{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,4]],"date-time":"2026-06-04T21:25:19Z","timestamp":1780608319616,"version":"3.54.1"},"reference-count":47,"publisher":"MDPI AG","issue":"6","license":[{"start":{"date-parts":[[2018,5,29]],"date-time":"2018-05-29T00:00:00Z","timestamp":1527552000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100003593","name":"Conselho Nacional de Desenvolvimento Cient\u00edfico e Tecnol\u00f3gico","doi-asserted-by":"publisher","award":["211546\/2013-5"],"award-info":[{"award-number":["211546\/2013-5"]}],"id":[{"id":"10.13039\/501100003593","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Remote Sensing"],"abstract":"<jats:p>The unicellular cyanobacterium Prochlorococcus is the most dominant resident of the subtropical gyres, which are considered to be the largest biomes on earth. In this study, the spatial and temporal variability in the global distribution of Prochlorococcus was estimated in the Atlantic Ocean using an empirical model based on data from 13 Atlantic Meridional Transect cruises. Our model uses satellite-derived sea surface temperature (SST), remote-sensing reflectance at 443 and 488 nm, and the water temperature at a depth of 200 m from Argo data. The model divides the population of Prochlorococcus into two groups: ProI, which dominates under high-light conditions associated with the surface, and ProII, which favors low light found near the deep chlorophyll maximum. ProI and ProII are then summed to provide vertical profiles of the concentration of Prochlorococcus cells. This model predicts that Prochlorococcus cells contribute 32 Mt of carbon biomass (7.4 \u00d7 1026 cells) to the Atlantic Ocean, concentrated mainly within the subtropical gyres (35%) and areas near the Equatorial Convergence Zone (30%). When projected globally, 3.4 \u00d7 1027 Prochlorococcus cells represent 171 Mt of carbon biomass, with 43% of this global biomass allocated to the upper ocean (0\u201345 m depth). Annual cell standing stocks were relatively stable between the years 2003 and 2014, and the contribution of the gyres varies seasonally as gyres expand and contract, tracking changes in light and temperature, with lowest cell abundances during the boreal and austral winter (1.4 \u00d7 1013 cells m\u22122), when surface cell concentrations were highest (9.8 \u00d7 104 cells mL\u22121), whereas the opposite scenario was observed in spring\u2013summer (2 \u00d7 1013 cells m\u22122). This model provides a three-dimensional view of the abundance of Prochlorococcus cells, revealing that Prochlorococcus contributes significantly to total phytoplankton biomass in the Atlantic Ocean, and can be applied using either in situ measurements at the sea surface (r2 = 0.83) or remote-sensing observables (r2 = 0.58).<\/jats:p>","DOI":"10.3390\/rs10060847","type":"journal-article","created":{"date-parts":[[2018,5,30]],"date-time":"2018-05-30T03:04:27Z","timestamp":1527649467000},"page":"847","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":23,"title":["Scratching Beneath the Surface: A Model to Predict the Vertical Distribution of Prochlorococcus Using Remote Sensing"],"prefix":"10.3390","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2136-2273","authenticated-orcid":false,"given":"Priscila K.","family":"Lange","sequence":"first","affiliation":[{"name":"Department of Earth Sciences, University of Oxford, South Parks Rd, OX1 3AN Oxford, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5134-8291","authenticated-orcid":false,"given":"Robert J. W.","family":"Brewin","sequence":"additional","affiliation":[{"name":"Plymouth Marine Laboratory, Prospect Place, The Hoe, PL1 3DH Plymouth, UK"},{"name":"National Centre for Earth Observation, Plymouth Marine Laboratory, PL1 3DH Plymouth, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Giorgio","family":"Dall\u2019Olmo","sequence":"additional","affiliation":[{"name":"Plymouth Marine Laboratory, Prospect Place, The Hoe, PL1 3DH Plymouth, UK"},{"name":"National Centre for Earth Observation, Plymouth Marine Laboratory, PL1 3DH Plymouth, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3695-5151","authenticated-orcid":false,"given":"Glen A.","family":"Tarran","sequence":"additional","affiliation":[{"name":"Plymouth Marine Laboratory, Prospect Place, The Hoe, PL1 3DH Plymouth, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shubha","family":"Sathyendranath","sequence":"additional","affiliation":[{"name":"Plymouth Marine Laboratory, Prospect Place, The Hoe, PL1 3DH Plymouth, UK"},{"name":"National Centre for Earth Observation, Plymouth Marine Laboratory, PL1 3DH Plymouth, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mikhail","family":"Zubkov","sequence":"additional","affiliation":[{"name":"National Oceanography Centre Southampton, Waterfront Campus, SO14 3ZH Southampton, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heather A.","family":"Bouman","sequence":"additional","affiliation":[{"name":"Department of Earth Sciences, University of Oxford, South Parks Rd, OX1 3AN Oxford, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2018,5,29]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"3153","DOI":"10.1016\/j.rse.2008.03.011","article-title":"Remote Sensing of Environment an absorption model to determine phytoplankton size classes from satellite ocean colour","volume":"112","author":"Hirata","year":"2008","journal-title":"Remote Sens. Environ."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1007\/s100219900068","article-title":"Minireviews: A Sea of Change: Biogeochemical Variability in the North Pacific Subtropical Gyre","volume":"2","author":"Karl","year":"1999","journal-title":"Ecosystems"},{"key":"ref_3","unstructured":"Longhurst, A. (1998). Ecological Geography of the Sea, Academic Press."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1641\/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2","article-title":"Terrestrial Ecoregions of the World: A New Map of Life on Earth","volume":"51","author":"Olson","year":"2001","journal-title":"Bioscience"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1038\/nclimate2838","article-title":"Revaluating ocean warming impacts on global phytoplankton","volume":"6","author":"Behrenfeld","year":"2015","journal-title":"Nat. Clim. Chang."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Westberry, T., Behrenfeld, M.J., Siegel, D.A., and Boss, E. (2008). Carbon-based primary productivity modeling with vertically resolved photoacclimation. Glob. Biogeochem. Cycles, 22.","DOI":"10.1029\/2007GB003078"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Aiken, J., Brewin, R.J.W., Dufois, F., Polimene, L., Hardman-Mountford, N.J., Jackson, T., Loveday, B., Hoya, S.M., Dall\u2019Olmo, G., and Stephens, J. (2016). A synthesis of the environmental response of the North and South Atlantic Sub-Tropical Gyres during two decades of AMT. Prog. Oceanogr.","DOI":"10.1016\/j.pocean.2016.08.004"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"791","DOI":"10.1139\/f82-108","article-title":"The deep chlorophyll maximum: Comparing vertical profiles of chlorophyll a","volume":"39","author":"Cullen","year":"1982","journal-title":"Can. J. Fish. Aquat. Sci."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Kirk, J.T.O. (2011). Light and Photosynthesis in Aquatic Ecosystems, Cambridge University Press. [3rd ed.].","DOI":"10.1017\/CBO9781139168212"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1111\/j.1365-2699.2005.01448.x","article-title":"Productivity and carbon fluxes of tropical savannas","volume":"33","author":"Grace","year":"2006","journal-title":"J. Biogeogr."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"340","DOI":"10.1038\/334340a0","article-title":"A novel free-living prochlorophyte abundant in the oceanic euphotic zone","volume":"334","author":"Chisholm","year":"1988","journal-title":"Nature"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1111\/j.1758-2229.2011.00241.x","article-title":"Water-column stratification governs the community structure of subtropical marine picophytoplankton","volume":"3","author":"Bouman","year":"2011","journal-title":"Environ. Microbiol. Rep."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1530","DOI":"10.1016\/j.dsr2.2006.05.005","article-title":"Prokaryoplankton standing stocks in oligotrophic gyre and equatorial provinces of the Atlantic Ocean: Evaluation of inter-annual variability","volume":"53","author":"Heywood","year":"2006","journal-title":"Deep Sea Res. II Top. Stud. Oceanogr."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1252","DOI":"10.1038\/ismej.2010.60","article-title":"Temporal dynamics of Prochlorococcus ecotypes in the Atlantic and Pacific oceans","volume":"4","author":"Malmstrom","year":"2010","journal-title":"ISME J."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"2588","DOI":"10.1111\/j.1462-2920.2007.01376.x","article-title":"Modelling the vertical distribution of Prochlorococcus and Synechococcus in the North Pacific Subtropical Ocean","volume":"9","author":"Rabouille","year":"2007","journal-title":"Environ. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1016\/S0079-6611(00)00008-2","article-title":"Picoplankton community structure on the Atlantic Meridional Transect: A comparison between seasons","volume":"45","author":"Zubkov","year":"2000","journal-title":"Prog. Oceanogr."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1339","DOI":"10.1016\/S0967-0637(98)00015-6","article-title":"Picoplanktonic community structure on an Atlantic transect from 50 degrees N to 50 degrees S","volume":"45","author":"Zubkov","year":"1998","journal-title":"Deep Sea Res. I Oceanogr. Res. Pap."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1132","DOI":"10.4319\/lo.1997.42.5_part_2.1132","article-title":"Bloom dynamics: Physiology, behavior, trophice: Ffects","volume":"42","author":"Smayda","year":"1997","journal-title":"Limonaology Oceanogr."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1721","DOI":"10.4319\/lo.2003.48.5.1721","article-title":"Elemental composition of marine Prochlorococcus and Synechococcus: Implications for the ecological stoichiometry of the sea","volume":"48","author":"Bertilsson","year":"2003","journal-title":"Limnol. Oceanogr."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"9824","DOI":"10.1073\/pnas.1307701110","article-title":"Present and future global distributions of the marine Cyanobacteria Prochlorococcus and Synechococcus","volume":"110","author":"Flombaum","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Williams, R.G., and Follows, M.J. (2011). Biological Fundamentals. Ocean Dynamics and the Carbon Cycle: Principles and Mechanisms, Cambridge University Press.","DOI":"10.1017\/CBO9780511977817"},{"key":"ref_22","unstructured":"(2014). NASA Moderate Resolution Imaging Spectroradiometer (MODIS-Aqua) Ocean Color Data, Goddard Space Flight Center Ocean Biology Processing Group."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Gordon, H.R., and Morel, A.Y. (1983). Remote Assessment of Ocean Color for Interpretation of Satellite Visible Imagery, Springer-Verlag New York.","DOI":"10.1007\/978-1-4684-6280-7"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"47","DOI":"10.5918\/jamstecr.8.47","article-title":"A monthly mean dataset of global oceanic temperature and salinity derived from Argo float observations","volume":"8","author":"Hosoda","year":"2008","journal-title":"JAMSTEC Rep. Res. Dev."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1737","DOI":"10.1126\/science.1118052","article-title":"Niche Partitioning Among Prochlorococcus Ecotypes Along Ocean-Scale Environmental Gradients","volume":"311","author":"Johnson","year":"2006","journal-title":"Science"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2205","DOI":"10.4319\/lo.2007.52.5.2205","article-title":"Influence of light and temperature on Prochlorococcus ecotype distributions in the Atlantic Ocean","volume":"52","author":"Zinser","year":"2007","journal-title":"Limnol. Oceanogr."},{"key":"ref_27","first-page":"687","article-title":"Photoinhibition of photosynthesis in natural assemblages of marine phytoplankton","volume":"38","author":"Platt","year":"1980","journal-title":"J. Mar. Res."},{"key":"ref_28","first-page":"1","article-title":"Mixed layer depth over the global ocean: An examination of profile data and a profile-based climatology","volume":"109","author":"Madec","year":"2004","journal-title":"J. Geophys. Res. Oceans"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1545","DOI":"10.4319\/lo.1989.34.8.1545","article-title":"Surface pigments, algal biomass profiles, and potential production of the euphotic layer: Relationships reinvestigated in view of remote-sensing applications","volume":"34","author":"Morel","year":"1989","journal-title":"Limnol. Oceanogr."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Venables, W.N., and Ripley, B.D. (2002). Modern Applied Statistics with S, Springer. [4th ed.].","DOI":"10.1007\/978-0-387-21706-2"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.rse.2013.09.016","article-title":"The Ocean Colour Climate Change Initiative: III. A round-robin comparison on in-water bio-optical algorithms","volume":"162","author":"Brewin","year":"2015","journal-title":"Remote Sens. Environ."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/0304-3800(94)00034-F","article-title":"A model comparison for daylength as a function of latitude and day of year","volume":"80","author":"Forsythe","year":"1995","journal-title":"Ecol. Model."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/0038-092X(69)90047-4","article-title":"The absorption of radiation in solar stills","volume":"12","author":"Cooper","year":"1969","journal-title":"Sol. Energy"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1038\/nrmicro3378","article-title":"Prochlorococcus: The structure and function of collective diversity","volume":"13","author":"Biller","year":"2014","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/j.dsr2.2003.08.002","article-title":"Subtropical gyre variability observed by ocean-color satellites","volume":"51","author":"McClain","year":"2004","journal-title":"Deep Sea Res. II Top. Stud. Oceanogr."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"845","DOI":"10.1357\/002224009792006124","article-title":"Plankton community composition, organic carbon and thorium-234 particle size distributions, and particle export in the Sargasso Sea","volume":"67","author":"Brew","year":"2009","journal-title":"J. Mar. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1","DOI":"10.4319\/lo.1997.42.1.0001","article-title":"Photosynthetic rates derived from satellite-based chlorophyll concentration","volume":"42","author":"Behrenfeld","year":"1997","journal-title":"Limnol. Oceanogr."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"956","DOI":"10.4319\/lo.1995.40.5.0956","article-title":"Relationship between the maximum quantum yield of carbon fixation and the minimum quantum yield of chlorophyll a in vivo fluorescence in the Gulf of St. Lawrence","volume":"40","author":"Babin","year":"1995","journal-title":"Limnol. Oceanogr."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"270","DOI":"10.3389\/fmars.2016.00270","article-title":"Comparison study of subtropical mode waters in the world ocean","volume":"3","author":"Tsubouchi","year":"2016","journal-title":"Front. Mar. Sci."},{"key":"ref_40","first-page":"345","article-title":"This Hemisphere","volume":"44","author":"Boggs","year":"1945","journal-title":"J. Geol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"3958","DOI":"10.1002\/2015JC010787","article-title":"What drives seasonal change in oligotrophic area in the subtropical North Atlantic?","volume":"120","author":"Dave","year":"2015","journal-title":"J. Geophys. Res. Oceans"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"L03618","DOI":"10.1029\/2007GL031745","article-title":"Ocean\u2019s least productive waters are expanding","volume":"35","author":"Polovina","year":"2008","journal-title":"Geophys. Res. Lett."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1983","DOI":"10.1016\/S0967-0645(00)00166-1","article-title":"Phytoplankton population dynamics at the Bermuda Atlantic Time-series station in the Sargasso Sea","volume":"48","author":"Durand","year":"2001","journal-title":"Deep Sea Res. II Top. Stud. Oceanogr."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1215\/21573689-2651533","article-title":"The impact of fine-scale turbulence on phytoplankton community structure","volume":"4","author":"Barton","year":"2014","journal-title":"Limnol. Oceanogr. Fluids Environ."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"37","DOI":"10.5194\/essd-4-37-2012","article-title":"Picophytoplankton biomass distribution in the global ocean","volume":"4","author":"Buitenhuis","year":"2012","journal-title":"Earth Syst. Sci. Data"},{"key":"ref_46","first-page":"285","article-title":"Prochlorococcus and synechococcus have evolved different adaptive mechanisms to cope with light and uv stress","volume":"3","author":"Six","year":"2012","journal-title":"Front. Microbiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1128\/MMBR.63.1.106-127.1999","article-title":"Prochlorococcus, a marine photosynthetic prokaryote of global significance","volume":"63","author":"Partensky","year":"1999","journal-title":"Microbiol. Mol. Biol. Rev."}],"container-title":["Remote Sensing"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-4292\/10\/6\/847\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T15:06:22Z","timestamp":1760195182000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-4292\/10\/6\/847"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,5,29]]},"references-count":47,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2018,6]]}},"alternative-id":["rs10060847"],"URL":"https:\/\/doi.org\/10.3390\/rs10060847","relation":{},"ISSN":["2072-4292"],"issn-type":[{"value":"2072-4292","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,5,29]]}}}