{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,12,31]],"date-time":"2025-12-31T10:10:41Z","timestamp":1767175841410,"version":"build-2238731810"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1010291","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,8,16]],"date-time":"2022-08-16T00:00:00Z","timestamp":1660608000000}}],"reference-count":73,"publisher":"Public Library of Science (PLoS)","issue":"7","license":[{"start":{"date-parts":[[2022,7,27]],"date-time":"2022-07-27T00:00:00Z","timestamp":1658880000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000270","name":"natural environment research council","doi-asserted-by":"publisher","award":["NE\/P012345\/1"],"award-info":[{"award-number":["NE\/P012345\/1"]}],"id":[{"id":"10.13039\/501100000270","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000275","name":"leverhulme trust","doi-asserted-by":"publisher","award":["RF-2020-653\\2"],"award-info":[{"award-number":["RF-2020-653\\2"]}],"id":[{"id":"10.13039\/501100000275","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000266","name":"engineering and physical sciences research council","doi-asserted-by":"publisher","award":["EP\/R029423\/1"],"award-info":[{"award-number":["EP\/R029423\/1"]}],"id":[{"id":"10.13039\/501100000266","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000266","name":"engineering and physical sciences research council","doi-asserted-by":"publisher","award":["EP\/R029326\/1"],"award-info":[{"award-number":["EP\/R029326\/1"]}],"id":[{"id":"10.13039\/501100000266","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003246","name":"nederlandse organisatie voor wetenschappelijk onderzoek","doi-asserted-by":"publisher","award":["OCENW.KLEIN.085"],"award-info":[{"award-number":["OCENW.KLEIN.085"]}],"id":[{"id":"10.13039\/501100003246","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>Microbes play a primary role in aquatic ecosystems and biogeochemical cycles. Spatial patchiness is a critical factor underlying these activities, influencing biological productivity, nutrient cycling and dynamics across trophic levels. Incorporating spatial dynamics into microbial models is a long-standing challenge, particularly where small-scale turbulence is involved. Here, we combine a fully 3D direct numerical simulation of convective mixed layer turbulence, with an individual-based microbial model to test the key hypothesis that the coupling of gyrotactic motility and turbulence drives intense microscale patchiness. The fluid model simulates turbulent convection caused by heat loss through the fluid surface, for example during the night, during autumnal or winter cooling or during a cold-air outbreak. We find that under such conditions, turbulence-driven patchiness is depth-structured and requires high motility: Near the fluid surface, intense convective turbulence overpowers motility, homogenising motile and non-motile microbes approximately equally. At greater depth, in conditions analogous to a thermocline, highly motile microbes can be over twice as patch-concentrated as non-motile microbes, and can substantially amplify their swimming velocity by efficiently exploiting fast-moving packets of fluid. Our results substantiate the predictions of earlier studies, and demonstrate that turbulence-driven patchiness is not a ubiquitous consequence of motility but rather a delicate balance of motility and turbulent intensity.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1010291","type":"journal-article","created":{"date-parts":[[2022,7,27]],"date-time":"2022-07-27T13:51:09Z","timestamp":1658929869000},"page":"e1010291","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Investigating microscale patchiness of motile microbes under turbulence in a simulated convective mixed layer"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7339-1845","authenticated-orcid":true,"given":"Alexander Kier","family":"Christensen","sequence":"first","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7526-6853","authenticated-orcid":true,"given":"Matthew D.","family":"Piggott","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2041-0704","authenticated-orcid":true,"given":"Erik","family":"van Sebille","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4840-5050","authenticated-orcid":true,"given":"Maarten","family":"van Reeuwijk","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8375-5684","authenticated-orcid":true,"given":"Samraat","family":"Pawar","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2022,7,27]]},"reference":[{"issue":"25","key":"pcbi.1010291.ref001","doi-asserted-by":"crossref","first-page":"6506","DOI":"10.1073\/pnas.1711842115","article-title":"The Biomass Distribution on Earth","volume":"115","author":"YM Bar-On","year":"2018","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"6266","key":"pcbi.1010291.ref002","doi-asserted-by":"crossref","first-page":"aac8455","DOI":"10.1126\/science.aac8455","article-title":"The Global Ocean Microbiome","volume":"350","author":"MA Moran","year":"2015","journal-title":"Science"},{"issue":"5879","key":"pcbi.1010291.ref003","doi-asserted-by":"crossref","first-page":"1034","DOI":"10.1126\/science.1153213","article-title":"The Microbial Engines That Drive Earth\u2019s Biogeochemical Cycles","volume":"320","author":"PG Falkowski","year":"2008","journal-title":"Science"},{"issue":"5374","key":"pcbi.1010291.ref004","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1126\/science.281.5374.237","article-title":"Primary Production of the Biosphere: Integrating Terrestrial and Oceanic Components","volume":"281","author":"CB Field","year":"1998","journal-title":"Science"},{"key":"pcbi.1010291.ref005","doi-asserted-by":"crossref","first-page":"257","DOI":"10.3354\/meps010257","article-title":"The Ecological Role of Water-Column Microbes in the Sea","volume":"10","author":"F Azam","year":"1983","journal-title":"Marine Ecology Progress Series"},{"key":"pcbi.1010291.ref006","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1007\/978-1-4684-7609-5_3","volume-title":"Advances in Microbial Ecology","author":"HW Ducklow","year":"1992"},{"issue":"6391","key":"pcbi.1010291.ref007","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1038\/359139a0","article-title":"Intense Hydrolytic Enzyme Activity on Marine Aggregates and Implications for Rapid Particle Dissolution","volume":"359","author":"DC Smith","year":"1992","journal-title":"Nature"},{"issue":"3","key":"pcbi.1010291.ref008","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1038\/ismej.2010.141","article-title":"Quantitative Analysis of a Deeply Sequenced Marine Microbial Metatranscriptome","volume":"5","author":"SM Gifford","year":"2011","journal-title":"The ISME Journal"},{"issue":"6","key":"pcbi.1010291.ref009","doi-asserted-by":"crossref","first-page":"1133","DOI":"10.4319\/lo.1998.43.6.1133","article-title":"Resource Competition in a Discrete Environment: Why Are Plankton Distributions Paradoxical?","volume":"43","author":"DA Siegel","year":"1998","journal-title":"Limnology and Oceanography"},{"key":"pcbi.1010291.ref010","first-page":"1","article-title":"The Plankton of the South Georgia Whaling Grounds and Adjacent Waters, 1926-1927","volume":"vol. 11","author":"AC Hardy","year":"1935","journal-title":"Discovery Reports"},{"issue":"Suppl.","key":"pcbi.1010291.ref011","first-page":"7","article-title":"A Further Example of the Patchiness of Plankton Distribution","volume":"3","author":"AC Hardy","year":"1955","journal-title":"Deep Sea Research"},{"issue":"04-06","key":"pcbi.1010291.ref012","first-page":"593","article-title":"The Variance Spectrum of Phytoplankton in a Turbulent Ocean","volume":"34","author":"KL Denman","year":"1976","journal-title":"Journal of Marine Research"},{"key":"pcbi.1010291.ref013","first-page":"125","article-title":"Effects of Physical Processes on Planktonic Ecosystems in Coastal Ocean","volume":"vol. 22","author":"KL Denman","year":"1984","journal-title":"Oceanography and Marine Biology\u2014An Annual Review"},{"issue":"2","key":"pcbi.1010291.ref014","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/S0079-6611(03)00085-5","article-title":"Phytoplankton Patchiness: The Role of Lateral Stirring and Mixing","volume":"57","author":"AP Martin","year":"2003","journal-title":"Progress in Oceanography"},{"issue":"2","key":"pcbi.1010291.ref015","first-page":"652","article-title":"Plankton Patchiness: Biology in the Physical Vernacular","volume":"37","author":"DL Mackas","year":"1985","journal-title":"Bulletin of Marine Science"},{"issue":"1","key":"pcbi.1010291.ref016","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3354\/ame035001","article-title":"Microscale Heterogeneity in the Activity of Coastal Bacterioplankton Communities","volume":"35","author":"JR Seymour","year":"2004","journal-title":"Aquatic Microbial Ecology"},{"issue":"3","key":"pcbi.1010291.ref017","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/j.jmarsys.2006.01.019","article-title":"Phytoplankton Patch Patterns: Seascape Anatomy in a Turbulent Ocean","volume":"69","author":"JG Mitchell","year":"2008","journal-title":"Journal of Marine Systems"},{"issue":"5","key":"pcbi.1010291.ref018","first-page":"197","article-title":"Microbial Micropatches within Microbial Hotspots","volume":"13","author":"LM Dann","year":"2018","journal-title":"PLOS ONE"},{"issue":"4","key":"pcbi.1010291.ref019","doi-asserted-by":"crossref","first-page":"613","DOI":"10.4319\/lo.1997.42.4.0613","article-title":"Spatially Explicit Simulations of a Microbial Food Web","volume":"42","author":"N Blackburn","year":"1997","journal-title":"Limnology and Oceanography"},{"issue":"6","key":"pcbi.1010291.ref020","doi-asserted-by":"crossref","first-page":"1309","DOI":"10.4319\/lo.2001.46.6.1309","article-title":"Marine Snow, Organic Solute Plumes, and Optimal Chemosensory Behavior of Bacteria","volume":"46","author":"T Ki\u00f8rboe","year":"2001","journal-title":"Limnology and Oceanography"},{"issue":"1","key":"pcbi.1010291.ref021","doi-asserted-by":"crossref","first-page":"109","DOI":"10.4319\/lo.2006.51.1.0109","article-title":"Individual Foraging Behaviors and Population Distributions of a Planktonic Predator Aggregating to Phytoplankton Thin Layers","volume":"51","author":"SM Deuer","year":"2006","journal-title":"Limnology and Oceanography"},{"issue":"6736","key":"pcbi.1010291.ref022","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1038\/21119","article-title":"Marine Viruses and Their Biogeochemical and Ecological Effects","volume":"399","author":"JA Fuhrman","year":"1999","journal-title":"Nature"},{"issue":"3","key":"pcbi.1010291.ref023","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1017\/S0025315406013464","article-title":"Microscale Patchiness of Virioplankton","volume":"86","author":"JR Seymour","year":"2006","journal-title":"Journal of the Marine Biological Association of the United Kingdom"},{"issue":"8","key":"pcbi.1010291.ref024","doi-asserted-by":"crossref","first-page":"1640","DOI":"10.4319\/lo.1992.37.8.1640","article-title":"Behavior of Acartia Tonsa in Patchy Food Environments","volume":"37","author":"P Tiselius","year":"1992","journal-title":"Limnology and Oceanography"},{"issue":"2","key":"pcbi.1010291.ref025","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1093\/plankt\/25.2.121","article-title":"Plankton Patchiness and Its Effect on Larger-Scale Productivity","volume":"25","author":"SJ Brentnall","year":"2003","journal-title":"Journal of Plankton Research"},{"issue":"2","key":"pcbi.1010291.ref026","doi-asserted-by":"crossref","first-page":"103","DOI":"10.3354\/ame026103","article-title":"Microscale Patchiness of Bacterioplankton Assemblage Richness in Seawater","volume":"26","author":"RA Long","year":"2001","journal-title":"Aquatic Microbial Ecology"},{"issue":"882","key":"pcbi.1010291.ref027","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1086\/282171","article-title":"The Paradox of the Plankton","volume":"95","author":"GE Hutchinson","year":"1961","journal-title":"The American Naturalist"},{"issue":"4","key":"pcbi.1010291.ref028","doi-asserted-by":"crossref","first-page":"1710","DOI":"10.1073\/pnas.67.4.1710","article-title":"Contemporaneous Disequilibrium, a New Hypothesis to Explain the \u201cParadox of the Plankton\u201d","volume":"67","author":"P Richerson","year":"1970","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"pcbi.1010291.ref029","first-page":"47","article-title":"Oceanic Turbulence at Millimeter Scales. In: Lectures on Plankton and Turbulence. No. 61 (Supl. 1) in Scientia Marina","author":"J Jim\u00e9nez","year":"1997","journal-title":"International Centre for Coastal Resources, Barcelona"},{"issue":"1","key":"pcbi.1010291.ref030","doi-asserted-by":"crossref","DOI":"10.1029\/2005GL024103","article-title":"Phytoplankton Microstructure in Fully Developed Oceanic Turbulence","volume":"33","author":"H Yamazaki","year":"2006","journal-title":"Geophysical Research Letters"},{"issue":"12","key":"pcbi.1010291.ref031","doi-asserted-by":"crossref","first-page":"1142","DOI":"10.1088\/0957-0233\/10\/12\/305","article-title":"Measurements of Plankton Distribution in the Ocean Using Submersible Holography","volume":"10","author":"E Malkiel","year":"1999","journal-title":"Measurement Science and Technology"},{"key":"pcbi.1010291.ref032","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms3148","article-title":"Turbulence Drives Microscale Patches of Motile Phytoplankton","volume":"4","author":"WM Durham","year":"2013","journal-title":"Nature Communications"},{"issue":"112","key":"pcbi.1010291.ref033","doi-asserted-by":"crossref","first-page":"20150791","DOI":"10.1098\/rsif.2015.0791","article-title":"Shear-Induced Orientational Dynamics and Spatial Heterogeneity in Suspensions of Motile Phytoplankton","volume":"12","author":"MT Barry","year":"2015","journal-title":"Journal of The Royal Society Interface"},{"issue":"6","key":"pcbi.1010291.ref034","doi-asserted-by":"crossref","DOI":"10.1103\/PhysRevE.93.063110","article-title":"Clustering of Particles in Turbulence Due to Phoresis","volume":"93","author":"L Schmidt","year":"2016","journal-title":"Physical Review E"},{"issue":"5999","key":"pcbi.1010291.ref035","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1038\/313218a0","article-title":"Hydrodynamic Focusing of Motile Algal Cells","volume":"313","author":"JO Kessler","year":"1985","journal-title":"Nature"},{"issue":"23","key":"pcbi.1010291.ref036","doi-asserted-by":"crossref","first-page":"238102","DOI":"10.1103\/PhysRevLett.106.238102","article-title":"Gyrotaxis in a Steady Vortical Flow","volume":"106","author":"WM Durham","year":"2011","journal-title":"Physical Review Letters"},{"key":"pcbi.1010291.ref037","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.mib.2015.03.003","article-title":"Microbes in Flow","volume":"25","author":"R Rusconi","year":"2015","journal-title":"Current Opinion in Microbiology"},{"issue":"4","key":"pcbi.1010291.ref038","doi-asserted-by":"crossref","first-page":"044502","DOI":"10.1103\/PhysRevLett.112.044502","article-title":"Turbulent Fluid Acceleration Generates Clusters of Gyrotactic Microorganisms","volume":"112","author":"F De Lillo","year":"2014","journal-title":"Physical Review Letters"},{"issue":"12","key":"pcbi.1010291.ref039","doi-asserted-by":"crossref","first-page":"124304","DOI":"10.1103\/PhysRevFluids.4.124304","article-title":"Role of Large-Scale Advection and Small-Scale Turbulence on Vertical Migration of Gyrotactic Swimmers","volume":"4","author":"C Marchioli","year":"2019","journal-title":"Physical Review Fluids"},{"key":"pcbi.1010291.ref040","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1017\/S0022112002001015","article-title":"Turbulent Convection Driven by Surface Cooling in Shallow Water","volume":"464","author":"O Zikanov","year":"2002","journal-title":"Journal of Fluid Mechanics"},{"issue":"13","key":"pcbi.1010291.ref041","doi-asserted-by":"crossref","first-page":"e2020GL086983","DOI":"10.1029\/2020GL086983","article-title":"Where Do Cold Air Outbreaks Occur, and How Have They Changed Over Time?","volume":"47","author":"ET Smith","year":"2020","journal-title":"Geophysical Research Letters"},{"key":"pcbi.1010291.ref042","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1016\/B0-12-227090-8\/00093-2","volume-title":"Encyclopedia of Atmospheric Sciences","author":"L Kantha","year":"2003"},{"issue":"5975","key":"pcbi.1010291.ref043","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1038\/310282a0","article-title":"Turbulence in an Oceanic Convective Mixed Layer","volume":"310","author":"TJ Shay","year":"1984","journal-title":"Nature"},{"issue":"1","key":"pcbi.1010291.ref044","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1029\/98RG02739","article-title":"Open-Ocean Convection: Observations, Theory, and Models","volume":"37","author":"J Marshall","year":"1999","journal-title":"Reviews of Geophysics"},{"issue":"9","key":"pcbi.1010291.ref045","article-title":"Lake-Size Dependency of Wind Shear and Convection as Controls on Gas Exchange","volume":"39","author":"JS Read","year":"2012","journal-title":"Geophysical Research Letters"},{"key":"pcbi.1010291.ref046","doi-asserted-by":"crossref","first-page":"2015","DOI":"10.1006\/rwos.2001.0118","article-title":"Open Ocean Convection","author":"A Soloviev","year":"2001","journal-title":"Encyclopedia of Ocean Sciences"},{"issue":"3","key":"pcbi.1010291.ref047","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1128\/aem.61.3.877-882.1995","article-title":"Long Lag Times and High Velocities in the Motility of Natural Assemblages of Marine Bacteria","volume":"61","author":"JG Mitchell","year":"1995","journal-title":"Applied and Environmental Microbiology"},{"issue":"2","key":"pcbi.1010291.ref048","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1007\/BF00347287","article-title":"Comparison of Sinking Velocity, Swimming Velocity, Rotation and Path Characteristics among Six Marine Dinoflagellate Species","volume":"113","author":"D Kamykowski","year":"1992","journal-title":"Marine Biology"},{"issue":"2","key":"pcbi.1010291.ref049","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1093\/plankt\/8.2.275","article-title":"The Temperature Acclimatized Swimming Speed of Selected Marine Dinoflagellates","volume":"8","author":"D Kamykowski","year":"1986","journal-title":"Journal of Plankton Research"},{"issue":"4","key":"pcbi.1010291.ref050","doi-asserted-by":"crossref","first-page":"792","DOI":"10.1128\/MMBR.00029-12","article-title":"Ecology and Physics of Bacterial Chemotaxis in the Ocean","volume":"76","author":"R Stocker","year":"2012","journal-title":"Microbiology and Molecular Biology Reviews"},{"issue":"6107","key":"pcbi.1010291.ref051","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1126\/science.1219417","article-title":"Trade-Offs of Chemotactic Foraging in Turbulent Water","volume":"338","author":"JR Taylor","year":"2012","journal-title":"Science"},{"issue":"7646","key":"pcbi.1010291.ref052","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1038\/nature21415","article-title":"Phytoplankton Can Actively Diversify Their Migration Strategy in Response to Turbulent Cues","volume":"543","author":"A Sengupta","year":"2017","journal-title":"Nature"},{"issue":"5","key":"pcbi.1010291.ref053","doi-asserted-by":"crossref","first-page":"965","DOI":"10.1111\/j.1529-8817.2007.00392.x","article-title":"Species-Specific Physiological Response of Dinoflagellates to Quantified Small-Scale Turbulence1","volume":"43","author":"E Berdalet","year":"2007","journal-title":"Journal of Phycology"},{"issue":"suppl.","key":"pcbi.1010291.ref054","first-page":"135","article-title":"Diel Vertical Migration by Dinoflagellates: Roles of Carbohydrate Metabolism and Behavioural Flexibility","volume":"27","author":"JJ Cullen","year":"1985","journal-title":"Contributions in Marine Science"},{"issue":"1","key":"pcbi.1010291.ref055","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1146\/annurev.fl.23.010191.002323","article-title":"Turbulent Mixing in Stratified Fluids","volume":"23","author":"HJS Fernando","year":"1991","journal-title":"Annual Review of Fluid Mechanics"},{"issue":"8","key":"pcbi.1010291.ref056","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.1175\/1520-0469(1970)027<1211:CVATSF>2.0.CO;2","article-title":"Convective Velocity and Temperature Scales for the Unstable Planetary Boundary Layer and for Rayleigh Convection","volume":"27","author":"JW Deardorff","year":"1970","journal-title":"Journal of the Atmospheric Sciences"},{"issue":"2","key":"pcbi.1010291.ref057","doi-asserted-by":"crossref","DOI":"10.1029\/2010GL045581","article-title":"Oceanic Convective Mixing and the Impact on Air-Sea Gas Transfer Velocity","volume":"38","author":"A Rutgersson","year":"2011","journal-title":"Geophysical Research Letters"},{"key":"pcbi.1010291.ref058","doi-asserted-by":"crossref","first-page":"500","DOI":"10.1017\/jfm.2014.640","article-title":"Energy Dispersion in Turbulent Jets. Part 1. Direct Simulation of Steady and Unsteady Jets","author":"J Craske","year":"2015","journal-title":"Journal of Fluid Mechanics"},{"issue":"1","key":"pcbi.1010291.ref059","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1016\/S0021-9991(03)00126-8","article-title":"Symmetry-Preserving Discretization of Turbulent Flow","volume":"187","author":"RWCP Verstappen","year":"2003","journal-title":"Journal of Computational Physics"},{"issue":"11","key":"pcbi.1010291.ref060","doi-asserted-by":"crossref","first-page":"5943","DOI":"10.1073\/pnas.1919332117","article-title":"Microbial Evolutionary Strategies in a Dynamic Ocean","volume":"117","author":"NG Walworth","year":"2020","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"5","key":"pcbi.1010291.ref061","doi-asserted-by":"crossref","first-page":"1374","DOI":"10.1038\/s41396-019-0355-6","article-title":"Live Cell Analysis at Sea Reveals Divergent Thermal Performance between Photosynthetic Ocean Microbial Eukaryote Populations","volume":"13","author":"AS McInnes","year":"2019","journal-title":"The ISME Journal"},{"issue":"5","key":"pcbi.1010291.ref062","doi-asserted-by":"crossref","first-page":"e2019JC015348","DOI":"10.1029\/2019JC015348","article-title":"Eddies in the Hawaiian Archipelago Region: Formation, Characterization, and Potential Implications on Larval Retention of Reef Fish","volume":"125","author":"D Lindo-Atichati","year":"2020","journal-title":"Journal of Geophysical Research: Oceans"},{"key":"pcbi.1010291.ref063","doi-asserted-by":"crossref","first-page":"109130","DOI":"10.1016\/j.ecolmodel.2020.109130","article-title":"Effects of Swimming Behaviour and Oceanography on Sea Turtle Hatchling Dispersal at the Intersection of Two Ocean Current Systems","volume":"431","author":"DZM Le Gouvello","year":"2020","journal-title":"Ecological Modelling"},{"issue":"4","key":"pcbi.1010291.ref064","doi-asserted-by":"crossref","first-page":"775","DOI":"10.1111\/evo.13946","article-title":"Phytoplankton Thermal Responses Adapt in the Absence of Hard Thermodynamic Constraints","volume":"74","author":"DG Kontopoulos","year":"2020","journal-title":"Evolution"},{"issue":"2","key":"pcbi.1010291.ref065","doi-asserted-by":"crossref","first-page":"e2019JC015583","DOI":"10.1029\/2019JC015583","article-title":"Influence of Barotropic Tidal Currents on Transport and Accumulation of Floating Microplastics in the Global Open Ocean","volume":"125","author":"MF Sterl","year":"2020","journal-title":"Journal of Geophysical Research: Oceans"},{"issue":"11","key":"pcbi.1010291.ref066","doi-asserted-by":"crossref","first-page":"115001","DOI":"10.1088\/2515-7620\/ab4e77","article-title":"Mixing of Passive Tracers at the Ocean Surface and Its Implications for Plastic Transport Modelling","volume":"1","author":"D Wichmann","year":"2019","journal-title":"Environmental Research Communications"},{"issue":"5","key":"pcbi.1010291.ref067","doi-asserted-by":"crossref","first-page":"1341","DOI":"10.5194\/os-15-1341-2019","article-title":"Basin-Scale Sources and Pathways of Microplastic That Ends up in the Gal\u00e1pagos Archipelago","volume":"15","author":"E van Sebille","year":"2019","journal-title":"Ocean Science"},{"key":"pcbi.1010291.ref068","first-page":"1","article-title":"Parcels v0.9: Prototyping a Lagrangian Ocean Analysis Framework for the Petascale Age","author":"M Lange","year":"2017","journal-title":"Geoscientific Model Development Discussions"},{"key":"pcbi.1010291.ref069","first-page":"1","article-title":"The Parcels v2.0 Lagrangian Framework: New Field Interpolation Schemes","author":"P Delandmeter","year":"2019","journal-title":"Geoscientific Model Development Discussions"},{"issue":"1","key":"pcbi.1010291.ref070","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1146\/annurev.fl.24.010192.001525","article-title":"Hydrodynamic Phenomena in Suspensions of Swimming Microorganisms","volume":"24","author":"TJ Pedley","year":"1992","journal-title":"Annual Review of Fluid Mechanics"},{"key":"pcbi.1010291.ref071","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1017\/S002211209400306X","article-title":"Biflagellate Gyrotaxis in a Shear Flow","volume":"281","author":"MS Jones","year":"1994","journal-title":"Journal of Fluid Mechanics"},{"issue":"4","key":"pcbi.1010291.ref072","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1006\/jtbi.1997.0421","article-title":"A Biased Random Walk Model for the Trajectories of Swimming Micro-Organisms","volume":"186","author":"NA Hill","year":"1997","journal-title":"Journal of Theoretical Biology"},{"issue":"5","key":"pcbi.1010291.ref073","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1007\/s40974-017-0071-x","article-title":"Swimming Characteristics of Gyrotactic Microorganisms in Low-Reynolds-Number Flow: Chlamydomonas Reinhardtii","volume":"2","author":"X Chen","year":"2017","journal-title":"Energy, Ecology and Environment"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1010291","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,8,16]],"date-time":"2022-08-16T00:00:00Z","timestamp":1660608000000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1010291","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,11,24]],"date-time":"2023-11-24T20:26:38Z","timestamp":1700857598000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1010291"}},"subtitle":[],"editor":[{"given":"Kiran Raosaheb","family":"Patil","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2022,7,27]]},"references-count":73,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2022,7,27]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1010291","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,7,27]]}}}