{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,7]],"date-time":"2026-03-07T18:47:52Z","timestamp":1772909272161,"version":"3.50.1"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1007890","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2020,6,5]],"date-time":"2020-06-05T00:00:00Z","timestamp":1591315200000}}],"reference-count":124,"publisher":"Public Library of Science (PLoS)","issue":"5","license":[{"start":{"date-parts":[[2020,5,26]],"date-time":"2020-05-26T00:00:00Z","timestamp":1590451200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100006146","name":"Office of Naval Reactors","doi-asserted-by":"publisher","award":["ONR N00014-17-1-2628"],"award-info":[{"award-number":["ONR N00014-17-1-2628"]}],"id":[{"id":"10.13039\/100006146","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["HL083464"],"award-info":[{"award-number":["HL083464"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"name":"NIH\/National Center for Advancing Translational Sciences Clinical and Translational Science","award":["TL1 TR00113"],"award-info":[{"award-number":["TL1 TR00113"]}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"DOI":"10.1371\/journal.pcbi.1007890","type":"journal-article","created":{"date-parts":[[2020,5,26]],"date-time":"2020-05-26T13:42:53Z","timestamp":1590500573000},"page":"e1007890","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":46,"title":["Non-uniform distribution of myosin-mediated forces governs red blood cell membrane curvature through tension modulation"],"prefix":"10.1371","volume":"16","author":[{"given":"Haleh","family":"Alimohamadi","sequence":"first","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7444-9327","authenticated-orcid":true,"given":"Alyson S.","family":"Smith","sequence":"additional","affiliation":[]},{"given":"Roberta B.","family":"Nowak","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3161-8802","authenticated-orcid":true,"given":"Velia M.","family":"Fowler","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5953-4347","authenticated-orcid":true,"given":"Padmini","family":"Rangamani","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2020,5,26]]},"reference":[{"key":"pcbi.1007890.ref001","doi-asserted-by":"crossref","first-page":"696","DOI":"10.1126\/science.8171320","article-title":"Engineering cell shape and function","volume":"264","author":"R Singhvi","year":"1994","journal-title":"Science"},{"key":"pcbi.1007890.ref002","doi-asserted-by":"crossref","first-page":"1356","DOI":"10.1016\/j.cell.2013.08.026","article-title":"Decoding information in cell shape","volume":"154","author":"P Rangamani","year":"2013","journal-title":"Cell"},{"key":"pcbi.1007890.ref003","doi-asserted-by":"crossref","first-page":"486","DOI":"10.1038\/nrm4012","article-title":"Forcing cells into shape: the mechanics of actomyosin contractility","volume":"16","author":"M Murrell","year":"2015","journal-title":"Nat Rev Mol Cell Biol"},{"key":"pcbi.1007890.ref004","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1016\/j.tcb.2005.11.003","article-title":"Dynamic modes of the cortical actomyosin gel during cell locomotion and division","volume":"16","author":"E Paluch","year":"2006","journal-title":"Trends Cell Biol"},{"key":"pcbi.1007890.ref005","doi-asserted-by":"crossref","first-page":"536","DOI":"10.1016\/j.tcb.2012.07.001","article-title":"Actin cortex mechanics and cellular morphogenesis","volume":"22","author":"G Salbreux","year":"2012","journal-title":"Trends Cell Biol"},{"key":"pcbi.1007890.ref006","doi-asserted-by":"crossref","first-page":"778","DOI":"10.1038\/nrm2786","article-title":"Non-muscle myosin II takes centre stage in cell adhesion and migration","volume":"10","author":"M Vicente-Manzanares","year":"2009","journal-title":"Nat Rev Mol Cell Biol"},{"key":"pcbi.1007890.ref007","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s00018-012-1002-9","article-title":"Nonmuscle myosin-2: mix and match","volume":"70","author":"SM Heissler","year":"2013","journal-title":"Cell Mol Life Sci"},{"key":"pcbi.1007890.ref008","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1242\/jcs.007112","article-title":"Nonmuscle myosin II moves in new directions","volume":"121","author":"MA Conti","year":"2008","journal-title":"J Cell Sci"},{"key":"pcbi.1007890.ref009","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1016\/j.tcb.2018.09.006","article-title":"Principles of Actomyosin Regulation In Vivo","volume":"29","author":"P Agarwal","year":"2019","journal-title":"Trends Cell Biol"},{"key":"pcbi.1007890.ref010","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1016\/j.ceb.2011.06.002","article-title":"Dynamics of actomyosin contractile activity during epithelial morphogenesis","volume":"23","author":"N Gorfinkiel","year":"2011","journal-title":"Curr Opin Cell Biol"},{"key":"pcbi.1007890.ref011","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1016\/j.ceb.2010.10.014","article-title":"Dynamics and regulation of contractile actin-myosin networks in morphogenesis","volume":"23","author":"KE Kasza","year":"2011","journal-title":"Curr Opin Cell Biol"},{"key":"pcbi.1007890.ref012","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1146\/annurev-cellbio-100109-104027","article-title":"Force generation, transmission, and integration during cell and tissue morphogenesis","volume":"27","author":"T Lecuit","year":"2011","journal-title":"Annu Rev Cell Dev Biol"},{"key":"pcbi.1007890.ref013","article-title":"Mechanisms of 3D cell migration","author":"KM Yamada","year":"2019","journal-title":"Nat Rev Mol Cell Biol"},{"key":"pcbi.1007890.ref014","doi-asserted-by":"crossref","first-page":"1420","DOI":"10.1172\/JCI39104","article-title":"The basics of epithelial-mesenchymal transition","volume":"119","author":"R Kalluri","year":"2009","journal-title":"J Clin Invest"},{"key":"pcbi.1007890.ref015","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1242\/bio.013409","article-title":"Changes in cell shape are correlated with metastatic potential in murine and human osteosarcomas","volume":"5","author":"SM Lyons","year":"2016","journal-title":"Biol Open"},{"key":"pcbi.1007890.ref016","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/j.actbio.2004.09.001","article-title":"Connections between single-cell biomechanics and human disease states: gastrointestinal cancer and malaria","volume":"1","author":"S Suresh","year":"2005","journal-title":"Acta Biomater"},{"key":"pcbi.1007890.ref017","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1016\/j.gene.2018.04.048","article-title":"MYH9: Structure, functions and role of non-muscle myosin IIA in human disease","volume":"664","author":"A Pecci","year":"2018","journal-title":"Gene"},{"key":"pcbi.1007890.ref018","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1016\/j.devcel.2018.11.011","article-title":"ARP3 Controls the Podocyte Architecture at the Kidney Filtration Barrier","volume":"47","author":"C Schell","year":"2018","journal-title":"Dev Cell"},{"key":"pcbi.1007890.ref019","doi-asserted-by":"crossref","first-page":"53","DOI":"10.3233\/JHD-180311","article-title":"Rac1 Activity Is Modulated by Huntingtin and Dysregulated in Models of Huntington\u2019s Disease","volume":"8","author":"A Tousley","year":"2019","journal-title":"J Huntingtons Dis"},{"key":"pcbi.1007890.ref020","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1038\/ncb3092","article-title":"Myosin II controls cellular branching morphogenesis and migration in three dimensions by minimizing cell-surface curvature","volume":"17","author":"H Elliott","year":"2015","journal-title":"Nat Cell Biol"},{"key":"pcbi.1007890.ref021","doi-asserted-by":"crossref","first-page":"382","DOI":"10.1557\/mrs2010.571","article-title":"Shape and Biomechanical Characteristics of Human Red Blood Cells in Health and Disease","volume":"35","author":"M Diez-Silva","year":"2010","journal-title":"MRS Bull"},{"key":"pcbi.1007890.ref022","volume-title":"Cell Shape: Determinants, Regulation, And Regulatory Role","author":"F Bronner","year":"2012"},{"key":"pcbi.1007890.ref023","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1146\/annurev.bb.23.060194.004035","article-title":"Mechanical properties of the red cell membrane in relation to molecular structure and genetic defects","volume":"23","author":"N Mohandas","year":"1994","journal-title":"Annu Rev Biophys Biomol Struct"},{"key":"pcbi.1007890.ref024","doi-asserted-by":"crossref","first-page":"3939","DOI":"10.1182\/blood-2008-07-161166","article-title":"Red cell membrane: past, present, and future","volume":"112","author":"N Mohandas","year":"2008","journal-title":"Blood"},{"key":"pcbi.1007890.ref025","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1146\/annurev.ph.49.030187.001141","article-title":"Red cell deformability and its relevance to blood flow","volume":"49","author":"S Chien","year":"1987","journal-title":"Annu Rev Physiol"},{"key":"pcbi.1007890.ref026","first-page":"39","volume-title":"Current Topics in Membranes","author":"VM Fowler","year":"2013"},{"key":"pcbi.1007890.ref027","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1042\/bj1980001","article-title":"The red cell membrane and its cytoskeleton","volume":"198","author":"WB Gratzer","year":"1981","journal-title":"Biochem J"},{"key":"pcbi.1007890.ref028","doi-asserted-by":"crossref","first-page":"206","DOI":"10.1097\/MOH.0000000000000227","article-title":"Feisty filaments: actin dynamics in the red blood cell membrane skeleton","volume":"23","author":"DS Gokhin","year":"2016","journal-title":"Curr Opin Hematol"},{"key":"pcbi.1007890.ref029","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1097\/00062752-200003000-00008","article-title":"New insights into erythrocyte membrane organization and microelasticity","volume":"7","author":"DE Discher","year":"2000","journal-title":"Curr Opin Hematol"},{"key":"pcbi.1007890.ref030","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1016\/S0022-5193(70)80032-7","article-title":"The minimum energy of bending as a possible explanation of the biconcave shape of the human red blood cell","volume":"26","author":"PB Canham","year":"1970","journal-title":"J Theor Biol"},{"key":"pcbi.1007890.ref031","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1515\/znc-1973-11-1209","article-title":"Elastic properties of lipid bilayers: theory and possible experiments","volume":"28","author":"W Helfrich","year":"1973","journal-title":"Z Naturforsch C"},{"key":"pcbi.1007890.ref032","doi-asserted-by":"crossref","first-page":"1335","DOI":"10.1051\/jphys:0197600370110133500","article-title":"The curvature elasticity of fluid membranes: A catalogue of vesicle shapes","volume":"37","author":"HJ Deuling","year":"1976","journal-title":"Journal de Physique"},{"key":"pcbi.1007890.ref033","doi-asserted-by":"crossref","first-page":"861","DOI":"10.1016\/S0006-3495(76)85736-0","article-title":"Red blood cell shapes as explained on the basis of curvature elasticity","volume":"16","author":"HJ Deuling","year":"1976","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref034","doi-asserted-by":"crossref","first-page":"4457","DOI":"10.1073\/pnas.71.11.4457","article-title":"Biological membranes as bilayer couples. A molecular mechanism of drug-erythrocyte interactions","volume":"71","author":"MP Sheetz","year":"1974","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref035","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1007\/s002490050140","article-title":"Amphiphile induced echinocyte-spheroechinocyte transformation of red blood cell shape","volume":"27","author":"A Igli\u010d","year":"1998","journal-title":"Eur Biophys J"},{"key":"pcbi.1007890.ref036","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0006-3495(81)84713-3","article-title":"Lateral organization of membranes and cell shapes","volume":"36","author":"VS Markin","year":"1981","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref037","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1051\/jphys:01986004703050700","article-title":"Curvature instability in membranes","volume":"47","author":"S Leibler","year":"1986","journal-title":"Journal de Physique"},{"key":"pcbi.1007890.ref038","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1209\/0295-5075\/19\/1\/010","article-title":"Equilibrium shape of two-component unilamellar membranes and vesicles","volume":"19","author":"D Andelman","year":"1992","journal-title":"EPL"},{"key":"pcbi.1007890.ref039","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1007\/BF00180372","article-title":"Shapes of bilayer vesicles with membrane embedded molecules","volume":"24","author":"V Kralj-Igli\u010d","year":"1996","journal-title":"Eur Biophys J"},{"key":"pcbi.1007890.ref040","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.febslet.2004.07.085","article-title":"Coupling between vesicle shape and the non-homogeneous lateral distribution of membrane constituents in Golgi bodies","volume":"574","author":"A Igli\u010d","year":"2004","journal-title":"FEBS Lett"},{"key":"pcbi.1007890.ref041","doi-asserted-by":"crossref","first-page":"5177","DOI":"10.1073\/pnas.1619264114","article-title":"Golgi apparatus self-organizes into the characteristic shape via postmitotic reassembly dynamics","volume":"114","author":"M Tachikawa","year":"2017","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref042","doi-asserted-by":"crossref","first-page":"062715","DOI":"10.1103\/PhysRevE.89.062715","article-title":"Anisotropic spontaneous curvatures in lipid membranes","volume":"89","author":"N Walani","year":"2014","journal-title":"Phys Rev E Stat Nonlin Soft Matter Phys"},{"key":"pcbi.1007890.ref043","doi-asserted-by":"crossref","first-page":"5319","DOI":"10.1039\/C8SM02356E","article-title":"Theoretical study of vesicle shapes driven by coupling curved proteins and active cytoskeletal forces","volume":"15","author":"M Fo\u0161nari\u010d","year":"2019","journal-title":"Soft Matter"},{"key":"pcbi.1007890.ref044","article-title":"Guided by curvature: shaping cells by coupling curved membrane proteins and cytoskeletal forces","volume":"373","author":"NS Gov","year":"2018","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"pcbi.1007890.ref045","doi-asserted-by":"crossref","first-page":"13356","DOI":"10.1073\/pnas.1311827110","article-title":"Lipid bilayer and cytoskeletal interactions in a red blood cell","volume":"110","author":"Z Peng","year":"2013","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref046","doi-asserted-by":"crossref","first-page":"1289","DOI":"10.1073\/pnas.0910785107","article-title":"Metabolic remodeling of the human red blood cell membrane","volume":"107","author":"Y Park","year":"2010","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref047","doi-asserted-by":"crossref","first-page":"4937","DOI":"10.1073\/pnas.0700257104","article-title":"Cytoskeletal dynamics of human erythrocyte","volume":"104","author":"J Li","year":"2007","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref048","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/S0021-9290(96)00100-5","article-title":"A possible mechanism determining the stability of spiculated red blood cells","volume":"30","author":"A Iglic","year":"1997","journal-title":"J Biomech"},{"key":"pcbi.1007890.ref049","doi-asserted-by":"crossref","first-page":"1756","DOI":"10.1016\/S0006-3495(02)75527-6","article-title":"Echinocyte shapes: bending, stretching, and shear determine spicule shape and spacing","volume":"82","author":"R Mukhopadhyay","year":"2002","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref050","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1083\/jcb.100.1.47","article-title":"Human erythrocyte myosin: identification and purification","volume":"100","author":"VM Fowler","year":"1985","journal-title":"J Cell Biol"},{"key":"pcbi.1007890.ref051","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1016\/S0021-9258(18)89689-2","article-title":"Myosin from human erythrocytes","volume":"260","author":"AJ Wong","year":"1985","journal-title":"J Biol Chem"},{"key":"pcbi.1007890.ref052","doi-asserted-by":"crossref","first-page":"1642","DOI":"10.1021\/bi00430a032","article-title":"Correlation of enzymatic properties and conformation of bovine erythrocyte myosin","volume":"28","author":"M Higashihara","year":"1989","journal-title":"Biochemistry"},{"key":"pcbi.1007890.ref053","doi-asserted-by":"crossref","first-page":"E4377","DOI":"10.1073\/pnas.1718285115","article-title":"Myosin IIA interacts with the spectrin-actin membrane skeleton to control red blood cell membrane curvature and deformability","volume":"115","author":"AS Smith","year":"2018","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref054","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1016\/S0006-3495(00)76295-3","article-title":"Effect of chain length and unsaturation on elasticity of lipid bilayers","volume":"79","author":"W Rawicz","year":"2000","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref055","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1007\/s002050050183","article-title":"Fluid Films with Curvature Elasticity","volume":"150","author":"DJ Steigmann","year":"1999","journal-title":"Arch Ration Mech Anal"},{"key":"pcbi.1007890.ref056","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1007\/s10237-012-0447-y","article-title":"Interaction between surface shape and intra-surface viscous flow on lipid membranes","volume":"12","author":"P Rangamani","year":"2013","journal-title":"Biomech Model Mechanobiol"},{"key":"pcbi.1007890.ref057","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1016\/j.bpj.2014.06.010","article-title":"Protein-induced membrane curvature alters local membrane tension","volume":"107","author":"P Rangamani","year":"2014","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref058","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1137\/0132063","article-title":"The Equations of Mechanical Equilibrium of a Model Membrane","volume":"32","author":"J Jenkins","year":"1977","journal-title":"SIAM J Appl Math"},{"key":"pcbi.1007890.ref059","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1007\/BF00275981","article-title":"Static equilibrium configurations of a model red blood cell","volume":"4","author":"JT Jenkins","year":"1977","journal-title":"J Math Biol"},{"key":"pcbi.1007890.ref060","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1016\/0021-9290(80)90184-0","article-title":"Equilibrium shapes of red blood cells in osmotic swelling","volume":"13","author":"BK Pai","year":"1980","journal-title":"J Biomech"},{"key":"pcbi.1007890.ref061","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1002\/9783527623372.ch2a","volume-title":"Soft Matter","author":"H. W. G Lim","year":"2008"},{"key":"pcbi.1007890.ref062","first-page":"171","article-title":"Red blood cell deformability, membrane material properties and shape: regulation by transmembrane, skeletal and cytosolic proteins and lipids","volume":"30","author":"N Mohandas","year":"1993","journal-title":"Semin Hematol"},{"key":"pcbi.1007890.ref063","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/S0006-3495(68)86484-7","article-title":"Theory of the sphering of red blood cells","volume":"8","author":"YC Fung","year":"1968","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref064","doi-asserted-by":"crossref","first-page":"34257","DOI":"10.1038\/srep34257","article-title":"Measuring cell surface area and deformability of individual human red blood cells over blood storage using quantitative phase imaging","volume":"6","author":"H Park","year":"2016","journal-title":"Sci Rep"},{"key":"pcbi.1007890.ref065","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1137\/0142026","article-title":"A Method for the Calculation of Vesicle Shapes","volume":"42","author":"J Luke","year":"1982","journal-title":"SIAM J Appl Math"},{"key":"pcbi.1007890.ref066","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1007\/BF00257107","article-title":"Membrane bending energy and shape determination of phospholipid vesicles and red blood cells","volume":"17","author":"S Svetina","year":"1989","journal-title":"Eur Biophys J"},{"key":"pcbi.1007890.ref067","doi-asserted-by":"crossref","first-page":"E1423","DOI":"10.1073\/pnas.1418491112","article-title":"Endocytic proteins drive vesicle growth via instability in high membrane tension environment","volume":"112","author":"N Walani","year":"2015","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref068","doi-asserted-by":"crossref","first-page":"238101","DOI":"10.1103\/PhysRevLett.88.238101","article-title":"Formation and interaction of membrane tubes","volume":"88","author":"I Der\u00e9nyi","year":"2002","journal-title":"Phys Rev Lett"},{"key":"pcbi.1007890.ref069","doi-asserted-by":"crossref","first-page":"120","DOI":"10.3390\/biom8040120","article-title":"Modeling Membrane Curvature Generation due to Membrane\u2013Protein Interactions","volume":"8","author":"H Alimohamadi","year":"2018","journal-title":"Biomolecules"},{"key":"pcbi.1007890.ref070","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/S0006-3495(64)86773-4","article-title":"MECHANICAL PROPERTIES OF THE RED CELL MEMBRANE. I. MEMBRANE STIFFNESS AND INTRACELLULAR PRESSURE","volume":"4","author":"RP Rand","year":"1964","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref071","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1007\/s10237-008-0143-0","article-title":"Modeling protein-mediated morphology in biomembranes","volume":"8","author":"A Agrawal","year":"2009","journal-title":"Biomech Model Mechanobiol"},{"key":"pcbi.1007890.ref072","author":"H Alimohamadi","journal-title":"Modeling membrane nanotube morphology: the role of heterogeneity in composition and material properties"},{"key":"pcbi.1007890.ref073","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1161\/01.RES.22.3.405","article-title":"Distribution of size and shape in populations of normal human red cells","volume":"22","author":"PB Canham","year":"1968","journal-title":"Circ Res"},{"key":"pcbi.1007890.ref074","doi-asserted-by":"crossref","first-page":"35","DOI":"10.3233\/BIR-1971-8106","article-title":"Comparative hemorheology\u2014hematological implications of species differences in blood viscosity","volume":"8","author":"S Chien","year":"1971","journal-title":"Biorheology"},{"key":"pcbi.1007890.ref075","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1016\/0026-2862(72)90069-6","article-title":"Improved measurements of the erythrocyte geometry","volume":"4","author":"E Evans","year":"1972","journal-title":"Microvasc Res"},{"key":"pcbi.1007890.ref076","doi-asserted-by":"crossref","first-page":"369","DOI":"10.3233\/BIR-1981-183-606","article-title":"High-resolution data on the geometry of red blood cells","volume":"18","author":"YC Fung","year":"1981","journal-title":"Biorheology"},{"key":"pcbi.1007890.ref077","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/0165-022X(85)90047-8","article-title":"Measurement of biophysical properties of red blood cells by resistance pulse spectroscopy: volume, shape, surface area, and deformability","volume":"11","author":"GV Richieri","year":"1985","journal-title":"J Biochem Biophys Methods"},{"key":"pcbi.1007890.ref078","doi-asserted-by":"crossref","first-page":"688","DOI":"10.1203\/00006450-199311000-00024","article-title":"Mechanical and geometrical properties of density-separated neonatal and adult erythrocytes","volume":"34","author":"O Linderkamp","year":"1993","journal-title":"Pediatr Res"},{"key":"pcbi.1007890.ref079","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1002\/cyto.990170402","article-title":"Optical and mathematical corrections of micropipette measurements of red blood cell geometry during anisotonic perifusion","volume":"17","author":"KG Engstr\u00f6m","year":"1994","journal-title":"Cytometry"},{"key":"pcbi.1007890.ref080","doi-asserted-by":"crossref","first-page":"1355","DOI":"10.1364\/AO.24.001355","article-title":"Flow-cytometric light scattering measurement of red blood cell volume and hemoglobin concentration","volume":"24","author":"DH Tycko","year":"1985","journal-title":"Appl Opt"},{"key":"pcbi.1007890.ref081","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1007\/978-1-4020-5502-7_8","volume-title":"OPTICS OF ERYTHROCYTES. Optics of Biological Particles","author":"P Tarasov","year":"2007"},{"key":"pcbi.1007890.ref082","article-title":"Computational study of scattering from healthy and diseased red blood cells","author":"\u00d6 Erg\u00fcl","year":"2010","journal-title":"J Biomed Opt"},{"key":"pcbi.1007890.ref083","first-page":"103","article-title":"Computational analysis of ultrastructural images of red blood cells","volume":"1","author":"M Das","year":"2015","journal-title":"Journal of Oncology Translational Research"},{"key":"pcbi.1007890.ref084","doi-asserted-by":"crossref","first-page":"81","DOI":"10.2170\/jjphysiol.5.81","article-title":"Contributions on the shapes of red blood corpuscles","volume":"5","author":"H Funaki","year":"1955","journal-title":"Jpn J Physiol"},{"key":"pcbi.1007890.ref085","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1006\/bulm.1998.0064","article-title":"Parametric-equation representation of biconcave erythrocytes","volume":"61","author":"PW Kuchel","year":"1999","journal-title":"Bull Math Biol"},{"key":"pcbi.1007890.ref086","unstructured":"Yurkin MA. Discrete dipole simulations of light scattering by blood cells. Universiteit van Amsterdam. 2007. Available: https:\/\/dare.uva.nl\/search?identifier=2df63a32-fc98-479c-8bfc-0cb95163574b"},{"key":"pcbi.1007890.ref087","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1117\/1.429883","article-title":"Scattering of light by a red blood cell","volume":"3","author":"AG Borovoi","year":"1998","journal-title":"J Biomed Opt"},{"key":"pcbi.1007890.ref088","doi-asserted-by":"crossref","first-page":"55001","DOI":"10.1117\/1.JBO.18.5.055001","article-title":"Modeling of light scattering by biconcave and deformed red blood cells with the invariant imbedding T-matrix method","volume":"18","author":"L Bi","year":"2013","journal-title":"J Biomed Opt"},{"key":"pcbi.1007890.ref089","doi-asserted-by":"crossref","first-page":"5249","DOI":"10.1364\/AO.44.005249","article-title":"Experimental and theoretical study of light scattering by individual mature red blood cells by use of scanning flow cytometry and a discrete dipole approximation","volume":"44","author":"MA Yurkin","year":"2005","journal-title":"Appl Opt"},{"key":"pcbi.1007890.ref090","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1016\/j.crme.2018.04.015","article-title":"Hybrid continuum\u2013coarse-grained modeling of erythrocytes","volume":"346","author":"J Lyu","year":"2018","journal-title":"Comptes Rendus M\u00e9canique"},{"key":"pcbi.1007890.ref091","doi-asserted-by":"crossref","DOI":"10.1201\/9781351074339","author":"EA Evans","year":"2018","journal-title":"Mechanics and Thermodynamics of Biomembranes: 0"},{"key":"pcbi.1007890.ref092","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.12088","article-title":"Mechanical sensitivity of Piezo1 ion channels can be tuned by cellular membrane tension","volume":"4","author":"AH Lewis","year":"2015","journal-title":"Elife"},{"key":"pcbi.1007890.ref093","doi-asserted-by":"crossref","first-page":"4134","DOI":"10.1529\/biophysj.107.117952","article-title":"Fluctuations of the red blood cell membrane: relation to mechanical properties and lack of ATP dependence","volume":"94","author":"J Evans","year":"2008","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref094","doi-asserted-by":"crossref","first-page":"1769","DOI":"10.1016\/j.cell.2018.09.054","article-title":"Cell Membranes Resist Flow","volume":"175","author":"Z Shi","year":"2018","journal-title":"Cell"},{"key":"pcbi.1007890.ref095","doi-asserted-by":"crossref","DOI":"10.1038\/nmeth.f.388","article-title":"The Airyscan detector from ZEISS: confocal imaging with improved signal-to-noise ratio and super-resolution","volume":"12","author":"J Huff","year":"2015","journal-title":"Nat Methods"},{"key":"pcbi.1007890.ref096","doi-asserted-by":"crossref","first-page":"218101","DOI":"10.1103\/PhysRevLett.97.218101","article-title":"Optical measurement of cell membrane tension","volume":"97","author":"G Popescu","year":"2006","journal-title":"Phys Rev Lett"},{"key":"pcbi.1007890.ref097","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.tcb.2012.09.006","article-title":"Use the force: membrane tension as an organizer of cell shape and motility","volume":"23","author":"A Diz-Mu\u00f1oz","year":"2013","journal-title":"Trends Cell Biol"},{"key":"pcbi.1007890.ref098","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1038\/378209a0","article-title":"Movement and force produced by a single myosin head","volume":"378","author":"JE Molloy","year":"1995","journal-title":"Nature"},{"key":"pcbi.1007890.ref099","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1080\/14786442708674422","article-title":"XXVI. Notice of some miscroscopic observations of the blood and animal tissues","volume":"2","author":"Lister JJ Hodgkin","year":"1827","journal-title":"Philos Mag"},{"key":"pcbi.1007890.ref100","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s00249-003-0337-4","article-title":"Elastic properties of the red blood cell membrane that determine echinocyte deformability","volume":"33","author":"D Kuzman","year":"2004","journal-title":"Eur Biophys J"},{"key":"pcbi.1007890.ref101","doi-asserted-by":"crossref","first-page":"031904","DOI":"10.1103\/PhysRevE.81.031904","article-title":"Multiscale simulation of erythrocyte membranes","volume":"81","author":"Z Peng","year":"2010","journal-title":"Phys Rev E Stat Nonlin Soft Matter Phys"},{"key":"pcbi.1007890.ref102","doi-asserted-by":"crossref","first-page":"7804","DOI":"10.1073\/pnas.1606751113","article-title":"Biomechanics of red blood cells in human spleen and consequences for physiology and disease","volume":"113","author":"IV Pivkin","year":"2016","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref103","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1017\/jfm.2014.14","article-title":"Erythrocyte responses in low-shear-rate flows: effects of non-biconcave stress-free state in the cytoskeleton","volume":"742","author":"Z Peng","year":"2014","journal-title":"J Fluid Mech"},{"key":"pcbi.1007890.ref104","doi-asserted-by":"crossref","first-page":"1859","DOI":"10.1529\/biophysj.104.045328","article-title":"Red blood cell membrane fluctuations and shape controlled by ATP-induced cytoskeletal defects","volume":"88","author":"NS Gov","year":"2005","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref105","doi-asserted-by":"crossref","first-page":"19742","DOI":"10.1038\/s41598-019-56128-0","article-title":"Normal red blood cells\u2019 shape stabilized by membrane's in-plane ordering","volume":"9","author":"L Mesarec","year":"2019","journal-title":"Sci Rep"},{"key":"pcbi.1007890.ref106","doi-asserted-by":"crossref","first-page":"1118","DOI":"10.1038\/s41557-018-0127-3","article-title":"A fluorescent membrane tension probe","volume":"10","author":"A Colom","year":"2018","journal-title":"Nat Chem"},{"key":"pcbi.1007890.ref107","doi-asserted-by":"crossref","first-page":"4107","DOI":"10.1529\/biophysj.104.056846","article-title":"Dynamics of myosin-driven skeletal muscle contraction: I. Steady-state force generation","volume":"88","author":"G Lan","year":"2005","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref108","doi-asserted-by":"crossref","first-page":"1348","DOI":"10.1016\/S0006-3495(96)79336-0","article-title":"What is the surface tension of a lipid bilayer membrane?","volume":"71","author":"F J\u00e4hnig","year":"1996","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref109","first-page":"175","article-title":"Membrane skeletal dynamics: role in modulation of red cell deformability, mobility of transmembrane proteins, and shape","volume":"20","author":"MP Sheetz","year":"1983","journal-title":"Semin Hematol"},{"key":"pcbi.1007890.ref110","doi-asserted-by":"crossref","first-page":"26326","DOI":"10.1074\/jbc.M110.123851","article-title":"Unconventional processive mechanics of non-muscle myosin IIB","volume":"285","author":"MF Norstrom","year":"2010","journal-title":"J Biol Chem"},{"key":"pcbi.1007890.ref111","doi-asserted-by":"crossref","first-page":"20554","DOI":"10.1038\/srep20554","article-title":"Load-dependent modulation of non-muscle myosin-2A function by tropomyosin 4.2","volume":"6","author":"N Hundt","year":"2016","journal-title":"Sci Rep"},{"key":"pcbi.1007890.ref112","doi-asserted-by":"crossref","first-page":"14847","DOI":"10.1073\/pnas.1615452113","article-title":"Biconcave shape of human red-blood-cell ghosts relies on density differences between the rim and dimple of the ghost\u2019s plasma membrane","volume":"113","author":"JF Hoffman","year":"2016","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref113","doi-asserted-by":"crossref","first-page":"1641","DOI":"10.1073\/pnas.1721463115","article-title":"Evidence that asymmetry of the membrane\/cytoskeletal complex in human red blood cell ghosts is responsible for their biconcave shape","volume":"115","author":"JF Hoffman","year":"2018","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1007890.ref114","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1016\/j.bpj.2018.11.3130","article-title":"A Model of Piezo1-Based Regulation of Red Blood Cell Volume","volume":"116","author":"S Svetina","year":"2019","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref115","doi-asserted-by":"crossref","first-page":"2024","DOI":"10.1091\/mbc.E18-02-0087","article-title":"The role of traction in membrane curvature generation","volume":"29","author":"H Alimohamadi","year":"2018","journal-title":"Mol Biol Cell"},{"key":"pcbi.1007890.ref116","doi-asserted-by":"crossref","first-page":"188101","DOI":"10.1103\/PhysRevLett.103.188101","article-title":"Why do red blood cells have asymmetric shapes even in a symmetric flow?","volume":"103","author":"B Kaoui","year":"2009","journal-title":"Phys Rev Lett"},{"key":"pcbi.1007890.ref117","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1039\/C9SM01494B","article-title":"A mechanical model reveals that non-axisymmetric buckling lowers the energy barrier associated with membrane neck constriction","volume":"16","author":"R Vasan","year":"2020","journal-title":"Soft Matter"},{"key":"pcbi.1007890.ref118","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1007\/s10237-015-0721-x","article-title":"A novel strain energy relationship for red blood cell membrane skeleton based on spectrin stiffness and its application to micropipette deformation","volume":"15","author":"S Svetina","year":"2016","journal-title":"Biomech Model Mechanobiol"},{"key":"pcbi.1007890.ref119","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1007\/s002490050140","article-title":"Amphiphile induced echinocyte\u2014spheroechinocyte red blood cell shape transformation","volume":"27","author":"A Iglic","year":"1998","journal-title":"Eur Biophys J"},{"key":"pcbi.1007890.ref120","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1016\/S0006-3495(95)79899-X","article-title":"Depletion of membrane skeleton in red blood cell vesicles","volume":"69","author":"A Iglic","year":"1995","journal-title":"Biophys J"},{"key":"pcbi.1007890.ref121","doi-asserted-by":"crossref","first-page":"051906","DOI":"10.1103\/PhysRevE.84.051906","article-title":"Computer simulation of cytoskeleton-induced blebbing in lipid membranes","volume":"84","author":"EJ Spangler","year":"2011","journal-title":"Phys Rev E Stat Nonlin Soft Matter Phys"},{"key":"pcbi.1007890.ref122","doi-asserted-by":"crossref","first-page":"1019","DOI":"10.1006\/bulm.1999.0128","article-title":"Membrane skeleton detachment in spherical and cylindrical microexovesicles","volume":"61","author":"H H\u00e4gerstrand","year":"1999","journal-title":"Bull Math Biol"},{"key":"pcbi.1007890.ref123","doi-asserted-by":"crossref","first-page":"3380","DOI":"10.1021\/jacs.8b13189","article-title":"Mechanosensitive Fluorescent Probes to Image Membrane Tension in Mitochondria, Endoplasmic Reticulum, and Lysosomes","volume":"141","author":"A Goujon","year":"2019","journal-title":"J Am Chem Soc"},{"key":"pcbi.1007890.ref124","doi-asserted-by":"crossref","first-page":"1714","DOI":"10.1016\/j.bpj.2019.09.034","article-title":"DLITE Uses Cell-Cell Interface Movement to Better Infer Cell-Cell Tensions","volume":"117","author":"R Vasan","year":"2019","journal-title":"Biophys J"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1007890","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2020,6,5]],"date-time":"2020-06-05T00:00:00Z","timestamp":1591315200000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1007890","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,3,17]],"date-time":"2021-03-17T00:40:04Z","timestamp":1615941604000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1007890"}},"subtitle":[],"editor":[{"given":"Arne","family":"Elofsson","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2020,5,26]]},"references-count":124,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2020,5,26]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1007890","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/668582","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,5,26]]}}}