{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:05:59Z","timestamp":1792785959533,"version":"4.5.0"},"reference-count":78,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T00:00:00Z","timestamp":1787788800000},"content-version":"vor","delay-in-days":238,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Department of Biology Education, Stockholm University","id":[{"id":"https:\/\/ror.org\/05f0yaq80","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Knut and Alice Wallenberg Foundation Fellowship and Prolongation","id":[{"id":"https:\/\/ror.org\/004hzzk67","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Swedish Research Councils VR","award":["2021-05563"],"award-info":[{"award-number":["2021-05563"]}],"id":[{"id":"https:\/\/ror.org\/03zttf063","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000066","name":"NIEHS","doi-asserted-by":"publisher","award":["1P01ES035542-01"],"award-info":[{"award-number":["1P01ES035542-01"]}],"id":[{"id":"10.13039\/100000066","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"NSF","doi-asserted-by":"publisher","award":["OCE-2414792"],"award-info":[{"award-number":["OCE-2414792"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001862","name":"Swedish Research Council FORMAS","doi-asserted-by":"publisher","award":["2022-01515"],"award-info":[{"award-number":["2022-01515"]}],"id":[{"id":"10.13039\/501100001862","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001862","name":"Swedish Research Council FORMAS","doi-asserted-by":"publisher","award":["FR2017-11-09"],"award-info":[{"award-number":["FR2017-11-09"]}],"id":[{"id":"10.13039\/501100001862","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,14]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Picocyanobacteria contribute significantly to carbon export but the mechanism by which their biomass is transported to depth is unclear. By analyzing 16S rRNA gene sequences from sediment trap material and size-fractionated water samples from different depths in the Baltic Sea, we show that picocyanobacterial contribution to phytoplankton biomass export was largely proportional (70%\u201390%) to their surface water abundances. However, we also found that both cell aggregation (enrichment in &amp;gt;10\u00a0\u03bcm fraction) and sedimentation (trap enrichment) varied among Synechococcus ASVs (amplicon sequence variants). A robust linear relationship was identified between ASV enrichment in traps and ASV enrichment in the &amp;gt;10\u00a0\u03bcm fraction, suggesting that cell aggregation is a direct determinant of Synechococcus sedimentation. Seasonal abundance patterns were similar among the dominant aggregate-forming ASVs but distinct from those of single-cell ASVs, suggesting aggregate\/colony formation being an important trait in determining distribution of Synechococcus strains and their response to environmental conditions. Correlating microscopy- and sequence-based abundances further allowed us to propose genotypes for the dominant Baltic Sea colonial picocyanobacterial morphotypes. Our results provide in situ data linking phenotypic characteristics of picocyanobacteria with sedimentation properties and several lines of evidence for a quantitative dependency between surface water picocyanobacterial aggregate\/colony formation and sinking. These findings are important for our understanding of mechanisms involved in export of pico-sized phytoplankton biomass from the photic zone to depth and emphasize the need to investigate occurrence and environmental regulation of cell aggregation in other picophytoplankton-dominated waters (e.g. open ocean gyres).<\/jats:p>","DOI":"10.1093\/ismejo\/wrag214","type":"journal-article","created":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T17:32:01Z","timestamp":1787851921000},"source":"Crossref","is-referenced-by-count":0,"title":["Differential aggregation properties among Baltic Sea picocyanobacterial strains impact their export"],"prefix":"10.1093","volume":"20","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7016-1473","authenticated-orcid":false,"given":"Martin","family":"Ekman","sequence":"first","affiliation":[{"name":"Department of Ecology, Environment and Plant Sciences (DEEP), Stockholm University , SE-106 91 Stockholm ,","place":["Sweden"]},{"name":"Department of Clinical Microbiology, Karolinska University Hospital , SE-14186 Stockholm ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elina","family":"Viinam\u00e4ki","sequence":"additional","affiliation":[{"name":"Department of Ecology, Environment and Plant Sciences (DEEP), Stockholm University , SE-106 91 Stockholm ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anders","family":"Verstraaten-Svensson","sequence":"additional","affiliation":[{"name":"Department of Ecology, Environment and Plant Sciences (DEEP), Stockholm University , SE-106 91 Stockholm ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3980-8181","authenticated-orcid":false,"given":"Ryan W","family":"Paerl","sequence":"additional","affiliation":[{"name":"Department of Marine, Earth, and Atmospheric Sciences, NC State University , Raleigh NC 27695 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3627-6899","authenticated-orcid":false,"given":"Anders F","family":"Andersson","sequence":"additional","affiliation":[{"name":"Department of Gene Technology, Science for Life Laboratory, KTH Royal Institute of Technology , SE-100 44 Stockholm ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Helena","family":"H\u00f6glander","sequence":"additional","affiliation":[{"name":"Department of Ecology, Environment and Plant Sciences (DEEP), Stockholm University , SE-106 91 Stockholm ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jakob","family":"Walve","sequence":"additional","affiliation":[{"name":"Department of Ecology, Environment and Plant Sciences (DEEP), Stockholm University , SE-106 91 Stockholm ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8696-1835","authenticated-orcid":false,"given":"Rachel A","family":"Foster","sequence":"additional","affiliation":[{"name":"Department of Ecology, Environment and Plant Sciences (DEEP), Stockholm University , SE-106 91 Stockholm ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,8,27]]},"reference":[{"key":"2026100200262314300_ref1","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1111\/jpy.13304","article-title":"An updated classification of cyanobacterial orders and families based on phylogenomic and polyphasic analysis","volume":"59","author":"Struneck\u00fd","year":"2023","journal-title":"J Phycol"},{"key":"2026100200262314300_ref2","doi-asserted-by":"publisher","first-page":"45","DOI":"10.4319\/lo.1990.35.1.0045","article-title":"Pigments, size, and distributions of Synechococcus in the North Atlantic and Pacific oceans","volume":"35","author":"Olson","year":"1990","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref3","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1128\/MMBR.00035-08","article-title":"Ecological genomics of marine Picocyanobacteria","volume":"73","author":"Scanlan","year":"2009","journal-title":"Microbiol Mol Biol Rev"},{"key":"2026100200262314300_ref4","doi-asserted-by":"publisher","first-page":"546","DOI":"10.3390\/microorganisms10030546","article-title":"The \u201cdark side\u201d of Picocyanobacteria: life as we do not know it (yet)","volume":"10","author":"Callieri","year":"2022","journal-title":"Microorganisms"},{"key":"2026100200262314300_ref5","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1016\/0198-0149(88)90126-4","article-title":"Primary production, sinking fluxes and the microbial food web","volume":"35","author":"Michaels","year":"1988","journal-title":"Deep Sea Res Part Oceanogr Res Pap"},{"key":"2026100200262314300_ref6","doi-asserted-by":"publisher","first-page":"838","DOI":"10.1126\/science.1133471","article-title":"Small phytoplankton and carbon export from the Surface Ocean","volume":"315","author":"Richardson","year":"2007","journal-title":"Science"},{"key":"2026100200262314300_ref7","doi-asserted-by":"publisher","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":"2026100200262314300_ref8","doi-asserted-by":"publisher","first-page":"717","DOI":"10.1038\/ngeo1265","article-title":"Assimilation of upwelled nitrate by small eukaryotes in the Sargasso Sea","volume":"4","author":"Fawcett","year":"2011","journal-title":"Nat Geosci"},{"key":"2026100200262314300_ref9","doi-asserted-by":"publisher","first-page":"203","DOI":"10.5194\/bg-8-203-2011","article-title":"Evidence for aggregation and export of cyanobacteria and nano-eukaryotes from the Sargasso Sea euphotic zone","volume":"8","author":"Lomas","year":"2011","journal-title":"Biogeosciences"},{"key":"2026100200262314300_ref10","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1016\/j.dsr2.2013.01.001","article-title":"DNA-based molecular fingerprinting of eukaryotic protists and cyanobacteria contributing to sinking particle flux at the Bermuda Atlantic time-series study","volume":"93","author":"Amacher","year":"2013","journal-title":"Deep Sea Res Part II Top Stud Oceanogr"},{"key":"2026100200262314300_ref11","doi-asserted-by":"publisher","first-page":"1309","DOI":"10.1002\/2015GB005134","article-title":"Small phytoplankton drive high summertime carbon and nutrient export in the Gulf of California and eastern tropical North Pacific","volume":"29","author":"Puigcorb\u00e9","year":"2015","journal-title":"Glob Biogeochem Cycles"},{"key":"2026100200262314300_ref12","doi-asserted-by":"publisher","first-page":"465","DOI":"10.1038\/nature16942","article-title":"Plankton networks driving carbon export in the oligotrophic ocean","volume":"532","author":"Guidi","year":"2016","journal-title":"Nature"},{"key":"2026100200262314300_ref13","doi-asserted-by":"publisher","first-page":"S448","DOI":"10.1002\/lno.10765","article-title":"Clade and strain specific contributions of Synechococcus and Prochlorococcus to carbon export in the Sargasso Sea","volume":"63","author":"De Martini","year":"2018","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref14","doi-asserted-by":"publisher","first-page":"6734","DOI":"10.1111\/1462-2920.15736","article-title":"Relating sinking and suspended microbial communities in the California current ecosystem: digestion resistance and the contributions of phytoplankton taxa to export","volume":"23","author":"Valencia","year":"2021","journal-title":"Environ Microbiol"},{"key":"2026100200262314300_ref15","doi-asserted-by":"publisher","first-page":"1211","DOI":"10.1038\/s41467-022-28842-3","article-title":"Adaptive carbon export response to warming in the Sargasso Sea","volume":"13","author":"Lomas","year":"2022","journal-title":"Nat Commun"},{"key":"2026100200262314300_ref16","doi-asserted-by":"publisher","first-page":"104708","DOI":"10.1016\/j.dsr2.2019.104708","article-title":"DNA metabarcoding reveals organisms contributing to particulate matter flux to abyssal depths in the north East Pacific Ocean","volume":"173","author":"Preston","year":"2020","journal-title":"Deep Sea Res Part II Top Stud Oceanogr"},{"key":"2026100200262314300_ref17","doi-asserted-by":"publisher","first-page":"103668","DOI":"10.1016\/j.dsr.2021.103668","article-title":"Microbial communities associated with sinking particles across an environmental gradient from coastal upwelling to the oligotrophic ocean","volume":"179","author":"Valencia","year":"2022","journal-title":"Deep Sea Res Part Oceanogr Res Pap"},{"key":"2026100200262314300_ref18","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/0025-3227(71)90070-3","article-title":"Normal and accelerated sinking of phytoplankton in the sea","volume":"11","author":"Smayda","year":"1971","journal-title":"Mar Geol"},{"key":"2026100200262314300_ref19","doi-asserted-by":"publisher","first-page":"24","DOI":"10.4319\/lo.1976.21.1.0024","article-title":"Particle retention efficiencies of an herbivorous copepod, Acartia clausi (adult and copepodite stages): effects on grazing1","volume":"21","author":"Nival","year":"1976","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref20","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1007\/978-94-007-3855-3_8","volume-title":"Ecology of Cyanobacteria II: Their Diversity in Space and Time","author":"Callieri","year":"2012"},{"key":"2026100200262314300_ref21","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1016\/j.hal.2006.07.006","article-title":"Phytoplankton vertical distributions and composition in Baltic Sea cyanobacterial blooms","volume":"6","author":"Hajdu","year":"2007","journal-title":"Harmful Algae"},{"key":"2026100200262314300_ref22","doi-asserted-by":"publisher","first-page":"311","DOI":"10.5194\/bgd-4-2667-2007","article-title":"Distribution of micro-organisms along a transect in the south-East Pacific Ocean (BIOSOPE cruise) using epifluorescence microscopy","volume":"5","author":"Masquelier","year":"2008","journal-title":"Biogeosciences"},{"key":"2026100200262314300_ref23","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1111\/1755-0998.13592","article-title":"A robust approach to estimate relative phytoplankton cell abundances from metagenomes","volume":"23","author":"Pierella Karlusich","year":"2023","journal-title":"Mol Ecol Resour"},{"key":"2026100200262314300_ref24","doi-asserted-by":"publisher","first-page":"805","DOI":"10.1002\/lno.10059","article-title":"Effects of clay minerals on the aggregation and subsequent settling of marine Synechococcus","volume":"60","author":"Deng","year":"2015","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref25","doi-asserted-by":"publisher","first-page":"39","DOI":"10.3354\/ame01803","article-title":"Effects of nutrient limitation on cell growth, TEP production and aggregate formation of marine Synechococcus","volume":"78","author":"Deng","year":"2016","journal-title":"Aquat Microb Ecol"},{"key":"2026100200262314300_ref26","doi-asserted-by":"publisher","first-page":"1864","DOI":"10.3389\/fmicb.2019.01864","article-title":"Heterotrophic bacteria enhance the aggregation of the marine Picocyanobacteria Prochlorococcus and Synechococcus","volume":"10","author":"Cruz","year":"2019","journal-title":"Front Microbiol"},{"key":"2026100200262314300_ref27","doi-asserted-by":"publisher","first-page":"1120","DOI":"10.1111\/1462-2920.13638","article-title":"Phenotypic plasticity in freshwater picocyanobacteria","volume":"19","author":"Huber","year":"2017","journal-title":"Environ Microbiol"},{"key":"2026100200262314300_ref28","doi-asserted-by":"publisher","DOI":"10.1093\/femsle\/fnx229","article-title":"Synechococcus plasticity under environmental changes","volume":"364","author":"Callieri","year":"2017","journal-title":"FEMS Microbiol Lett"},{"key":"2026100200262314300_ref29","doi-asserted-by":"publisher","first-page":"2611","DOI":"10.1038\/ismej.2017.114","article-title":"High abundance and expression of transposases in bacteria from the Baltic Sea","volume":"11","author":"Vigil-Stenman","year":"2017","journal-title":"ISME J"},{"key":"2026100200262314300_ref30","doi-asserted-by":"publisher","first-page":"1076","DOI":"10.1038\/ismej.2014.198","article-title":"Genomes and gene expression across light and productivity gradients in eastern subtropical Pacific microbial communities","volume":"9","author":"Dupont","year":"2015","journal-title":"ISME J"},{"key":"2026100200262314300_ref31","doi-asserted-by":"publisher","first-page":"1841","DOI":"10.1016\/S0967-0637(99)00032-1","article-title":"Representing phytoplankton aggregates in biogeochemical models","volume":"46","author":"Kriest","year":"1999","journal-title":"Deep Sea Res Part Oceanogr Res Pap"},{"key":"2026100200262314300_ref32","doi-asserted-by":"publisher","first-page":"1695","DOI":"10.1016\/j.csr.2003.06.001","article-title":"BASIC: Baltic Sea cyanobacteria. An investigation of the structure and dynamics of water blooms of cyanobacteria in the Baltic Sea\u2014responses to a changing environment","volume":"23","author":"Stal","year":"2003","journal-title":"Cont Shelf Res"},{"key":"2026100200262314300_ref33","doi-asserted-by":"publisher","first-page":"14330","DOI":"10.1038\/s41598-022-18454-8","article-title":"Seasonal dynamics in picocyanobacterial abundance and clade composition at coastal and offshore stations in the Baltic Sea","volume":"12","author":"Zufia","year":"2022","journal-title":"Sci Rep"},{"key":"2026100200262314300_ref34","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1016\/S0924-7963(00)00016-6","article-title":"Nitrogen fixation by diazotrophic cyanobacteria in the Baltic Sea and transfer of the newly fixed nitrogen to picoplankton organisms","volume":"25","author":"Ohlendieck","year":"2000","journal-title":"J Mar Syst"},{"key":"2026100200262314300_ref35","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1579\/0044-7447-32.8.577","article-title":"Assessing Secchi and photic zone depth in the Baltic Sea from satellite data","volume":"32","author":"Kratzer","year":"2003","journal-title":"Ambio"},{"key":"2026100200262314300_ref36","doi-asserted-by":"publisher","first-page":"2397","DOI":"10.5194\/bg-6-2397-2009","article-title":"Dissolved iron (II) in the Baltic Sea surface water and implications for cyanobacterial bloom development","volume":"6","author":"Breitbarth","year":"2009","journal-title":"Biogeosciences"},{"key":"2026100200262314300_ref37","volume-title":"Guidelines for Monitoring of Phytoplankton Composition, Abundance and Biomass (Version 2017-07-31)","author":"HELCOM"},{"key":"2026100200262314300_ref38","doi-asserted-by":"publisher","first-page":"1955","DOI":"10.1002\/lno.12636","article-title":"Colony-forming and single-cell picocyanobacteria nitrogen acquisition strategies and carbon fixation in the brackish Baltic Sea","volume":"69","author":"Laber","year":"2024","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref39","doi-asserted-by":"publisher","first-page":"205","DOI":"10.3354\/meps031205","article-title":"A new sediment trap system","volume":"31","author":"Larsson","year":"1986","journal-title":"Mar Ecol Prog Ser"},{"key":"2026100200262314300_ref40","doi-asserted-by":"publisher","first-page":"429","DOI":"10.1016\/0272-7714(90)90007-E","article-title":"Two hydrodynamically stable self-suspended buoyant sediment traps","volume":"30","author":"Broman","year":"1990","journal-title":"Estuar Coast Shelf Sci"},{"key":"2026100200262314300_ref41","article-title":"Biovolumes and size-classes of phytoplankton in the Baltic Sea","volume":"106","author":"Olenina I","year":"2006","journal-title":"HELCOM Balt Sea Env Proc"},{"key":"2026100200262314300_ref42","doi-asserted-by":"publisher","first-page":"22026","DOI":"10.1038\/s41598-020-79157-6","article-title":"Picophytoplankton dynamics in a large temperate estuary and impacts of extreme storm events","volume":"10","author":"Paerl","year":"2020","journal-title":"Sci Rep"},{"key":"2026100200262314300_ref43","doi-asserted-by":"publisher","first-page":"1571","DOI":"10.1038\/ismej.2011.41","article-title":"Transitions in bacterial communities along the 2000 km salinity gradient of the Baltic Sea","volume":"5","author":"Herlemann","year":"2011","journal-title":"ISME J"},{"key":"2026100200262314300_ref44","doi-asserted-by":"publisher","first-page":"5116","DOI":"10.1128\/AEM.01403-14","article-title":"DegePrime, a program for degenerate primer design for broad-taxonomic-range PCR in microbial ecology studies","volume":"80","author":"Hugerth","year":"2014","journal-title":"Appl Environ Microbiol"},{"key":"2026100200262314300_ref45","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1038\/s41597-023-02825-5","article-title":"A comprehensive dataset on spatiotemporal variation of microbial plankton communities in the Baltic Sea","volume":"11","author":"Latz","year":"2024","journal-title":"Sci Data"},{"key":"2026100200262314300_ref46","doi-asserted-by":"publisher","first-page":"276","DOI":"10.1038\/s41587-020-0439-x","article-title":"The nf-core framework for community-curated bioinformatics pipelines","volume":"38","author":"Ewels","year":"2020","journal-title":"Nat Biotechnol"},{"key":"2026100200262314300_ref47","doi-asserted-by":"publisher","first-page":"550420","DOI":"10.3389\/fmicb.2020.550420","article-title":"Interpretations of environmental microbial community studies are biased by the selected 16S rRNA (gene) amplicon sequencing pipeline","volume":"11","author":"Straub","year":"2020","journal-title":"Front Microbiol"},{"key":"2026100200262314300_ref48","doi-asserted-by":"publisher","first-page":"1435","DOI":"10.1111\/1755-0998.12401","article-title":"Phyto REF: a reference database of the plastidial 16S rRNA gene of photosynthetic eukaryotes with curated taxonomy","volume":"15","author":"Decelle","year":"2015","journal-title":"Mol Ecol Resour"},{"key":"2026100200262314300_ref49","doi-asserted-by":"publisher","first-page":"923","DOI":"10.3389\/fmicb.2017.00923","article-title":"Identification of cyanobacteria in a eutrophic coastal lagoon on the southern Baltic coast","volume":"8","author":"Albrecht","year":"2017","journal-title":"Front Microbiol"},{"key":"2026100200262314300_ref50","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13059-015-0834-7","article-title":"Metagenome-assembled genomes uncover a global brackish microbiome","volume":"16","author":"Hugerth","year":"2015","journal-title":"Genome Biol"},{"key":"2026100200262314300_ref51","doi-asserted-by":"publisher","first-page":"1892","DOI":"10.1038\/ismej.2014.35","article-title":"Picocyanobacteria containing a novel pigment gene cluster dominate the brackish water Baltic Sea","volume":"8","author":"Larsson","year":"2014","journal-title":"ISME J"},{"key":"2026100200262314300_ref52","doi-asserted-by":"publisher","first-page":"1166","DOI":"10.4319\/lo.1999.44.4.1166","article-title":"Blooms of the picoplanktonic cyanobacterium Synechococcus in Florida bay, a subtropical inner-shelf lagoon","volume":"44","author":"Phlips","year":"1999","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref53","doi-asserted-by":"publisher","first-page":"786590","DOI":"10.3389\/fmicb.2021.786590","article-title":"Seasonality of coastal Picophytoplankton growth, nutrient limitation, and biomass contribution","volume":"12","author":"Alegria Zufia","year":"2021","journal-title":"Front Microbiol"},{"key":"2026100200262314300_ref54","doi-asserted-by":"publisher","first-page":"1405","DOI":"10.1093\/plankt\/19.10.1405","article-title":"Cyanobacterial picoplankton from the Central Baltic Sea: cell size classification by image-analyzed fluorescence microscopy","volume":"19","author":"Albertano","year":"1997","journal-title":"J Plankton Res"},{"key":"2026100200262314300_ref55","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1007\/s10750-012-1278-7","article-title":"Occurrence of cyanobacteria and cyanotoxin in the southern Baltic proper. Filamentous cyanobacteria versus single-celled picocyanobacteria","volume":"701","author":"Mazur-Marzec","year":"2013","journal-title":"Hydrobiologia"},{"key":"2026100200262314300_ref56","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1016\/j.hal.2013.10.016","article-title":"Environmental factors controlling colony formation in blooms of the cyanobacteria Microcystis spp. in Lake Taihu, China","volume":"31","author":"Ma","year":"2014","journal-title":"Harmful Algae"},{"key":"2026100200262314300_ref57","doi-asserted-by":"publisher","first-page":"165","DOI":"10.3354\/ame018165","article-title":"Nutrient control of cyanobacterial blooms in the Baltic Sea","volume":"18","author":"Stal","year":"1999","journal-title":"Aquat Microb Ecol"},{"key":"2026100200262314300_ref58","doi-asserted-by":"publisher","first-page":"1960","DOI":"10.1038\/s41396-019-0386-z","article-title":"Untangling hidden nutrient dynamics: rapid ammonium cycling and single-cell ammonium assimilation in marine plankton communities","volume":"13","author":"Klawonn","year":"2019","journal-title":"ISME J"},{"key":"2026100200262314300_ref59","doi-asserted-by":"publisher","first-page":"57","DOI":"10.3354\/ame01130","article-title":"Blooms of Baltic Sea Aphanizomenon sp. (cyanobacteria) collapse after internal phosphorus depletion","volume":"49","author":"Walve","year":"2007","journal-title":"Aquat Microb Ecol"},{"key":"2026100200262314300_ref60","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1016\/j.marchem.2013.11.002","article-title":"Trace metals and nutrients in Baltic Sea cyanobacteria: internal and external fractions and potential use in nitrogen fixation","volume":"158","author":"Walve","year":"2014","journal-title":"Mar Chem"},{"key":"2026100200262314300_ref61","doi-asserted-by":"publisher","first-page":"15","DOI":"10.3354\/meps266015","article-title":"Phosphorus dynamics during the transition from nitrogen to phosphate limitation in the Central Baltic Sea","volume":"266","author":"Nausch","year":"2004","journal-title":"Mar Ecol Prog Ser"},{"key":"2026100200262314300_ref62","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1016\/j.ecss.2015.07.013","article-title":"Key role of phosphorus and nitrogen in regulating cyanobacterial community composition in the northern Baltic Sea","volume":"164","author":"Andersson","year":"2015","journal-title":"Estuar Coast Shelf Sci"},{"key":"2026100200262314300_ref63","doi-asserted-by":"publisher","first-page":"e2023GB007756","DOI":"10.1029\/2023GB007756","article-title":"Biodiversity and stoichiometric plasticity increase Pico-phytoplankton contributions to marine net primary productivity and the biological pump","volume":"37","author":"Letscher","year":"2023","journal-title":"Glob Biogeochem Cycles"},{"key":"2026100200262314300_ref64","doi-asserted-by":"publisher","first-page":"924","DOI":"10.4319\/lo.1993.38.5.0924","article-title":"Phylogenetic diversity of aggregate-attached vs. free-living marine bacterial assemblages","volume":"38","author":"DeLong","year":"1993","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref65","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1038\/ismej.2013.144","article-title":"Metagenomic analysis of size-fractionated picoplankton in a marine oxygen minimum zone","volume":"8","author":"Ganesh","year":"2014","journal-title":"ISME J"},{"key":"2026100200262314300_ref66","doi-asserted-by":"publisher","first-page":"469","DOI":"10.3389\/fmicb.2015.00469","article-title":"Microbial community structure and function on sinking particles in the North Pacific subtropical gyre","volume":"6","author":"Fontanez","year":"2015","journal-title":"Front Microbiol"},{"key":"2026100200262314300_ref67","doi-asserted-by":"publisher","first-page":"689","DOI":"10.1007\/BF00355486","article-title":"Marine snow studies in the Northeast Atlantic Ocean: distribution, composition and role as a food source for migrating plankton","volume":"116","author":"Lampitt","year":"1993","journal-title":"Mar Biol"},{"key":"2026100200262314300_ref68","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/S0924-7963(99)00056-1","article-title":"Seasonal stages of phytoplankton community structure and sinking loss in the Gulf of Riga","volume":"23","author":"Olli","year":"1999","journal-title":"J Mar Syst"},{"key":"2026100200262314300_ref69","doi-asserted-by":"publisher","first-page":"87","DOI":"10.4319\/lo.2000.45.1.0087","article-title":"Mass sedimentation of picoplankton embedded in organic aggregates","volume":"45","author":"Waite","year":"2000","journal-title":"Limnol Oceanogr"},{"key":"2026100200262314300_ref70","doi-asserted-by":"publisher","first-page":"11","DOI":"10.3354\/meps08648","article-title":"Autotrophic picoplankton in mesozooplankton guts: evidence of aggregate feeding in the mesopelagic zone and export of small phytoplankton","volume":"412","author":"Wilson","year":"2010","journal-title":"Mar Ecol Prog Ser"},{"key":"2026100200262314300_ref71","doi-asserted-by":"publisher","first-page":"e79230","DOI":"10.1371\/journal.pone.0079230","article-title":"Mesozooplankton grazing on Picocyanobacteria in the Baltic Sea as inferred from molecular diet analysis","volume":"8","author":"Motwani","year":"2013","journal-title":"PLoS One"},{"key":"2026100200262314300_ref72","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1016\/0198-0149(89)90131-3","article-title":"Direct observations of the mass flocculation of diatom blooms: characteristics, settling velocities and formation of diatom aggregates","volume":"36","author":"Alldredge","year":"1989","journal-title":"Deep Sea Res Part Oceanogr Res Pap"},{"key":"2026100200262314300_ref73","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1017\/S0967026202003657","article-title":"Diatom aggregation in the sea: mechanisms and ecological implications","volume":"37","author":"Thornton","year":"2002","journal-title":"Eur J Phycol"},{"key":"2026100200262314300_ref74","doi-asserted-by":"publisher","first-page":"4155","DOI":"10.1016\/S0967-0645(01)00084-4","article-title":"Aggregation, sedimentation and biodegradability of phytoplankton-derived material during spring in the Ross Sea, Antarctica","volume":"48","author":"Becquevort","year":"2001","journal-title":"Deep Sea Res Part II Top Stud Oceanogr"},{"key":"2026100200262314300_ref75","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1046\/j.1365-2435.1998.00233.x","article-title":"The twelfth Tansley lecture. Small is beautiful: the picophytoplankton","volume":"12","author":"Raven","year":"1998","journal-title":"Funct Ecol"},{"key":"2026100200262314300_ref76","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1127\/archiv-hydrobiol\/131\/1994\/175","article-title":"Sedimentation of vernal and late summer phytoplankton communities in the coastal Baltic Sea","volume":"131","author":"Heiskanen","year":"1994","journal-title":"Arch F\u00fcr Hydrobiol"},{"key":"2026100200262314300_ref77","doi-asserted-by":"publisher","first-page":"404","DOI":"10.3389\/fmars.2020.00404","article-title":"Seasonal variability in benthic\u2013pelagic coupling: quantifying organic matter inputs to the seafloor and benthic macrofauna using a multi-marker approach","volume":"7","author":"Rodil","year":"2020","journal-title":"Front Mar Sci"},{"key":"2026100200262314300_ref78","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1007\/BF00386803","article-title":"Sinking rates of natural copepod fecal pellets","volume":"51","author":"Small","year":"1979","journal-title":"Mar Biol"}],"container-title":["The ISME Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/ismej\/advance-article-pdf\/doi\/10.1093\/ismejo\/wrag214\/70821003\/wrag214.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismej\/article-pdf\/20\/1\/wrag214\/70821003\/wrag214.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismej\/article-pdf\/20\/1\/wrag214\/70821003\/wrag214.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T04:26:28Z","timestamp":1790915188000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/ismej\/article\/doi\/10.1093\/ismejo\/wrag214\/8772006"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1]]},"references-count":78,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,1,14]]}},"URL":"https:\/\/doi.org\/10.1093\/ismejo\/wrag214","relation":{},"ISSN":["1751-7362","1751-7370"],"issn-type":[{"value":"1751-7362","type":"print"},{"value":"1751-7370","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,1]]},"published":{"date-parts":[[2026,1]]},"article-number":"wrag214"}}