{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T21:43:53Z","timestamp":1792100633256,"version":"4.4.0"},"reference-count":77,"publisher":"Elsevier BV","issue":"9","license":[{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T00:00:00Z","timestamp":1787184000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000054","name":"National Cancer Institute","doi-asserted-by":"publisher","award":["R01CA279129"],"award-info":[{"award-number":["R01CA279129"]}],"id":[{"id":"10.13039\/100000054","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000054","name":"National Cancer Institute","doi-asserted-by":"publisher","award":["P30CA014236"],"award-info":[{"award-number":["P30CA014236"]}],"id":[{"id":"10.13039\/100000054","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Cell Reports"],"published-print":{"date-parts":[[2026,9]]},"DOI":"10.1016\/j.celrep.2026.117962","type":"journal-article","created":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T20:15:33Z","timestamp":1789071333000},"page":"117962","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["SERINC2-mediated phosphatidylserine remodeling activates protrusion-localized AXL signaling to promote brain metastasis"],"prefix":"10.1016","volume":"45","author":[{"given":"Zhibo","family":"Yan","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maja","family":"Beus","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhannan","family":"Han","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lihong","family":"Pan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yihui","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Junqi","family":"Lu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Loren B.","family":"Weidenhammer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liang","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Spencer","family":"Rosario","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingyun","family":"Lee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cristina M.","family":"Furdui","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hui-Wen","family":"Lo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eugene S.","family":"Kandel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Erik J.","family":"Soderblom","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weili","family":"Ma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qing","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sergei V.","family":"Kotenko","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Raymond B.","family":"Birge","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7868-0518","authenticated-orcid":false,"given":"Mikhail A.","family":"Nikiforov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.celrep.2026.117962_bib1","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1016\/j.tcb.2017.03.003","article-title":"Tumor Cell Invadopodia: Invasive Protrusions that Orchestrate Metastasis","volume":"27","author":"Eddy","year":"2017","journal-title":"Trends Cell Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib2","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1007\/s10585-006-9014-1","article-title":"Invadopodia: specialized cell structures for cancer invasion","volume":"23","author":"Weaver","year":"2006","journal-title":"Clin. Exp. Metastasis"},{"key":"10.1016\/j.celrep.2026.117962_bib3","doi-asserted-by":"crossref","DOI":"10.1111\/boc.70018","article-title":"Protrusion-Derived Extracellular Vesicles (PD-EVs) and Their Diverse Origins: Key Players in Cellular Communication, Cancer Progression, and T Cell Modulation","volume":"117","author":"Duro","year":"2025","journal-title":"Biol. Cell"},{"key":"10.1016\/j.celrep.2026.117962_bib4","doi-asserted-by":"crossref","first-page":"1689","DOI":"10.1016\/j.cell.2023.02.002","article-title":"Cancer neuroscience: State of the field, emerging directions","volume":"186","author":"Winkler","year":"2023","journal-title":"Cell"},{"key":"10.1016\/j.celrep.2026.117962_bib5","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1126\/science.1152066","article-title":"Membrane phosphatidylserine regulates surface charge and protein localization","volume":"319","author":"Yeung","year":"2008","journal-title":"Science"},{"key":"10.1016\/j.celrep.2026.117962_bib6","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1016\/j.ejcb.2016.06.006","article-title":"A new front in cell invasion: The invadopodial membrane","volume":"95","author":"Hastie","year":"2016","journal-title":"Eur. J. Cell Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib7","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.semcdb.2019.11.017","article-title":"Filopodia and their links with membrane traffic and cell adhesion","volume":"102","author":"Gallop","year":"2020","journal-title":"Semin. Cell Dev. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib8","doi-asserted-by":"crossref","DOI":"10.3390\/cancers14030466","article-title":"AXL Receptor Tyrosine Kinase as a Promising Therapeutic Target Directing Multiple Aspects of Cancer Progression and Metastasis","volume":"14","author":"Goyette","year":"2022","journal-title":"Cancers (Basel)"},{"key":"10.1016\/j.celrep.2026.117962_bib9","doi-asserted-by":"crossref","first-page":"5712","DOI":"10.1038\/ncomms6712","article-title":"Low MITF\/AXL ratio predicts early resistance to multiple targeted drugs in melanoma","volume":"5","author":"M\u00fcller","year":"2014","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2026.117962_bib10","doi-asserted-by":"crossref","first-page":"7304","DOI":"10.1038\/s41467-022-34899-x","article-title":"Decoding molecular programs in melanoma brain metastases","volume":"13","author":"Radke","year":"2022","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2026.117962_bib11","doi-asserted-by":"crossref","first-page":"1313","DOI":"10.1158\/1078-0432.CCR-15-1458","article-title":"Molecular Pathways: AXL, a Membrane Receptor Mediator of Resistance to Therapy","volume":"22","author":"Scaltriti","year":"2016","journal-title":"Clin. Cancer Res."},{"key":"10.1016\/j.celrep.2026.117962_bib12","doi-asserted-by":"crossref","first-page":"592","DOI":"10.1093\/neuros\/nyaa471","article-title":"Differences in Genomic Alterations Between Brain Metastases and Primary Tumors","volume":"88","author":"Dono","year":"2021","journal-title":"Neurosurgery"},{"key":"10.1016\/j.celrep.2026.117962_bib13","doi-asserted-by":"crossref","first-page":"1693","DOI":"10.1016\/j.ccell.2024.08.015","article-title":"Distinct tumor architectures and microenvironments for the initiation of breast cancer metastasis in the brain","volume":"42","author":"Gan","year":"2024","journal-title":"Cancer Cell"},{"key":"10.1016\/j.celrep.2026.117962_bib14","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1158\/0008-5472.CAN-23-1459","article-title":"AXL\/WRNIP1 Mediates Replication Stress Response and Promotes Therapy Resistance and Metachronous Metastasis in HER2+ Breast Cancer","volume":"84","author":"Marquez-Palencia","year":"2024","journal-title":"Cancer Res."},{"key":"10.1016\/j.celrep.2026.117962_bib15","doi-asserted-by":"crossref","first-page":"3421","DOI":"10.1016\/j.celrep.2019.11.018","article-title":"A TAZ-AXL-ABL2 Feed-Forward Signaling Axis Promotes Lung Adenocarcinoma Brain Metastasis","volume":"29","author":"Hoj","year":"2019","journal-title":"Cell Rep."},{"key":"10.1016\/j.celrep.2026.117962_bib16","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2024596118","article-title":"The GAS6-AXL signaling pathway triggers actin remodeling that drives membrane ruffling, macropinocytosis, and cancer-cell invasion","volume":"118","author":"Zd\u017calik-Bielecka","year":"2021","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.celrep.2026.117962_bib17","doi-asserted-by":"crossref","first-page":"2634","DOI":"10.1158\/0008-5472.CAN-18-2316","article-title":"Cross-Talk between Receptor Tyrosine Kinases AXL and ERBB3 Regulates Invadopodia Formation in Melanoma Cells","volume":"79","author":"Revach","year":"2019","journal-title":"Cancer Res."},{"key":"10.1016\/j.celrep.2026.117962_bib18","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1146\/annurev.biophys.093008.131234","article-title":"The distribution and function of phosphatidylserine in cellular membranes","volume":"39","author":"Leventis","year":"2010","journal-title":"Annu. Rev. Biophys."},{"key":"10.1016\/j.celrep.2026.117962_bib19","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1186\/s12964-019-0438-z","article-title":"Distribution, dynamics and functional roles of phosphatidylserine within the cell","volume":"17","author":"Kay","year":"2019","journal-title":"Cell Commun. Signal."},{"key":"10.1016\/j.celrep.2026.117962_bib20","doi-asserted-by":"crossref","DOI":"10.3390\/biom10060928","article-title":"ORP5 and ORP8: Sterol Sensors and Phospholipid Transfer Proteins at Membrane Contact Sites?","volume":"10","author":"Santos","year":"2020","journal-title":"Biomolecules"},{"key":"10.1016\/j.celrep.2026.117962_bib21","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.coi.2019.11.009","article-title":"Flippase and scramblase for phosphatidylserine exposure","volume":"62","author":"Nagata","year":"2020","journal-title":"Curr. Opin. Immunol."},{"key":"10.1016\/j.celrep.2026.117962_bib22","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1186\/s12964-025-02090-6","article-title":"Dys-regulated phosphatidylserine externalization as a cell intrinsic immune escape mechanism in cancer","volume":"23","author":"Pulica","year":"2025","journal-title":"Cell Commun. Signal."},{"key":"10.1016\/j.celrep.2026.117962_bib23","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.tcb.2006.02.003","article-title":"Phosphatidylserine recognition by phagocytes: a view to a kill","volume":"16","author":"Wu","year":"2006","journal-title":"Trends Cell Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib24","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1007\/978-3-031-92785-0_2","article-title":"Efferocytosis in Homeostasis","volume":"1481","author":"Mauer\u00f6der","year":"2025","journal-title":"Adv. Exp. Med. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib25","article-title":"Loss of lipid asymmetry facilitates plasma membrane blebbing by decreasing membrane lipid packing","volume":"122","author":"Wang","year":"2025","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.celrep.2026.117962_bib26","doi-asserted-by":"crossref","DOI":"10.1101\/cshperspect.a009076","article-title":"Biology of the TAM receptors","volume":"5","author":"Lemke","year":"2013","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib27","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1093\/neuonc\/noae219","article-title":"The immune landscape in brain metastasis","volume":"27","author":"Schreurs","year":"2025","journal-title":"Neuro Oncol."},{"key":"10.1016\/j.celrep.2026.117962_bib28","doi-asserted-by":"crossref","first-page":"30022","DOI":"10.1074\/jbc.271.47.30022","article-title":"Identification of the product of growth arrest-specific gene 6 as a common ligand for Axl, Sky, and Mer receptor tyrosine kinases","volume":"271","author":"Nagata","year":"1996","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.celrep.2026.117962_bib29","article-title":"The Emerging Role of the Serine Incorporator Protein Family in Regulating Viral Infection","volume":"10","author":"Xu","year":"2022","journal-title":"Front. Cell Dev. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib30","doi-asserted-by":"crossref","first-page":"35776","DOI":"10.1074\/jbc.M505712200","article-title":"Serinc, an activity-regulated protein family, incorporates serine into membrane lipid synthesis","volume":"280","author":"Inuzuka","year":"2005","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.celrep.2026.117962_bib31","article-title":"Microglia contribute to bipolar depression through Serinc2-dependent phospholipid synthesis","volume":"122","author":"Wang","year":"2025","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.celrep.2026.117962_bib32","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/j.molimm.2016.12.007","article-title":"Disruption of Serinc1, which facilitates serine-derived lipid synthesis, fails to alter macrophage function, lymphocyte proliferation or autoimmune disease susceptibility","volume":"82","author":"Chu","year":"2017","journal-title":"Mol. Immunol."},{"key":"10.1016\/j.celrep.2026.117962_bib33","doi-asserted-by":"crossref","first-page":"13702","DOI":"10.1074\/jbc.M117.797332","article-title":"The host-cell restriction factor SERINC5 restricts HIV-1 infectivity without altering the lipid composition and organization of viral particles","volume":"292","author":"Trautz","year":"2017","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.celrep.2026.117962_bib34","doi-asserted-by":"crossref","first-page":"4368","DOI":"10.1038\/s41467-023-39262-2","article-title":"Antiviral HIV-1 SERINC restriction factors disrupt virus membrane asymmetry","volume":"14","author":"Leonhardt","year":"2023","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2026.117962_bib35","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1038\/s41594-019-0357-0","article-title":"A bipartite structural organization defines the SERINC family of HIV-1 restriction factors","volume":"27","author":"Pye","year":"2020","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib36","doi-asserted-by":"crossref","DOI":"10.3390\/biom14050570","article-title":"Disruption of Transmembrane Phosphatidylserine Asymmetry by HIV-1 Incorporated SERINC5 Is Not Responsible for Virus Restriction","volume":"14","author":"Raghunath","year":"2024","journal-title":"Biomolecules"},{"key":"10.1016\/j.celrep.2026.117962_bib37","doi-asserted-by":"crossref","first-page":"970","DOI":"10.1016\/j.chembiol.2022.01.007","article-title":"Phosphorylation of guanosine monophosphate reductase triggers a GTP-dependent switch from pro- to anti-oncogenic function of EPHA4","volume":"29","author":"Wolff","year":"2022","journal-title":"Cell Chem. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib38","doi-asserted-by":"crossref","first-page":"1720","DOI":"10.1158\/2159-8290.CD-19-0270","article-title":"Polyunsaturated Fatty Acids from Astrocytes Activate PPAR\u03b3 Signaling in Cancer Cells to Promote Brain Metastasis","volume":"9","author":"Zou","year":"2019","journal-title":"Cancer Discov."},{"key":"10.1016\/j.celrep.2026.117962_bib39","doi-asserted-by":"crossref","first-page":"1005","DOI":"10.1038\/nature08021","article-title":"Genes that mediate breast cancer metastasis to the brain","volume":"459","author":"Bos","year":"2009","journal-title":"Nature"},{"key":"10.1016\/j.celrep.2026.117962_bib40","doi-asserted-by":"crossref","first-page":"6091","DOI":"10.1038\/s41467-021-26324-6","article-title":"Regulation of local GTP availability controls RAC1 activity and cell invasion","volume":"12","author":"Bianchi-Smiraglia","year":"2021","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2026.117962_bib41","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1091\/mbc.12.2.265","article-title":"Integrin-mediated adhesion regulates cell polarity and membrane protrusion through the Rho family of GTPases","volume":"12","author":"Cox","year":"2001","journal-title":"Mol. Biol. Cell"},{"key":"10.1016\/j.celrep.2026.117962_bib42","doi-asserted-by":"crossref","first-page":"R362","DOI":"10.1016\/j.cub.2008.02.028","article-title":"Invadopodia","volume":"18","author":"Weaver","year":"2008","journal-title":"Curr. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib43","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1016\/j.celrep.2013.09.015","article-title":"A purine nucleotide biosynthesis enzyme guanosine monophosphate reductase is a suppressor of melanoma invasion","volume":"5","author":"Wawrzyniak","year":"2013","journal-title":"Cell Rep."},{"key":"10.1016\/j.celrep.2026.117962_bib44","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/0022-1759(95)00072-I","article-title":"A novel assay for apoptosis. Flow cytometric detection of phosphatidylserine expression on early apoptotic cells using fluorescein labelled Annexin V","volume":"184","author":"Vermes","year":"1995","journal-title":"J. Immunol. Methods"},{"key":"10.1016\/j.celrep.2026.117962_bib45","doi-asserted-by":"crossref","first-page":"4577","DOI":"10.1093\/emboj\/19.17.4577","article-title":"Localization of phosphatidylinositol 3-phosphate in yeast and mammalian cells","volume":"19","author":"Gillooly","year":"2000","journal-title":"Embo j"},{"key":"10.1016\/j.celrep.2026.117962_bib46","doi-asserted-by":"crossref","first-page":"880","DOI":"10.1038\/nbt.4201","article-title":"Efficient proximity labeling in living cells and organisms with TurboID","volume":"36","author":"Branon","year":"2018","journal-title":"Nat. Biotechnol."},{"key":"10.1016\/j.celrep.2026.117962_bib47","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1038\/s41568-018-0038-z","article-title":"Every step of the way: integrins in cancer progression and metastasis","volume":"18","author":"Hamidi","year":"2018","journal-title":"Nat. Rev. Cancer"},{"key":"10.1016\/j.celrep.2026.117962_bib48","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1146\/annurev-biochem-061516-044932","article-title":"Endoplasmic Reticulum-Plasma Membrane Contact Sites","volume":"86","author":"Saheki","year":"2017","journal-title":"Annu. Rev. Biochem."},{"key":"10.1016\/j.celrep.2026.117962_bib49","doi-asserted-by":"crossref","first-page":"1235","DOI":"10.1016\/j.cub.2005.05.055","article-title":"STIM is a Ca2+ sensor essential for Ca2+-store-depletion-triggered Ca2+ influx","volume":"15","author":"Liou","year":"2005","journal-title":"Curr. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib50","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1038\/nature04702","article-title":"A mutation in Orai1 causes immune deficiency by abrogating CRAC channel function","volume":"441","author":"Feske","year":"2006","journal-title":"Nature"},{"key":"10.1016\/j.celrep.2026.117962_bib51","first-page":"11","article-title":"Junctophilins: a novel family of junctional membrane complex proteins","volume":"6","author":"Takeshima","year":"2000","journal-title":"Mol. Cell"},{"key":"10.1016\/j.celrep.2026.117962_bib52","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1016\/j.celrep.2013.09.038","article-title":"Feedback regulation of receptor-induced Ca2+ signaling mediated by E-Syt1 and Nir2 at endoplasmic reticulum-plasma membrane junctions","volume":"5","author":"Chang","year":"2013","journal-title":"Cell Rep."},{"key":"10.1016\/j.celrep.2026.117962_bib53","doi-asserted-by":"crossref","first-page":"19133","DOI":"10.1074\/jbc.272.31.19133","article-title":"Cloning of a Chinese hamster ovary (CHO) cDNA encoding phosphatidylserine synthase (PSS) II, overexpression of which suppresses the phosphatidylserine biosynthetic defect of a PSS I-lacking mutant of CHO-K1 cells","volume":"272","author":"Kuge","year":"1997","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.celrep.2026.117962_bib54","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1126\/science.aab1370","article-title":"INTRACELLULAR TRANSPORT. PI4P\/phosphatidylserine countertransport at ORP5- and ORP8-mediated ER-plasma membrane contacts","volume":"349","author":"Chung","year":"2015","journal-title":"Science"},{"key":"10.1016\/j.celrep.2026.117962_bib55","first-page":"1","article-title":"OSBP-Related Protein Family in Lipid Transport Over Membrane Contact Sites","volume":"8","author":"Olkkonen","year":"2015","journal-title":"Lipid Insights"},{"key":"10.1016\/j.celrep.2026.117962_bib56","doi-asserted-by":"crossref","first-page":"552","DOI":"10.1038\/nature13269","article-title":"Structure of a lipid-bound extended synaptotagmin indicates a role in lipid transfer","volume":"510","author":"Schauder","year":"2014","journal-title":"Nature"},{"key":"10.1016\/j.celrep.2026.117962_bib57","article-title":"Endoplasmic Reticulum-Plasma Membrane Contact Sites: Regulators, Mechanisms, and Physiological Functions","volume":"9","author":"Li","year":"2021","journal-title":"Front. Cell Dev. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib58","doi-asserted-by":"crossref","first-page":"757","DOI":"10.1038\/s41467-017-00861-5","article-title":"ORP5 and ORP8 bind phosphatidylinositol-4, 5-biphosphate (PtdIns(4,5)P (2)) and regulate its level at the plasma membrane","volume":"8","author":"Ghai","year":"2017","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2026.117962_bib59","doi-asserted-by":"crossref","first-page":"623","DOI":"10.1038\/373623a0","article-title":"Axl receptor tyrosine kinase stimulated by the vitamin K-dependent protein encoded by growth-arrest-specific gene 6","volume":"373","author":"Varnum","year":"1995","journal-title":"Nature"},{"key":"10.1016\/j.celrep.2026.117962_bib60","doi-asserted-by":"crossref","first-page":"738","DOI":"10.1016\/j.tibs.2017.06.004","article-title":"Phosphatidylserine Is the Signal for TAM Receptors and Their Ligands","volume":"42","author":"Lemke","year":"2017","journal-title":"Trends Biochem. Sci."},{"key":"10.1016\/j.celrep.2026.117962_bib61","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1186\/s12943-019-1090-3","article-title":"AXL receptor tyrosine kinase as a promising anti-cancer approach: functions, molecular mechanisms and clinical applications","volume":"18","author":"Zhu","year":"2019","journal-title":"Mol. Cancer"},{"key":"10.1016\/j.celrep.2026.117962_bib62","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1083\/jcb.201211138","article-title":"Extracellular vesicles: exosomes, microvesicles, and friends","volume":"200","author":"Raposo","year":"2013","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib63","doi-asserted-by":"crossref","DOI":"10.3390\/cancers13164234","article-title":"Quantitative Tyrosine Phosphoproteome Profiling of AXL Receptor Tyrosine Kinase Signaling Network","volume":"13","author":"Wu","year":"2021","journal-title":"Cancers (Basel)"},{"key":"10.1016\/j.celrep.2026.117962_bib64","doi-asserted-by":"crossref","DOI":"10.3390\/cancers17050755","article-title":"Targeting STAT3 for Cancer Therapy: Focusing on Y705, S727, or Dual Inhibition?","volume":"17","author":"Berkley","year":"2025","journal-title":"Cancers (Basel)"},{"key":"10.1016\/j.celrep.2026.117962_bib65","doi-asserted-by":"crossref","first-page":"442","DOI":"10.3389\/fimmu.2014.00442","article-title":"Tetraspanins in extracellular vesicle formation and function","volume":"5","author":"Andreu","year":"2014","journal-title":"Front. Immunol."},{"key":"10.1016\/j.celrep.2026.117962_bib66","doi-asserted-by":"crossref","first-page":"1875","DOI":"10.1016\/j.cub.2009.09.059","article-title":"ARF6-regulated shedding of tumor cell-derived plasma membrane microvesicles","volume":"19","author":"Muralidharan-Chari","year":"2009","journal-title":"Curr. Biol."},{"key":"10.1016\/j.celrep.2026.117962_bib67","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1016\/j.cell.2019.02.029","article-title":"Reassessment of Exosome Composition","volume":"177","author":"Jeppesen","year":"2019","journal-title":"Cell"},{"key":"10.1016\/j.celrep.2026.117962_bib68","doi-asserted-by":"crossref","first-page":"25750","DOI":"10.1074\/jbc.M114.569020","article-title":"Receptor tyrosine kinases, TYRO3, AXL, and MER, demonstrate distinct patterns and complex regulation of ligand-induced activation","volume":"289","author":"Tsou","year":"2014","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.celrep.2026.117962_bib69","doi-asserted-by":"crossref","first-page":"1808","DOI":"10.1126\/science.1281555","article-title":"Activation of transcription by IFN-gamma: tyrosine phosphorylation of a 91-kD DNA binding protein","volume":"258","author":"Shuai","year":"1992","journal-title":"Science"},{"key":"10.1016\/j.celrep.2026.117962_bib70","doi-asserted-by":"crossref","first-page":"962","DOI":"10.1038\/cdd.2016.11","article-title":"Phosphatidylserine is a global immunosuppressive signal in efferocytosis, infectious disease, and cancer","volume":"23","author":"Birge","year":"2016","journal-title":"Cell Death Differ."},{"key":"10.1016\/j.celrep.2026.117962_bib71","doi-asserted-by":"crossref","DOI":"10.1172\/JCI163446","article-title":"Macrophages and microglia in glioblastoma: heterogeneity, plasticity, and therapy","volume":"133","author":"Khan","year":"2023","journal-title":"J. Clin. Investig."},{"key":"10.1016\/j.celrep.2026.117962_bib72","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.immuni.2014.06.008","article-title":"Macrophage activation and polarization: nomenclature and experimental guidelines","volume":"41","author":"Murray","year":"2014","journal-title":"Immunity"},{"key":"10.1016\/j.celrep.2026.117962_bib73","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1152\/physrev.00011.2010","article-title":"Physiology of microglia","volume":"91","author":"Kettenmann","year":"2011","journal-title":"Physiol. Rev."},{"key":"10.1016\/j.celrep.2026.117962_bib74","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1146\/annurev-bioeng-110220-115419","article-title":"Regulation of Tumor Invasion by the Physical Microenvironment: Lessons from Breast and Brain Cancer","volume":"24","author":"Beeghly","year":"2022","journal-title":"Annu. Rev. Biomed. Eng."},{"key":"10.1016\/j.celrep.2026.117962_bib75","article-title":"The biomechanical signature of tumor invasion","volume":"13","author":"Liu","year":"2026","journal-title":"Genes Dis."},{"key":"10.1016\/j.celrep.2026.117962_bib76","doi-asserted-by":"crossref","first-page":"2586","DOI":"10.1016\/j.cell.2025.02.034","article-title":"Cell membranes sustain phospholipid imbalance via cholesterol asymmetry","volume":"188","author":"Doktorova","year":"2025","journal-title":"Cell"},{"key":"10.1016\/j.celrep.2026.117962_bib77","article-title":"Trogocytosis of cancer-associated fibroblasts promotes pancreatic cancer growth and immune suppression via phospholipid scramblase anoctamin 6 (ANO6)","author":"Ogier","year":"2023","journal-title":"bioRxiv"}],"container-title":["Cell Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2211124726010405?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2211124726010405?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T20:45:31Z","timestamp":1792097131000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S2211124726010405"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9]]},"references-count":77,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2026,9]]}},"alternative-id":["S2211124726010405"],"URL":"https:\/\/doi.org\/10.1016\/j.celrep.2026.117962","relation":{},"ISSN":["2211-1247"],"issn-type":[{"value":"2211-1247","type":"print"}],"subject":[],"published":{"date-parts":[[2026,9]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"SERINC2-mediated phosphatidylserine remodeling activates protrusion-localized AXL signaling to promote brain metastasis","name":"articletitle","label":"Article Title"},{"value":"Cell Reports","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.celrep.2026.117962","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Author(s). Published by Elsevier Inc.","name":"copyright","label":"Copyright"}],"article-number":"117962"}}