{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T10:26:49Z","timestamp":1787567209204,"version":"build-2736575974"},"reference-count":65,"publisher":"eLife Sciences Publications, Ltd","license":[{"start":{"date-parts":[[2026,8,5]],"date-time":"2026-08-05T00:00:00Z","timestamp":1785888000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,8,5]],"date-time":"2026-08-05T00:00:00Z","timestamp":1785888000000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,8,5]],"date-time":"2026-08-05T00:00:00Z","timestamp":1785888000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01AI158963"],"award-info":[{"award-number":["R01AI158963"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["R01AI158963"],"award-info":[{"award-number":["R01AI158963"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01AI097274"],"award-info":[{"award-number":["R01AI097274"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["R01AI097274"],"award-info":[{"award-number":["R01AI097274"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["P30GM140963"],"award-info":[{"award-number":["P30GM140963"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["P30GM140963"],"award-info":[{"award-number":["P30GM140963"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["P20GM103474"],"award-info":[{"award-number":["P20GM103474"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["P20GM103474"],"award-info":[{"award-number":["P20GM103474"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"name":"UM Center for Environmental Health Sciences","award":["S10OD025019-01"],"award-info":[{"award-number":["S10OD025019-01"]}]},{"award":["S10OD025019-01"],"award-info":[{"award-number":["S10OD025019-01"]}],"id":[{"id":"https:\/\/ror.org\/04rq5mt64","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":["elifesciences.org"],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>Whether infection of cells by individual virions occurs randomly, or if there is some form(s) of competition or cooperativity between individual virions, remains largely unknown for most virus\u2013cell associations. Here, we studied cooperativity\/competition for three different strains of human cytomegalovirus (HCMV) on two different cell types (fibroblasts and epithelial cells). By titrating viral inocula concentrations in small steps over several orders of magnitude, and by using flow cytometry to precisely measure the frequency of infected cells, we found that for most virus\u2013cell associations, the frequency of cell infection increases faster than linear with an increasing inoculum concentration, indicating cooperativity between individual infecting virions. Mathematical modeling suggests that this apparent cooperativity cannot be explained by heterogeneity in either the infectivity of the individual virions or the resistance of individual cells to infection, or by simple aggregation\/clumping of viral particles. Stochastic simulations of two additional alternative models that allow for (1) reduction in cell resistance to infection when exposed to multiple virions, or (2) compensation in infectivity of poorly infectious virions when coinfecting cells with more infectious virions, resulted in apparent viral cooperativity. Analysis of other published datasets suggests the presence of apparent viral cooperativity for HIV and vaccinia virus, infecting CRFK or HeLa cells, respectively, but not for tobacco mosaic virus forming plaques on plant leaves. We thus (1) propose a methodology to rigorously evaluate apparent cooperativity of viruses infecting target cells, and (2) demonstrate that knowing the degree of virus cooperativity for any given virus\u2013cell combination is important for an accurate quantification of multiplicity of infection.<\/jats:p>","DOI":"10.7554\/elife.108921","type":"journal-article","created":{"date-parts":[[2025,11,14]],"date-time":"2025-11-14T17:30:13Z","timestamp":1763141413000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.108921","source":"Crossref","is-referenced-by-count":0,"title":["Apparent cooperativity between human CMV virions introduces errors in conventional methods of calculating multiplicity of infection"],"prefix":"10.7554","volume":"14","author":[{"given":"Christopher","family":"Peterson","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0078xmk34","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Biological Studies, College of Humanities and Sciences, University of Montana","place":["Missoula, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0698-8318","authenticated-orcid":true,"given":"Joshua","family":"Miller","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0524sp257","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Geographical Sciences, University of Bristol","place":["Bristol, United Kingdom"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"given":"Brent","family":"Ryckman","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0078xmk34","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Biological Studies, College of Humanities and Sciences, University of Montana","place":["Missoula, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"corresponding-author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6572-1691","authenticated-orcid":true,"given":"Vitaly V","family":"Ganusov","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/00wbskb04","id-type":"ROR","asserted-by":"publisher"}],"name":"Host-Pathogen Interactions program, Texas Biomedical Research Institute","place":["San Antonio, United States"]},{"id":[{"id":"https:\/\/ror.org\/020f3ap87","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Microbiology, University of Tennessee","place":["Knoxville, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"corresponding-author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]}],"member":"4374","published-online":{"date-parts":[[2026,8,5]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","DOI":"10.1128\/jvi.01273-23","type":"journal-article","article-title":"All the single cells: if you like it then you should put some virus on it","volume":"98","author":"Adams","year":"2024","journal-title":"Journal of Virology"},{"key":"bib2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1008181","type":"journal-article","article-title":"Experimental determination of the force of malaria infection reveals a non-linear relationship to mosquito sporozoite loads","volume":"16","author":"Aleshnick","year":"2020","journal-title":"PLOS Pathogens"},{"key":"bib3","doi-asserted-by":"publisher","first-page":"6773","DOI":"10.1128\/JVI.00649-15","type":"journal-article","article-title":"Particle-to-PFU ratio of Ebola virus influences disease course and survival in cynomolgus macaques","volume":"89","author":"Alfson","year":"2015","journal-title":"Journal of Virology"},{"key":"bib4","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.abd4942","type":"journal-article","article-title":"Cooperative nature of viral replication","volume":"6","author":"Andreu-Moreno","year":"2020","journal-title":"Science Advances"},{"key":"bib5","doi-asserted-by":"publisher","first-page":"4465","DOI":"10.1128\/jvi.74.10.4465-4473.2000","type":"journal-article","article-title":"Expression of herpes simplex virus ICP47 and human cytomegalovirus US11 prevents recognition of transgene products by CD8(+) cytotoxic T lymphocytes","volume":"74","author":"Berger","year":"2000","journal-title":"Journal of Virology"},{"key":"bib6","doi-asserted-by":"publisher","DOI":"10.1101\/2024.12.09.627549","type":"preprint","article-title":"What Challenges Remain in Harmonizing Cytomegalovirus Viral Load Quantification across Laboratories?","volume-title":"bioRxiv","author":"Boutolleau","year":"2024"},{"key":"bib7","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1002\/rmv.655","type":"journal-article","article-title":"Review of cytomegalovirus seroprevalence and demographic characteristics associated with infection","volume":"20","author":"Cannon","year":"2010","journal-title":"Reviews in Medical Virology"},{"key":"bib8","doi-asserted-by":"publisher","first-page":"6917","DOI":"10.1128\/JVI.00081-09","type":"journal-article","article-title":"Enumeration of an extremely high particle-to-PFU ratio for Varicella-zoster virus","volume":"83","author":"Carpenter","year":"2009","journal-title":"Journal of Virology"},{"key":"bib9","doi-asserted-by":"publisher","first-page":"863","DOI":"10.1128\/JVI.73.2.863-870.1999","type":"journal-article","article-title":"Transcriptional regulation of the human cytomegalovirus US11 early gene","volume":"73","author":"Chau","year":"1999","journal-title":"Journal of Virology"},{"key":"bib10","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1009480","type":"journal-article","article-title":"Time to revisit the endpoint dilution assay and to replace the TCID50 as a measure of a virus sample\u2019s infection concentration","volume":"17","author":"Cresta","year":"2021","journal-title":"PLOS Computational Biology"},{"key":"bib11","type":"web-resource","article-title":"How many spheres can fit inside this larger sphere?","author":"D\u2019Aurizio","year":"2019","unstructured":"D\u2019Aurizio J. 2019. How many spheres can fit inside this larger sphere?. https:\/\/math.stackexchange.com\/questions\/3375911\/how-many-spheres-can-fit-inside-this-larger-sphere [Accessed July 16, 2026]."},{"key":"bib12","doi-asserted-by":"publisher","first-page":"336","DOI":"10.1002\/bdra.23601","type":"journal-article","article-title":"Cytomegalovirus infection in pregnancy","volume":"109","author":"Davis","year":"2017","journal-title":"Birth Defects Research"},{"key":"bib13","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1084\/jem.99.2.167","type":"journal-article","article-title":"Plaque formation and isolation of pure lines with poliomyelitis viruses","volume":"99","author":"Dulbecco","year":"1954","journal-title":"The Journal of Experimental Medicine"},{"key":"bib14","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1002\/biot.201400429","type":"journal-article","article-title":"Defective interfering viruses and their impact on vaccines and viral vectors","volume":"10","author":"Frensing","year":"2015","journal-title":"Biotechnology Journal"},{"key":"bib15","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1128\/jb.88.2.433-439.1964","type":"journal-article","article-title":"Relative plaque-forming, cell-infecting, and interfering qualities of vaccinia virus","volume":"88","author":"Galasso","year":"1964","journal-title":"Journal of Bacteriology"},{"key":"bib16","doi-asserted-by":"publisher","first-page":"1138","DOI":"10.1128\/jb.90.4.1138-1142.1965","type":"journal-article","article-title":"Effect of particle aggregation on the survival of irradiated vaccinia virus","volume":"90","author":"Galasso","year":"1965","journal-title":"Journal of Bacteriology"},{"key":"bib17","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms7022","type":"journal-article","article-title":"Sorting of small infectious virus particles by flow virometry reveals distinct infectivity profiles","volume":"6","author":"Gaudin","year":"2015","journal-title":"Nature Communications"},{"key":"bib18","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1159\/000150509","type":"journal-article","article-title":"Structure and assembly of the virion","volume":"39","author":"Gibson","year":"1996","journal-title":"Intervirology"},{"key":"bib19","doi-asserted-by":"publisher","first-page":"22","DOI":"10.3748\/wjg.v13.i1.22","type":"journal-article","article-title":"Viral and cellular determinants involved in hepadnaviral entry","volume":"13","author":"Glebe","year":"2007","journal-title":"World Journal of Gastroenterology"},{"key":"bib20","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1093\/oxfordjournals.aje.a113662","type":"journal-article","article-title":"Estimation of risk due to low doses of microorganisms: a comparison of alternative methodologies","volume":"118","author":"Haas","year":"1983","journal-title":"American Journal of Epidemiology"},{"key":"bib21","doi-asserted-by":"publisher","DOI":"10.1002\/9781118910030","type":"book","volume-title":"Quantitative Microbial Risk Assessment","author":"Haas","year":"2014"},{"key":"bib22","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1016\/j.bbi.2023.12.033","type":"journal-article","article-title":"Intermittent cytomegalovirus infection alters neurobiological metabolism and induces cognitive deficits in mice","volume":"117","author":"Harrison","year":"2024","journal-title":"Brain, Behavior, and Immunity"},{"key":"bib23","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1093\/cid\/ciae535","type":"journal-article","article-title":"Current and future strategies for the prevention and treatment of cytomegalovirus infections in transplantation","volume":"81","author":"Heldman","year":"2025","journal-title":"Clinical Infectious Diseases"},{"key":"bib24","doi-asserted-by":"publisher","first-page":"620","DOI":"10.1016\/0042-6822(73)90063-9","type":"journal-article","article-title":"Mechanism of influenza recombination II Virus aggregation and its effect on plaque formation by so-called noninfective virus","volume":"56","author":"Hirst","year":"1973","journal-title":"Virology"},{"key":"bib25","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1016\/0042-6822(83)90122-8","type":"journal-article","article-title":"Isolation and characterization of a noninfectious virion-like particle released from cells infected with human strains of cytomegalovirus","volume":"130","author":"Irmiere","year":"1983","journal-title":"Virology"},{"key":"bib26","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/s0065-3527(08)60597-7","type":"journal-article","article-title":"Particle counts and infectivity titrations for animal viruses","volume":"4","author":"Isaacs","year":"1957","journal-title":"Advances in Virus Research"},{"key":"bib27","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1099\/00221287-4-1-53","type":"journal-article","article-title":"Interpreting relationships between the concentrations of plant viruses and numbers of local lesions","volume":"4","author":"Kleczkowski","year":"1950","journal-title":"Journal of General Microbiology"},{"key":"bib28","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1016\/j.chom.2016.07.011","type":"journal-article","article-title":"A multicomponent animal virus isolated from mosquitoes","volume":"20","author":"Ladner","year":"2016","journal-title":"Cell Host & Microbe"},{"key":"bib29","doi-asserted-by":"publisher","first-page":"4471","DOI":"10.1073\/pnas.1419875112","type":"journal-article","article-title":"A viral regulator of glycoprotein complexes contributes to human cytomegalovirus cell tropism","volume":"112","author":"Li","year":"2015","journal-title":"PNAS"},{"key":"bib30","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1038\/s41579-018-0115-z","type":"journal-article","article-title":"Host and viral determinants of influenza A virus species specificity","volume":"17","author":"Long","year":"2019","journal-title":"Nature Reviews. Microbiology"},{"key":"bib31","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1016\/s0092-1157(85)80022-6","type":"journal-article","article-title":"The standardization of infectivity titrations of poliovaccines--a WHO collaborative study","volume":"13","author":"Magrath","year":"1985","journal-title":"Journal of Biological Standardization"},{"key":"bib32","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1016\/j.mex.2015.02.003","type":"journal-article","article-title":"A method based on light scattering to estimate the concentration of virus particles without the need for virus particle standards","volume":"2","author":"Makra","year":"2015","journal-title":"MethodsX"},{"key":"bib33","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.00571-18","type":"journal-article","article-title":"Real-time visualization and quantification of human cytomegalovirus replication in living cells using the ANCHOR DNA labeling technology","volume":"92","author":"Mariam\u00e9","year":"2018","journal-title":"Journal of Virology"},{"key":"bib34","doi-asserted-by":"publisher","first-page":"4112","DOI":"10.1128\/JVI.02489-08","type":"journal-article","article-title":"Essential role for either TRS1 or IRS1 in human cytomegalovirus replication","volume":"83","author":"Marshall","year":"2009","journal-title":"Journal of Virology"},{"key":"bib35","doi-asserted-by":"publisher","DOI":"10.1186\/s12985-021-01566-4","type":"journal-article","article-title":"The basic reproductive number and particle-to-plaque ratio: comparison of these two parameters of viral infectivity","volume":"18","author":"McCormick","year":"2021","journal-title":"Virology Journal"},{"key":"bib36","type":"software","article-title":"HCMV_Dose_Infection","volume-title":"Software Heritage","author":"Miller","year":"2026","unstructured":"Miller J. 2026. HCMV_Dose_Infection. Software Heritage. swh:1:rev:e55178c9fa9581d24f0d51030d5ee6cab7a6b752. https:\/\/archive.softwareheritage.org\/swh:1:dir:1f8a6c138e28c58c5907bc3c85e3de724f1b9074;origin=https:\/\/github.com\/Joshua-Miller-161\/HCMV_Dose_Infection;visit=swh:1:snp:4710ad3083d45b6085554e0e7793ce249153b81c;anchor=swh:1:rev:e55178c9fa9581d24f0d51030d5ee6cab7a6b752."},{"key":"bib37","volume-title":"Cytomegaloviruses and Their Replication","author":"Mocarski","year":"1996","type":"book"},{"key":"bib38","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2201787119","type":"journal-article","article-title":"Computational modeling of protracted HCMV replication using genome substrates and protein temporal profiles","volume":"119","author":"Monti","year":"2022","journal-title":"PNAS"},{"key":"bib39","doi-asserted-by":"publisher","DOI":"10.1080\/22221751.2024.2447614","type":"journal-article","article-title":"Replication kinetics, pathogenicity and virus-induced cellular responses of cattle-origin influenza A(H5N1) isolates from Texas, United States","volume":"14","author":"Mostafa","year":"2025","journal-title":"Emerging Microbes & Infections"},{"key":"bib40","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.3390\/v7031189","type":"journal-article","article-title":"Simple mathematical models do not accurately predict early SIV dynamics","volume":"7","author":"Noecker","year":"2015","journal-title":"Viruses"},{"key":"bib41","doi-asserted-by":"publisher","DOI":"10.3390\/v16030440","type":"journal-article","article-title":"Quantifying plant viruses: evolution from bioassay to infectivity dilution curves along the model of tobamoviruses","volume":"16","author":"Nourinejhad Zarghani","year":"2024","journal-title":"Viruses"},{"key":"bib42","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2408078121","type":"journal-article","article-title":"The Pentamer glycoprotein complex inhibits viral Immediate Early transcription during human cytomegalovirus infections","volume":"121","author":"Ohman","year":"2024","journal-title":"PNAS"},{"key":"bib43","doi-asserted-by":"publisher","DOI":"10.3390\/v14020438","type":"journal-article","article-title":"Viral aggregation: the knowns and unknowns","volume":"14","author":"Pradhan","year":"2022","journal-title":"Viruses"},{"key":"bib44","type":"web-resource","article-title":"The Purity of Plaques","author":"Racaniello","year":"2017","unstructured":"Racaniello V. 2017. The Purity of Plaques. https:\/\/virology.ws\/2017\/03\/30\/the-purity-ofplaques\/ [Accessed July 16, 2026]."},{"key":"bib45","doi-asserted-by":"publisher","DOI":"10.1002\/alz.14401","type":"journal-article","article-title":"Alzheimer\u2019s disease-associated CD83(+) microglia are linked with increased immunoglobulin G4 and human cytomegalovirus in the gut, vagal nerve, and brain","volume":"21","author":"Readhead","year":"2025","journal-title":"Alzheimer\u2019s & Dementia"},{"key":"bib46","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.32303","type":"journal-article","article-title":"Extreme heterogeneity of influenza virus infection in single cells","volume":"7","author":"Russell","year":"2018","journal-title":"eLife"},{"key":"bib47","doi-asserted-by":"publisher","first-page":"14118","DOI":"10.1073\/pnas.0804365105","type":"journal-article","article-title":"HCMV gH\/gL\/UL128\u2013131 interferes with virus entry into epithelial cells: Evidence for cell type-specific receptors","volume":"105","author":"Ryckman","year":"2008","journal-title":"PNAS"},{"key":"bib48","doi-asserted-by":"publisher","first-page":"9512","DOI":"10.1128\/JVI.78.17.9512-9523.2004","type":"journal-article","article-title":"Human cytomegalovirus tegument proteins ppUL82 (pp71) and ppUL35 interact and cooperatively activate the major immediate-early enhancer","volume":"78","author":"Schierling","year":"2004","journal-title":"Journal of Virology"},{"key":"bib49","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.00034-20","type":"journal-article","article-title":"Specialization for cell-free or cell-to-cell spread of BAC-cloned human cytomegalovirus strains is determined by factors beyond the UL128-131 and RL13 Loci","volume":"94","author":"Schultz","year":"2020","journal-title":"Journal of Virology"},{"key":"bib50","doi-asserted-by":"publisher","first-page":"863","DOI":"10.1088\/0022-3727\/2\/6\/311","type":"journal-article","article-title":"The density of random close packing of spheres","volume":"2","author":"Scott","year":"1969","journal-title":"Journal of Physics D"},{"key":"bib51","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.43599","type":"journal-article","article-title":"A multicellular way of life for a multipartite virus","volume":"8","author":"Sicard","year":"2019","journal-title":"eLife"},{"key":"bib52","doi-asserted-by":"publisher","first-page":"359","DOI":"10.1099\/vir.0.83286-0","type":"journal-article","article-title":"Cloning and sequencing of a highly productive, endotheliotropic virus strain derived from human cytomegalovirus TB40\/E","volume":"89","author":"Sinzger","year":"2008","journal-title":"The Journal of General Virology"},{"key":"bib53","doi-asserted-by":"publisher","DOI":"10.3390\/v16121854","type":"journal-article","article-title":"Virulence and replicative fitness of HIV-1 transmitted\/founder (T\/F) viruses harbouring drug resistance-associated mutation","volume":"16","author":"Sonawane","year":"2024","journal-title":"Viruses"},{"key":"bib54","doi-asserted-by":"publisher","first-page":"9593","DOI":"10.1021\/acs.analchem.4c01209","type":"journal-article","article-title":"Multiangle light scattering as a lentivirus purification process analytical technology","volume":"96","author":"Sripada","year":"2024","journal-title":"Analytical Chemistry"},{"key":"bib55","doi-asserted-by":"publisher","first-page":"3191","DOI":"10.1172\/JCI42955","type":"journal-article","article-title":"Reconstruction of the complete human cytomegalovirus genome in a BAC reveals RL13 to be a potent inhibitor of replication","volume":"120","author":"Stanton","year":"2010","journal-title":"The Journal of Clinical Investigation"},{"key":"bib56","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1007\/s12551-016-0218-6","type":"journal-article","article-title":"Dynamic light scattering: a practical guide and applications in biomedical sciences","volume":"8","author":"Stetefeld","year":"2016","journal-title":"Biophysical Reviews"},{"key":"bib57","doi-asserted-by":"publisher","first-page":"4219","DOI":"10.1021\/nl3018109","type":"journal-article","article-title":"Cooperative vaccinia infection demonstrated at the single-cell level using FluidFM","volume":"12","author":"Stiefel","year":"2012","journal-title":"Nano Letters"},{"key":"bib58","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1111\/j.1600-0668.2009.00621.x","type":"journal-article","article-title":"Review and comparison between the Wells-Riley and dose-response approaches to risk assessment of infectious respiratory diseases","volume":"20","author":"Sze To","year":"2010","journal-title":"Indoor Air"},{"key":"bib59","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.59988","type":"journal-article","article-title":"Mutational resilience of antiviral restriction favors primate TRIM5\u03b1 in host-virus evolutionary arms races","volume":"9","author":"Tenthorey","year":"2020","journal-title":"eLife"},{"key":"bib60","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1016\/0025-5564(75)90037-1","type":"journal-article","article-title":"Some classes of hit-theory models","volume":"23","author":"Turner","year":"1975","journal-title":"Mathematical Biosciences"},{"key":"bib61","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1128\/jb.91.1.213-220.1966","type":"journal-article","article-title":"Interactions of human cytomegalovirus with human fibroblasts","volume":"91","author":"Vonka","year":"1966","journal-title":"Journal of Bacteriology"},{"key":"bib62","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1128\/br.24.1.151-166.1960","type":"journal-article","article-title":"Viral interference. some considerations of basic mechanisms and their potential relationship to host resistance","volume":"24","author":"Wagner","year":"1960","journal-title":"Bacteriological Reviews"},{"key":"bib63","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1016\/S0092-8674(00)81054-5","type":"journal-article","article-title":"The human cytomegalovirus US11 gene product dislocates mhc class i heavy chains from the endoplasmic reticulum to the cytosol","volume":"84","author":"Wiertz","year":"1996","journal-title":"Cell"},{"key":"bib64","doi-asserted-by":"publisher","first-page":"8999","DOI":"10.1128\/JVI.01325-15","type":"journal-article","article-title":"Human cytomegalovirus gH\/gL\/gO promotes the fusion step of entry into all cell types, whereas gH\/gL\/UL128-131 broadens virus tropism through a distinct mechanism","volume":"89","author":"Zhou","year":"2015","journal-title":"Journal of Virology"},{"key":"bib65","doi-asserted-by":"publisher","DOI":"10.1002\/rmv.2034","type":"journal-article","article-title":"Estimation of the worldwide seroprevalence of cytomegalovirus: A systematic review and meta-analysis","volume":"29","author":"Zuhair","year":"2019","journal-title":"Reviews in Medical Virology"}],"container-title":["eLife"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/cdn.elifesciences.org\/articles\/108921\/elife-108921-v1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/cdn.elifesciences.org\/articles\/108921\/elife-108921-v1.xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/elifesciences.org\/articles\/108921","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T08:54:25Z","timestamp":1787561665000},"score":1,"resource":{"primary":{"URL":"https:\/\/elifesciences.org\/articles\/108921"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,5]]},"references-count":65,"alternative-id":["10.7554\/eLife.108921"],"URL":"https:\/\/doi.org\/10.7554\/elife.108921","archive":["CLOCKSS"],"relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2025.04.23.650360","asserted-by":"subject"}],"is-same-as":[{"id-type":"doi","id":"10.7554\/eLife.108921.3","asserted-by":"subject"}],"is-version-of":[{"id-type":"doi","id":"10.7554\/eLife.108921.1","asserted-by":"subject"},{"id-type":"doi","id":"10.7554\/eLife.108921.2","asserted-by":"subject"}],"is-supplemented-by":[{"id-type":"uri","id":"https:\/\/archive.softwareheritage.org\/swh:1:dir:1f8a6c138e28c58c5907bc3c85e3de724f1b9074;origin=https:\/\/github.com\/Joshua-Miller-161\/HCMV_Dose_Infection;visit=swh:1:snp:4710ad3083d45b6085554e0e7793ce249153b81c;anchor=swh:1:rev:e55178c9fa9581d24f0d51030d5ee6cab7a6b752","asserted-by":"subject"}]},"ISSN":["2050-084X"],"issn-type":[{"value":"2050-084X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,5]]},"assertion":[{"value":"single anonymised","name":"peer_review_transparency","label":"Peer review transparency","group":{"name":"peer_review_taxonomy"}},{"value":"other reviewer(s), editor","name":"peer_review_interaction","label":"Peer review interaction","group":{"name":"peer_review_taxonomy"}},{"value":"review summaries, review reports, author\/editor communication, reviewer identities reviewer opt in, editor identities","name":"peer_review_published","label":"Peer review published","group":{"name":"peer_review_taxonomy"}},{"value":"open (sign in with ORCID iD required)","name":"post_publication_commenting","label":"Post publication commenting","group":{"name":"post_publication_commenting"}}],"article-number":"RP108921"}}