,1,27]]},"reference":[{"key":"2026012709571377900_bib1","doi-asserted-by":"publisher","first-page":"144","DOI":"10.1016\/j.cub.2013.11.048","article-title":"Coordination of Rho family GTPase activities to orchestrate cytoskeleton responses during cell wound repair","volume":"24","author":"Abreu-Blanco","year":"2014","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib2","doi-asserted-by":"publisher","first-page":"1661","DOI":"10.1101\/gr.3726705","article-title":"Drosophila melanogaster: A case study of a model genomic sequence and its consequences","volume":"15","author":"Ashburner","year":"2005","journal-title":"Genome Res."},{"key":"2026012709571377900_bib3","doi-asserted-by":"publisher","first-page":"575","DOI":"10.1083\/jcb.200407124","article-title":"DRhoGEF2 regulates actin organization and contractility in the Drosophila blastoderm embryo","volume":"168","author":"Barmchi","year":"2005","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib4","doi-asserted-by":"publisher","first-page":"5393","DOI":"10.1242\/jcs.02640","article-title":"Involvement of Rac in actin cytoskeleton rearrangements induced by MIM-B","volume":"118","author":"Bompard","year":"2005","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib5","doi-asserted-by":"publisher","first-page":"1628","DOI":"10.1016\/j.cub.2014.05.069","article-title":"Cellular control of cortical actin nucleation","volume":"24","author":"Bovellan","year":"2014","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib6","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1016\/s1534-5807(02)00117-x","article-title":"Drosophila\u2019s insulin\/PI3-kinase pathway coordinates cellular metabolism with nutritional conditions","volume":"2","author":"Britton","year":"2002","journal-title":"Dev. Cell."},{"key":"2026012709571377900_bib7","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.202104046","article-title":"WASP integrates substrate topology and cell polarity to guide neutrophil migration","volume":"221","author":"Brunetti","year":"2022","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib8","doi-asserted-by":"publisher","first-page":"579","DOI":"10.1016\/j.cub.2014.01.072","article-title":"Homeostatic actin cytoskeleton networks are regulated by assembly factor competition for monomers","volume":"24","author":"Burke","year":"2014","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib9","doi-asserted-by":"publisher","first-page":"770","DOI":"10.1016\/j.cub.2010.02.060","article-title":"Cortical actin dynamics facilitate early-stage centrosome separation","volume":"20","author":"Cao","year":"2010","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib10","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1042\/BJ20120329","article-title":"Dimerization is necessary for MIM-mediated membrane deformation and endocytosis","volume":"446","author":"Cao","year":"2012","journal-title":"Biochem. J."},{"key":"2026012709571377900_bib11","doi-asserted-by":"publisher","first-page":"jcs186254","DOI":"10.1242\/jcs.186254","article-title":"The actin cortex at a glance","volume":"131","author":"Chugh","year":"2018","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib12","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1083\/jcb.201305021","article-title":"Cell biology in neuroscience: Mechanisms of cell migration in the nervous system","volume":"202","author":"Cooper","year":"2013","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib13","doi-asserted-by":"publisher","first-page":"2037","DOI":"10.1016\/j.cub.2012.08.046","article-title":"RhoGEF and positioning of rappaport-like furrows in the early Drosophila embryo","volume":"22","author":"Crest","year":"2012","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib14","doi-asserted-by":"publisher","first-page":"1781","DOI":"10.1038\/onc.2011.376","article-title":"Mtss1 regulates epidermal growth factor signaling in head and neck squamous carcinoma cells","volume":"31","author":"Dawson","year":"2012","journal-title":"Oncogene"},{"key":"2026012709571377900_bib15","doi-asserted-by":"publisher","first-page":"iyad184","DOI":"10.1093\/genetics\/iyad184","article-title":"Adhesion and polarity protein distribution-regulates hexagon dominated plasma membrane organization in Drosophila blastoderm embryos","volume":"225","author":"Dey","year":"2023","journal-title":"Genetics"},{"key":"2026012709571377900_bib16","doi-asserted-by":"publisher","first-page":"jcs240168","DOI":"10.1242\/jcs.240168","article-title":"DE-cadherin and Myosin II balance regulates furrow length for onset of polygon shape in syncytial embryos","volume":"133","author":"Dey","year":"2020","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib17","doi-asserted-by":"publisher","first-page":"790","DOI":"10.1038\/nature00859","article-title":"Mechanism of regulation of WAVE1-induced actin nucleation by Rac1 and Nck","volume":"418","author":"Eden","year":"2002","journal-title":"Nature"},{"key":"2026012709571377900_bib18","doi-asserted-by":"publisher","first-page":"1540","DOI":"10.1016\/j.yexcr.2008.01.016","article-title":"RhoA\/ROCK-mediated switching between Cdc42- and Rac1-dependent protrusion in MTLn3 carcinoma cells","volume":"314","author":"El-Sibai","year":"2008","journal-title":"Exp. Cell Res."},{"key":"2026012709571377900_bib19","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/ncomms3244","article-title":"Tubular endocytosis drives remodelling of the apical surface during epithelial morphogenesis in Drosophila","volume":"4","author":"Fabrowski","year":"2013","journal-title":"Nat. Commun."},{"key":"2026012709571377900_bib20","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1242\/jcs.61.1.31","article-title":"Studies of nuclear and cytoplasmic behaviour during the five mitotic cycles that precede gastrulation in Drosophila embryogenesis","volume":"61","author":"Foe","year":"1983","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib21","doi-asserted-by":"publisher","first-page":"1767","DOI":"10.1242\/dev.127.9.1767","article-title":"Microtubules and mitotic cycle phase modulate spatiotemporal distributions of F-actin and myosin II in Drosophila syncytial blastoderm embryos","volume":"127","author":"Foe","year":"2000","journal-title":"Development"},{"key":"2026012709571377900_bib22","doi-asserted-by":"publisher","first-page":"1429","DOI":"10.1016\/j.cub.2009.07.058","article-title":"Drosophila Cip4\/Toca-1 integrates membrane trafficking and actin dynamics through WASP and SCAR\/WAVE","volume":"19","author":"Fricke","year":"2009","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib23","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1016\/j.cell.2009.04.010","article-title":"The BAR domain superfamily: Membrane-molding macromolecules","volume":"137","author":"Frost","year":"2009","journal-title":"Cell"},{"key":"2026012709571377900_bib24","doi-asserted-by":"publisher","first-page":"10900","DOI":"10.1038\/s41467-025-66351-1","article-title":"Efficient cellular transformation via protein delivery through the protrusion-derived extracellular vesicles","volume":"16","author":"Fujioka","year":"2025","journal-title":"Nat. Commun."},{"key":"2026012709571377900_bib25","doi-asserted-by":"publisher","first-page":"1009","DOI":"10.1242\/dev.01669","article-title":"RhoGEF2 and the formin Dia control the formation of the furrow canal by directed actin assembly during Drosophila cellularisation","volume":"132","author":"Grosshans","year":"2005","journal-title":"Development"},{"key":"2026012709571377900_bib26","doi-asserted-by":"publisher","first-page":"1955","DOI":"10.1016\/j.cellsig.2013.04.009","article-title":"Signaling networks of Rho GTPases in cell motility","volume":"25","author":"Hanna","year":"2013","journal-title":"Cell. Signal."},{"key":"2026012709571377900_bib27","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1016\/j.ydbio.2022.08.004","article-title":"Sponge\/DOCK-dependent regulation of F-actin networks directing cortical cap behaviors and syncytial furrow ingression","volume":"491","author":"Henry","year":"2022","journal-title":"Dev. Biol."},{"key":"2026012709571377900_bib28","doi-asserted-by":"publisher","first-page":"2316","DOI":"10.1242\/dev.120998","article-title":"A rapid, membrane-dependent pathway directs furrow formation through RalA in the early Drosophila embryo","volume":"142","author":"Holly","year":"2015","journal-title":"Development"},{"key":"2026012709571377900_bib29","doi-asserted-by":"publisher","first-page":"5138","DOI":"10.1091\/mbc.e09-02-0144","article-title":"Exploring the roles of diaphanous and enabled activity in shaping the balance between filopodia and lamellipodia","volume":"20","author":"Homem","year":"2009","journal-title":"Mol. Biol. Cell"},{"key":"2026012709571377900_bib30","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1016\/s0092-8674(02)01044-9","article-title":"Endocytosis: Driving membranes around the Bend","volume":"111","author":"Hurley","year":"2002","journal-title":"Cell"},{"key":"2026012709571377900_bib31","doi-asserted-by":"publisher","first-page":"4195","DOI":"10.1083\/jcb.201903152","article-title":"Par-1 controls the composition and growth of cortical actin caps during embryo cleavage","volume":"218","author":"Jiang","year":"2019","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib32","doi-asserted-by":"publisher","first-page":"3578","DOI":"10.1038\/s41467-022-31207-5","article-title":"Branched actin networks are organized for asymmetric force production during clathrin-mediated endocytosis in mammalian cells","volume":"13","author":"Jin","year":"2022","journal-title":"Nat. Commun."},{"key":"2026012709571377900_bib33","doi-asserted-by":"publisher","first-page":"893","DOI":"10.1021\/acssynbio.9b00516","article-title":"Light-inducible generation of membrane curvature in live cells with engineered BAR domain proteins","volume":"9","author":"Jones","year":"2020","journal-title":"ACS Synth. Biol."},{"key":"2026012709571377900_bib34","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/j.celrep.2018.06.013","article-title":"MTSS1 regulation of actin-nucleating formin DAAM1 in dendritic filopodia determines final dendritic configuration of Purkinje cells","volume":"24","author":"Kawabata Galbraith","year":"2018","journal-title":"Cell Rep."},{"key":"2026012709571377900_bib35","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1038\/nrm3861","article-title":"Steering cell migration: Lamellipodium dynamics and the regulation of directional persistence","volume":"15","author":"Krause","year":"2014","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012709571377900_bib36","doi-asserted-by":"publisher","first-page":"1639","DOI":"10.1016\/j.cub.2008.09.063","article-title":"Drosophila Cip4 and WASp define a branch of the Cdc42-Par6-aPKC pathway regulating E-cadherin endocytosis","volume":"18","author":"Leibfried","year":"2008","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib37","doi-asserted-by":"publisher","first-page":"1475","DOI":"10.1242\/jcs.198937","article-title":"Missing-in-metastasis protein downregulates CXCR4 by promoting ubiquitylation and interaction with small Rab GTPases","volume":"130","author":"Li","year":"2017","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib38","doi-asserted-by":"publisher","first-page":"6494","DOI":"10.1074\/jbc.RA118.006071","article-title":"Differential interactions of missing in metastasis and insulin receptor tyrosine kinase substrate with RAB proteins in the endocytosis of CXCR4","volume":"294","author":"Li","year":"2019","journal-title":"J. Biol. Chem."},{"key":"2026012709571377900_bib39","doi-asserted-by":"publisher","first-page":"2059","DOI":"10.1038\/sj.onc.1208412","article-title":"Differential regulation of cortactin and N-WASP-mediated actin polymerization by missing in metastasis (MIM) protein","volume":"24","author":"Lin","year":"2005","journal-title":"Oncogene"},{"key":"2026012709571377900_bib40","doi-asserted-by":"publisher","first-page":"1787","DOI":"10.1101\/gad.8.15.1787","article-title":"Distinct morphogenetic functions of similar small GTPases: Drosophila Drac1 is involved in axonal outgrowth and myoblast fusion","volume":"8","author":"Luo","year":"1994","journal-title":"Genes Dev."},{"key":"2026012709571377900_bib41","doi-asserted-by":"publisher","first-page":"1350","DOI":"10.1080\/15384101.2017.1325045","article-title":"Daam1 regulates fascin for actin assembly in mouse oocyte meiosis","volume":"16","author":"Lu","year":"2017","journal-title":"Cell Cycle"},{"key":"2026012709571377900_bib42","doi-asserted-by":"publisher","first-page":"569","DOI":"10.1007\/s00109-007-0207-0","article-title":"MIM: A multifunctional scaffold protein","volume":"85","author":"Machesky","year":"2007","journal-title":"J. Mol. Med."},{"key":"2026012709571377900_bib43","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1083\/jcb.201603077","article-title":"RhoA GTPase inhibition organizes contraction during epithelial morphogenesis","volume":"214","author":"Mason","year":"2016","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib44","doi-asserted-by":"publisher","first-page":"953","DOI":"10.1083\/jcb.200609176","article-title":"Missing-in-metastasis and IRSp53 deform PI(4,5)P2-rich membranes by an inverse BAR domain-like mechanism","volume":"176","author":"Mattila","year":"2007","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib45","doi-asserted-by":"publisher","first-page":"8452","DOI":"10.1074\/jbc.M212113200","article-title":"Mouse MIM, a tissue-specific regulator of cytoskeletal dynamics, interacts with ATP-actin monomers through its C-terminal WH2 domain","volume":"278","author":"Mattila","year":"2003","journal-title":"J. Biol. Chem."},{"key":"2026012709571377900_bib46","doi-asserted-by":"publisher","first-page":"892","DOI":"10.1242\/dev.128876","article-title":"Rab8 directs furrow ingression and membrane addition during epithelial formation in Drosophila melanogaster","volume":"143","author":"Mavor","year":"2016","journal-title":"Development"},{"key":"2026012709571377900_bib47","doi-asserted-by":"publisher","first-page":"dev201547","DOI":"10.1242\/dev.201547","article-title":"A Rab39-Klp98A-Rab35 endocytic recycling pathway is essential for rapid Golgi-dependent furrow ingression","volume":"150","author":"Miao","year":"2023","journal-title":"Development"},{"key":"2026012709571377900_bib48","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1101\/sqb.1985.050.01.012","article-title":"Studies on the cytoplasmic organization of early Drosophila embryos","volume":"50","author":"Miller","year":"1985","journal-title":"Cold Spring Harb. Symp. Quant. Biol."},{"key":"2026012709571377900_bib49","doi-asserted-by":"publisher","first-page":"101736","DOI":"10.1016\/j.xpro.2022.101736","article-title":"Imaging and quantification of apical microvilli in the syncytial blastoderm of Drosophila embryos","volume":"3","author":"Mitra","year":"2022","journal-title":"STAR Protoc."},{"key":"2026012709571377900_bib50","doi-asserted-by":"publisher","first-page":"4763","DOI":"10.1242\/dev.098509","article-title":"Par3 controls neural crest migration by promoting microtubule catastrophe during contact inhibition of locomotion","volume":"140","author":"Moore","year":"2013","journal-title":"Development"},{"key":"2026012709571377900_bib51","doi-asserted-by":"publisher","first-page":"884","DOI":"10.1016\/j.cell.2018.06.032","article-title":"Systematic nanoscale analysis of endocytosis links efficient vesicle formation to patterned actin nucleation","volume":"174","author":"Mund","year":"2018","journal-title":"Cell"},{"key":"2026012709571377900_bib52","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1038\/nature14603","article-title":"A self-organized biomechanical network drives shape changes during tissue morphogenesis","volume":"524","author":"Munjal","year":"2015","journal-title":"Nature"},{"key":"2026012709571377900_bib53","doi-asserted-by":"publisher","first-page":"842","DOI":"10.1016\/j.devcel.2021.02.029","article-title":"Filopodium-derived vesicles produced by MIM enhance the migration of recipient cells","volume":"56","author":"Nishimura","year":"2021","journal-title":"Dev. Cell"},{"key":"2026012709571377900_bib54","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1186\/1478-811X-8-23","article-title":"Rac and Rho GTPases in cancer cell motility control","volume":"8","author":"Parri","year":"2010","journal-title":"Cell Commun. Signal."},{"key":"2026012709571377900_bib55","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1016\/j.ydbio.2013.03.012","article-title":"WAVE\/SCAR promotes endocytosis and early endosome morphology in polarized C. elegans epithelia","volume":"377","author":"Patel","year":"2013","journal-title":"Dev. Biol."},{"key":"2026012709571377900_bib56","doi-asserted-by":"publisher","first-page":"1848","DOI":"10.1016\/j.cub.2003.10.023","article-title":"Trafficking through Rab11 endosomes is required for cellularization during Drosophila embryogenesis","volume":"13","author":"Pelissier","year":"2003","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib57","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.202003086","article-title":"The WAVE complex associates with sites of saddle membrane curvature","volume":"220","author":"Pipathsouk","year":"2021","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib58","doi-asserted-by":"publisher","first-page":"F111","DOI":"10.1083\/jcb.150.5.f111","article-title":"Molecular links between endocytosis and the actin cytoskeleton","volume":"150","author":"Qualmann","year":"2000","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib59","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1083\/jcb.200910136","article-title":"I-BAR protein antagonism of endocytosis mediates directional sensing during guided cell migration","volume":"189","author":"Quinones","year":"2010","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib60","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1083\/jcb.200305115","article-title":"Actin cytoskeleton remodeling during early Drosophila furrow formation requires recycling endosomal components nuclear-fallout and Rab11","volume":"163","author":"Riggs","year":"2003","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib61","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1242\/bio.20149936","article-title":"Dynamin regulates metaphase furrow formation and plasma membrane compartmentalization in the syncytial Drosophila embryo","volume":"4","author":"Rikhy","year":"2015","journal-title":"Biol. Open"},{"key":"2026012709571377900_bib62","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.48093","article-title":"B cells extract antigens at Arp2\/3-generated actin foci interspersed with linear filaments","volume":"8","author":"Roper","year":"2019","journal-title":"Elife"},{"key":"2026012709571377900_bib63","doi-asserted-by":"publisher","first-page":"604","DOI":"10.1016\/j.devcel.2015.01.012","article-title":"Ect2\/Pbl acts via Rho and polarity proteins to direct the assembly of an isotropic actomyosin cortex upon mitotic entry","volume":"32","author":"Rosa","year":"2015","journal-title":"Dev. Cell"},{"key":"2026012709571377900_bib64","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1080\/19490992.2015.1090670","article-title":"Competition and collaboration between different actin assembly pathways allows for homeostatic control of the actin cytoskeleton","volume":"5","author":"Rotty","year":"2014","journal-title":"Bioarchitecture"},{"key":"2026012709571377900_bib65","doi-asserted-by":"publisher","first-page":"644","DOI":"10.1016\/j.devcel.2015.04.014","article-title":"MIM-induced membrane bending promotes dendritic spine initiation","volume":"33","author":"Saarikangas","year":"2015","journal-title":"Dev. Cell"},{"key":"2026012709571377900_bib66","doi-asserted-by":"publisher","first-page":"1245","DOI":"10.1242\/jcs.082610","article-title":"Missing-in-metastasis MIM\/MTSS1 promotes actin assembly at intercellular junctions and is required for integrity of kidney epithelia","volume":"124","author":"Saarikangas","year":"2011","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib67","doi-asserted-by":"publisher","first-page":"1835","DOI":"10.1038\/s41467-018-03955-w","article-title":"Small GTPases and BAR domain proteins regulate branched actin polymerisation for clathrin and dynamin-independent endocytosis","volume":"9","author":"Sathe","year":"2018","journal-title":"Nat. Commun."},{"key":"2026012709571377900_bib68","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.63535","article-title":"Spatiotemporal recruitment of RhoGTPase protein GRAF inhibits actomyosin ring constriction in Drosophila cellularization","volume":"10","author":"Sharma","year":"2021","journal-title":"Elife"},{"key":"2026012709571377900_bib69","doi-asserted-by":"publisher","first-page":"jcs235846","DOI":"10.1242\/jcs.235846","article-title":"F-BAR domain protein Syndapin regulates actomyosin dynamics during apical cap remodeling in syncytial Drosophila embryos","volume":"133","author":"Sherlekar","year":"2020","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib70","doi-asserted-by":"publisher","first-page":"2064","DOI":"10.1091\/mbc.E15-09-0656","article-title":"Syndapin promotes pseudocleavage furrow formation by actin organization in the syncytial Drosophila embryo","volume":"27","author":"Sherlekar","year":"2016","journal-title":"Mol. Biol. Cell"},{"key":"2026012709571377900_bib71","doi-asserted-by":"publisher","first-page":"1917","DOI":"10.1091\/mbc.E12-01-0006","article-title":"Arp2\/3 mediates early endosome dynamics necessary for the maintenance of PAR asymmetry in Caenorhabditis elegans","volume":"23","author":"Shivas","year":"2012","journal-title":"Mol. Biol. Cell"},{"key":"2026012709571377900_bib94","doi-asserted-by":"publisher","first-page":"1047","DOI":"10.1016\/j.cell.2009.04.013","article-title":"Cyfip1 is a putative invasion suppressor in epithelial cancers","volume":"137","author":"Silva","year":"2009","journal-title":"Cell"},{"key":"2026012709571377900_bib72","doi-asserted-by":"publisher","first-page":"705","DOI":"10.1016\/s0960-9822(02)00807-2","article-title":"Arp2\/3-dependent pseudocleavage [correction of psuedocleavage] furrow assembly in syncytial Drosophila embryos","volume":"12","author":"Stevenson","year":"2002","journal-title":"Curr. Biol."},{"key":"2026012709571377900_bib73","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1083\/jcb.200912134","article-title":"Clasp-mediated microtubule bundling regulates persistent motility and contact repulsion in Drosophila macrophages in vivo","volume":"189","author":"Stramer","year":"2010","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib74","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41467-018-06738-5","article-title":"Mechanochemical feedback control of dynamin independent endocytosis modulates membrane tension in adherent cells","volume":"9","author":"Thottacherry","year":"2018","journal-title":"Nat. Commun."},{"key":"2026012709571377900_bib75","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.celrep.2019.06.012","article-title":"Activation of Arp2\/3 by WASp is essential for the endocytosis of Delta only during cytokinesis in Drosophila","volume":"28","author":"Trylinski","year":"2019","journal-title":"Cell Rep."},{"key":"2026012709571377900_bib76","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/0012-1606(76)90070-1","article-title":"Scanning electron microscopy of Drosophila embryogenesis. 1. The structure of the egg envelopes and the formation of the cellular blastoderm","volume":"50","author":"Turner","year":"1976","journal-title":"Dev. Biol."},{"key":"2026012709571377900_bib77","doi-asserted-by":"publisher","first-page":"2093","DOI":"10.1016\/j.febslet.2008.04.039","article-title":"Rho GTPases in cancer cell biology","volume":"582","author":"Vega","year":"2008","journal-title":"FEBS Lett."},{"key":"2026012709571377900_bib78","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1534\/genetics.110.120998","article-title":"Genetic evidence for antagonism between Pak protein kinase and Rho1 small GTPase signaling in regulation of the actin cytoskeleton during Drosophila oogenesis","volume":"187","author":"Vlachos","year":"2011","journal-title":"Genetics"},{"key":"2026012709571377900_bib79","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1038\/s41421-022-00507-x","article-title":"MTSS1 curtails lung adenocarcinoma immune evasion by promoting AIP4-mediated PD-L1 monoubiquitination and lysosomal degradation","volume":"9","author":"Wang","year":"2023","journal-title":"Cell Discov."},{"key":"2026012709571377900_bib80","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1242\/jcs.1986.supplement_5.20","article-title":"The cytoskeleton of the early Drosophila embryo","volume":"5","author":"Warn","year":"1986","journal-title":"J. Cell Sci. Suppl."},{"key":"2026012709571377900_bib81","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1042\/BJ20021962","article-title":"MIM-B, a putative metastasis suppressor protein, binds to actin and to protein tyrosine phosphatase delta","volume":"371","author":"Woodings","year":"2003","journal-title":"Biochem. J."},{"key":"2026012709571377900_bib82","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1083\/jcb.200103048","article-title":"RhoA is required for monocyte tail retraction during transendothelial migration","volume":"154","author":"Worthylake","year":"2001","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib83","doi-asserted-by":"publisher","DOI":"10.1101\/2024.07.08.600855","article-title":"The WAVE complex forms linear arrays at negative membrane curvature to instruct lamellipodia formation","author":"Wu","year":"2024","journal-title":"bioRxiv"},{"key":"2026012709571377900_bib84","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1007174","article-title":"Differentially-dimensioned furrow formation by zygotic gene expression and the MBT","volume":"14","author":"Xie","year":"2018","journal-title":"PLoS Genet."},{"key":"2026012709571377900_bib85","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.63046","article-title":"Combinatorial deployment of F-actin regulators to build complex 3D actin structures in vivo","volume":"10","author":"Xie","year":"2021","journal-title":"Elife"},{"key":"2026012709571377900_bib86","doi-asserted-by":"publisher","first-page":"14929","DOI":"10.1074\/jbc.M309408200","article-title":"A novel actin bundling\/filopodium-forming domain conserved in insulin receptor tyrosine kinase substrate p53 and missing in metastasis protein","volume":"279","author":"Yamagishi","year":"2004","journal-title":"J. Biol. Chem."},{"key":"2026012709571377900_bib87","doi-asserted-by":"publisher","first-page":"1796","DOI":"10.1242\/jcs.118422","article-title":"The F-BAR protein Cip4\/Toca-1 antagonizes the formin Diaphanous in membrane stabilization and compartmentalization","volume":"126","author":"Yan","year":"2013","journal-title":"J. Cell Sci."},{"key":"2026012709571377900_bib88","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0020845","article-title":"Murine missing in metastasis (MIM) mediates cell polarity and regulates the motility response to growth factors","volume":"6","author":"Yu","year":"2011","journal-title":"PLoS One"},{"key":"2026012709571377900_bib89","doi-asserted-by":"publisher","first-page":"689","DOI":"10.1083\/jcb.200109057","article-title":"SCAR is a primary regulator of Arp2\/3-dependent morphological events in Drosophila","volume":"156","author":"Zallen","year":"2002","journal-title":"J. Cell Biol."},{"key":"2026012709571377900_bib90","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1016\/j.devcel.2018.04.024","article-title":"Collision of expanding actin caps with actomyosin borders for cortical bending and mitotic rounding in a syncytium","volume":"45","author":"Zhang","year":"2018","journal-title":"Dev. Cell"},{"key":"2026012709571377900_bib91","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/j.ceb.2010.10.005","article-title":"I-BAR domain proteins: Linking actin and plasma membrane dynamics","volume":"23","author":"Zhao","year":"2011","journal-title":"Curr. Opin. Cell Biol."},{"key":"2026012709571377900_bib92","doi-asserted-by":"publisher","first-page":"96635","DOI":"10.1039\/c6ra21530k","article-title":"Downregulation of MIM protein inhibits the cellular endocytosis process of magnetic nanoparticles in macrophages","volume":"6","author":"Zhao","year":"2016","journal-title":"RSC Adv."},{"key":"2026012709571377900_bib93","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1016\/j.bbagen.2018.12.002","article-title":"Missing-in-metastasis protein promotes internalization of magnetic nanoparticles via association with clathrin light chain and Rab7","volume":"1863","author":"Zhao","year":"2019","journal-title":"Biochim. Biophys. Acta Gen. Subj."}],"container-title":["Journal of Cell Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/rupress.org\/jcb\/article-pdf\/doi\/10.1083\/jcb.202502184\/2025533\/jcb_202502184.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/rupress.org\/jcb\/article-pdf\/doi\/10.1083\/jcb.202502184\/2025533\/jcb_202502184.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,27]],"date-time":"2026-01-27T14:57:31Z","timestamp":1769525851000},"score":1,"resource":{"primary":{"URL":"https:\/\/rupress.org\/jcb\/article\/225\/4\/e202502184\/281439\/MIM-triggers-formin-to-Arp2-3-based-actin-assembly"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,27]]},"references-count":94,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2026,4,6]]}},"URL":"https:\/\/doi.org\/10.1083\/jcb.202502184","relation":{},"ISSN":["0021-9525","1540-8140"],"issn-type":[{"value":"0021-9525","type":"print"},{"value":"1540-8140","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,27]]},"article-number":"e202502184"}}