{"id":14,"date":"2019-09-09T18:53:43","date_gmt":"2019-09-09T18:53:43","guid":{"rendered":"https:\/\/lab.research.sickkids.ca\/freeman\/?page_id=14"},"modified":"2026-02-02T18:37:55","modified_gmt":"2026-02-02T18:37:55","slug":"publications","status":"publish","type":"page","link":"https:\/\/lab.research.sickkids.ca\/freeman\/publications\/","title":{"rendered":"PUBLICATIONS"},"content":{"rendered":"<div class=\"wpb-content-wrapper\"><p>[vc_row][vc_column]<div class=\"dt-fancy-separator h3-size style-double title-color\" style=\"width: 100%;\"><div class=\"dt-fancy-title\"><span class=\"separator-holder separator-left\"><\/span>Publications<span class=\"separator-holder separator-right\"><\/span><\/div><\/div>[vc_column_text]<\/p>\n<p><span style=\"color: #3366ff\"><strong>2026<\/strong><\/span><\/p>\n<p>Le T, Li YM, Saric A, Holthuis JCM, Sergio Grinstein, <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.molbiolcell.org\/doi\/abs\/10.1091\/mbc.E25-05-0245\">Lipid scrambling via TMEM16F mediates the formation and release of apoptotic vesicles.<\/a> <em>bioRxiv<\/em>, 2026<\/p>\n<p><span style=\"color: #3366ff\"><strong>2025<\/strong><\/span><\/p>\n<p><strong>Freeman SA<\/strong>, Grinstein S. <a href=\"https:\/\/rupress.org\/jcb\/article\/225\/1\/e202511084\/278534\">Is the Parkinson\u2019s-associated protein TMEM175 a proton channel: Yay or nay?<\/a> <em>Journal of Cell Biology<\/em>, 2025<\/p>\n<p>Wyss LC, <strong>Freeman SA<\/strong> (2025). <a href=\"https:\/\/onlinelibrary.wiley.com\/doi\/full\/10.1002\/pgr2.70040\">The glycocalyx: barriers and opportunities at cell-cell encounters.<\/a>\u00a0Proteoglycan Research, 2025<\/p>\n<p>Kavianpour AA, Ghasempour S, Meyer KJ, Le T, Cai R, Marques PE, Nodwell JR, <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.cell.com\/current-biology\/abstract\/S0960-9822(25)00952-2\">Phosphatidylethanolamine is a phagocytic ligand implicated in the binding and removal of apoptotic and bacterial extracellular vesicles.<\/a> Current Biology, 2025<\/p>\n<p>de Paz Linares G, Mossemann J, Sayed B, <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.molbiolcell.org\/doi\/full\/10.1091\/mbc.E24-11-0511\">The scaffolding and activation of NLRP3 on acidic vesicles depends on their biophysical properties and is independent of intermediate filaments<\/a>.\u00a0 Molecular Biology of the Cell, 2025<\/p>\n<p>Wu JZ, Pemberton JG, Morioka S, Sasaki J, Bablani, P, Sasaki T, Balla T, Grinstein S, <strong>Freeman SA<\/strong>. <a href=\"https:\/\/rupress.org\/jcb\/article-abstract\/224\/6\/e202408174\/277345\/Sorting-nexin-10-regulates-lysosomal-ionic\">Sorting nexin 10 regulates lysosomal ionic homeostasis via CIC-7 by controlling PI(3,5)P2<\/a>.\u00a0 Journal of Cell Biology, v2025<\/p>\n<p>Scott O, Saran E, <strong>Freeman SA<\/strong>. <a class=\"gsc_oci_title_link\" href=\"https:\/\/www.embopress.org\/doi\/abs\/10.1038\/s44319-025-00405-9\" data-clk=\"hl=en&amp;sa=T&amp;ei=wTPcZ8D_E4CxieoPk8a88Q4\">The spectrum of lysosomal stress and damage responses: from mechanosensing to inflammation<\/a>. <em>EMBO Reports<\/em>, 2025<\/p>\n<p>Skrtic M, Yusuf B, Patel S, Reddy EC, Ting KKY, Cybulsky MI, <strong>Freeman SA<\/strong>, Robinson LA. <a class=\"gsc_a_at\" href=\"https:\/\/scholar.google.ca\/citations?view_op=view_citation&amp;hl=en&amp;user=RUUm0V0AAAAJ&amp;sortby=pubdate&amp;citation_for_view=RUUm0V0AAAAJ:nb7KW1ujOQ8C\">The neurorepellent SLIT2 inhibits LPS-induced proinflammatory signaling in macrophages<\/a>. <em>Journal of Immunology<\/em>, 2025<\/p>\n<p><span style=\"color: #3366ff\"><strong>2024<\/strong><\/span><\/p>\n<p class=\"gsc_oci_value\">Cai R, Scott O, Ye G, Le T, Saran E, Kwon W, Inpanathan S, Sayed BA, Botelho RJ, Saric A, Uderhardt S, <strong>Freeman SA<\/strong>. <a class=\"gsc_oci_title_link\" href=\"https:\/\/www.nature.com\/articles\/s41556-024-01459-y\" data-clk=\"hl=en&amp;sa=T&amp;ei=shWZZo_jD92xy9YPzOP1kQ4\">Pressure sensing of lysosomes enables control of TFEB responses in macrophages. <\/a><em>Nature Cell Biology<\/em>, 2024<\/p>\n<p>Le T, Ferling I, Qin L, Nabaile C, Assuncao L, Roskelley CD, Grinstein S, and <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.cell.com\/developmental-cell\/abstract\/S1534-5807(24)00040-6\">Redistribution of the glycocalyx exposes phagocytic determinants on apoptotic cells<\/a>. <em>Developmental Cell<\/em>, 2024<\/p>\n<p>Chadwick S, Barreda D, Wu JZ, Ye G, Yusuf B, Ren D, and <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.pnas.org\/doi\/10.1073\/pnas.2309465121?doi=10.1073%2Fpnas.2309465121\">Two pore channels regulate endomembrane tension to enable remodeling and resolution of phagolysosomes<\/a>. <em>PNAS<\/em>, 2024<\/p>\n<p>Barreda D, Grinstein S, and <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0171933523000973\">Target lysis by cholesterol extraction is a rate limiting step in the resolution of phagolysosomes<\/a>. <em>European Journal of Cell Biology<\/em>, 2024<\/p>\n<p>Dissanayake D, Firouzabady A, Massumi M, de Paz Linares GA, Marshall C, <strong>Freeman SA<\/strong>, Laxer RM, Yeung RSM. <a class=\"gsc_oci_title_link\" href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0006497124015179\" data-clk=\"hl=en&amp;sa=T&amp;ei=s2NoZsSxPOaAy9YP3p6ByAg\">Interleukin-1 mediated hyperinflammation in XIAP-deficiency is associated with defective autophagy.<\/a> <em>Blood<\/em>, 2024<\/p>\n<div id=\"gsc_oci_table\">\n<div class=\"gs_scl\">\n<p class=\"gsc_oci_value\">Lu C-Y, Wu JW, Yao HHY, Liu RJY, Li L, Pluthero FG, <strong>Freeman SA<\/strong>, Kahr WHA. <a class=\"gsc_oci_title_link\" href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S1538783624002472\" data-clk=\"hl=en&amp;sa=T&amp;ei=95pHZuesLPGDy9YPlOu1CA\">Acidification of \u03b1-granules in megakaryocytes by vacuolar-type ATPase is essential for organelle biogenesis<\/a>. <em>Journal of Thrombosis and Haemostasis<\/em>, 2024<\/p>\n<p class=\"gsc_oci_value\"><span style=\"color: #3366ff\"><strong>2023<\/strong><\/span><\/p>\n<\/div>\n<\/div>\n<p>Mylvaganam S and <strong>Freeman SA<\/strong>. <a href=\"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/imr.13260\">The resolution of phagosomes<\/a>. <em>Immunological Reviews<\/em>, 2023<\/p>\n<p>de Paz Linares GA, <strong>Freeman SA<\/strong>, Cai R. <a class=\"gsc_a_at\" href=\"https:\/\/scholar.google.com\/citations?view_op=view_citation&amp;hl=en&amp;user=RUUm0V0AAAAJ&amp;sortby=pubdate&amp;authuser=1&amp;citation_for_view=RUUm0V0AAAAJ:pqnbT2bcN3wC\">Using Ion Substitution and Fluid Indicators to Monitor Macropinosome Dynamics in Live Cells<\/a>. <em>Phagocytosis and Phagosomes: Methods and Protocols, <\/em>2023<\/p>\n<p>Wu J, Zeziulia M, Kwon W, Jentsch TJ, Grinstein S, <strong>Freeman SA<\/strong>. <a class=\"gsc_oci_title_link\" href=\"https:\/\/rupress.org\/jcb\/article-abstract\/222\/6\/e202208155\/213990\" data-clk=\"hl=en&amp;sa=T&amp;ei=s5uIZNg_y_aaAcbztIgO&amp;authuser=1\">ClC-7 drives intraphagosomal chloride accumulation to support hydrolase activity and phagosome resolution<\/a>. <em>Journal of Cell Biology, <\/em>2023<\/p>\n<p class=\"gs_gray\">Hirata Y, Cai R, Steinberg BE, Volchuk A, Saito Y, Matsuzawa A, Grinstein S, <strong>Freeman SA<\/strong>. <a class=\"gsc_a_at\" href=\"https:\/\/www.cell.com\/current-biology\/pdf\/S0960-9822(23)00200-2.pdf\">Lipid peroxidation increases membrane tension, Piezo1 gating and cation permeability to execute ferroptosis<\/a>. <em>Current Biology<\/em>, 2023<\/p>\n<p class=\"gs_gray\">Teoh CW, Riedl Khursigara M, Ortiz-Sandoval CG, Park JW, Li J, Bohorquez-Hernandez A, Bruno V, Bowen EE, <strong>Freeman SA<\/strong>, Robinson LA, Licht C.\u00a0<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC9968885\/\">The loss of glycocalyx integrity impairs complement factor H binding and contributes to cyclosporine-induced endothelial cell injury<\/a>. <em>Frontiers in Medicine<\/em>, 2023<\/p>\n<p class=\"gs_gray\"><strong>Freeman SA<\/strong>, Grinstein S, Orlowski J. <a class=\"gsc_a_at\" href=\"https:\/\/scholar.google.ca\/citations?view_op=view_citation&amp;hl=en&amp;user=RUUm0V0AAAAJ&amp;sortby=pubdate&amp;authuser=1&amp;citation_for_view=RUUm0V0AAAAJ:vV6vV6tmYwMC\">Determinants, maintenance, and function of organellar pH<\/a>.\u00a0<em>Physiological Reviews<\/em>, 2023<\/p>\n<p class=\"gs_gray\">Ghasempour S, and <strong>Freeman SA<\/strong>. <a href=\"https:\/\/febs.onlinelibrary.wiley.com\/doi\/10.1111\/febs.16236\">The glycocalyx and immune evasion in cancer<\/a>. <em>The FEBS Journal<\/em>, 2023<\/p>\n<p><span style=\"color: #3366ff\"><strong>2022<\/strong><\/span><\/p>\n<p>Yazdani S, Bilan PJ, Jaldin-Fincati JR, Pang J, Ceban F, Saran E, Brumell JH, <strong>Freeman SA<\/strong>, Klip A. \u00a0<a href=\"https:\/\/www.molbiolcell.org\/doi\/abs\/10.1091\/mbc.E22-04-0146\">Dynamic glucose uptake, storage and release by human microvascular endothelial cells<\/a>. <em>MBoC<\/em>, 2022<\/p>\n<p>Mylvaganam S, Plumb J, Yusuf B, Li R, Lu CY, Robinson LA, <strong>Freeman SA<\/strong>, Grinstein S. \u00a0<a href=\"https:\/\/www.nature.com\/articles\/s41556-022-00953-5\">The spectrin cytoskeleton integrates endothelial mechanoresponses<\/a>. <em>Nature Cell Biology<\/em>, 2022<\/p>\n<p><span style=\"color: #3366ff\"><strong>2021<\/strong><\/span><\/p>\n<p>Leung G, Zhou Y, Ostrowski P, Mylvaganam S, Boroumand P, Mulder DJ, Guo C, Muise AM, <strong>Freeman SA<\/strong>. \u00a0<a href=\"https:\/\/insight.jci.org\/articles\/view\/149376\">ARPC1B binds WASP to control actin polymerization and curtail tonic signaling in B cells<\/a>. <em>JCI Insight<\/em>, 2021<\/p>\n<p>Arif AA, Huang YH, <strong>Freeman SA<\/strong>, Atif J, Dean P, Lai JCY, Blanchet MR, Wiegand KC, McNagny KM, Underhill TM, Gold MR, Johnson P, Roskelley CD. <a href=\"https:\/\/mcr.aacrjournals.org\/content\/early\/2021\/10\/16\/1541-7786.MCR-21-0009.abstract\">Inflammation-induced metastatic colonization of the lung is facilitated by hepatocyte growth factor-secreting monocyte-derived macrophages.<\/a>\u00a0 <em>Molecular Cancer Research<\/em>, 2021<\/p>\n<p>Saric A,\u00a0<strong>Freeman SA<\/strong>, Williamson C, Jarnik M, Guardia C, Fernandopulle M, Gershlick D, Bonifacino J. <a href=\"https:\/\/www.nature.com\/articles\/s41467-021-24709-1#citeas\">SNX19 Restricts endolysosome motility through contacts with endoplasmic reticulum<\/a>.\u00a0<em>Nature Communications<\/em> 12(4552), 2021<\/p>\n<p>Mylvaganam S, <strong>Freeman SA<\/strong>, Grinstein S. <a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0960982221000695?via%3Dihub\">The cytoskeleton in phagocytosis and macropinocytosis<\/a>. <i>Current Biology <\/i><i>31<\/i>(10), 619\u2013632, 2021<\/p>\n<p>Wang L, Aschenbrenner D, Zeng Z, Cao X, Mayr D, Mehta M, Capitani M, Warner N, Pan J, Wang L, Li Q, Zuo T, Cohen-Kedar S, Lu J, Ardy RC, Mulder DJ, Dissanayake D, Peng K, Huang Z, Li X, Wang Y, Wang X, Li S, Bullers S, Gammage AN, Warnatz K, Schiefer AI, Krivan G, Goda V, Kahr WHA, Lemaire M; Genomics England Research Consortium, Lu CY, Siddiqui I, Surette MG, Kotlarz D, Engelhardt KR, Griffin HR, Rottapel R, Decaluwe H, Laxer RM, Proietti M, Hambleton S, Elcombe S, Guo CH, Grimbacher B, Dotan I, Ng SC, <strong>Freeman SA<\/strong>, Snapper SB, Klein C, Boztug K, Huang Y, Li D, Uhlig HH, Muise AM.\u00a0<a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/33782605\/\">Gain-of-function variants in SYK cause immune dysregulation and systemic inflammation in humans and mice<\/a>. <i>Nature Genetics<\/i>\u00a053,4: 500-510, 2021<\/p>\n<p>Chadwick SR, Grinstein S,\u00a0<strong>Freeman SA<\/strong>.\u00a0 <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/33706237\/\">From the inside out: Ion fluxes at the centre of endocytic traffic<\/a>.\u00a0\u00a0<em>Current Opinion in Cell Biology<\/em> 71, 77-86, 2021<\/p>\n<p>Saric A and <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2020.611326\/full\">Endomembrane Tension and Trafficking<\/a>.\u00a0<i>Frontiers in Cell and Developmental Biology.<\/i> 8:611326, 2021<\/p>\n<p>Saric A and <strong>Freeman SA<\/strong>. <a href=\"https:\/\/rdcu.be\/cd2AC\">Solutes as controllers of endomembrane dynamics<\/a>.\u00a0<i>Nature Reviews Molecular Cell Biology<\/i>,\u00a02021<\/p>\n<p>Chadwick SR, Wu JZ, <strong>Freeman SA<\/strong>\u00a0\u00a0<a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/33385319\/\">Solute Transport Controls Membrane Tension and Organellar Volume.<\/a>\u00a0<em> Cellular Physiology &amp; Biochemistry<\/em>. 55(S1):1-24, 2021<\/p>\n<p><strong>Freeman SA<\/strong> and Grinstein S.\u00a0<a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/34152790\/\">Promoters and Antagonists of Phagocytosis: A Plastic and Tunable Response<\/a>.\u00a0<em> Annual Review of Cell and Developmental Biology,\u00a0<\/em>volume 37, 2021<\/p>\n<p><span style=\"color: #3366ff\"><strong>2020<\/strong><\/span><\/p>\n<div class=\"gs_gray\">\n<p>Kwon W and <strong>Freeman SA<\/strong>.\u00a0<a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/33281830\/\">Phagocytosis by the Retinal Pigment Epithelium: Recognition, Resolution, Recycling.<\/a>\u00a0 <em>Frontiers in Immunology<\/em>. 11:604205,\u00a02020<\/p>\n<p>Imbert PRC, Saric A, Pedram K, Bertozzi CR, Grinstein S, <strong>Freeman SA<\/strong>. <a class=\"gsc_a_at\" href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0960982220314950\" data-href=\"\/citations?view_op=view_citation&amp;hl=en&amp;user=RUUm0V0AAAAJ&amp;sortby=pubdate&amp;citation_for_view=RUUm0V0AAAAJ:iH-uZ7U-co4C\">An Acquired and Endogenous Glycocalyx Forms a Bidirectional \u201cDon\u2019t Eat\u201d and \u201cDon\u2019t Eat Me\u201d Barrier to Phagocytosis.<\/a>\u00a0<em>Current Biology<\/em>, 2020<\/p>\n<p>Westman J, <strong>Freeman SA<\/strong>, Grinstein S. <a href=\"https:\/\/www.sciencedirect.com\/science\/article\/abs\/pii\/S0143416020301111\">Unconventional role of lysosomes in phagocytosis.<\/a><em> Cell Calcium 91, <\/em>102269, 2020<\/p>\n<p>Mercer J, Lee JE, Saphire EO, <strong>Freeman SA<\/strong>.\u00a0<a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0092867420308102\">SnapShot: Enveloped Virus Entry.<\/a><em> Cell <\/em>182 (3), 786-786. e1, 2020<\/p>\n<p>Mylvaganam S, Riedl M, Vega A, Collins RF, Jaqaman K, Grinstein S, <strong>Freeman SA<\/strong>.\u00a0<a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S2211124720307798\">Stabilization of Endothelial Receptor Arrays by a Polarized Spectrin Cytoskeleton Facilitates Rolling and Adhesion of Leukocytes.<\/a><em> Cell Reports <\/em>31 (12), 107798, 2020<\/p>\n<p><strong>Freeman SA<\/strong>, Uderhardt S, Saric A, Collins RF, Buckley CM, Mylvaganam S, Boroumand P, Plumb J, Germain RN, Ren D, Grinstein S.\u00a0<a href=\"https:\/\/science.sciencemag.org\/content\/367\/6475\/301.abstract\">Lipid-gated monovalent ion fluxes regulate endocytic traffic and support immune surveillance.<\/a><em> Science <\/em>367 (6475), 301-305, 2020<\/p>\n<p><strong>Freeman SA<\/strong> and Grinstein S.\u00a0<a href=\"https:\/\/www.sciencedirect.com\/science\/article\/abs\/pii\/S0960982219315179\">Phagocytosis: mechanosensing, traction forces, and a molecular clutch<em>.<\/em><\/a><em> Current Biology<\/em> 30 (1),<em> R24-R26,<\/em> 2020<\/p>\n<p><span style=\"color: #3366ff\"><strong>2019<\/strong><\/span><\/p>\n<p>Marques PE, Nyegaard S, Collins RF, Troise F, <strong>Freeman SA<\/strong>, Trimble WS, Grinstein S. <a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/31231038\">Multimerization and Retention of the Scavenger Receptor SR-B1 in the Plasma Membrane.<\/a> <em>Developmental Cell<\/em><em>.<\/em> 50 (3), 283-295, 2019<\/p>\n<p>Ostrowski PP, <strong>Freeman SA<\/strong>, Fairn G, Grinstein S. <a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/31231039\">Dynamic Podosome-Like Structures in Nascent Phagosomes Are Coordinated by Phosphoinositides<\/a>. <em>Developmental Cell<\/em><em>.<\/em>\u00a050 (4), 397-410, 2019<\/p>\n<p><span style=\"color: #3366ff\"><strong>2018<\/strong><\/span><\/p>\n<p><strong>Freeman SA<\/strong> and Grinstein S.\u00a0<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/30159984\">Resolution of macropinosomes, phagosomes and autolysosomes: Osmotically driven shrinkage enables tubulation and vesiculation.<\/a>\u00a0<em>Traffic<\/em>. 19 (12), 965-974, 2018<\/p>\n<p>Mylvaganam SM, Grinstein S, <strong>Freeman SA<\/strong>.\u00a0<a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/30209546\">Picket-fences in the plasma membrane: functions in immune cells and phagocytosis.<\/a> <em>Semin Immunopathol<\/em>. 40 (6), 605-615, 2018<\/p>\n<p>Schlam D, Grinstein S, <strong>Freeman SA<\/strong>. <a href=\"https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/30062408\">Screening for Rho GTPase Modulators in Actin-Dependent Processes Exemplified by Phagocytosis.<\/a><em> Methods Mol Biol<\/em>. 107-127, 2018<\/p>\n<p>Vega AR, Grinstein S, <strong>Freeman SA<\/strong>, Jaqaman K. <a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0006349518301322\">Multistep Track Segmentation and Motion Classification for Transient Mobility Analysis.<\/a> <em>Biophysical Journal. <\/em>114(5): 1018-1025, 2018<\/p>\n<p><strong>Freeman SA<\/strong>, Vega AR, Riedl M, Collins RF, Ostrowski PP, Woods EC, Bertozzi CR, Tammi MI, Lidke DS, Johnson P, Mayor S, Jaqaman K, Grinstein S.\u00a0<a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/29328918\/\">Transmembrane pickets connect cyto-and pericellular skeletons forming barriers to receptor engagement.<\/a> <em>Cell. <\/em>172(1): 305-317, 2016<\/p>\n<p><span style=\"color: #3366ff\"><strong>2017<\/strong><\/span><\/p>\n<p>Marques PE, Grinstein S, <strong>Freeman SA<\/strong>. <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/28475901\/\">SnapShot: Macropinocytosis.<\/a> <em>Cell.\u00a0<\/em>169(4): 766, 2017<\/p>\n<p>Wang JC, Lee JY-J, Christian S, Dang-Lawson M, Pritchard C, <strong>Freeman SA<\/strong>, Gold MR. <a href=\"https:\/\/jcs.biologists.org\/content\/130\/6\/1094\">The Rap1\u2013cofilin-1 pathway coordinates actin reorganization and MTOC polarization at the B cell immune synapse.<\/a> <em>Journal of Cell Science. 130(6): <\/em>1094-1109,\u00a02017<\/p>\n<p><strong>Freeman SA<\/strong>, Christian S<strong>,<\/strong> Austin P, Iu I, Graves ML, Huang L, Tang S, Coombs D, Gold MR, Roskelley CD. <a href=\"https:\/\/jcs.biologists.org\/content\/130\/1\/152\">Applied stretch initiates directional invasion through the action of Rap1 GTPase as a tension sensor.<\/a> <em>Journal of Cell Science. <\/em>130(1): 152-163, 2017<\/p>\n<p>Sharfe N, Karanxha A, Dadi H, Merico D, Chitayat D, Herbrick J, <strong>Freeman SA<\/strong>, Grinstein S, Roifman CM. <a href=\"https:\/\/www.jacionline.org\/article\/S0091-6749(17)31814-6\/fulltext\">Dual loss of p110\u03b4\u00a0PI3-kinase and SKAP (KNSTRN) expression leads to combined immunodeficiency and multisystem syndromic features.<\/a> <em>Journal of Allergy and Clinical Immunology. <\/em>142(2): 618-629, 2017<\/p>\n<p>Hua R, Cheng D, Coyaud \u00c9, <strong>Freeman SA<\/strong>, Di Pietro E, Wang Y, Vissa A, Yip CM, Fairn GD,\u00a0 Braverman N, Brumell JH, Trimble WS, Raught B, Peter K Kim.\u00a0<a href=\"https:\/\/rupress.org\/jcb\/article\/216\/2\/367\/46157\/VAPs-and-ACBD5-tether-peroxisomes-to-the-ER-for\">VAPs and ACBD5 tether peroxisomes to the ER for peroxisome maintenance and lipid homeostasis.<\/a> <em>J Cell Biol. 216(8): <\/em>367-377,\u00a02017<\/p>\n<p><span style=\"color: #3366ff\"><strong>2016<\/strong><\/span><\/p>\n<p>Saric A, Grinstein S, <strong>Freeman SA<\/strong>.\u00a0<a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/28237098\/\">Measurement of Autolysosomal pH by Dual-Wavelength Ratio Imaging.<\/a> <em>Methods in Enzymology. 588: <\/em>15-29, 2016<\/p>\n<p><strong>Freeman SA<\/strong> and Grinstein S. <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/27997839\/\">Phagocytosis: How Macrophages Tune Their Non-professional Counterparts.<\/a> <em>Current Biology. 26(24): <\/em>1279-1282, 2016<\/p>\n<p>Wong HS, Jaumouill\u00e9 V, <strong>Freeman SA<\/strong>,\u00a0Doodnauth SA, Schlam D, Canton J, Mukovozov IM, Saric A, Grinstein S, Robinson LA. <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/26997267\/\">Chemokine signaling enhances CD36 responsiveness toward oxidized low-density lipoproteins and accelerates foam cell formation.<\/a> <em>Cell Reports. <\/em>14(12): 2859-71, 2016<\/p>\n<p>Ostrowski PP, Grinstein S, <strong>Freeman SA<\/strong>. <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/27459066\/\">Diffusion barriers, mechanical forces, and the biophysics of phagocytosis.<\/a> <em>Developmental Cell. <\/em>38(2): 135-146, 2016<\/p>\n<p>Oudhoff MJ, Braam MJS, <strong>Freeman SA<\/strong>, Wong D, Rattray DG, Wang J, Antignano F, Snyder K, Refaeli I, Hughes MR, McNagny KM, Gold MR, Arrowsmith CH, Sato T, Rossi FMV, Tatlock JH, Owen DR, Brown PJ, Zaph C. <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/27046831\/\">SETD7 controls intestinal regeneration and tumorigenesis by regulating Wnt\/\u03b2-catenin and hippo\/YAP signaling.<\/a> <em>Developmental Cell. <\/em>37(1): 47-57, 2016<\/p>\n<p>Schlam D, Canton J, Carre\u00f1o M, Kopinski H, <strong>Freeman SA<\/strong>, Grinstein S, Fairn GD. <a href=\"https:\/\/mbio.asm.org\/content\/7\/2\/e02242-15.abstract\">Gliotoxin Suppresses Macrophage Immune Function by Subverting Phosphatidylinositol 3, 4, 5-Trisphosphate Homeostasis.<\/a> <em>MBio. <\/em>7(2): 02242-15, 2016<\/p>\n<p><strong>Freeman SA<\/strong>, Goyette J, Furuya W, Woods EC, Bertozzi CR, Bergmeier W, Hinz B, van der Merwe PA, Das R, Grinstein S. <a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0092867415015676\">Integrins form an expanding diffusional barrier that coordinates phagocytosis.<\/a> <em>Cell<\/em>. 164(1): 128-140, 2016<\/p>\n<p><span style=\"color: #3366ff\"><strong>2015<\/strong><\/span><\/p>\n<p><strong>Freeman SA<\/strong>, Jaumouill\u00e9 V, Choi K, Hsu BE, Wong HS, Abraham L, Graves ML, Coombs D, Roskelley CD, Das R, Grinstein S, Gold MR.\u00a0<a href=\"https:\/\/www.nature.com\/articles\/ncomms7168\">Toll-like receptor ligands sensitize B-cell receptor signalling by reducing actin-dependent spatial confinement of the receptor.<\/a> <em>Nature Communications. <\/em>6: 6168, 2015<\/p>\n<p>Schlam D, Bagshaw R, <strong>Freeman SA<\/strong>, Collins RF, Fairn G, Grinstein S. <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/26465210\/\">Phosphoinositide 3-kinase enables phagocytosis of large particles by terminating actin assembly through Rac\/Cdc42 GTPase-activating proteins<\/a>. <em>Nature Communications. <\/em>6: 8623, 2015<\/p>\n<p><span style=\"color: #3366ff\"><strong>&#8211; 2014<\/strong><\/span><\/p>\n<p>Barsyte-Lovejoy, Li F, Oudhoff MJ, Tatlock JH, Dong A, Zeng H, Wu H, <strong>Freeman SA<\/strong>, Schapira M, Senisterra GA, Kuznetsova E, Marcellus R, Allali-Hassani A, Kennedy S, Lambert JP, Couzens AL, Aman A, Gingras AC, Al-Awar R, Fish PV, Gerstenberger BS, Roberts L, Benn CL, Grimley RL, Braam MJ, Rossi FM, Sudol M, Brown PJ, Bunnage ME, Owen DR, Zaph C, Vedadi M, Arrowsmith CH. <a href=\"https:\/\/www.pnas.org\/content\/111\/35\/12853\">(R)-PFI-2 is a potent and selective inhibitor of SETD7 methyltransferase activity in cells.<\/a> <em>Proceedings of the National Academy of Sciences. <\/em>111(35): 12853-12858, 2014<\/p>\n<p><strong>Freeman SA<\/strong> and Grinstein S.\u00a0<a href=\"https:\/\/www.researchgate.net\/publication\/266914701_Phagocytosis_Receptors_signal_integration_and_the_cytoskeleton\">Phagocytosis: receptors, signal integration, and the cytoskeleton.<\/a><em> Immunological Reviews. <\/em>262(1): 193-215, 2014<\/p>\n<p>Oudhoff MJ, <strong>Freeman SA<\/strong>, Couzens AL, Antignano F, Kuznetsova E, Min PH, Northrop JP, Lehnertz B, Barsyte-Lovejoy D, Vedadi M, Arrowsmith CH, Nishina H, Gold MR, Rossi FMV, Gingras A, Zaph C. <a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S1534580713003195\">Control of the hippo pathway by set7-dependent methylation of yap.<\/a> <em>Developmental Cell. <\/em>26(2): 188-194, 2013<\/p>\n<p>Austin P, <strong>Freeman SA<\/strong>, Gray CA, Gold MR, Vogl AW, Andersen RJ, Roberge M, Roskelley CD. <a href=\"https:\/\/mcr.aacrjournals.org\/content\/11\/5\/530\">The invasion inhibitor sarasinoside A1 reverses mesenchymal tumor transformation in an E-cadherin-independent manner.<\/a> <em>Molecular Cancer Research. <\/em>11(5): 530-40, 2013<\/p>\n<p>Maltby S, <strong>Freeman SA<\/strong>, Gold MJ, Baker JHE, Minchinton AI, Gold MR, Roskelley CD, McNagny KM. <a href=\"https:\/\/journals.plos.org\/plosone\/article?id=10.1371\/journal.pone.0018160\">Opposing Roles for CD34 in B16 Melanoma Tumor Growth Alter Early Stage Vasculature and Late Stage Immune Cell Infiltration.<\/a><em> PloS one. <\/em>6(4): 18160, 2011<\/p>\n<p><strong>Freeman SA<\/strong>, Lei V, Dang-Lawson M, Mizuno K, Roskelley CD, Gold MR. <a href=\"https:\/\/www.jimmunol.org\/content\/early\/2011\/11\/06\/jimmunol.1102233\">Cofilin-Mediated F-Actin Severing Is Regulated by the Rap GTPase and Controls the Cytoskeletal Dynamics That Drive Lymphocyte Spreading and BCR Microcluster Formation.<\/a> <em>The Journal of Immunology. <\/em>187(11): 5887-5900, 2011<\/p>\n<p>Davies AH, Barrett I, Pambid MR, Hu K, Stratford AL, F<strong>reeman SA<\/strong>, Berquin IM, Pelech S, Hieter P, Maxwell C, Dunn SE. <a href=\"https:\/\/www.nature.com\/articles\/onc201182\">YB-1 evokes susceptibility to cancer through cytokinesis failure, mitotic dysfunction and HER2 amplification.<\/a> <em>Oncogene. <\/em>30(34): 3649-60, 2011<\/p>\n<p><strong>Freeman SA<\/strong>, McLeod SJ, Dukowski J, Austin P, Lee CCY, Millen-Martin B, Kubes P, McCafferty D, Gold MR, Roskelley CD.<a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/20484042\/\"> Preventing the activation or cycling of the Rap1 GTPase alters adhesion and cytoskeletal dynamics and blocks metastatic melanoma cell extravasation into the lungs.<\/a> <em>Cancer Research. <\/em>70(11): 4590-601, 2010<\/p>\n<p>Lin KBL, <strong>Freeman SA<\/strong>, Gold MR. <a href=\"https:\/\/www.tandfonline.com\/doi\/full\/10.4161\/cam.4.3.11114\">Rap GTPase-mediated adhesion and migration: A target for limiting the dissemination of B-cell lymphomas?.<\/a> <em>Cell adhesion &amp; migration. <\/em>4(3):327-32, 2010<\/p>\n<p>Lin KBL, Tan P, <strong>Freeman SA<\/strong>, Lam M, McNagny KM, Gold MR. <a href=\"https:\/\/pubmed.ncbi.nlm.nih.gov\/19838206\/\">The Rap GTPases regulate the migration, invasiveness and in vivo dissemination of B-cell lymphomas.<\/a><em> Oncogene. <\/em>29(4): 608-615, 2009<\/p>\n<p>Lin KBL, <strong>Freeman SA,<\/strong> Zabetian S, Brugger H, Weber M, Lei V, Dang-Lawson M, Kathy WK,\u00a0 Santamaria R, Batista FD, Gold MR. <a href=\"https:\/\/www.sciencedirect.com\/science\/article\/pii\/S1074761307005845\">The rap GTPases regulate B cell morphology, immune-synapse formation, and signaling by particulate B cell receptor ligands.<\/a>\u00a0 <em>Immunity. <\/em>28(1): 75-87,\u00a02008<\/p>\n<p>Badour K, McGavin MKH, Zhang J, <strong>Freeman SA<\/strong>, Vieira C, Filipp D, Julius M, Mills GB, Siminovitch KA. <a href=\"https:\/\/www.pnas.org\/content\/104\/5\/1593\">Interaction of the Wiskott-Aldrich syndrome protein with sorting nexin 9 is required for CD28 endocytosis and cosignaling in T cells.<\/a> <em>Proceedings of the National Academy of Sciences<\/em><em>. <\/em>104(5): 1593-1598, 2007<\/p>\n<p>Leng Y, Zhang J, Badour K, Arpaia E, <strong>Freeman SA<\/strong>, Cheung P, Siu M, Siminovitch K.<a href=\"https:\/\/www.pnas.org\/content\/102\/4\/1098\"> Abelson-interactor-1 promotes WAVE2 membrane translocation and Abelson-mediated tyrosine phosphorylation required for WAVE2 activation.<\/a>\u00a0 <em>Proceedings of the National Academy of Sciences<\/em><em>. <\/em>102(4): 1098-1103, 2005<\/p>\n<p>[vc_row][vc_column width=&#8221;1\/1&#8243;][vc_column_text]<\/p>\n<\/div>\n<p>[\/vc_column_text][\/vc_column][\/vc_row][vc_row][vc_column][vc_column_text]<\/p>\n<p>[\/vc_column_text][\/vc_column][\/vc_row]<\/p>\n<\/div>","protected":false},"excerpt":{"rendered":"<p>[vc_row][vc_column][vc_column_text] 2026 Le T, Li YM, Saric A, Holthuis JCM, Sergio Grinstein, Freeman SA. Lipid scrambling via TMEM16F mediates the formation and release of apoptotic vesicles. bioRxiv, 2026 2025 Freeman SA, Grinstein S. Is the Parkinson\u2019s-associated protein TMEM175 a proton channel: Yay or nay? Journal of Cell Biology, 2025 Wyss LC, Freeman SA (2025). The&hellip;<\/p>\n","protected":false},"author":119,"featured_media":196,"parent":0,"menu_order":0,"comment_status":"closed","ping_status":"closed","template":"","meta":{"footnotes":""},"class_list":["post-14","page","type-page","status-publish","has-post-thumbnail","hentry","description-off"],"yoast_head":"<!-- This site is optimized with the Yoast SEO Premium plugin v27.0 (Yoast SEO v27.0) - https:\/\/yoast.com\/product\/yoast-seo-premium-wordpress\/ -->\n<title>PUBLICATIONS - Freeman Lab<\/title>\n<meta name=\"robots\" content=\"index, follow, max-snippet:-1, max-image-preview:large, max-video-preview:-1\" \/>\n<link rel=\"canonical\" href=\"https:\/\/lab.research.sickkids.ca\/freeman\/publications\/\" \/>\n<meta property=\"og:locale\" content=\"en_US\" \/>\n<meta property=\"og:type\" content=\"article\" \/>\n<meta property=\"og:title\" content=\"PUBLICATIONS\" \/>\n<meta property=\"og:description\" content=\"[vc_row][vc_column][vc_column_text] 2026 Le T, Li YM, Saric A, Holthuis JCM, Sergio Grinstein, Freeman SA. 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