{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,3]],"date-time":"2025-10-03T13:07:40Z","timestamp":1759496860818,"version":"3.37.3"},"reference-count":62,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,7,21]],"date-time":"2021-07-21T00:00:00Z","timestamp":1626825600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,7,21]],"date-time":"2021-07-21T00:00:00Z","timestamp":1626825600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"the Joint Usage\/Research Center on Tropical Disease, Institute of Tropical Medicine, Nagasaki University","award":["#2020-Ippan-05","#2020-Ippan-05"],"award-info":[{"award-number":["#2020-Ippan-05","#2020-Ippan-05"]}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP17H03935"],"award-info":[{"award-number":["JP17H03935"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100008884","name":"Nakatani Foundation for Advancement of Measuring Technologies in Biomedical Engineering","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100008884","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007449","name":"Takeda Science Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100007449","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007428","name":"The Naito Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100007428","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100008664","name":"ONO Medical Research Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100008664","id-type":"DOI","asserted-by":"crossref"}]},{"name":"The NOVARTIS Foundation (Japan) for the Promotion of Science"},{"DOI":"10.13039\/100008732","name":"The Uehara Memorial Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100008732","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Lipid rafts, sterol-rich and sphingolipid-rich microdomains on the plasma membrane are important in processes like cell signaling, adhesion, and protein and lipid transport. The virulence of many eukaryotic parasites is related to raft microdomains on the cell membrane. In the malaria parasite<jats:italic>Plasmodium falciparum<\/jats:italic>, glycosylphosphatidylinositol-anchored proteins, which are important for invasion and are possible targets for vaccine development, are localized in the raft. However, rafts are poorly understood. We used quick-freezing and freeze-fracture immuno-electron microscopy to examine the localization of monosialotetrahexosylganglioside (GM1) and monosialodihexosylganglioside (GM3), putative raft microdomain components in<jats:italic>P. falciparum<\/jats:italic>and infected erythrocytes. This method immobilizes molecules in situ, minimizing artifacts. GM3 was localized in the exoplasmic (EF) and cytoplasmic leaflets (PF) of the parasite and the parasitophorous vacuole (PV) membranes, but solely in the EF of the infected erythrocyte membrane, as in the case for uninfected erythrocytes. Phosphatidylinositol 4,5-bisphosphate (PtdIns(4,5)P<jats:sub>2<\/jats:sub>) was localized solely in the PF of erythrocyte, parasite, and PV membranes. This is the first time that GM3, the major component of raft microdomains, was found in the PF of a biological membrane. The unique localization of raft microdomains may be due to<jats:italic>P. falciparum<\/jats:italic>lipid metabolism and its unique biological processes, like protein transport from the parasite to infected erythrocytes.<\/jats:p>","DOI":"10.1038\/s41598-021-94037-3","type":"journal-article","created":{"date-parts":[[2021,7,21]],"date-time":"2021-07-21T10:07:53Z","timestamp":1626862073000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Glycosphingolipid GM3 is localized in both exoplasmic and cytoplasmic leaflets of Plasmodium falciparum malaria parasite plasma membrane"],"prefix":"10.1038","volume":"11","author":[{"given":"Shiomi","family":"Koudatsu","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tatsunori","family":"Masatani","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rikako","family":"Konishi","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Masahito","family":"Asada","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hassan","family":"Hakimi","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuna","family":"Kurokawa","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kanna","family":"Tomioku","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Osamu","family":"Kaneko","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2990-2134","authenticated-orcid":false,"given":"Akikazu","family":"Fujita","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2021,7,21]]},"reference":[{"key":"94037_CR1","doi-asserted-by":"publisher","first-page":"569","DOI":"10.1038\/42408","volume":"387","author":"K Simons","year":"1997","unstructured":"Simons, K. & Ikonen, E. Functional rafts in cell membranes. Nature 387, 569\u2013572. https:\/\/doi.org\/10.1038\/42408 (1997).","journal-title":"Nature"},{"key":"94037_CR2","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1016\/0092-8674(92)90189-j","volume":"68","author":"DA Brown","year":"1992","unstructured":"Brown, D. A. & Rose, J. K. Sorting of GPI-anchored proteins to glycolipid-enriched membrane subdomains during transport to the apical cell surface. Cell 68, 533\u2013544. https:\/\/doi.org\/10.1016\/0092-8674(92)90189-j (1992).","journal-title":"Cell"},{"key":"94037_CR3","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1006\/jsbi.2000.4266","volume":"131","author":"MC Giocondi","year":"2000","unstructured":"Giocondi, M. C., Vie, V., Lesniewska, E., Goudonnet, J. P. & Le Grimellec, C. In situ imaging of detergent-resistant membranes by atomic force microscopy. J. Struct. Biol. 131, 38\u201343. https:\/\/doi.org\/10.1006\/jsbi.2000.4266 (2000).","journal-title":"J. Struct. Biol."},{"key":"94037_CR4","doi-asserted-by":"publisher","first-page":"2693","DOI":"10.1016\/S0006-3495(02)75278-8","volume":"83","author":"H Heerklotz","year":"2002","unstructured":"Heerklotz, H. Triton promotes domain formation in lipid raft mixtures. Biophys. J. 83, 2693\u20132701 (2002).","journal-title":"Biophys. J."},{"key":"94037_CR5","doi-asserted-by":"publisher","first-page":"2112","DOI":"10.1091\/mbc.E07-01-0071","volume":"18","author":"A Fujita","year":"2007","unstructured":"Fujita, A. et al. Gangliosides GM1 and GM3 in the living cell membrane form clusters susceptible to cholesterol depletion and chilling. Mol. Biol. Cell. 18, 2112\u20132122. https:\/\/doi.org\/10.1091\/mbc.E07-01-0071 (2007).","journal-title":"Mol. Biol. Cell."},{"key":"94037_CR6","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1007\/s00418-007-0341-3","volume":"128","author":"A Fujita","year":"2007","unstructured":"Fujita, A. & Fujimoto, T. Quantitative retention of membrane lipids in the freeze-fracture replica. Histochem. Cell. Biol. 128, 385\u2013389. https:\/\/doi.org\/10.1007\/s00418-007-0341-3 (2007).","journal-title":"Histochem. Cell. Biol."},{"key":"94037_CR7","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1038\/nprot.2010.20","volume":"5","author":"A Fujita","year":"2010","unstructured":"Fujita, A., Cheng, J. & Fujimoto, T. Quantitative electron microscopy for the nanoscale analysis of membrane lipid distribution. Nat. Protoc. 5, 661\u2013669. https:\/\/doi.org\/10.1038\/nprot.2010.20 (2010).","journal-title":"Nat. Protoc."},{"key":"94037_CR8","doi-asserted-by":"publisher","first-page":"2453","DOI":"10.1242\/jcs.109.10.2453","volume":"109","author":"K Fujimoto","year":"1996","unstructured":"Fujimoto, K., Umeda, M. & Fujimoto, T. Transmembrane phospholipid distribution revealed by freeze-fracture replica labeling. J. Cell. Sci. 109, 2453\u20132460 (1996).","journal-title":"J. Cell. Sci."},{"key":"94037_CR9","doi-asserted-by":"publisher","first-page":"5164","DOI":"10.1073\/pnas.0834959100","volume":"100","author":"VK Goel","year":"2003","unstructured":"Goel, V. K. et al. Band 3 is a host receptor binding merozoite surface protein 1 during the Plasmodium falciparum invasion of erythrocytes. Proc. Natl. Acad. Sci. USA 100, 5164\u20135169. https:\/\/doi.org\/10.1073\/pnas.0834959100 (2003).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"94037_CR10","doi-asserted-by":"publisher","first-page":"40169","DOI":"10.1074\/jbc.M509631200","volume":"280","author":"PR Sanders","year":"2005","unstructured":"Sanders, P. R. et al. Distinct protein classes including novel merozoite surface antigens in Raft-like membranes of Plasmodium falciparum. J. Biol. Chem. 280, 40169\u201340176. https:\/\/doi.org\/10.1074\/jbc.M509631200 (2005).","journal-title":"J. Biol. Chem."},{"key":"94037_CR11","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1016\/j.ijpara.2016.10.002","volume":"47","author":"SA Arredondo","year":"2017","unstructured":"Arredondo, S. A. & Kappe, S. H. I. The s48\/45 six-cysteine proteins: Mediators of interaction throughout the Plasmodium life cycle. Int. J. Parasitol. 47, 409\u2013423. https:\/\/doi.org\/10.1016\/j.ijpara.2016.10.002 (2017).","journal-title":"Int. J. Parasitol."},{"key":"94037_CR12","doi-asserted-by":"publisher","first-page":"1920","DOI":"10.1182\/blood-2003-09-3165","volume":"103","author":"SC Murphy","year":"2004","unstructured":"Murphy, S. C. et al. Erythrocyte detergent-resistant membrane proteins: Their characterization and selective uptake during malarial infection. Blood 103, 1920\u20131928. https:\/\/doi.org\/10.1182\/blood-2003-09-3165 (2004).","journal-title":"Blood"},{"key":"94037_CR13","doi-asserted-by":"publisher","first-page":"29319","DOI":"10.1074\/jbc.M101268200","volume":"276","author":"BU Samuel","year":"2001","unstructured":"Samuel, B. U. et al. The role of cholesterol and glycosylphosphatidylinositol-anchored proteins of erythrocyte rafts in regulating raft protein content and malarial infection. J. Biol. Chem. 276, 29319\u201329329. https:\/\/doi.org\/10.1074\/jbc.M101268200 (2001).","journal-title":"J. Biol. Chem."},{"key":"94037_CR14","doi-asserted-by":"publisher","first-page":"3556","DOI":"10.1093\/emboj\/19.14.3556","volume":"19","author":"S Lauer","year":"2000","unstructured":"Lauer, S. et al. Vacuolar uptake of host components, and a role for cholesterol and sphingomyelin in malarial infection. EMBO J. 19, 3556\u20133564. https:\/\/doi.org\/10.1093\/emboj\/19.14.3556 (2000).","journal-title":"EMBO J."},{"key":"94037_CR15","doi-asserted-by":"publisher","first-page":"388","DOI":"10.1016\/j.bbalip.2009.01.008","volume":"1791","author":"A Fujita","year":"2009","unstructured":"Fujita, A., Cheng, J. & Fujimoto, T. Segregation of GM1 and GM3 clusters in the cell membrane depends on the intact actin cytoskeleton. Biochim. Biophys. Acta 1791, 388\u2013396 (2009).","journal-title":"Biochim. Biophys. Acta"},{"key":"94037_CR16","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1038\/35036052","volume":"1","author":"K Simons","year":"2000","unstructured":"Simons, K. & Toomre, D. Lipid rafts and signal transduction. Nat. Rev. Mol. Cell. Biol. 1, 31\u201339 (2000).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"94037_CR17","doi-asserted-by":"publisher","first-page":"9256","DOI":"10.1073\/pnas.0900216106","volume":"106","author":"A Fujita","year":"2009","unstructured":"Fujita, A., Cheng, J., Tauchi-Sato, K., Takenawa, T. & Fujimoto, T. A distinct pool of phosphatidylinositol 4,5-bisphosphate in caveolae revealed by a nanoscale labeling technique. Proc. Natl. Acad. Sci. USA 106, 9256\u20139261. https:\/\/doi.org\/10.1073\/pnas.0900216106 (2009).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"94037_CR18","doi-asserted-by":"publisher","first-page":"2132","DOI":"10.1182\/blood-2007-04-083873","volume":"110","author":"SC Murphy","year":"2007","unstructured":"Murphy, S. C. et al. Cytoplasmic remodeling of erythrocyte raft lipids during infection by the human malaria parasite Plasmodium falciparum. Blood 110, 2132\u20132139. https:\/\/doi.org\/10.1182\/blood-2007-04-083873 (2007).","journal-title":"Blood"},{"key":"94037_CR19","doi-asserted-by":"publisher","first-page":"10635","DOI":"10.1073\/pnas.91.22.10635","volume":"91","author":"X Han","year":"1994","unstructured":"Han, X. & Gross, R. W. Electrospray ionization mass spectroscopic analysis of human erythrocyte plasma membrane phospholipids. Proc. Natl. Acad. Sci. USA 91, 10635\u201310639. https:\/\/doi.org\/10.1073\/pnas.91.22.10635 (1994).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"94037_CR20","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1016\/0162-0134(94)00010-8","volume":"57","author":"J Bramham","year":"1995","unstructured":"Bramham, J. & Riddell, F. G. The effect of lithium therapy upon the composition of the human erythrocyte membrane. J. Inorg. Biochem. 57, 23\u201332. https:\/\/doi.org\/10.1016\/0162-0134(94)00010-8 (1995).","journal-title":"J. Inorg. Biochem."},{"key":"94037_CR21","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1126\/science.803712","volume":"187","author":"JA Dvorak","year":"1975","unstructured":"Dvorak, J. A., Miller, L. H., Whitehouse, W. C. & Shiroishi, T. Invasion of erythrocytes by malaria merozoites. Science 187, 748\u2013750. https:\/\/doi.org\/10.1126\/science.803712 (1975).","journal-title":"Science"},{"key":"94037_CR22","doi-asserted-by":"publisher","first-page":"291","DOI":"10.1042\/bst0220291","volume":"22","author":"AA Holder","year":"1994","unstructured":"Holder, A. A. et al. Malaria parasites and erythrocyte invasion. Biochem. Soc. Trans. 22, 291\u2013295. https:\/\/doi.org\/10.1042\/bst0220291 (1994).","journal-title":"Biochem. Soc. Trans."},{"key":"94037_CR23","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/s0014-4894(02)00143-1","volume":"102","author":"E Nagao","year":"2002","unstructured":"Nagao, E., Seydel, K. B. & Dvorak, J. A. Detergent-resistant erythrocyte membrane rafts are modified by a Plasmodium falciparum infection. Exp. Parasitol. 102, 57\u201359. https:\/\/doi.org\/10.1016\/s0014-4894(02)00143-1 (2002).","journal-title":"Exp. Parasitol."},{"key":"94037_CR24","doi-asserted-by":"publisher","first-page":"1141","DOI":"10.1182\/blood.v97.4.1141","volume":"97","author":"U Salzer","year":"2001","unstructured":"Salzer, U. & Prohaska, R. Stomatin, flotillin-1, and flotillin-2 are major integral proteins of erythrocyte lipid rafts. Blood 97, 1141\u20131143. https:\/\/doi.org\/10.1182\/blood.v97.4.1141 (2001).","journal-title":"Blood"},{"key":"94037_CR25","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1007\/BF02703669","volume":"30","author":"A Ciana","year":"2005","unstructured":"Ciana, A., Balduini, C. & Minetti, G. Detergent-resistant membranes in human erythrocytes and their connection to the membrane-skeleton. J. Biosci. 30, 317\u2013328. https:\/\/doi.org\/10.1007\/BF02703669 (2005).","journal-title":"J. Biosci."},{"key":"94037_CR26","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1111\/j.2517-6161.1977.tb01615.x","volume":"39","author":"BD Ripley","year":"1977","unstructured":"Ripley, B. D. Modeling spatial patterns. J. R. Stat. Soc. Ser. B 39, 172\u2013212 (1977).","journal-title":"J. R. Stat. Soc. Ser. B"},{"key":"94037_CR27","doi-asserted-by":"crossref","first-page":"368","DOI":"10.1111\/j.2517-6161.1979.tb01091.x","volume":"41","author":"BD Ripley","year":"1979","unstructured":"Ripley, B. D. Tests of randomness for spatial point patterns. J. R. Stat. Soc. Ser. B 41, 368\u2013374 (1979).","journal-title":"J. R. Stat. Soc. Ser. B"},{"key":"94037_CR28","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1083\/jcb.200209091","volume":"160","author":"IA Prior","year":"2003","unstructured":"Prior, I. A., Muncke, C., Parton, R. G. & Hancock, J. F. Direct visualization of Ras proteins in spatially distinct cell surface microdomains. J. Cell. Biol. 160, 165\u2013170 (2003).","journal-title":"J. Cell. Biol."},{"key":"94037_CR29","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1016\/j.tracli.2008.03.014","volume":"15","author":"L Tilley","year":"2008","unstructured":"Tilley, L. & Hanssen, E. A 3D view of the host cell compartment in P. falciparum-infected erythrocytes. Transfus Clin. Biol. 15, 72\u201381. https:\/\/doi.org\/10.1016\/j.tracli.2008.03.014 (2008).","journal-title":"Transfus Clin. Biol."},{"key":"94037_CR30","doi-asserted-by":"publisher","first-page":"4650","DOI":"10.1073\/pnas.76.9.4650","volume":"76","author":"A Kilejian","year":"1979","unstructured":"Kilejian, A. Characterization of a protein correlated with the production of knob-like protrusions on membranes of erythrocytes infected with Plasmodium falciparum. Proc. Natl. Acad. Sci. USA 76, 4650\u20134653. https:\/\/doi.org\/10.1073\/pnas.76.9.4650 (1979).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"94037_CR31","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1006\/jsbi.2000.4236","volume":"130","author":"E Nagao","year":"2000","unstructured":"Nagao, E., Kaneko, O. & Dvorak, J. A. Plasmodium falciparum-infected erythrocytes: Qualitative and quantitative analyses of parasite-induced knobs by atomic force microscopy. J. Struct. Biol. 130, 34\u201344. https:\/\/doi.org\/10.1006\/jsbi.2000.4236 (2000).","journal-title":"J. Struct. Biol."},{"key":"94037_CR32","doi-asserted-by":"publisher","first-page":"1100","DOI":"10.1177\/28.10.6775025","volume":"28","author":"GA Ackerman","year":"1980","unstructured":"Ackerman, G. A., Wolken, K. W. & Gelder, F. B. Surface distribution of monosialoganglioside GM1 on human blood cells and the effect of exogenous GM1 and neuraminidase on cholera toxin surface labelling. A quantitative immunocytochemical study. J. Histochem. Cytochem. 28, 1100\u20131112. https:\/\/doi.org\/10.1177\/28.10.6775025 (1980).","journal-title":"J. Histochem. Cytochem."},{"key":"94037_CR33","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1016\/j.tcb.2011.12.002","volume":"22","author":"NE Ziolkowska","year":"2012","unstructured":"Ziolkowska, N. E., Christiano, R. & Walther, T. C. Organized living: Formation mechanisms and functions of plasma membrane domains in yeast. Trends Cell. Biol. 22, 151\u2013158. https:\/\/doi.org\/10.1016\/j.tcb.2011.12.002 (2012).","journal-title":"Trends Cell. Biol."},{"key":"94037_CR34","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1083\/jcb.143.2.501","volume":"143","author":"P Varnai","year":"1998","unstructured":"Varnai, P. & Balla, T. Visualization of phosphoinositides that bind pleckstrin homology domains: Calcium- and agonist-induced dynamic changes and relationship to myo-[3H]inositol-labeled phosphoinositide pools. J. Cell Biol. 143, 501\u2013510 (1998).","journal-title":"J. Cell Biol."},{"key":"94037_CR35","doi-asserted-by":"publisher","first-page":"1019","DOI":"10.1152\/physrev.00028.2012","volume":"93","author":"T Balla","year":"2013","unstructured":"Balla, T. Phosphoinositides: Tiny lipids with giant impact on cell regulation. Physiol. Rev. 93, 1019\u20131137 (2013).","journal-title":"Physiol. Rev."},{"key":"94037_CR36","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/S0308-2261(21)00460-4","volume":"14","author":"SL Schrier","year":"1985","unstructured":"Schrier, S. L. Red cell membrane biology: Introduction. Clin. Haematol. 14, 1\u201312 (1985).","journal-title":"Clin. Haematol."},{"key":"94037_CR37","doi-asserted-by":"publisher","first-page":"1112","DOI":"10.1182\/blood.V74.3.1112.1112","volume":"74","author":"JA Chasis","year":"1989","unstructured":"Chasis, J. A., Prenant, M., Leung, A. & Mohandas, N. Membrane assembly and remodeling during reticulocyte maturation. Blood 74, 1112\u20131120 (1989).","journal-title":"Blood"},{"key":"94037_CR38","doi-asserted-by":"publisher","first-page":"755","DOI":"10.1016\/j.cell.2006.02.006","volume":"124","author":"AF Cowman","year":"2006","unstructured":"Cowman, A. F. & Crabb, B. S. Invasion of red blood cells by malaria parasites. Cell 124, 755\u2013766. https:\/\/doi.org\/10.1016\/j.cell.2006.02.006 (2006).","journal-title":"Cell"},{"key":"94037_CR39","doi-asserted-by":"publisher","first-page":"5197","DOI":"10.1074\/jbc.M509807200","volume":"281","author":"J Baum","year":"2006","unstructured":"Baum, J. et al. A conserved molecular motor drives cell invasion and gliding motility across malaria life cycle stages and other apicomplexan parasites. J. Biol. Chem. 281, 5197\u20135208. https:\/\/doi.org\/10.1074\/jbc.M509807200 (2006).","journal-title":"J. Biol. Chem."},{"key":"94037_CR40","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1016\/j.mib.2009.06.008","volume":"12","author":"W Daher","year":"2009","unstructured":"Daher, W. & Soldati-Favre, D. Mechanisms controlling glideosome function in apicomplexans. Curr. Opin. Microbiol. 12, 408\u2013414. https:\/\/doi.org\/10.1016\/j.mib.2009.06.008 (2009).","journal-title":"Curr. Opin. Microbiol."},{"key":"94037_CR41","doi-asserted-by":"publisher","first-page":"3039","DOI":"10.1091\/mbc.e07-01-0040","volume":"18","author":"TM Johnson","year":"2007","unstructured":"Johnson, T. M., Rajfur, Z., Jacobson, K. & Beckers, C. J. Immobilization of the type XIV myosin complex in Toxoplasma gondii. Mol. Biol. Cell 18, 3039\u20133046. https:\/\/doi.org\/10.1091\/mbc.e07-01-0040 (2007).","journal-title":"Mol. Biol. Cell"},{"key":"94037_CR42","doi-asserted-by":"publisher","first-page":"148","DOI":"10.1016\/j.molbiopara.2007.04.013","volume":"154","author":"PR Sanders","year":"2007","unstructured":"Sanders, P. R. et al. Identification of protein complexes in detergent-resistant membranes of Plasmodium falciparum schizonts. Mol. Biochem. Parasitol. 154, 148\u2013157. https:\/\/doi.org\/10.1016\/j.molbiopara.2007.04.013 (2007).","journal-title":"Mol. Biochem. Parasitol."},{"key":"94037_CR43","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1016\/j.chom.2012.06.005","volume":"12","author":"ML Jones","year":"2012","unstructured":"Jones, M. L., Collins, M. O., Goulding, D., Choudhary, J. S. & Rayner, J. C. Analysis of protein palmitoylation reveals a pervasive role in Plasmodium development and pathogenesis. Cell Host Microbe 12, 246\u2013258. https:\/\/doi.org\/10.1016\/j.chom.2012.06.005 (2012).","journal-title":"Cell Host Microbe"},{"key":"94037_CR44","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/j.molbiopara.2006.04.008","volume":"149","author":"RR Rees-Channer","year":"2006","unstructured":"Rees-Channer, R. R. et al. Dual acylation of the 45 kDa gliding-associated protein (GAP45) in Plasmodium falciparum merozoites. Mol. Biochem. Parasitol. 149, 113\u2013116. https:\/\/doi.org\/10.1016\/j.molbiopara.2006.04.008 (2006).","journal-title":"Mol. Biochem. Parasitol."},{"key":"94037_CR45","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0033845","volume":"7","author":"MA Ridzuan","year":"2012","unstructured":"Ridzuan, M. A. et al. Subcellular location, phosphorylation and assembly into the motor complex of GAP45 during Plasmodium falciparum schizont development. PLoS ONE 7, e33845. https:\/\/doi.org\/10.1371\/journal.pone.0033845 (2012).","journal-title":"PLoS ONE"},{"key":"94037_CR46","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1016\/j.chom.2010.09.002","volume":"8","author":"K Frenal","year":"2010","unstructured":"Frenal, K. et al. Functional dissection of the apicomplexan glideosome molecular architecture. Cell Host Microbe 8, 343\u2013357. https:\/\/doi.org\/10.1016\/j.chom.2010.09.002 (2010).","journal-title":"Cell Host Microbe"},{"key":"94037_CR47","doi-asserted-by":"publisher","first-page":"22050","DOI":"10.1073\/pnas.1016184107","volume":"107","author":"I Levental","year":"2010","unstructured":"Levental, I., Lingwood, D., Grzybek, M., Coskun, U. & Simons, K. Palmitoylation regulates raft affinity for the majority of integral raft proteins. Proc. Natl. Acad. Sci. USA 107, 22050\u201322054. https:\/\/doi.org\/10.1073\/pnas.1016184107 (2010).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"94037_CR48","doi-asserted-by":"publisher","first-page":"2204","DOI":"10.1111\/j.1432-1033.2004.04150.x","volume":"271","author":"AS Couto","year":"2004","unstructured":"Couto, A. S. et al. Glycosphingolipids in Plasmodium falciparum. Presence of an active glucosylceramide synthase. Eur. J. Biochem. 271, 2204\u20132214. https:\/\/doi.org\/10.1111\/j.1432-1033.2004.04150.x (2004).","journal-title":"Eur. J. Biochem."},{"key":"94037_CR49","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1515\/bchm2.1984.365.1.185","volume":"365","author":"R Schauer","year":"1984","unstructured":"Schauer, R., Wember, M. & Howard, R. J. Malaria parasites do not contain or synthesize sialic acids. Hoppe Seylers Z. Physiol. Chem. 365, 185\u2013194. https:\/\/doi.org\/10.1515\/bchm2.1984.365.1.185 (1984).","journal-title":"Hoppe Seylers Z. Physiol. Chem."},{"key":"94037_CR50","doi-asserted-by":"publisher","first-page":"2183","DOI":"10.1083\/jcb.108.6.2183","volume":"108","author":"K Haldar","year":"1989","unstructured":"Haldar, K., de Amorim, A. F. & Cross, G. A. Transport of fluorescent phospholipid analogues from the erythrocyte membrane to the parasite in Plasmodium falciparum-infected cells. J. Cell. Biol. 108, 2183\u20132192. https:\/\/doi.org\/10.1083\/jcb.108.6.2183 (1989).","journal-title":"J. Cell. Biol."},{"key":"94037_CR51","doi-asserted-by":"publisher","first-page":"13106","DOI":"10.1021\/bi020385t","volume":"41","author":"X Buton","year":"2002","unstructured":"Buton, X. et al. Transbilayer movement of monohexosylsphingolipids in endoplasmic reticulum and Golgi membranes. Biochemistry 41, 13106\u201313115. https:\/\/doi.org\/10.1021\/bi020385t (2002).","journal-title":"Biochemistry"},{"key":"94037_CR52","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1016\/s0962-8924(98)01249-5","volume":"8","author":"S Ichikawa","year":"1998","unstructured":"Ichikawa, S. & Hirabayashi, Y. Glucosylceramide synthase and glycosphingolipid synthesis. Trends Cell. Biol. 8, 198\u2013202. https:\/\/doi.org\/10.1016\/s0962-8924(98)01249-5 (1998).","journal-title":"Trends Cell. Biol."},{"key":"94037_CR53","doi-asserted-by":"publisher","first-page":"13305","DOI":"10.1074\/jbc.M113.457937","volume":"288","author":"J Suzuki","year":"2013","unstructured":"Suzuki, J. et al. Calcium-dependent phospholipid scramblase activity of TMEM16 protein family members. J Biol Chem 288, 13305\u201313316. https:\/\/doi.org\/10.1074\/jbc.M113.457937 (2013).","journal-title":"J Biol Chem"},{"key":"94037_CR54","doi-asserted-by":"publisher","DOI":"10.1101\/2020.1106.1122.165258","author":"S Haase","year":"2020","unstructured":"Haase, S. et al. Identification and characterisation of a phospholipid scramblase in the malaria parasite Plasmodium falciparum. bioRxiv https:\/\/doi.org\/10.1101\/2020.1106.1122.165258 (2020).","journal-title":"bioRxiv"},{"key":"94037_CR55","doi-asserted-by":"publisher","first-page":"1146","DOI":"10.1016\/S0021-9258(20)71220-2","volume":"260","author":"RG Sleight","year":"1985","unstructured":"Sleight, R. G. & Pagano, R. E. Transbilayer movement of a fluorescent phosphatidylethanolamine analogue across the plasma membranes of cultured mammalian cells. J. Biol. Chem. 260, 1146\u20131154 (1985).","journal-title":"J. Biol. Chem."},{"key":"94037_CR56","doi-asserted-by":"publisher","first-page":"5890","DOI":"10.1016\/S0021-9258(18)45658-X","volume":"262","author":"OC Martin","year":"1987","unstructured":"Martin, O. C. & Pagano, R. E. Transbilayer movement of fluorescent analogs of phosphatidylserine and phosphatidylethanolamine at the plasma membrane of cultured cells. Evidence for a protein-mediated and ATP-dependent process(es). J. Biol. Chem. 262, 5890\u20135898 (1987).","journal-title":"J. Biol. Chem."},{"key":"94037_CR57","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1212\/WNL.47.1.237","volume":"47","author":"S Kusunoki","year":"1996","unstructured":"Kusunoki, S. et al. GM1b is a new member of antigen for serum antibody in Guillain-Barre syndrome. Neurology 47, 237\u2013242 (1996).","journal-title":"Neurology"},{"key":"94037_CR58","doi-asserted-by":"publisher","first-page":"595","DOI":"10.1177\/002215549704500411","volume":"45","author":"T Fujimoto","year":"1997","unstructured":"Fujimoto, T. & Fujimoto, K. Metal sandwich method to quick-freeze monolayer cultured cells for freeze-fracture. J. Histochem. Cytochem. 45, 595\u2013598 (1997).","journal-title":"J. Histochem. Cytochem."},{"key":"94037_CR59","doi-asserted-by":"publisher","first-page":"1298","DOI":"10.1016\/j.bbamem.2016.03.011","volume":"1858","author":"A Yoshida","year":"2016","unstructured":"Yoshida, A., Shigekuni, M., Tanabe, K. & Fujita, A. Nanoscale analysis reveals agonist-sensitive and heterogeneous pools of phosphatidylinositol 4-phosphate in the plasma membrane. Biochim. Biophys. Acta 1858, 1298\u20131305. https:\/\/doi.org\/10.1016\/j.bbamem.2016.03.011 (2016).","journal-title":"Biochim. Biophys. Acta"},{"key":"94037_CR60","doi-asserted-by":"publisher","first-page":"3443","DOI":"10.1242\/jcs.108.11.3443","volume":"108","author":"K Fujimoto","year":"1995","unstructured":"Fujimoto, K. Freeze-fracture replica electron microscopy combined with SDS digestion for cytochemical labeling of integral membrane proteins. Application to the immunogold labeling of intercellular junctional complexes. J. Cell. Sci. 108, 3443\u20133449 (1995).","journal-title":"J. Cell. Sci."},{"key":"94037_CR61","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1136\/jcp.20.4.603","volume":"20","author":"AJ Salsbury","year":"1967","unstructured":"Salsbury, A. J. & Clarke, J. A. New method for detecting changes in the surface appearance of human red blood cells. J. Clin. Pathol. 20, 603\u2013610 (1967).","journal-title":"J. Clin. Pathol."},{"key":"94037_CR62","first-page":"507","volume":"77","author":"JG White","year":"1974","unstructured":"White, J. G. Effects of an ionophore, A23187, on the surface morphology of normal erythrocytes. Am. J. Pathol. 77, 507\u2013518 (1974).","journal-title":"Am. J. Pathol."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-94037-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-94037-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-94037-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,9,4]],"date-time":"2024-09-04T18:02:15Z","timestamp":1725472935000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-94037-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,7,21]]},"references-count":62,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["94037"],"URL":"https:\/\/doi.org\/10.1038\/s41598-021-94037-3","relation":{},"ISSN":["2045-2322"],"issn-type":[{"type":"electronic","value":"2045-2322"}],"subject":[],"published":{"date-parts":[[2021,7,21]]},"assertion":[{"value":"11 April 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"28 June 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"21 July 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"14890"}}