{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T04:18:34Z","timestamp":1779337114169,"version":"3.51.4"},"reference-count":67,"publisher":"Public Library of Science (PLoS)","issue":"10","license":[{"start":{"date-parts":[[2015,10,27]],"date-time":"2015-10-27T00:00:00Z","timestamp":1445904000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"DOI":"10.1371\/journal.pcbi.1004450","type":"journal-article","created":{"date-parts":[[2015,10,27]],"date-time":"2015-10-27T18:37:35Z","timestamp":1445971055000},"page":"e1004450","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":46,"title":["Parsimonious Model of Vascular Patterning Links Transverse Hormone Fluxes to Lateral Root Initiation: Auxin Leads the Way, while Cytokinin Levels Out"],"prefix":"10.1371","volume":"11","author":[{"given":"Sedeer","family":"el-Showk","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hanna","family":"Help-Rinta-Rahko","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tiina","family":"Blomster","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Riccardo","family":"Siligato","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Athanasius F. M.","family":"Mar\u00e9e","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ari Pekka","family":"M\u00e4h\u00f6nen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ver\u00f4nica A.","family":"Grieneisen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2015,10,27]]},"reference":[{"key":"ref1","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1242\/dev.119.1.71","article-title":"Cellular organisation of the Arabidopsis thaliana root","volume":"119","author":"L Dolan","year":"1993","journal-title":"Development"},{"issue":"23","key":"ref2","doi-asserted-by":"crossref","first-page":"2938","DOI":"10.1101\/gad.189200","article-title":"A novel two-component hybrid molecule regulates vascular morphogenesis of the Arabidopsis root","volume":"14","author":"AP M\u00e4h\u00f6nen","year":"2000","journal-title":"Genes & Development"},{"issue":"11","key":"ref3","doi-asserted-by":"crossref","first-page":"917","DOI":"10.1016\/j.cub.2011.04.017","article-title":"A mutually inhibitory interaction between auxin and cytokinin specifies vascular pattern in roots","volume":"21","author":"A Bishopp","year":"2011","journal-title":"Current Biology"},{"issue":"2","key":"ref4","doi-asserted-by":"crossref","first-page":"857","DOI":"10.1073\/pnas.1221766111","article-title":"Integration of hormonal signaling networks and mobile microRNAs is required for vascular patterning in Arabidopsis roots","volume":"111","author":"D Muraro","year":"2014","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"5","key":"ref5","doi-asserted-by":"crossref","first-page":"1002","DOI":"10.1016\/j.cell.2012.07.017","article-title":"A bistable circuit involving SCARECROW-RETINOBLASTOMA integrates cues to inform asymmetric stem cell division","volume":"150","author":"A Cruz-Ram\u00edrez","year":"2012","journal-title":"Cell"},{"issue":"9","key":"ref6","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1016\/S1360-1385(97)84623-7","article-title":"Auxin-cytokinin interactions in higher plants: old problems and new tools","volume":"2","author":"C Coenen","year":"1997","journal-title":"Trends in Plant Science"},{"key":"ref7","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1146\/annurev-cellbio-101011-155741","article-title":"Hormonal interactions in the regulation of plant development","volume":"28","author":"M Vanstraelen","year":"2012","journal-title":"Annual Review of Cell and Developmental Biology"},{"issue":"7","key":"ref8","doi-asserted-by":"crossref","first-page":"1373","DOI":"10.1242\/dev.086371","article-title":"Crossing paths: cytokinin signalling and crosstalk","volume":"140","author":"S El-Showk","year":"2013","journal-title":"Development"},{"issue":"7301","key":"ref9","doi-asserted-by":"crossref","first-page":"1089","DOI":"10.1038\/nature09126","article-title":"Hormonal control of the shoot stem-cell niche","volume":"465","author":"Z Zhao","year":"2010","journal-title":"Nature"},{"issue":"5906","key":"ref10","doi-asserted-by":"crossref","first-page":"1380","DOI":"10.1126\/science.1164147","article-title":"A genetic framework for the control of cell division and differentiation in the root meristem","volume":"322","author":"R Dello Ioio","year":"2008","journal-title":"Science"},{"issue":"4","key":"ref11","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1111\/j.1365-313X.2005.02548.x","article-title":"Hormonally controlled expression of the Arabidopsis MAX4 shoot branching regulatory gene","volume":"44","author":"K Bainbridge","year":"2005","journal-title":"The Plant Journal"},{"issue":"2","key":"ref12","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1046\/j.1365-313x.2000.00862.x","article-title":"The hormonal regulation of axillary bud growth in Arabidopsis","volume":"24","author":"SP Chatfield","year":"2000","journal-title":"The Plant Journal"},{"issue":"12","key":"ref13","doi-asserted-by":"crossref","first-page":"3889","DOI":"10.1105\/tpc.107.055863","article-title":"Cytokinins act directly on lateral root founder cells to inhibit root initiation","volume":"19","author":"L Laplaze","year":"2007","journal-title":"The Plant Cell"},{"issue":"40","key":"ref14","doi-asserted-by":"crossref","first-page":"41866","DOI":"10.1074\/jbc.M406337200","article-title":"Arabidopsis CYP735A1 and CYP735A2 encode cytokinin hydroxylases that catalyze the biosynthesis of trans-Zeatin","volume":"279","author":"K Takei","year":"2004","journal-title":"The Journal of Biological Chemistry"},{"issue":"1","key":"ref15","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1046\/j.1365-313X.2003.01945.x","article-title":"Expression of cytokinin biosynthetic isopentenyltransferase genes in Arabidopsis: tissue specificity and regulation by auxin, cytokinin, and nitrate","volume":"37","author":"K Miyawaki","year":"2004","journal-title":"The Plant Journal"},{"issue":"9","key":"ref16","doi-asserted-by":"crossref","first-page":"2956","DOI":"10.1105\/tpc.110.074856","article-title":"Cytokinin regulation of auxin synthesis in Arabidopsis involves a homeostatic feedback loop regulated via auxin and cytokinin signal transduction","volume":"22","author":"B Jones","year":"2010","journal-title":"The Plant Cell"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1055\/s-2006-923928","article-title":"New insights into the biology of cytokinin degradation","volume":"8","author":"T Werner","year":"2006","journal-title":"Plant Biology"},{"issue":"7290","key":"ref18","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1038\/nature08836","article-title":"MONOPTEROS controls embryonic root initiation by regulating a mobile transcription factor","volume":"464","author":"A Schlereth","year":"2010","journal-title":"Nature"},{"issue":"7198","key":"ref19","doi-asserted-by":"crossref","first-page":"1094","DOI":"10.1038\/nature06943","article-title":"Cytokinin and auxin interaction in root stem-cell specification during early embryogenesis","volume":"453","author":"B M\u00fcller","year":"2008","journal-title":"Nature"},{"issue":"2","key":"ref20","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1093\/pcp\/pcl063","article-title":"ARR1 directly activates cytokinin response genes that encode proteins with diverse regulatory functions","volume":"48","author":"M Taniguchi","year":"2007","journal-title":"Plant and Cell Physiology"},{"issue":"11","key":"ref21","doi-asserted-by":"crossref","first-page":"4284","DOI":"10.1073\/pnas.0900060106","article-title":"Cytokinin regulates root meristem activity via modulation of the polar auxin transport","volume":"106","author":"K R\u016f\u017ei\u010dka","year":"2009","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"5775","key":"ref22","doi-asserted-by":"crossref","first-page":"914","DOI":"10.1126\/science.1123542","article-title":"PIN proteins perform a rate-limiting function in cellular auxin efflux","volume":"312","author":"J Petr\u00e1sek","year":"2006","journal-title":"Science"},{"issue":"8","key":"ref23","doi-asserted-by":"crossref","first-page":"382","DOI":"10.1016\/j.tplants.2006.06.002","article-title":"Auxin transport: a field in flux","volume":"11","author":"EM Kramer","year":"2006","journal-title":"Trends in Plant Science"},{"issue":"2","key":"ref24","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1016\/j.devcel.2013.12.008","article-title":"Auxin-callose-mediated plasmodesmatal gating is essential for tropic auxin gradient formation and signaling","volume":"28","author":"X Han","year":"2014","journal-title":"Developmental Cell"},{"issue":"1","key":"ref25","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1105\/tpc.106.040782","article-title":"Interactions among PIN-FORMED and P-glycoprotein auxin transporters in Arabidopsis","volume":"19","author":"JJ Blakeslee","year":"2007","journal-title":"The Plant Cell"},{"issue":"20","key":"ref26","doi-asserted-by":"crossref","first-page":"3345","DOI":"10.1242\/dev.021071","article-title":"Interaction of PIN and PGP transport mechanisms in auxin distribution-dependent development","volume":"135","author":"J Mravec","year":"2008","journal-title":"Development"},{"issue":"2","key":"ref27","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1111\/j.1365-313X.2005.02519.x","article-title":"Cellular efflux of auxin catalyzed by the Arabidopsis MDR\/PGP transporter AtPGP1","volume":"44","author":"M Geisler","year":"2005","journal-title":"The Plant Journal"},{"issue":"11","key":"ref28","doi-asserted-by":"crossref","first-page":"2922","DOI":"10.1105\/tpc.105.035816","article-title":"PGP4, an ATP Binding Cassette P-glycoprotein, catalyzes auxin transport in Arabidopsis thaliana roots","volume":"17","author":"K Terasaka","year":"2005","journal-title":"The Plant Cell"},{"issue":"370","key":"ref29","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1093\/jexbot\/53.370.835","article-title":"The Arabidopsis dual-affinity nitrate transporter gene AtNRT1.1 (CHL1) is regulated by auxin in both shoots and roots","volume":"53","author":"FQ Guo","year":"2002","journal-title":"Journal of Experimental Botany"},{"issue":"6","key":"ref30","doi-asserted-by":"crossref","first-page":"927","DOI":"10.1016\/j.devcel.2010.05.008","article-title":"Nitrate-regulated auxin transport by NRT1.1 defines a mechanism for nutrient sensing in plants","volume":"18","author":"G Krouk","year":"2010","journal-title":"Developmental Cell"},{"issue":"7165","key":"ref31","doi-asserted-by":"crossref","first-page":"1008","DOI":"10.1038\/nature06215","article-title":"Auxin transport is sufficient to generate a maximum and gradient guiding root growth","volume":"2007","author":"VA Grieneisen","journal-title":"Nature"},{"issue":"7021","key":"ref32","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1038\/nature03184","article-title":"The PIN auxin efflux facilitator network controls growth and patterning in Arabidopsis roots","volume":"433","author":"I Blilou","year":"2005","journal-title":"Nature"},{"issue":"5775","key":"ref33","doi-asserted-by":"crossref","first-page":"883","DOI":"10.1126\/science.1121356","article-title":"Polar PIN localization directs auxin flow in plants","volume":"312","author":"J Wisniewska","year":"2006","journal-title":"Science"},{"issue":"11","key":"ref34","doi-asserted-by":"crossref","first-page":"1116","DOI":"10.1016\/j.cub.2006.04.030","article-title":"Cytokinins regulate a bidirectional phosphorelay network in Arabidopsis","volume":"16","author":"AP M\u00e4h\u00f6nen","year":"2006","journal-title":"Current Biology"},{"issue":"5757","key":"ref35","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1126\/science.1118875","article-title":"Cytokinin signaling and its inhibitor AHP6 regulate cell fate during vascular development","volume":"311","author":"AP M\u00e4h\u00f6nen","year":"2006","journal-title":"Science"},{"issue":"7296","key":"ref36","doi-asserted-by":"crossref","first-page":"316","DOI":"10.1038\/nature08977","article-title":"Cell signalling by microRNA165\/6 directs gene dose-dependent root cell fate","volume":"465","author":"A Carlsbecker","year":"2010","journal-title":"Nature"},{"issue":"32","key":"ref37","doi-asserted-by":"crossref","first-page":"13630","DOI":"10.1073\/pnas.0906416106","article-title":"Brassinosteroid signaling and auxin transport are required to establish the periodic pattern of Arabidopsis shoot vascular bundles","volume":"106","author":"M Iba\u00f1es","year":"2009","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"10","key":"ref38","doi-asserted-by":"crossref","first-page":"4002","DOI":"10.1073\/pnas.1200636109","article-title":"Cytokinin signaling as a positional cue for patterning the apical-basal axis of the growing Arabidopsis shoot meristem","volume":"109","author":"VS Chickarmane","year":"2012","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"12","key":"ref39","doi-asserted-by":"crossref","first-page":"e307","DOI":"10.1371\/journal.pbio.0060307","article-title":"Root system architecture from coupling cell shape to auxin transport","volume":"6","author":"M Laskowski","year":"2008","journal-title":"PLoS Biology"},{"issue":"6","key":"ref40","doi-asserted-by":"crossref","first-page":"788","DOI":"10.1093\/pcp\/pcj043","article-title":"Expression profiling of auxin-treated Arabidopsis roots: toward a molecular analysis of lateral root emergence","volume":"47","author":"M Laskowski","year":"2006","journal-title":"Plant and Cell Physiology"},{"issue":"6197","key":"ref41","doi-asserted-by":"crossref","first-page":"1255215","DOI":"10.1126\/science.1255215","article-title":"Integration of growth and patterning during vascular tissue formation in Arabidopsis","volume":"345","author":"B De Rybel","year":"2014","journal-title":"Science"},{"issue":"1","key":"ref42","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1137\/0103003","article-title":"The numerical solution of parabolic and elliptic differential equations","volume":"3","author":"DW Peaceman","year":"1955","journal-title":"Journal of the Society for Industrial and Applied Mathematics"},{"key":"ref43","doi-asserted-by":"crossref","first-page":"3069","DOI":"10.1105\/tpc.104.024737","article-title":"Structure-function analysis of the presumptive Arabidopsis auxin permease AUX1","volume":"16","author":"R Swarup","year":"2004","journal-title":"The Plant Cell"},{"issue":"6","key":"ref44","doi-asserted-by":"crossref","first-page":"810","DOI":"10.1101\/gad.462608","article-title":"Auxin influx carriers stabilize phyllotactic patterning","volume":"22","author":"K Bainbridge","year":"2008","journal-title":"Genes & Development"},{"issue":"3","key":"ref45","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1023\/A:1005961303167","article-title":"The Arabidopsis AUX1 gene: a model system to study mRNA processing in plants","volume":"36","author":"A Marchant","year":"1998","journal-title":"Plant Molecular Biology"},{"issue":"8","key":"ref46","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1038\/ncb1754","article-title":"The auxin influx carrier LAX3 promotes lateral root emergence","volume":"10","author":"K Swarup","year":"2008","journal-title":"Nature Cell Biology"},{"issue":"3","key":"ref47","doi-asserted-by":"crossref","first-page":"862","DOI":"10.1105\/tpc.113.119495","article-title":"Systems analysis of auxin transport in the Arabidopsis root apex","volume":"26","author":"LR Band","year":"2014","journal-title":"The Plant Cell"},{"issue":"7","key":"ref48","doi-asserted-by":"crossref","first-page":"2874","DOI":"10.1105\/tpc.112.097766","article-title":"AUX\/LAX genes encode a family of auxin influx transporters that perform distinct functions during Arabidopsis development","volume":"24","author":"B P\u00e9ret","year":"2012","journal-title":"The Plant Cell"},{"issue":"4","key":"ref49","doi-asserted-by":"crossref","first-page":"e1005183","DOI":"10.1371\/journal.pgen.1005183","article-title":"Auxin influx carriers control vascular paterning and xylem differentiation in Arabidopsis thaliana","volume":"11","author":"N F\u00e0bregas","year":"2015","journal-title":"PLoS Genetics"},{"issue":"3","key":"ref50","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1242\/dev.125.3.521","article-title":"The HOBBIT gene is required for formation of the root meristem in the Arabidopsis embryo","volume":"125","author":"V Willemsen","year":"1998","journal-title":"Development"},{"issue":"23","key":"ref51","doi-asserted-by":"crossref","first-page":"6903","DOI":"10.1093\/emboj\/17.23.6903","article-title":"AtPIN2 defines a locus of Arabidopsis for root gravitropism control","volume":"17","author":"A M\u00fcller","year":"1998","journal-title":"The EMBO Journal"},{"issue":"11","key":"ref52","doi-asserted-by":"crossref","first-page":"927","DOI":"10.1016\/j.cub.2011.04.049","article-title":"Phloem-transported cytokinin regulates polar auxin transport and maintains vascular pattern in the root meristem","volume":"21","author":"A Bishopp","year":"2011","journal-title":"Current Biology"},{"issue":"4","key":"ref53","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1104\/pp.106.083790","article-title":"How far can a molecule of weak acid travel in the apoplast or xylem?","volume":"141","author":"EM Kramer","year":"2006","journal-title":"Plant Physiology"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1186\/1752-0509-6-37","article-title":"Morphogengineering roots: comparing mechanisms of morphogen gradient formation","volume":"6","author":"VA Grieneisen","year":"2012","journal-title":"BMC Systems Biology"},{"key":"ref55","first-page":"1","article-title":"A bHLH complex controls embryonic vascular tissue establishment and indeterminate growth in Arabidopsis","volume":"24","author":"B De Rybel","year":"2013","journal-title":"Developmental Cell"},{"issue":"1","key":"ref56","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1111\/j.1744-7909.2010.00898.x","article-title":"Metabolism and long-distance translocation of cytokinins","volume":"52","author":"T Kudo","year":"2010","journal-title":"Journal of Integrative Plant Biology"},{"issue":"3","key":"ref57","doi-asserted-by":"crossref","first-page":"1066","DOI":"10.1104\/pp.112.211763","article-title":"A robust and sensitive synthetic sensor to monitor the transcriptional output of the cytokining signaling network in planta","volume":"161","author":"E Zurcher","year":"2013","journal-title":"Plant Physiology"},{"issue":"4","key":"ref58","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1242\/dev.02753","article-title":"Auxin-dependent regulation of lateral root positioning in the basal meristem of Arabidopsis","volume":"134","author":"I De Smet","year":"2007","journal-title":"Development"},{"issue":"21","key":"ref59","doi-asserted-by":"crossref","first-page":"4301","DOI":"10.1242\/dev.090548","article-title":"To branch or not to branch: the role of pre-patterning in lateral root formation","volume":"140","author":"JM Van Norman","year":"2013","journal-title":"Development"},{"issue":"25","key":"ref60","doi-asserted-by":"crossref","first-page":"8790","DOI":"10.1073\/pnas.0712307105","article-title":"Auxin acts as a local morphogenetic trigger to specify lateral root founder cells","volume":"105","author":"JG Dubrovsky","year":"2008","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"5","key":"ref61","doi-asserted-by":"crossref","first-page":"903","DOI":"10.1093\/aob\/mcj604","article-title":"Lateral root initiation in Arabidopsis: developmental window, spatial patterning, density and predictability","volume":"97","author":"JG Dubrovsky","year":"2006","journal-title":"Annals of botany"},{"issue":"1","key":"ref62","doi-asserted-by":"crossref","DOI":"10.1142\/S0219720013400106","article-title":"Mathematical modeling of auxin transport in protoxylem and protophloem of Arabidopsis thaliana root tips","volume":"11","author":"ES Novoselova","year":"2013","journal-title":"Journal of Bioinformatics and Computational Biology"},{"issue":"1","key":"ref63","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1093\/jxb\/erm171","article-title":"Auxin fluxes in the root apex co-regulate gravitropism and lateral root initiation","volume":"59","author":"M Lucas","year":"2008","journal-title":"Journal of Experimental Botany"},{"issue":"4","key":"ref64","doi-asserted-by":"crossref","first-page":"743","DOI":"10.1007\/s00285-011-0472-y","article-title":"Multiscale modelling of auxin transport in the plant-root elongation zone","volume":"65","author":"LR Band","year":"2012","journal-title":"Journal of Mathematical biology"},{"issue":"7525","key":"ref65","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1038\/nature13663","article-title":"PLETHORA gradient formation mechanism separates auxin responses","volume":"515","author":"AP M\u00e4h\u00f6nen","year":"2014","journal-title":"Nature"},{"issue":"6","key":"ref66","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1016\/j.devcel.2014.05.006","article-title":"D6 PROTEIN KINASE activates auxin transport-dependent growth and PIN-FORMED phosphorylation at the plasma membrane","volume":"29","author":"ICR Barbosa","year":"2014","journal-title":"Developmental Cell"},{"issue":"20","key":"ref67","doi-asserted-by":"crossref","first-page":"1979","DOI":"10.1016\/j.cub.2013.08.008","article-title":"Cytokinin induces cell division in the quiescent center of the Arabidopsis root apical meristem","volume":"23","author":"W Zhang","year":"2013","journal-title":"Current Biology"}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pcbi.1004450","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,5,24]],"date-time":"2022-05-24T15:11:49Z","timestamp":1653405109000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1004450"}},"subtitle":[],"editor":[{"given":"Tom","family":"Beeckman","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2015,10,27]]},"references-count":67,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2015,10,27]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1004450","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2015,10,27]]}}}