Articles | Volume 22, issue 23
https://doi.org/10.5194/bg-22-7563-2025
© Author(s) 2025. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Special issue:
https://doi.org/10.5194/bg-22-7563-2025
© Author(s) 2025. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Research article
| 
03 Dec 2025
Research article |
| 03 Dec 2025
| 03 Dec 2025
Biogeochemical dynamics of the sea-surface microlayer in a multidisciplinary mesocosm study
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Mariana Ribas-Ribas
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Leonie Jaeger
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Carola Lehners
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Lisa Gassen
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Edgar Fernando Cortés-Espinoza
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Jochen Wollschläger
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Claudia Thölen
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Hannelore Waska
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Jasper Zöbelein
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Thorsten Brinkhoff
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Isha Athale
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
Rüdiger Röttgers
Helmholtz-Zentrum Hereon, Institute of Carbon Cycles, Geesthacht, Germany
Michael Novak
Helmholtz-Zentrum Hereon, Institute of Carbon Cycles, Geesthacht, Germany
Anja Engel
GEOMAR Helmholtz Centre for Ocean Research Kiel, Kiel, Germany
Theresa Barthelmeß
GEOMAR Helmholtz Centre for Ocean Research Kiel, Kiel, Germany
Josefine Karnatz
GEOMAR Helmholtz Centre for Ocean Research Kiel, Kiel, Germany
Thomas Reinthaler
Department of Functional and Evolutionary Ecology, University of Vienna, Vienna, Austria
Dmytro Spriahailo
Department of Functional and Evolutionary Ecology, University of Vienna, Vienna, Austria
Gernot Friedrichs
Institute of Physical Chemistry, Kiel University, Kiel, Germany
Falko Asmussen Schäfer
Institute of Physical Chemistry, Kiel University, Kiel, Germany
Oliver Wurl
Institute for Chemistry and Biology of the Marine Environment (ICBM), School of Mathematics and Science, Carl von Ossietzky Universität Oldenburg, Oldenburg, Germany
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This paper presents the fourth version of the ESA Ocean Colour Climate Change Initiative global in situ bio-optical database. The updated compilation combines observations from many international data sources collected between 1996 and 2025 and includes major expansions in coverage and variables. The dataset provides a consistent reference for validating satellite ocean-colour products and supports algorithm development and long-term climate studies.
Samuel Mintah Ayim and Oliver Wurl
EGUsphere, https://doi.org/10.5194/egusphere-2026-4866, https://doi.org/10.5194/egusphere-2026-4866, 2026
This preprint is open for discussion and under review for Ocean Science (OS).
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We investigated whether small-scale changes in seawater properties at the ocean surface reflect those below. An autonomous surface vehicle measured water density at the ultra-thin skin layer and one meter deep in the German Bight. Skin density patterns frequently differed from the underlying water and were driven more by salinity than temperature. This suggests that satellite observations of the ocean skin may not accurately represent underlying water dynamics in complex coastal waters.
Sophia Hirschmann, Anabel von Jackowski, Benjamin Pontiller, Katja Metfies, Vasily Povazhnyi, Stefan Neuhaus, Eva-Maria Nöthig, and Anja Engel
EGUsphere, https://doi.org/10.5194/egusphere-2026-4192, https://doi.org/10.5194/egusphere-2026-4192, 2026
This preprint is open for discussion and under review for Biogeosciences (BG).
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Michelle Albinus, Thomas H. Badewien, Lisa Gassen, Oliver Wurl, and Jens Meyerjürgens
Ocean Sci., 22, 2143–2160, https://doi.org/10.5194/os-22-2143-2026, https://doi.org/10.5194/os-22-2143-2026, 2026
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Biogeosciences, 23, 4145–4169, https://doi.org/10.5194/bg-23-4145-2026, https://doi.org/10.5194/bg-23-4145-2026, 2026
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Biogeosciences, 23, 3697–3721, https://doi.org/10.5194/bg-23-3697-2026, https://doi.org/10.5194/bg-23-3697-2026, 2026
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Biogeosciences, 23, 3517–3539, https://doi.org/10.5194/bg-23-3517-2026, https://doi.org/10.5194/bg-23-3517-2026, 2026
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Anja Engel, Gernot Friedrichs, Kerstin E. Krall, and Bernd Jähne
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Martin Hieronymi, Daniel Behr, Shun Bi, and Rüdiger Röttgers
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Lin Yang, Bin Yang, Jing Zhang, Anja Engel, and Gui-Peng Yang
Biogeosciences, 23, 1261–1278, https://doi.org/10.5194/bg-23-1261-2026, https://doi.org/10.5194/bg-23-1261-2026, 2026
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Carbon monoxide (CO), chromophoric dissolved organic matter (CDOM), and fluorescent dissolved organic matter (FDOM) were more frequently enriched in the higher temperature and salinity off-shore regions. Marine-humic like CDOM tends to inhibit the sea-to-air flux of CO in the sea-surface microlayer (SML). The enrichment and photochemical process of CO in the SML were more active during the daytime. The photochemical production and microbial consumption rates of CO in the SML were more active than in the subsurface layer (SSW).
Carsten Rauch, Lisa Deyle, Leonie Jaeger, Edgar Fernando Cortés-Espinoza, Mariana Ribas-Ribas, Josefine Karnatz, Anja Engel, and Oliver Wurl
Ocean Sci., 22, 403–426, https://doi.org/10.5194/os-22-403-2026, https://doi.org/10.5194/os-22-403-2026, 2026
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Microsensors measuring oxygen and temperature were used to gain high-resolution profiles across the surface of a water basin, in which an algal bloom was induced. These novel data show that the oxygen at the sea surface is highly influenced by algal blooms, while the temperature is only indirectly affected by them. Since algal blooms occur globally, this has considerable implications for calculating global air-sea exchanges of gases or heat, especially under low-wind conditions.
Lina A. Holthusen, Hermann W. Bange, Thomas H. Badewien, Julia C. Muchowski, Tina Santl-Temkiv, Jennie Spicker Schmidt, Oliver Wurl, and Damian L. Arévalo-Martínez
The Cryosphere, 20, 535–550, https://doi.org/10.5194/tc-20-535-2026, https://doi.org/10.5194/tc-20-535-2026, 2026
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Ander López-Puertas, Oliver Wurl, Sanja Frka, and Mariana Ribas-Ribas
Ocean Sci., 21, 3471–3485, https://doi.org/10.5194/os-21-3471-2025, https://doi.org/10.5194/os-21-3471-2025, 2025
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We studied how daily cycles affect inorganic carbon variables in the ocean's surface microlayer. Using data from three full days and nights off the Croatian coast, we found that thermohaline properties and key indicators like pH and pCO₂ change significantly from day to night. Ignoring nighttime conditions may lead to global carbon budget errors and highlights the need for continuous ocean observations.
Edgar Fernando Cortés-Espinoza, Alisa Wüst, Ander Lopéz-Puertas, Oliver Wurl, José Martín Hernández-Ayón, Hannelore Waska, and Mariana Ribas-Ribas
EGUsphere, https://doi.org/10.5194/egusphere-2025-5265, https://doi.org/10.5194/egusphere-2025-5265, 2025
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In a mesocosm experiment, we measured carbon system variables to examine how organic matter accumulation increases organic alkalinity and alters seawater pH in the sea-surface microlayer and underlying water during an induced phytoplankton bloom. Organic alkalinity was consistently higher in the sea-surface microlayer, and during bloom peak its pH effect extended into the underlying water, reducing and sometimes reversing surface–subsurface pH differences.
Lisa Gassen, Samuel M. Ayim, Leonie Jaeger, Jens Meyerjürgens, Mariana Ribas-Ribas, and Oliver Wurl
Ocean Sci., 21, 2787–2804, https://doi.org/10.5194/os-21-2787-2025, https://doi.org/10.5194/os-21-2787-2025, 2025
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This study investigates how abrupt weather changes, such as shifts in air temperature, wind speed and precipitation, impact temperature and salinity in the ocean’s skin layer (upper first millimetre). Two events in the harbour of Bremerhaven and one event in the North Sea revealed that the skin layer reacts instantly, with greater temperature changes than those at a depth of 100 cm, underscoring its key role in air-sea interactions and climate dynamics.
Vlad A. Macovei, Louise C. V. Rewrie, Rüdiger Röttgers, and Yoana G. Voynova
Biogeosciences, 22, 3375–3396, https://doi.org/10.5194/bg-22-3375-2025, https://doi.org/10.5194/bg-22-3375-2025, 2025
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We found that biogeochemical variability at the land–sea interface (LSI) in two major temperate estuaries is modulated by the 14 d spring–neap tidal cycle, with large effects on dissolved inorganic and organic carbon concentrations and distribution. As this effect increases the strength of the carbon source to the atmosphere by up to 74 % during spring tide, it should be accounted for in regional models, which aim to resolve biogeochemical processing at the LSI.
Anisbel Leon-Marcos, Moritz Zeising, Manuela van Pinxteren, Sebastian Zeppenfeld, Astrid Bracher, Elena Barbaro, Anja Engel, Matteo Feltracco, Ina Tegen, and Bernd Heinold
Geosci. Model Dev., 18, 4183–4213, https://doi.org/10.5194/gmd-18-4183-2025, https://doi.org/10.5194/gmd-18-4183-2025, 2025
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This study represents the primary marine organic aerosol (PMOA) emissions, focusing on their sea–atmosphere transfer. Using the FESOM2.1–REcoM3 model, concentrations of key organic biomolecules were estimated and integrated into the ECHAM6.3–HAM2.3 aerosol–climate model. Results highlight the influence of marine biological activity and surface winds on PMOA emissions, with reasonably good agreement with observations improving aerosol representation in the southern oceans.
Lisa Deyle, Thomas H. Badewien, Oliver Wurl, and Jens Meyerjürgens
Earth Syst. Sci. Data, 16, 2099–2112, https://doi.org/10.5194/essd-16-2099-2024, https://doi.org/10.5194/essd-16-2099-2024, 2024
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A dataset from the North Sea of 85 surface drifters from 2017–2021 is presented. Surface drifters enable the analysis of ocean currents by determining the velocities of surface currents and tidal effects. The entire North Sea has not been studied using drifters before, but the analysis of ocean currents is essential, e.g., to understand the pathways of plastic. The results show that there are strong tidal effects in the shallow North Sea area and strong surface currents in the deep areas.
Karine Sellegri, Theresa Barthelmeß, Jonathan Trueblood, Antonia Cristi, Evelyn Freney, Clémence Rose, Neill Barr, Mike Harvey, Karl Safi, Stacy Deppeler, Karen Thompson, Wayne Dillon, Anja Engel, and Cliff Law
Atmos. Chem. Phys., 23, 12949–12964, https://doi.org/10.5194/acp-23-12949-2023, https://doi.org/10.5194/acp-23-12949-2023, 2023
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The amount of sea spray emitted to the atmosphere depends on the ocean temperature, but this dependency is not well understood, especially when ocean biology is involved. In this study, we show that sea spray emissions are increased by up to a factor of 4 at low seawater temperatures compared to moderate temperatures, and we quantify the temperature dependence as a function of the ocean biogeochemistry.
Shungudzemwoyo P. Garaba, Michelle Albinus, Guido Bonthond, Sabine Flöder, Mario L. M. Miranda, Sven Rohde, Joanne Y. L. Yong, and Jochen Wollschläger
Earth Syst. Sci. Data, 15, 4163–4179, https://doi.org/10.5194/essd-15-4163-2023, https://doi.org/10.5194/essd-15-4163-2023, 2023
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These high-quality data document a harmful algal bloom dominated by Nodularia spumigena, a cyanobacterium that has been recurring in waters around the world, using advanced water observation technologies. We also showcase the benefits of experiments of opportunity and the issues with obtaining synoptic spatio-temporal data for monitoring water quality. The dataset can be leveraged to gain more knowledge on related blooms, develop detection algorithms and optimize future monitoring efforts.
Manon Rocco, Erin Dunne, Alexia Saint-Macary, Maija Peltola, Theresa Barthelmeß, Neill Barr, Karl Safi, Andrew Marriner, Stacy Deppeler, James Harnwell, Anja Engel, Aurélie Colomb, Alfonso Saiz-Lopez, Mike Harvey, Cliff S. Law, and Karine Sellegri
EGUsphere, https://doi.org/10.5194/egusphere-2023-516, https://doi.org/10.5194/egusphere-2023-516, 2023
Preprint archived
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During the Sea2cloud campaign in the Southern Pacific Ocean, we measured air-sea emissions from phytopankton of two key atmospheric compounds: DMS and MeSH. These compounds are well-known to play a great role in atmospheric chemistry and climate. We see in this paper that these compounds are most emited by the nanophytoplankton population. We provide here parameters for climate models to predict future trends of the emissions of these compounds and their roles and impacts on the global warming.
Damian L. Arévalo-Martínez, Amir Haroon, Hermann W. Bange, Ercan Erkul, Marion Jegen, Nils Moosdorf, Jens Schneider von Deimling, Christian Berndt, Michael Ernst Böttcher, Jasper Hoffmann, Volker Liebetrau, Ulf Mallast, Gudrun Massmann, Aaron Micallef, Holly A. Michael, Hendrik Paasche, Wolfgang Rabbel, Isaac Santos, Jan Scholten, Katrin Schwalenberg, Beata Szymczycha, Ariel T. Thomas, Joonas J. Virtasalo, Hannelore Waska, and Bradley A. Weymer
Biogeosciences, 20, 647–662, https://doi.org/10.5194/bg-20-647-2023, https://doi.org/10.5194/bg-20-647-2023, 2023
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Groundwater flows at the land–ocean transition and the extent of freshened groundwater below the seafloor are increasingly relevant in marine sciences, both because they are a highly uncertain term of biogeochemical budgets and due to the emerging interest in the latter as a resource. Here, we discuss our perspectives on future research directions to better understand land–ocean connectivity through groundwater and its potential responses to natural and human-induced environmental changes.
Alexia D. Saint-Macary, Andrew Marriner, Theresa Barthelmeß, Stacy Deppeler, Karl Safi, Rafael Costa Santana, Mike Harvey, and Cliff S. Law
Ocean Sci., 19, 1–15, https://doi.org/10.5194/os-19-1-2023, https://doi.org/10.5194/os-19-1-2023, 2023
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The uppermost oceanic layer was sampled to determine what can explain a potential dimethyl sulfide (DMS) enrichment in this environment. A novel sampling method was used, and the results showed that DMS was not as enriched as expected. Our results showed that the phytoplanktonic composition influenced the DMS concentration, confirming results from another study in this oceanic region. However, additional factors are required to observe a DMS enrichment in the uppermost oceanic layer.
André Valente, Shubha Sathyendranath, Vanda Brotas, Steve Groom, Michael Grant, Thomas Jackson, Andrei Chuprin, Malcolm Taberner, Ruth Airs, David Antoine, Robert Arnone, William M. Balch, Kathryn Barker, Ray Barlow, Simon Bélanger, Jean-François Berthon, Şükrü Beşiktepe, Yngve Borsheim, Astrid Bracher, Vittorio Brando, Robert J. W. Brewin, Elisabetta Canuti, Francisco P. Chavez, Andrés Cianca, Hervé Claustre, Lesley Clementson, Richard Crout, Afonso Ferreira, Scott Freeman, Robert Frouin, Carlos García-Soto, Stuart W. Gibb, Ralf Goericke, Richard Gould, Nathalie Guillocheau, Stanford B. Hooker, Chuamin Hu, Mati Kahru, Milton Kampel, Holger Klein, Susanne Kratzer, Raphael Kudela, Jesus Ledesma, Steven Lohrenz, Hubert Loisel, Antonio Mannino, Victor Martinez-Vicente, Patricia Matrai, David McKee, Brian G. Mitchell, Tiffany Moisan, Enrique Montes, Frank Muller-Karger, Aimee Neeley, Michael Novak, Leonie O'Dowd, Michael Ondrusek, Trevor Platt, Alex J. Poulton, Michel Repecaud, Rüdiger Röttgers, Thomas Schroeder, Timothy Smyth, Denise Smythe-Wright, Heidi M. Sosik, Crystal Thomas, Rob Thomas, Gavin Tilstone, Andreia Tracana, Michael Twardowski, Vincenzo Vellucci, Kenneth Voss, Jeremy Werdell, Marcel Wernand, Bozena Wojtasiewicz, Simon Wright, and Giuseppe Zibordi
Earth Syst. Sci. Data, 14, 5737–5770, https://doi.org/10.5194/essd-14-5737-2022, https://doi.org/10.5194/essd-14-5737-2022, 2022
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A compiled set of in situ data is vital to evaluate the quality of ocean-colour satellite data records. Here we describe the global compilation of bio-optical in situ data (spanning from 1997 to 2021) used for the validation of the ocean-colour products from the ESA Ocean Colour Climate Change Initiative (OC-CCI). The compilation merges and harmonizes several in situ data sources into a simple format that could be used directly for the evaluation of satellite-derived ocean-colour data.
Lin Yang, Jing Zhang, Anja Engel, and Gui-Peng Yang
Biogeosciences, 19, 5251–5268, https://doi.org/10.5194/bg-19-5251-2022, https://doi.org/10.5194/bg-19-5251-2022, 2022
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Enrichment factors of dissolved organic matter (DOM) in the eastern marginal seas of China exhibited a significant spatio-temporal variation. Photochemical and enrichment processes co-regulated DOM enrichment in the sea-surface microlayer (SML). Autochthonous DOM was more frequently enriched in the SML than terrestrial DOM. DOM in the sub-surface water exhibited higher aromaticity than that in the SML.
Quentin Devresse, Kevin W. Becker, Arne Bendinger, Johannes Hahn, and Anja Engel
Biogeosciences, 19, 5199–5219, https://doi.org/10.5194/bg-19-5199-2022, https://doi.org/10.5194/bg-19-5199-2022, 2022
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Eddies are ubiquitous in the ocean and alter physical, chemical, and biological processes. However, how they affect organic carbon production and consumption is largely unknown. Here we show how an eddy triggers a cascade effect on biomass production and metabolic activities of phyto- and bacterioplankton. Our results may contribute to the improvement of biogeochemical models used to estimate carbon fluxes in the ocean.
Theresa Barthelmeß and Anja Engel
Biogeosciences, 19, 4965–4992, https://doi.org/10.5194/bg-19-4965-2022, https://doi.org/10.5194/bg-19-4965-2022, 2022
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Greenhouse gases released by human activity cause a global rise in mean temperatures. While scientists can predict how much of these gases accumulate in the atmosphere based on not only human-derived sources but also oceanic sinks, it is rather difficult to predict the major influence of coastal ecosystems. We provide a detailed study on the occurrence, composition, and controls of substances that suppress gas exchange. We thus help to determine what controls coastal greenhouse gas fluxes.
Manuela van Pinxteren, Tiera-Brandy Robinson, Sebastian Zeppenfeld, Xianda Gong, Enno Bahlmann, Khanneh Wadinga Fomba, Nadja Triesch, Frank Stratmann, Oliver Wurl, Anja Engel, Heike Wex, and Hartmut Herrmann
Atmos. Chem. Phys., 22, 5725–5742, https://doi.org/10.5194/acp-22-5725-2022, https://doi.org/10.5194/acp-22-5725-2022, 2022
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A class of marine particles (transparent exopolymer particles, TEPs) that is ubiquitously found in the world oceans was measured for the first time in ambient marine aerosol particles and marine cloud waters in the tropical Atlantic Ocean. TEPs are likely to have good properties for influencing clouds. We show that TEPs are transferred from the ocean to the marine atmosphere via sea-spray formation and our results suggest that they can also form directly in aerosol particles and in cloud water.
Tristan Petit, Børge Hamre, Håkon Sandven, Rüdiger Röttgers, Piotr Kowalczuk, Monika Zablocka, and Mats A. Granskog
Ocean Sci., 18, 455–468, https://doi.org/10.5194/os-18-455-2022, https://doi.org/10.5194/os-18-455-2022, 2022
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We provide the first insights on bio-optical processes in Storfjorden (Svalbard). Information on factors controlling light propagation in the water column in this arctic fjord becomes crucial in times of rapid sea ice decline. We find a significant contribution of dissolved matter to light absorption and a subsurface absorption maximum linked to phytoplankton production. Dense bottom waters from sea ice formation carry elevated levels of dissolved and particulate matter.
France Van Wambeke, Vincent Taillandier, Karine Desboeufs, Elvira Pulido-Villena, Julie Dinasquet, Anja Engel, Emilio Marañón, Céline Ridame, and Cécile Guieu
Biogeosciences, 18, 5699–5717, https://doi.org/10.5194/bg-18-5699-2021, https://doi.org/10.5194/bg-18-5699-2021, 2021
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Simultaneous in situ measurements of (dry and wet) atmospheric deposition and biogeochemical stocks and fluxes in the sunlit waters of the open Mediterranean Sea revealed complex physical and biological processes occurring within the mixed layer. Nitrogen (N) budgets were computed to compare the sources and sinks of N in the mixed layer. The transitory effect observed after a wet dust deposition impacted the microbial food web down to the deep chlorophyll maximum.
Frédéric Gazeau, France Van Wambeke, Emilio Marañón, Maria Pérez-Lorenzo, Samir Alliouane, Christian Stolpe, Thierry Blasco, Nathalie Leblond, Birthe Zäncker, Anja Engel, Barbara Marie, Julie Dinasquet, and Cécile Guieu
Biogeosciences, 18, 5423–5446, https://doi.org/10.5194/bg-18-5423-2021, https://doi.org/10.5194/bg-18-5423-2021, 2021
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Our study shows that the impact of dust deposition on primary production depends on the initial composition and metabolic state of the tested community and is constrained by the amount of nutrients added, to sustain both the fast response of heterotrophic prokaryotes and the delayed one of phytoplankton. Under future environmental conditions, heterotrophic metabolism will be more impacted than primary production, therefore reducing the capacity of surface waters to sequester anthropogenic CO2.
Cited articles
Balch, W. M.: The ecology, biogeochemistry, and optical properties of coccolithophores, Annu. Rev. Mar. Sci., 10, 71–98, https://doi.org/10.1146/annurev-marine-121916-063319, 2018.
Bauer, H.-U., Herrmann, M., and Villmann, T.: Neural maps and topographic vector quantization, Neural. Netw., 12, 659–676, https://doi.org/10.1016/S0893-6080(99)00027-1,1999.
Barthelmeß, T. and Engel, A.: How biogenic polymers control surfactant dynamics in the surface microlayer: insights from a coastal Baltic Sea study, Biogeosciences, 19, 4965–4992, https://doi.org/10.5194/bg-19-4965-2022, 2022.
Barthelmeß, T., Schütte, F., and Engel, A.: Variability of the sea surface microlayer across a filament's edge and potential influences on gas exchange, Front. Mar. Sci., 8, 718384, https://doi.org/10.3389/fmars.2021.718384, 2021.
Behrenfeld, M. J., Boss, E., Siegel, D. A., and Shea, D. M.: Carbon-based ocean productivity and phytoplankton physiology from space, Global Biogeochem. Cycles., 19, 1–14, https://doi.org/10.1029/2004GB002299, 2005.
Bibi, R., Kang, H. Y., Kim, D., Jang, J., Kundu, G. K., Kim, Y. K., and Kang, C.-K.: Dominance of autochthonous phytoplankton-derived particulate organic matter in a low-turbidity temperate estuarine embayment, Gwangyang Bay, Korea, Front. Mar. Sci., 7, 580260, https://doi.org/10.3389/fmars.2020.580260, 2020.
Bibi, R., Ribas-Ribas, M., Jaeger, L., Lehners, C., Gassen, L., Cortés, E., Wollschläger, J., Thölen, C., Waska, H., Zöbelein, J., Brinkhoff, T., Athale, I., Röttgers, R., Novak, M., Engel, A., Barthelmeß, T., Karnatz, J., Reinthaler, T., Spriahailo, D., Friedrichs, G., Schäfer, F., and Wurl, O.: Physical, chemical, and biogeochemical parameters from a mesocosm experiment at the Sea Surface Facility (SURF), Wilhelmshaven, Germany, spring 2023, PANGAEA [dataset], https://doi.org/10.1594/PANGAEA.984101, 2025.
Brockmann, U. H., Huhnerfuss, H., Kattner, G., Broecker, H. C., and Hentzschel, G.: Artificial surface films in the sea area near Sylt 1, Limnol. Oceanogr., 27, 1050–1058, https://doi.org/10.4319/lo.1982.27.6.1050, 1982.
Broecker, H.-C., Petermann, J., and Siems, W.: The influence of wind on CO2-exchange in a wind-wave tunnel, including the effects of monolayers, J. Mar. Res., 36, 595–610, 1978.
Brussaard, C. P., Payet, J. P., Winter, C., and Weinbauer, M. G.: Quantification of aquatic viruses by flow cytometry, Man. Aquat. Viral Ecol., 11, 102–109, 2010.
Brussaard, C. P. D., Gast, G. J., van Duyl, F. C., and Riegman, R.: Impact of phytoplankton bloom magnitude on a pelagic microbial food web, Mar. Ecol. Prog. Ser., 144, 211–221, https://doi.org/10.3354/meps144211, 1996.
Buchan, A., LeCleir, G. R., Gulvik, C. A., and González, J. M.: Master recyclers: features and functions of bacteria associated with phytoplankton blooms, Nat. Rev. Microbiol., 12, 686–698, https://doi.org/10.1038/nrmicro3326, 2014.
Burson, A., Stomp, M., Akil, L., Brussaard, C. P., and Huisman, J.: Unbalanced reduction of nutrient loads has created an offshore gradient from phosphorus to nitrogen limitation in the North Sea: Offshore gradient from P to N limitation, Limnol. Oceanogr., 61, 869–888, https://doi.org/10.1002/lno.10257,2016.
Bunse, C., Bertos-Fortis, M., Sassenhagen, I., Sildever, S., Sjöqvist, C., Godhe, A., Gross, S., Kremp, A., Lips, I., Lundholm, N., Rengefors, K., Sefbom, J., Pinhassi, J., and Legrand, C.: Spatio-temporal interdependence of bacteria and phytoplankton during a Baltic Sea Spring bloom, Front. Microbiol., 7, 517, https://doi.org/10.3389/fmicb.2016.00517, 2016.
Carlson, D. J.: A field evaluation of plate and screen microlayer sampling techniques, Mar. Chem., 11, 189–208, https://doi.org/10.1016/0304-4203(82)90015-9, 1982.
Charrad, M., Ghazzali, N., Boiteau, V., and Niknafs, A.: NbClust: an R package for determining the relevant number of clusters in a data set, Journal of Statistical Software, 61, 1–36, https://doi.org/10.18637/jss.v061.i06, 2014.
Cifuentes, L. A., Sharp, J. H., and Fogel, M. L.: Stable carbon and nitrogen isotope biogeochemistry in the Delaware estuary, Limnol. Oceanogr., 33, 1102–1115, https://doi.org/10.4319/lo.1988.33.5.1102,1988.
Ciuraru, R., Fine, L., van Pinxteren, M., D'Anna, B., Herrmann, H., and George, C.: Unravelling new processes at interfaces: photochemical isoprene production at the sea surface, Environ. Sci. Technol., 49, 13199–13205, https://doi.org/10.1021/acs.est.5b02388, 2015.
Clark, C. D. and Zika, R. G.: Marine organic photochemistry: from the sea surface to marine aerosols, Mar. Chem., 1–33, https://doi.org/10.1007/10683826_1, 2000.
Cloern, J. E., Grenz, C., and Vidergar-Lucas, L.: An empirical model of the phytoplankton chlorophyll: carbon ratio-the conversion factor between productivity and growth rate, Limnol. Oceanogr., 40, 1313–1321, https://doi.org/10.4319/lo.1995.40.7.1313, 1995.
Ćosović, B.: Direct determination of surface-active substances in natural waters, Mar. Chem., 22, 363–373, https://doi.org/10.1016/0304-4203(87)90020-X, 1987.
Cunliffe, M. and Murrell, J. C.: The sea-surface microlayer is a gelatinous biofilm, ISME J., 3, 1001–1003, https://doi.org/10.1038/ismej.2009.69, 2009.
Cunliffe, M. and Murrell, J. C.: Eukarya 18S rRNA gene diversity in the sea surface microlayer: implications for the structure of the neustonic microbial loop, ISME J., 4, 455–458, https://doi.org/10.1038/ismej.2009.133, 2010.
Cunliffe, M., Whiteley, A. S., Newbold, L., Oliver, A., Schäfer, H., and Murrell, J. C.: Comparison of bacterioneuston and bacterioplankton dynamics during a phytoplankton bloom in a fjord mesocosm, Appl. Environ. Microbiol., 75, 7173–7181, https://doi.org/10.1128/AEM.01374-09, 2009.
Cunliffe, M., Upstill-Goddard, R. C., and Murrell, J. C.: Microbiology of aquatic surface microlayers, FEMS Microbiol. Reviews, 35, 233–246, https://doi.org/10.1111/j.1574-6976.2010.00246.x, 2011.
Cunliffe, M., Engel, A., Frka, S., Gašparović, B., Guitart, C., Murrell, J. C., Salter, M., Stolle, C., Upstill-Goddard, R., and Wurl, O.: Sea surface microlayers: A unified physicochemical and biological perspective of the air–ocean interface, Prog. Oceanogr., 109, 104–116, https://doi.org/10.1016/j.pocean.2012.08.004, 2013.
Doane, T. A. and Horwáth, W. R.: Spectrophotometric determination of nitrate with a single reagent, Anal. Lett., 36, 2713–2722, https://doi.org/10.1081/AL-120024647, 2003.
Ellison, G. B., Tuck, A. F., and Vaida, V.: Atmospheric processing of organic aerosols, J. Geophys. Res. Atmos., 104, 11633–11641, https://doi.org/10.1029/1999JD900073, 1999.
Engel, A. and Galgani, L.: The organic sea-surface microlayer in the upwelling region off the coast of Peru and potential implications for air–sea exchange processes, Biogeosciences, 13, 989–1007, https://doi.org/10.5194/bg-13-989-2016, 2016.
Engel, A., Sperling, M., Sun, C., Grosse, J., and Friedrichs, G.: Organic matter in the surface microlayer: Insights from a wind wave channel experiment, Front. Mar. Sci., 5, 182, https://doi.org/10.3389/fmars.2018.00182, 2018.
Fanning, K. A. and Pilson, M.: On the spectrophotometric determination of dissolved silica in natural waters, Anal. Chem., 45, 136–140, https://doi.org/10.1021/ac60323a021, 1973.
Ferderer, A., Schulz, K. G., Riebesell, U., Baker, K. G., Chase, Z., and Bach, L. T.: Investigating the effect of silicate- and calcium-based ocean alkalinity enhancement on diatom silicification, Biogeosciences, 21, 2777–2794, https://doi.org/10.5194/bg-21-2777-2024, 2024.
Finkel, Z. V.: Light absorption and size scaling of light-limited metabolism in marine diatoms, Limnol. Oceanogr., 46, 86–94, https://doi.org/10.4319/lo.2001.46.1.0086, 2001.
Franklin, M. P., McDonald, I. R., Bourne, D. G., Owens, N. J., Upstill-Goddard, R. C., and Murrell, J. C.: Bacterial diversity in the bacterioneuston (sea surface microlayer): the bacterioneuston through the looking glass, Environ. Microbiol., 7, 723–736, https://doi.org/10.1111/j.1462-2920.2004.00736.x, 2005.
Gade, M., Byfield, V., Ermakov, S., Lavrova, O., and Mitnik, L.: Slicks as indicators for marine processes, Oceanogr., 26, 138–149, https://doi.org/10.5670/oceanog.2013.39, 2013.
Galgani, L., Stolle, C., Endres, S., Schulz, K. G., and Engel, A.: Effects of ocean acidification on the biogenic composition of the sea-surface microlayer: Results from a mesocosm study, J. Geophys. Res.-Oceans., 119, 7911–7924, https://doi.org/10.1002/2014JC010188, 2014.
Galili, T.: dendextend: an R package for visualizing, adjusting and comparing trees of hierarchical clustering, Bioinformatics, 31, 3718–3720, https://doi.org/10.1093/bioinformatics/btv428, 2015.
Garland, J. L.: Analytical approaches to the characterization of samples of microbial communities using patterns of potential C source utilization, Soil. Biolog. Biochem., 28, 213–221, https://doi.org/10.1016/0038-0717(95)00112-3, 1996.
Garland, J. L. and Mills, A. L.: Classification and characterization of heterotrophic microbial communities on the basis of patterns of community-level sole-carbon-source utilization, Appl. Environ. Microbiol., 57, 2351–2359, 10.1128/aem.57.8.2351-2359.1991, 1991.
Gassen, L., Esters, L., Ribas-Ribas, M., and Wurl, O.: The impact of rainfall on the sea surface salinity: a mesocosm study, Sci Rep., 14, 6353, https://doi.org/10.1038/s41598-024-56915-4, 2024.
Geider, R. J.: Light and temperature dependence of the carbon to chlorophyll a ratio in microalgae and cyanobacteria: implications for physiology and growth of phytoplankton, New Phytol., 106, 1–34, https://doi.org/10.1111/j.1469-8137.1987.tb04788.x, 1987.
Geider, R. J., MacIntyre, H. L., and Kana, T. M.: Dynamic model of phytoplankton growth and acclimation: responses of the balanced growth rate and the chlorophyll a: carbon ratio to light, nutrient limitation, and temperature, Mar. Ecol. Prog. Ser., 148, 187–200, https://doi.org/10.3354/meps148187, 1997.
Geider, R. J., MacIntyre, H. L., and Kana, T. M.: A dynamic regulatory model of phytoplanktonic acclimation to light, nutrients, and temperature, Limnol. Oceanogr., 43, 679–694, https://doi.org/10.4319/lo.1998.43.4.0679, 1998.
Goldman, J. C., McCarthy, J. J., and Peavey, D. G.: Growth rate influence on the chemical composition of phytoplankton in oceanic waters, Nature, 279, 210–215, https://doi.org/10.1038/279210a0, 1979.
Gordon, H. R., Smyth, T. J., Balch, W. M., Boynton, G. C., and Tarran, G. A.: Light scattering by coccoliths detached from Emiliania huxleyi, Appl. Optics, 48, 6059–6073, 10.1364/AO.48.006059, 2009.
González-Gil, R., Banas, N. S., Bresnan, E., and Heath, M. R.: The onset of the spring phytoplankton bloom in the coastal North Sea supports the Disturbance Recovery Hypothesis, Biogeosciences, 19, 2417–2426, https://doi.org/10.5194/bg-19-2417-2022, 2022.
Grizzetti, B., Bouraoui, F., and Aloe, A.: Changes of nitrogen and phosphorus loads to European seas, Glob. Change Biol., 18, 769–782, https://doi.org/10.1111/j.1365-2486.2011.02576.x, 2012.
Guo, X., Wang, Z.-H., Zhao, J.-G., Xiao, L., and Jiang, T.: Effects of inorganic nutrients on the phytoplankton community in the sea surface microlayer of Daya Bay, South China Sea, J. Sea Res., 156, 101830, https://doi.org/10.1016/j.seares.2019.101830, 2020.
Hardy, J. T.: The sea surface microlayer: biology, chemistry and anthropogenic enrichment, Prog. Oceanogr., 11, 307–328, https://doi.org/10.1016/0079-6611(82)90001-5, 1982.
Harvey, G. W. and Burzell, L. A.: A simple microlayer method for small samples 1, Limnol. Oceanogr., 17, 156–157, https://doi.org/10.4319/lo.1972.17.1.0156, 1972.
Hyun, M. J., Choi, D. H., Lee, H., Won, J., Kim, G.-U., Lee, Y., Jeong, J.-Y., Ra, K., Yang, W., Lee, J., Jeong, J., Lee, C. M., and Noh, J. H.: Phytoplankton Spring succession pattern in the Yellow Sea surveyed at Socheongcho Ocean Research Station, Front. Mar. Sci., 10, 1280612, https://doi.org/10.3389/fmars.2023.1280612, 2023.
Jakobsen, H. H. and Markager, S.: Carbon-to-chlorophyll ratio for phytoplankton in temperate coastal waters: seasonal patterns and relationship to nutrients, Limnol. Oceanogr., 61, 1853–1868, https://doi.org/10.1002/lno.10338, 2016.
Jarvis, N., Garrett, W., Scheiman, M., and Timmons, C.: Surface chemical characterization of surface-active material in seawater, Limnol. Oceanogr., 12, 88–96, https://doi.org/10.4319/lo.1967.12.1.0088, 1967.
Jenkinson, I. R., Seuront, L., Ding, H., and Elias, F.: Biological modification of mechanical properties of the sea surface microlayer, influencing waves, ripples, foam and air–sea fluxes, Elem. Sci. Anth., 6, 26, https://doi.org/10.1525/elementa.283, 2018.
Kattner, G.: Storage of dissolved inorganic nutrients in seawater: poisoning with mercuric chloride, Mar. Chem., 67, 61–66, https://doi.org/10.1016/S0304-4203(99)00049-3, 1999.
Kurata, N., Vella, K., Hamilton, B., Shivji, M., Soloviev, A., Matt, S., Tartar, A., and Perrie, W.: Surfactant-associated bacteria in the near-surface layer of the ocean, Sci. Rep., 6, 19123, https://doi.org/10.1038/srep19123, 2016.
Kohonen, T.: Self-Organizing Maps, 3rd edn., Springer, Berlin, https://doi.org/10.1007/978-3-642-56927-2, 2001.
Kohonen, T.: Essentials of the self-organizing map, Neural. Netw., 37, 52–65, https://doi.org/10.1016/j.neunet.2012.09.018, 2013.
Krause, J. W., Schulz, I. K., Rowe, K. A., Dobbins, W., Winding, M. H., Sejr, M. K., Duarte, C. M., and Agustí, S.: Silicic acid limitation drives bloom termination and potential carbon sequestration in an Arctic bloom, Sci. Rep., 9, 8149, https://doi.org/10.1038/s41598-019-44587-4, 2019.
Kuznetsova, M., Lee, C., Aller, J., and Frew, N.: Enrichment of amino acids in the sea surface microlayer at coastal and open ocean sites in the North Atlantic Ocean, Limnol. Oceanogr., 49, 1605–1619, https://doi.org/10.4319/lo.2004.49.5.1605, 2004.
Laskov, C., Herzog, C., Lewandowski, J., and Hupfer, M.: Miniaturized photometrical methods for the rapid analysis of phosphate, ammonium, ferrous iron, and sulfate in pore water of freshwater sediments, Limnol. Oceanogr. Methods., 5, 63–71, https://doi.org/10.4319/lom.2007.5.63, 2007.
Laxague, N. J., Zappa, C. J., Soumya, S., and Wurl, O.: The suppression of ocean waves by biogenic slicks, J. R. Soc. Interface, 21, 20240385, https://doi.org/10.1098/rsif.2024.0385, 2024.
Laß, K. and Friedrichs, G.: Revealing structural properties of the marine nanolayer from vibrational sum frequency generation spectra, J. Geophys. Res., 116, https://doi.org/10.1029/2010JC006609, 2011.
Laß, K., Bange, H. W., and Friedrichs, G.: Seasonal signatures in SFG vibrational spectra of the sea surface nanolayer at Boknis Eck Time Series Station (SW Baltic Sea), Biogeosciences, 10, 5325–5334, https://doi.org/10.5194/bg-10-5325-2013, 2013.
Legendre, P. and Legendre, L.: Numerical Ecology, 3rd edn., vol. 24, Elsevier, Amsterdam, ISBN 978-0-444-53868-0, 2012.
Lessard, E. J., Merico, A., and Tyrrell, T.: Nitrate: phosphate ratios and Emiliania huxleyi blooms, Limnol. Oceanogr., 50, 1020-1024, https://doi.org/10.4319/lo.2005.50.3.1020, 2005.
Liénart, C., Susperregui, N., Rouaud, V., Cavalheiro, J., David, V., Del Amo, Y., Duran, R., Lauga, B., Monperrus, M., Pigot, T., Bichon, S., Charlier, K., and Savoye, N.: Dynamics of particulate organic matter in a coastal system characterized by the occurrence of marine mucilage-A stable isotope study, J. Sea Res., 116, 12–22, https://doi.org/10.1016/j.seares.2016.08.001, 2016.
Logue, J. B., Mouquet, N., Peter, H., and Hillebrand, H.: Empirical approaches to metacommunities: a review and comparison with theory, Trends Ecol. Evol., 26, 482–491, https://doi.org/10.1016/j.tree.2011.04.009, 2011.
Ly, J., Philippart, C. J. M., and Kromkamp, J. C.: Phosphorus limitation during a phytoplankton spring bloom in the western Dutch Wadden Sea, J. Sea Res., 88, 109–120, https://doi.org/10.1016/j.seares.2013.12.010, 2014.
Marzetz, V., Spijkerman, E., Striebel, M., and Wacker, A.: Phytoplankton community responses to interactions between light intensity, light variations, and phosphorus supply, Front. Environ. Sci., 8, 539733, https://doi.org/10.3389/fenvs.2020.539733, 2020.
Miranda, K. M., Espey, M. G., and Wink, D. A.: A rapid, simple spectrophotometric method for simultaneous detection of nitrate and nitrite, Nitric Oxide, 5, 62–71, https://doi.org/10.1006/niox.2000.0319, 2001.
Moore, C. M., Mills, M. M., Langlois, R., Milne, A., Achterberg, E. P., La Roche, J., and Geider, R. J.: Relative influence of nitrogen and phosphorus availability on phytoplankton physiology and productivity in the oligotrophic subtropical North Atlantic Ocean, Limnol. Oceanogr., 53, 291–305, https://doi.org/10.4319/lo.2008.53.1.0291, 2008.
Moore, C. M., Mills, M. M., Arrigo, K. R., Berman-Frank, I., Bopp, L., Boyd, P. W., Galbraith, E. D., Geider, R. J., Guieu, C., Jaccard, S. L., Jickells, T. D., La Roche, J., Lenton, T. M., Mahowald, N. M., Marañón, E., Marinov, I., Moore, J. K., Nakatsuka, T., Oschlies, A., Saito, M. A., Thingstad, T. F., Tsuda, A., and Ulloa, O.: Processes and patterns of oceanic nutrient limitation, Nat. Geosci., 6, 701–710, https://doi.org/10.1038/ngeo1765, 2013.
Mustaffa, N. I. H., Ribas-Ribas, M., and Wurl, O.: High-resolution variability of the enrichment of fluorescence dissolved organic matter in the sea surface microlayer of an upwelling region, Elem. Sci. Anth., 5, 52, https://doi.org/10.1525/elementa.242, 2017.
Mustaffa, N. I. H., Badewien, T. H., Ribas-Ribas, M., and Wurl, O.: High-resolution observations on enrichment processes in the sea-surface microlayer, Sci. Rep., 8, 13122, https://doi.org/10.1038/s41598-018-31465-8, 2018.
Mustaffa, N. I. H., Ribas-Ribas, M., Banko-Kubis, H. M., and Wurl, O.: Global reduction of in situ CO2 transfer velocity by natural surfactants in the sea-surface microlayer, Proc. Math. Phys. Eng. Sci., 476, 20190763, https://doi.org/10.1098/rspa.2019.0763, 2020a.
Mustaffa, N. I. H., Kallajoki, L., Hillebrand, H., Wurl, O., and Striebel, M.: Sea surface phytoplankton community response to nutrient and light changes, Mar. Biol., 167, 1–15, https://doi.org/10.1007/s00227-020-03738-2, 2020b.
Nakajima, R., Tsuchiya, K., Nakatomi, N., Yoshida, T., Tada, Y., Konno, F., Toda, T., Kuwahara, V. S., Hamasaki, K., and Othman, B. H. R.: Enrichment of microbial abundance in the sea-surface microlayer over a coral reef: implications for biogeochemical cycles in reef ecosystems, Mar. Ecol. Prog. Ser., 490, 11–22, https://doi.org/10.3354/meps10481, 2013.
Neukermans, G. and Fournier, G.: Optical modeling of spectral backscattering and remote sensing reflectance from Emiliania huxleyi blooms, Front. Mar. Sci., 5, 146, https://doi.org/10.3389/fmars.2018.00146, 2018.
Novak, G. A. and Bertram, T. H.: Reactive VOC production from photochemical and heterogeneous reactions occurring at the air–ocean interface, Acc. Chem. Res., 53, 1014–1023, 10.1021/acs.accounts.0c00095, 2020.
Obernosterer, I., Catala, P., Lami, R., Caparros, J., Ras, J., Bricaud, A., Dupuy, C., van Wambeke, F., and Lebaron, P.: Biochemical characteristics and bacterial community structure of the sea surface microlayer in the South Pacific Ocean, Biogeosciences, 5, 693–705, https://doi.org/10.5194/bg-5-693-2008, 2008.
Park, Y. S., Céréghino, R., Compin, A., and Lek, S.: Applications of artificial neural networks for patterning and predicting aquatic insect species richness in running waters, Ecol. Model., 160, 265–280, https://doi.org/10.1016/S0304-3800(02)00258-2, 2003.
Passow, U.: Transparent exopolymer particles (TEP) in aquatic environments, Prog. Oceanogr., 55, 287–333, 2002.
Pereira, R., Ashton, I., Sabbaghzadeh, B., Shutler, J. D., and Upstill-Goddard, R. C.: Reduced air–sea CO2 exchange in the Atlantic Ocean due to biological surfactants, Nature Geosci., 11, 492–496, https://doi.org/10.1038/s41561-018-0136-2, 2018.
Pinhassi, J., Sala, M. M., Havskum, H., Peters, F., Guadayol, O., Malits, A., and Marrasé, C.: Changes in bacterioplankton composition under different phytoplankton regimens, Appl. Environ. Microbiol., 70, 6753–6766, https://doi.org/10.1128/AEM.70.11.6753-6766.2004, 2004.
Pérez, M. T., Pausz, C., and Herndl, G. J.: Major shift in bacterioplankton utilization of enantiomeric amino acids between surface waters and the ocean's interior, Limnol. Oceanogr., 48, 755–763, https://doi.org/10.4319/lo.2003.48.2.0755, 2003.
Putland, J. and Iverson, R.: Phytoplankton biomass in a subtropical estuary: distribution, size composition, and carbon: chlorophyll ratios, Estuar. Coasts., 30, 878–885, https://doi.org/10.1007/BF02841341, 2007.
Rahlff, J., Esser, S. P., Plewka, J., Heinrichs, M. E., Soares, A., Scarchilli, C., Grigioni, P., Wex, H., Giebel, H.-A., and Probst, A. J.: Marine viruses disperse bidirectionally along the natural water cycle, Nat. Commun., 14, 6354, https://doi.org/10.1038/s41467-023-42125-5, 2023.
Rauch, C., Deyle, L., Jaeger, L., Cortés-Espinoza, E. F., Ribas-Ribas, M., Karnatz, J., Engel, A., and Wurl, O.: Phytoplankton blooms affect microscale gradients of oxygen and temperature across the sea surface microlayer, EGUsphere [preprint], https://doi.org/10.5194/egusphere-2025-4833, 2025.
Reinthaler, T., Sintes, E., and Herndl, G. J.: Dissolved organic matter and bacterial production and respiration in the sea-surface microlayer of the open Atlantic and the western Mediterranean Sea, Limnol. Oceanogr., 53, 122–136, https://doi.org/10.4319/lo.2008.53.1.0122, 2008.
Ribas-Ribas, M., Mustaffa, N. I. H., Rahlff, J., Stolle, C., and Wurl, O.: Sea Surface Scanner (S3): A catamaran for high-resolution measurements of biogeochemical properties of the sea surface microlayer, Atmos. Oceanic. Technol., 34, 1433–1448, https://doi.org/10.1175/JTECH-D-17-0017.1, 2017.
Ribas-Ribas, M., Helleis, F., Rahlff, J., and Wurl, O.: Air-sea CO2-exchange in a large annular wind-wave tank and the effects of surfactants, Front. Mar. Sci., 5, 457, https://doi.org/10.3389/fmars.2018.00457, 2018.
Riemann, L., Steward, G. F., and Azam, F.: Dynamics of bacterial community composition and activity during a mesocosm diatom bloom, Appl. Environ. Microbiol., 2, 578–87, https://doi.org/10.1128/AEM.66.2.578-587.2000, 2000.
Romano, J.-C.: Sea-surface slick occurrence in the open sea (Mediterranean, Red Sea, Indian Ocean) in relation to wind speed, Deep-Sea Res. Pt. I., 43, 411–423, https://doi.org/10.1016/0967-0637(96)00024-6, 1996.
R Core Team: R: A language and environment for statistical computing, R Foundation for Statistical Computing: Vienna, Austria, https://www.R-project.org/ (last access: 1 December 2025), 2020.
Sabbaghzadeh, B., Upstill-Goddard, R., Beale, R., Pereira, R., and Nightingale, P. D.: The Atlantic Ocean surface microlayer from 50° N to 50° S is ubiquitously enriched in surfactants at wind speeds up to 13 m s−1, Geophys. Res. Lett., 44, 2852–2858, https://doi.org/10.1002/2017GL072988, 2017.
Salter, M., Upstill-Goddard, R., Nightingale, P., Archer, S., Blomquist, B., Ho, D., Huebert, B., Schlosser, P., and Yang, M.: Impact of an artificial surfactant release on air-sea gas fluxes during Deep Ocean Gas Exchange Experiment II, J. Geophys. Res.- Oceans, 116, https://doi.org/10.1029/2011JC007023, 2011.
Sathyendranath, S., Stuart, V., Nair, A., Oka, K., Nakane, T., Bouman, H., Forget, M.-H., Maass, H., and Platt, T.: Carbon-to-chlorophyll ratio and growth rate of phytoplankton in the sea, Mar. Ecol. Prog. Ser., 383, 73–84, https://doi.org/10.3354/meps07998, 2009.
Savoye, N., Aminot, A., Tréguer, P., Fontugne, M., Naulet, N., and Kérouel, R.: Dynamics of particulate organic matter δ15N and δ13C during spring phytoplankton blooms in a macrotidal ecosystem (Bay of Seine, France), Mar. Ecol. Prog. Ser., 255, 27–41, https://doi.org/10.3354/meps255027, 2003.
Schnetger, B. and Lehners, C.: Determination of nitrate plus nitrite in small volume marine water samples using vanadium(III) chloride as a reduction agent, Mar. Chem., 160, 91–98, https://doi.org/10.1016/j.marchem.2014.01.010, 2014.
Sharoni, S. and Halevy, I.: Nutrient ratios in marine particulate organic matter are predicted by the population structure of well-adapted phytoplankton, Sci. Adv., 6, https://doi.org/10.1126/sciadv.aaw9371, 2020.
Sharp, J. H.: Improved analysis for “particulate” organic carbon and nitrogen from seawater, Limnol. Oceanogr., 19, 984–989, https://doi.org/10.4319/lo.1974.19.6.0984, 1974.
Sieburth, J. M., Willis, P.-J., Johnson, K. M., Burney, C. M., Lavoie, D. M., Hinga, K. R., Caron, D. A., French III, F. W., Johnson, P. W., and Davis, P. G.: Dissolved organic matter and heterotrophic microneuston in the surface microlayers of the North Atlantic, Sci., 194, 1415–1418, https://doi.org/10.1126/science.194.4272.1415, 1976.
Stolle, C., Labrenz, M., Meeske, C., and Jürgens, K.: Bacterioneuston community structure in the southern Baltic sea and its dependence on meteorological conditions, Appl. Environ. Microbiol., 77, 3726–3733, https://doi.org/10.1128/AEM.00042-11, 2011.
Smetacek, V. and Cloern, J. E.: Oceans. On phytoplankton trends, Science, 319, 1346–1348, https://doi.org/10.1126/science.1151330, 2008.
Sugimura, Y. and Suzuki, Y.: A high-temperature catalytic oxidation method for the determination of non-volatile dissolved organic carbon in seawater by direct injection of a liquid sample, Mar. Chem., 24, 105–131, https://doi.org/10.1016/0304-4203(88)90043-6, 1988.
Teeling, H., Fuchs, B. M., Becher, D., Klockow, C., Gardebrecht, A., Bennke, C. M., Kassabgy, M., Huang, S., Mann, A. J., Waldmann, J., Weber, M., Klindworth, A., Otto, A., Lange, J., Bernhardt, J., Reinsch, C., Hecker, M., Peplies, J., Bockelmann, F. D., Callies, U., Gerdts, G., Wichels, A., Wiltshire, K. H., Glöckner, F. O., Schweder, T., and Amann, R.: Substrate-controlled succession of marine bacterioplankton populations induced by a phytoplankton bloom, Science, 336, 608–611. https://doi.org/10.1126/science.1218344, 2012.
Thompson, P. A., Guo, M. X., and Harrison, P. J.: Effects of variation in temperature. I. On the biochemical composition of eight species of marine phytoplankton, J. Phycol., 28, 481–488, https://doi.org/10.1111/j.0022-3646.1992.00488.x, 1992.
Thornton, D. C.: Dissolved organic matter (DOM) release by phytoplankton in the contemporary and future ocean, Eur. J. Phycol., 49, 20–46, https://doi.org/10.1080/09670262.2013.875596, 2014.
Tyrrell, T., Holligan, P., and Mobley, C.: Optical impacts of oceanic coccolithophore blooms, J. Geophys. Res.-Oceans, 104, 3223–3241, https://doi.org/10.1029/1998JC900052,1999.
Van Pinxteren, M., Barthel, S., Fomba, K. W., Müller, K., Von Tümpling, W., and Herrmann, H.: The influence of environmental drivers on the enrichment of organic carbon in the sea surface microlayer and in submicron aerosol particles–measurements from the Atlantic Ocean, Elem. Sci. Anth., 5, 35, https://doi.org/10.1525/elementa.225, 2017.
Vesanto, J. and Alhoniemi, E.: Clustering of the self-organizing map, IEEE Trans. Neural. Netw., 11, 586–600, 10.1109/72.846731, 2000.
Wang, X. J., Behrenfeld, M., Le Borgne, R., Murtugudde, R., and Boss, E.: Regulation of phytoplankton carbon to chlorophyll ratio by light, nutrients and temperature in the Equatorial Pacific Ocean: a basin-scale model, Biogeosciences, 6, 391–404, https://doi.org/10.5194/bg-6-391-2009, 2009.
Wang, Z.-H., Song, S.-H., and Qi, Y.-Z.: A comparative study of phytoneuston and the phytoplankton community structure in Daya Bay, South China Sea, J. Sea Res., 85, 474–482, 2014.
Wear, E. K., Carlson, C. A., James, A. K., Brzezinski, M. A., Windecker, L. A., and Nelson, C. E.: Synchronous shifts in dissolved organic carbon bioavailability and bacterial community responses over the course of an upwelling-driven phytoplankton bloom, Limnol. Oceanogr., 60, 657–677, https://doi.org/10.1002/lno.10042, 2015.
Wentzky, V. C., Seifert, L. I., Rinke, K., and Weitere, M.: Seasonal succession of functional traits in phytoplankton communities and their interaction with trophic state, J. Ecol., 108, 1649–1663, https://doi.org/10.1111/1365-2745.13395, 2020.
Wiltshire, K. H., Kraberg, A. C., Bartsch, I., Boersma, M., Franke, H. D., Freund, J., Gebühr, C., Gerdts, G., Stockmann, K., and Wichels, A.: Resilience of North Sea phytoplankton spring bloom dynamics: An analysis of long-term data at Helgoland Roads, Limnol. Oceanogr., 53, 1294–1302, https://doi.org/10.4319/lo.2008.53.4.1294, 2008.
Wurl, O. and Holmes, M.: The gelatinous nature of the sea-surface microlayer, Mar. Chem., 110, 89–97, https://doi.org/10.1016/j.marchem.2008.02.009, 2008.
Wurl, O., Miller, L., Röttgers, R., and Vagle, S.: The distribution and fate of surface-active substances in the sea-surface microlayer and water column, Mar. Chem., 115, 1–9, https://doi.org/10.1016/j.marchem.2009.04.007, 2009.
Wurl, O., Wurl, E., Miller, L., Johnson, K., and Vagle, S.: Formation and global distribution of sea-surface microlayers, Biogeosciences, 8, 121–135, https://doi.org/10.5194/bg-8-121-2011, 2011.
Wurl, O., Stolle, C., Van Thuoc, C., Thu, P. T., and Mari, X.: Biofilm-like properties of the sea surface and predicted effects on air–sea CO2 exchange, Progress. Oceanogr., 144, 15–24, https://doi.org/10.1016/j.pocean.2016.03.002, 2016.
Wurl, O., Ekau, W., Landing, W. M., and Zappa, C. J.: Sea surface microlayer in a changing ocean–A perspective, Elem. Sci. Anth., 5, 31, https://doi.org/10.1525/elementa.228, 2017.
Zäncker, B., Bracher, A., Röttgers, R., and Engel, A.: Variations of the organic matter composition in the sea surface microlayer: a comparison between open ocean, coastal, and upwelling sites off the Peruvian coast, Front. Microbiol., 8, 2369, https://doi.org/10.3389/fmicb.2017.02369, 2017.
Zapata, M., Rodríguez, F., and Garrido, J. L.: Separation of chlorophylls and carotenoids from marine phytoplankton: a new HPLC method using a reversed phase C8 column and pyridine-containing mobile phases, Mar. Ecol. Prog. Ser., 195, 29–45, https://doi.org/10.3354/meps195029, 2000.
Ẑutić, V., Ćosović, B., Marčenko, E., Bihari, N., and Kršinić, F.: Surfactant production by marine phytoplankton, Mar. Chem., 10, 505–520, https://doi.org/10.1016/0304-4203(81)90004-9,1981.
Short summary
A multidisciplinary mesocosm study was conducted to investigate biogeochemical processes and their relationships in the sea-surface microlayer and underlying water during an induced phytoplankton bloom. Phytoplankton-derived organic matter, fuelled microbial activity and biofilm formation, supporting high bacterial abundance. Distinct temporal patterns in biogeochemical parameters and greater variability in the sea-surface microlayer highlight its influence on air–sea interactions.
A multidisciplinary mesocosm study was conducted to investigate biogeochemical processes and...
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