{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T03:05:49Z","timestamp":1789009549247,"version":"build-2803163510"},"reference-count":55,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,8,4]],"date-time":"2021-08-04T00:00:00Z","timestamp":1628035200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,8,4]],"date-time":"2021-08-04T00:00:00Z","timestamp":1628035200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Oligomerization of Pr55\n                    <jats:sup>Gag<\/jats:sup>\n                    is a critical step of the late stage of the HIV life cycle. It has been known that the binding of IP6, an abundant endogenous cyclitol molecule at the MA domain, has been linked to the oligomerization of Pr55\n                    <jats:sup>Gag<\/jats:sup>\n                    . However, the exact binding site of IP6 on MA remains unknown and the structural details of this interaction are missing. Here, we present three high-resolution crystal structures of the MA domain in complex with IP6 molecules to reveal its binding mode. Additionally, extensive Differential Scanning Fluorimetry analysis combined with cryo- and ambient-temperature\u00a0X-ray crystallography and GNM-based transfer entropy calculations identify the key residues that participate in IP6 binding. Our data provide novel insights about the multilayered HIV-1 virion assembly process that involves the interplay of IP6 with PIP2, a phosphoinositide essential for the binding of Pr55\n                    <jats:sup>Gag<\/jats:sup>\n                    to membrane. IP6 and PIP2 have neighboring alternate binding sites within the same highly basic region (residues 18\u201333). This indicates that IP6 and PIP2 bindings are not mutually exclusive and may play a key role in coordinating virion particles\u2019 membrane localization. Based on our three different IP6-MA complex crystal structures, we propose a new model that involves IP6 coordination of the oligomerization of outer MA and inner CA domain\u2019s 2D layers during assembly and budding.\n                  <\/jats:p>","DOI":"10.1038\/s41598-021-95236-8","type":"journal-article","created":{"date-parts":[[2021,8,4]],"date-time":"2021-08-04T06:03:09Z","timestamp":1628056989000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":12,"title":["Structural insight into host plasma membrane association and assembly of HIV-1 matrix protein"],"prefix":"10.1038","volume":"11","author":[{"given":"Halilibrahim","family":"Ciftci","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hiroshi","family":"Tateishi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kotaro","family":"Koiwai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryoko","family":"Koga","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kensaku","family":"Anraku","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kazuaki","family":"Monde","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"\u00c7a\u011fda\u015f","family":"Da\u011f","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ebru","family":"Destan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Busra","family":"Yuksel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Esra","family":"Ayan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gunseli","family":"Yildirim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Merve","family":"Yigin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"F. Betul","family":"Ertem","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alaleh","family":"Shafiei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Omur","family":"Guven","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sabri O.","family":"Besler","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Raymond G.","family":"Sierra","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chun Hong","family":"Yoon","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhen","family":"Su","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mengling","family":"Liang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Burcin","family":"Acar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Turkan","family":"Haliloglu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Masami","family":"Otsuka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fumiaki","family":"Yumoto","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mikako","family":"Fujita","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Toshiya","family":"Senda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hasan","family":"DeMirci","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,8,4]]},"reference":[{"key":"95236_CR1","doi-asserted-by":"crossref","unstructured":"Chinchar, V. G. Replication of Viruses. Encycl. Virol. 1471\u20131478 (1999).","DOI":"10.1006\/rwvi.1999.0245"},{"key":"95236_CR2","unstructured":"Walker, B. & Mcmichael, A. The T-Cell Response to HIV. 1\u201319 (2012)."},{"key":"95236_CR3","doi-asserted-by":"publisher","first-page":"484","DOI":"10.1038\/nrmicro3490","volume":"13","author":"EO Freed","year":"2015","unstructured":"Freed, E. O. HIV-1 assembly, release and maturation. Nat. Rev. Microbiol. 13, 484\u2013496. https:\/\/doi.org\/10.1038\/nrmicro3490 (2015).","journal-title":"Nat. Rev. Microbiol."},{"key":"95236_CR4","doi-asserted-by":"crossref","unstructured":"Sundquist, W. I. & Kra, H. HIV-1 Assembly, Budding, and Maturation. 1\u201324 (2012).","DOI":"10.1101\/cshperspect.a015420"},{"key":"95236_CR5","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fmicb.2012.00055","volume":"3","author":"RH Ghanam","year":"2012","unstructured":"Ghanam, R. H., Samal, A. B., Fernandez, T. F. & Saad, J. S. Role of the HIV-1 matrix protein in Gag intracellular trafficking and targeting to the plasma membrane for virus assembly. Front. Microbiol. 3, 1\u201315. https:\/\/doi.org\/10.3389\/fmicb.2012.00055 (2012).","journal-title":"Front. Microbiol."},{"key":"95236_CR6","doi-asserted-by":"publisher","first-page":"2113","DOI":"10.1016\/j.jmb.2018.04.042","volume":"430","author":"CR Gaines","year":"2018","unstructured":"Gaines, C. R. et al. HIV-1 matrix protein interactions with tRNA: implications for membrane targeting. J. Mol. Biol. 430, 2113\u20132127. https:\/\/doi.org\/10.1016\/j.jmb.2018.04.042 (2018).","journal-title":"J. Mol. Biol."},{"key":"95236_CR7","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1038\/s41586-018-0396-4","volume":"560","author":"RA Dick","year":"2018","unstructured":"Dick, R. A. et al. Inositol phosphates are assembly co-factors for HIV-1. Nature 560, 509\u2013512. https:\/\/doi.org\/10.1038\/s41586-018-0396-4 (2018).","journal-title":"Nature"},{"key":"95236_CR8","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1006\/jmbi.1994.1719","volume":"244","author":"MA Massiah","year":"1994","unstructured":"Massiah, M. A. et al. Three-dimensional structure of the human immunodeficiency virus type 1 matrix protein. J. Mol. Biol. 244, 198\u2013223. https:\/\/doi.org\/10.1006\/jmbi.1994.1719 (1994).","journal-title":"J. Mol. Biol."},{"key":"95236_CR9","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1073\/pnas.0305665101","volume":"101","author":"C Tang","year":"2004","unstructured":"Tang, C. et al. Entropic switch regulates myristate exposure in the HIV-1 matrix protein. Proc. Natl. Acad. Sci. U. S. A. 101, 517\u2013522. https:\/\/doi.org\/10.1073\/pnas.0305665101 (2004).","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"95236_CR10","doi-asserted-by":"publisher","first-page":"14889","DOI":"10.1073\/pnas.0405596101","volume":"101","author":"A Ono","year":"2004","unstructured":"Ono, A., Ablan, S. D., Lockett, S. J., Nagashima, K. & Freed, E. O. Phosphatidylinositol (4,5) bisphosphate regulates HIV-1 Gag targeting to the plasma membrane.\u00a0Proc. Natl. Acad. Sci. U. S. A. 101, 14889\u201314894. https:\/\/doi.org\/10.1073\/pnas.0405596101\u00a0(2004).\u00a0","journal-title":"Proc. Natl. Acad. Sci. U. S. A"},{"key":"95236_CR11","doi-asserted-by":"publisher","first-page":"3949","DOI":"10.1128\/jvi.69.6.3949-3954.1995","volume":"69","author":"EO Freed","year":"1995","unstructured":"Freed, E. O., Englund, G. & Martin, M. A. Role of the basic domain of human immunodeficiency virus type 1 matrix in macrophage infection. J. Virol. 69, 3949\u20133954. https:\/\/doi.org\/10.1128\/jvi.69.6.3949-3954.1995 (1995).","journal-title":"J. Virol."},{"key":"95236_CR12","doi-asserted-by":"publisher","first-page":"1521","DOI":"10.1016\/j.str.2005.07.010","volume":"13","author":"PS Murray","year":"2005","unstructured":"Murray, P. S. et al. Retroviral matrix domains share electrostatic homology: Models for membrane binding function throughout the viral life cycle. Structure 13, 1521\u20131531. https:\/\/doi.org\/10.1016\/j.str.2005.07.010 (2005).","journal-title":"Structure"},{"key":"95236_CR13","doi-asserted-by":"publisher","first-page":"5657","DOI":"10.1128\/jvi.00509-16","volume":"90","author":"A Alfadhli","year":"2016","unstructured":"Alfadhli, A. et al. Trimer enhancement mutation effects on HIV-1 matrix protein binding activities. J. Virol. 90, 5657\u20135664. https:\/\/doi.org\/10.1128\/jvi.00509-16 (2016).","journal-title":"J. Virol."},{"key":"95236_CR14","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1016\/j.virol.2014.08.017","volume":"468","author":"B Sanford","year":"2014","unstructured":"Sanford, B. et al. Deletions in the fifth alpha helix of HIV-1 matrix block virus release. Virology 468, 293\u2013302. https:\/\/doi.org\/10.1016\/j.virol.2014.08.017 (2014).","journal-title":"Virology"},{"key":"95236_CR15","doi-asserted-by":"publisher","first-page":"5311","DOI":"10.1128\/jvi.68.8.5311-5320.1994","volume":"68","author":"EO Freed","year":"1994","unstructured":"Freed, E. O., Orenstein, J. M., Buckler-White, A. J. & Martin, M. A. Single amino acid changes in the human immunodeficiency virus type 1 matrix protein block virus particle production. J. Virol. 68, 5311\u20135320. https:\/\/doi.org\/10.1128\/jvi.68.8.5311-5320.1994 (1994).","journal-title":"J. Virol."},{"key":"95236_CR16","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1006\/viro.1998.9398","volume":"251","author":"EO Freed","year":"1998","unstructured":"Freed, E. O. HIV-1 gag proteins: diverse functions in the virus life cycle. Virology 251, 1\u201315. https:\/\/doi.org\/10.1006\/viro.1998.9398 (1998).","journal-title":"Virology"},{"key":"95236_CR17","doi-asserted-by":"publisher","first-page":"2556","DOI":"10.1128\/jvi.68.4.2556-2569.1994","volume":"68","author":"W Zhou","year":"1994","unstructured":"Zhou, W., Parent, L. J., Wills, J. W. & Resh, M. D. Identification of a membrane-binding domain within the amino-terminal region of human immunodeficiency virus type 1 Gag protein which interacts with acidic phospholipids. J. Virol. 68, 2556\u20132569. https:\/\/doi.org\/10.1128\/jvi.68.4.2556-2569.1994 (1994).","journal-title":"J. Virol."},{"key":"95236_CR18","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-017-09129-w","volume":"7","author":"H Tateishi","year":"2017","unstructured":"Tateishi, H. et al. A clue to unprecedented strategy to HIV eradication: \u2018Lock-in and apoptosis\u2019. Sci. Rep. 7, 1\u20138. https:\/\/doi.org\/10.1038\/s41598-017-09129-w (2017).","journal-title":"Sci. Rep."},{"key":"95236_CR19","doi-asserted-by":"publisher","first-page":"11364","DOI":"10.1073\/pnas.0602818103","volume":"103","author":"JS Saad","year":"2006","unstructured":"Saad, J. S. et al. Structural basis for targeting HIV-1 Gag proteins to the plasma membrane for virus assembly. Proc. Natl. Acad. Sci. U. S. A. 103, 11364\u201311369. https:\/\/doi.org\/10.1073\/pnas.0602818103 (2006).","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"95236_CR20","doi-asserted-by":"publisher","first-page":"10875","DOI":"10.1073\/pnas.191224698","volume":"98","author":"S Campbell","year":"2001","unstructured":"Campbell, S. et al. Modulation of HIV-like particle assembly in vitro by inositol phosphates. Proc. Natl. Acad. Sci. U. S. A. 98, 10875\u201310879. https:\/\/doi.org\/10.1073\/pnas.191224698 (2001).","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"95236_CR21","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1042\/BJ20081417","volume":"416","author":"AJ Letcher","year":"2008","unstructured":"Letcher, A. J., Schell, M. J. & Irvine, R. F. Do mammals make all their own inositol hexakisphosphate?. Biochem. J. 416, 263\u2013270. https:\/\/doi.org\/10.1042\/BJ20081417 (2008).","journal-title":"Biochem. J."},{"key":"95236_CR22","doi-asserted-by":"publisher","first-page":"3983","DOI":"10.1016\/j.celrep.2019.11.050","volume":"29","author":"DL Mallery","year":"2019","unstructured":"Mallery, D. L. et al. Cellular IP6 levels limit HIV production while viruses that cannot efficiently package IP6 are attenuated for infection and replication. Cell Rep. 29, 3983-3996.e4. https:\/\/doi.org\/10.1016\/j.celrep.2019.11.050 (2019).","journal-title":"Cell Rep."},{"key":"95236_CR23","first-page":"799","volume":"365","author":"AKD Siddhartha","year":"2007","unstructured":"Siddhartha, A. K. D. et al. Interactions between HIV-1 gag molecules in solution: an inositol phosphate-mediated switch. Nat. Institutes Heal. 365, 799\u2013811 (2007).","journal-title":"Nat. Institutes Heal."},{"key":"95236_CR24","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1159\/000331995","volume":"55","author":"S Sierra-Arag\u00f3n","year":"2012","unstructured":"Sierra-Arag\u00f3n, S. & Walter, H. Targets for inhibition of HIV replication: entry, enzyme action, release and maturation. Intervirology 55, 84\u201397. https:\/\/doi.org\/10.1159\/000331995 (2012).","journal-title":"Intervirology"},{"key":"95236_CR25","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1128\/JVI.01079-19","volume":"93","author":"A Alfadhli","year":"2019","unstructured":"Alfadhli, A. et al. Analysis of HIV-1 . J. Virol. 93, 1\u201310 (2019).","journal-title":"J. Virol."},{"key":"95236_CR26","doi-asserted-by":"publisher","first-page":"1058","DOI":"10.1107\/S0907444906023821","volume":"62","author":"M Hiraki","year":"2006","unstructured":"Hiraki, M. et al. Development of an automated large-scale protein-crystallization and monitoring system for high-throughput protein-structure analyses. Acta Crystallogr. Sect. D Biol. Crystallogr. 62, 1058\u20131065. https:\/\/doi.org\/10.1107\/S0907444906023821 (2006).","journal-title":"Acta Crystallogr. Sect. D Biol. Crystallogr."},{"issue":"Pt 2","key":"95236_CR27","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1107\/S0907444909047337","volume":"66","author":"W Kabsch","year":"2010","unstructured":"Kabsch, W. XDS. Acta Crystallogr. Sect. D Biol. Crystallogr. 66(Pt 2), 125\u2013132. https:\/\/doi.org\/10.1107\/S0907444909047337 (2010).","journal-title":"Acta Crystallogr. Sect. D Biol. Crystallogr."},{"key":"95236_CR28","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1107\/S0907444905036693","volume":"62","author":"P Evans","year":"2006","unstructured":"Evans, P. Scaling and assessment of data quality. Acta Crystallogr. Sect. D Biol. Crystallogr. 62, 72\u201382. https:\/\/doi.org\/10.1107\/S0907444905036693 (2006).","journal-title":"Acta Crystallogr. Sect. D Biol. Crystallogr."},{"key":"95236_CR29","doi-asserted-by":"publisher","first-page":"1204","DOI":"10.1107\/S0907444913000061","volume":"69","author":"PR Evans","year":"2013","unstructured":"Evans, P. R. & Murshudov, G. N. How good are my data and what is the resolution?. Acta Crystallogr. Sect. D Biol. Crystallogr. 69, 1204\u20131214. https:\/\/doi.org\/10.1107\/S0907444913000061 (2013).","journal-title":"Acta Crystallogr. Sect. D Biol. Crystallogr."},{"key":"95236_CR30","doi-asserted-by":"publisher","first-page":"1022","DOI":"10.1107\/S0021889897006766","volume":"30","author":"A Vagin","year":"1997","unstructured":"Vagin, A. & Teplyakov, A. MOLREP: an automated program for molecular replacement. J. Appl. Crystallogr. 30, 1022\u20131025. https:\/\/doi.org\/10.1107\/S0021889897006766 (1997).","journal-title":"J. Appl. Crystallogr."},{"key":"95236_CR31","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1107\/S0907444996012255","volume":"53","author":"GN Murshudov","year":"1997","unstructured":"Murshudov, G. N., Vagin, A. A. & Dodson, E. J. Refinement of macromolecular structures by the maximum-likelihood method. Acta Crystallogr. Sect. D Biol. Crystallogr. 53, 240\u2013255. https:\/\/doi.org\/10.1107\/S0907444996012255 (1997).","journal-title":"Acta Crystallogr. Sect. D Biol. Crystallogr."},{"key":"95236_CR32","doi-asserted-by":"publisher","first-page":"352","DOI":"10.1107\/S0907444912001308","volume":"68","author":"PV Afonine","year":"2012","unstructured":"Afonine, P. V. et al. Towards automated crystallographic structure refinement with phenix.refine. Acta Crystallogr. Sect. D Biol. Crystallogr. 68, 352\u2013367. https:\/\/doi.org\/10.1107\/S0907444912001308 (2012).","journal-title":"Acta Crystallogr. Sect. D Biol. Crystallogr."},{"key":"95236_CR33","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1107\/S0907444910007493","volume":"66","author":"P Emsley","year":"2010","unstructured":"Emsley, P., Lohkamp, B., Scott, W. G. & Cowtan, K. Features and development of Coot. Acta Crystallogr. Sect. D Biol. Crystallogr. 66, 486\u2013501. https:\/\/doi.org\/10.1107\/S0907444910007493 (2010).","journal-title":"Acta Crystallogr. Sect. D Biol. Crystallogr."},{"key":"95236_CR34","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1016\/j.jmb.2007.05.022","volume":"372","author":"E Krissinel","year":"2007","unstructured":"Krissinel, E. & Henrick, K. Inference of macromolecular assemblies from crystalline state. J. Mol. Biol. 372, 774\u2013797. https:\/\/doi.org\/10.1016\/j.jmb.2007.05.022 (2007).","journal-title":"J. Mol. Biol."},{"key":"95236_CR35","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3390\/ijms20071675","volume":"20","author":"HI Ciftci","year":"2019","unstructured":"Ciftci, H. I. et al. Serial femtosecond X-ray diffraction of HIV-1 gag MA-IP6 microcrystals at ambient temperature. Int. J. Mol. Sci. 20, 1\u201314. https:\/\/doi.org\/10.3390\/ijms20071675 (2019).","journal-title":"Int. J. Mol. Sci."},{"key":"95236_CR36","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1016\/j.nima.2013.01.057","volume":"718","author":"S Herrmann","year":"2013","unstructured":"Herrmann, S. et al. CSPAD-140k: A versatile detector for LCLS experiments. Nucl. Instruments Methods Phys. Res. Sect. A Accel. Spectrometers, Detect. Assoc. Equip. 718, 550\u2013553. https:\/\/doi.org\/10.1016\/j.nima.2013.01.057 (2013).","journal-title":"Nucl. Instruments Methods Phys. Res. Sect. A Accel. Spectrometers, Detect. Assoc. Equip."},{"key":"95236_CR37","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1038\/nmeth.3667","volume":"13","author":"RG Sierra","year":"2016","unstructured":"Sierra, R. G. et al. Concentric-flow electrokinetic injector enables serial crystallography of ribosome and photosystem-II. Nat. Methods 13, 59\u201362. https:\/\/doi.org\/10.1038\/nmeth.3667 (2016).","journal-title":"Nat. Methods"},{"key":"95236_CR38","doi-asserted-by":"publisher","first-page":"672","DOI":"10.1107\/S1600576716004349","volume":"49","author":"D Damiani","year":"2016","unstructured":"Damiani, D. et al. Linac coherent light source data analysis using psana. J. Appl. Crystallogr. 49, 672\u2013679. https:\/\/doi.org\/10.1107\/S1600576716004349 (2016).","journal-title":"J. Appl. Crystallogr."},{"key":"95236_CR39","doi-asserted-by":"publisher","DOI":"10.1186\/s40679-016-0037-7","author":"J Thayer","year":"2017","unstructured":"Thayer, J. et al. Data systems for the Linac coherent light source. Adv. Struct. Chem. Imaging https:\/\/doi.org\/10.1186\/s40679-016-0037-7 (2017).","journal-title":"Adv. Struct. Chem. Imaging"},{"key":"95236_CR40","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1107\/S0021889812002312","volume":"45","author":"TA White","year":"2012","unstructured":"White, T. A. et al. CrystFEL: A software suite for snapshot serial crystallography. J. Appl. Crystallogr. 45, 335\u2013341. https:\/\/doi.org\/10.1107\/S0021889812002312 (2012).","journal-title":"J. Appl. Crystallogr."},{"key":"95236_CR41","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1107\/S205979831801238X","volume":"75","author":"TA White","year":"2019","unstructured":"White, T. A. Processing serial crystallography data with crystFEL: a step-by-step guide. Acta Crystallogr. Sect. D Struct. Biol. 75, 219\u2013233. https:\/\/doi.org\/10.1107\/S205979831801238X (2019).","journal-title":"Acta Crystallogr. Sect. D Struct. Biol."},{"key":"95236_CR42","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.35335","author":"DL Mallery","year":"2018","unstructured":"Mallery, D. L. et al. IP6 is an HIV pocket factor that prevents capsid collapse and promotes DNA synthesis. Elife https:\/\/doi.org\/10.7554\/eLife.35335 (2018).","journal-title":"Elife"},{"key":"95236_CR43","doi-asserted-by":"publisher","DOI":"10.1002\/0471140864.ps2809s79","author":"K Huynh","year":"2015","unstructured":"Huynh, K. & Partch, C. L. Analysis of protein ligand-receptor binding by photoaffinity cross-linking. Curr. Protoc. Protein Sci. https:\/\/doi.org\/10.1002\/0471140864.ps2809s79 (2015).","journal-title":"Curr. Protoc. Protein Sci."},{"key":"95236_CR44","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1016\/S1359-0278(97)00024-2","volume":"2","author":"I Bahar","year":"1997","unstructured":"Bahar, I., Atilgan, A. R. & Erman, B. Direct evaluation of thermal fluctuations in proteins using a single-parameter harmonic potential. Fold. Des. 2, 173\u2013181. https:\/\/doi.org\/10.1016\/S1359-0278(97)00024-2 (1997).","journal-title":"Fold. Des."},{"key":"95236_CR45","doi-asserted-by":"publisher","first-page":"3090","DOI":"10.1103\/PhysRevLett.79.3090","volume":"79","author":"T Haliloglu","year":"1997","unstructured":"Haliloglu, T., Bahar, I. & Erman, B. Gaussian dynamics of folded proteins. Phys. Rev. Lett. 79, 3090\u20133093. https:\/\/doi.org\/10.1103\/PhysRevLett.79.3090 (1997).","journal-title":"Phys. Rev. Lett."},{"key":"95236_CR46","doi-asserted-by":"publisher","first-page":"1056","DOI":"10.1002\/prot.25272","volume":"85","author":"A Hacisuleyman","year":"2017","unstructured":"Hacisuleyman, A. & Erman, B. Causality, transfer entropy, and allosteric communication landscapes in proteins with harmonic interactions. Proteins Struct. Funct. Bioinforma. 85, 1056\u20131064. https:\/\/doi.org\/10.1002\/prot.25272 (2017).","journal-title":"Proteins Struct. Funct. Bioinforma."},{"key":"95236_CR47","doi-asserted-by":"publisher","first-page":"651","DOI":"10.1016\/j.str.2020.03.014","volume":"28","author":"B Acar","year":"2020","unstructured":"Acar, B. et al. Distinct allosteric networks underlie mechanistic speciation of ABC transporters. Structure 28, 651-663.e5. https:\/\/doi.org\/10.1016\/j.str.2020.03.014 (2020).","journal-title":"Structure"},{"key":"95236_CR48","doi-asserted-by":"publisher","first-page":"1101","DOI":"10.1016\/j.bpj.2015.06.004","volume":"109","author":"I Bahar","year":"2015","unstructured":"Bahar, I., Cheng, M. H., Lee, J. Y., Kaya, C. & Zhang, S. Structure-encoded global motions and their role in mediating protein-substrate interactions. Biophys. J. 109, 1101\u20131109. https:\/\/doi.org\/10.1016\/j.bpj.2015.06.004 (2015).","journal-title":"Biophys. J."},{"key":"95236_CR49","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1111\/j.1348-0421.2008.00008.x","volume":"52","author":"T Murakami","year":"2008","unstructured":"Murakami, T. Roles of the interactions between Env and Gag proteins in the HIV-1 replication cycle. Microbiol. Immunol. 52, 287\u2013295. https:\/\/doi.org\/10.1111\/j.1348-0421.2008.00008.x (2008).","journal-title":"Microbiol. Immunol."},{"key":"95236_CR50","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1080\/15476286.2018.1481696","volume":"15","author":"N Dubois","year":"2018","unstructured":"Dubois, N. et al. The C-terminal p6 domain of the HIV-1 Pr55Gag precursor is required for specific binding to the genomic RNA. RNA Biol. 15, 923\u2013936. https:\/\/doi.org\/10.1080\/15476286.2018.1481696 (2018).","journal-title":"RNA Biol."},{"key":"95236_CR51","doi-asserted-by":"publisher","first-page":"5006","DOI":"10.1039\/c4ob00350k","volume":"12","author":"H Tateishi","year":"2014","unstructured":"Tateishi, H. et al. Design and synthesis of lipid-coupled inositol 1,2,3,4,5,6-hexakisphosphate derivatives exhibiting high-affinity binding for the HIV-1 MA domain. Org. Biomol. Chem. 12, 5006\u20135022. https:\/\/doi.org\/10.1039\/c4ob00350k (2014).","journal-title":"Org. Biomol. Chem."},{"key":"95236_CR52","doi-asserted-by":"publisher","first-page":"1637","DOI":"10.1016\/j.jmb.2016.03.005","volume":"428","author":"PY Mercredi","year":"2016","unstructured":"Mercredi, P. Y. et al. Structural and molecular determinants of membrane binding by the HIV-1 matrix protein. J. Mol. Biol. 428, 1637\u20131655. https:\/\/doi.org\/10.1016\/j.jmb.2016.03.005 (2016).","journal-title":"J. Mol. Biol."},{"key":"95236_CR53","doi-asserted-by":"publisher","first-page":"434","DOI":"10.1016\/j.jmb.2008.07.027","volume":"382","author":"JS Saad","year":"2008","unstructured":"Saad, J. S. et al. Structure of the myristylated human immunodeficiency virus type 2 Matrix protein and the role of phosphatidylinositol-(4,5)- bisphosphate in membrane targeting. J. Mol. Biol. 382, 434\u2013447. https:\/\/doi.org\/10.1016\/j.jmb.2008.07.027 (2008).","journal-title":"J. Mol. Biol."},{"key":"95236_CR54","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1016\/j.jmb.2011.08.038","volume":"413","author":"LS Ehrlich","year":"2011","unstructured":"Ehrlich, L. S., Medina, G. N. & Carter, C. A. ESCRT machinery potentiates HIV-1 utilization of the PI(4,5)P 2-PLC-IP3R-Ca2+ signaling cascade. J. Mol. Biol. 413, 347\u2013358. https:\/\/doi.org\/10.1016\/j.jmb.2011.08.038 (2011).","journal-title":"J. Mol. Biol."},{"key":"95236_CR55","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.sbi.2008.02.001","volume":"18","author":"BK Ganser-Pornillos","year":"2008","unstructured":"Ganser-Pornillos, B. K., Yeager, M. & Sundquist, W. I. The structural biology of HIV assembly. Curr. Opin. Cell Biol. 18, 1\u201324. https:\/\/doi.org\/10.1016\/j.sbi.2008.02.001 (2008).","journal-title":"Curr. Opin. Cell Biol."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-95236-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-95236-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-95236-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,3]],"date-time":"2022-12-03T07:08:33Z","timestamp":1670051313000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-021-95236-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,8,4]]},"references-count":55,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["95236"],"URL":"https:\/\/doi.org\/10.1038\/s41598-021-95236-8","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-467430\/v1","asserted-by":"object"}]},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,8,4]]},"assertion":[{"value":"27 April 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"15 July 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"4 August 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":"15819"}}