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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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.
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We investigated how organic molecules in the ocean’s surface layer accumulate and respond to wind. Using a large wind-wave tank filled with seawater, we found that natural molecules produced by marine microbes gather at the surface under light winds, slowing the exchange of gases such as carbon dioxide. When winds increase, this layer rapidly breaks down. These findings suggest that marine life can influence how the ocean and atmosphere interact, particularly in calm conditions.
Lisa Gassen, Samuel M. Ayim, Leonie Jaeger, Jens Meyerjürgens, Mariana Ribas-Ribas, and Oliver Wurl
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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.
Lin Yang, Peiyi Bian, Jing Zhang, Anja Engel, Bin Yang, and Gui-Peng Yang
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CO, CDOM, and 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 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 SSW.
Lisa Deyle, Thomas H. Badewien, Oliver Wurl, and Jens Meyerjürgens
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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
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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.
Evelyn Freney, Karine Sellegri, Alessia Nicosia, Leah R. Williams, Matteo Rinaldi, Jonathan T. Trueblood, André S. H. Prévôt, Melilotus Thyssen, Gérald Grégori, Nils Haëntjens, Julie Dinasquet, Ingrid Obernosterer, France Van Wambeke, Anja Engel, Birthe Zäncker, Karine Desboeufs, Eija Asmi, Hilkka Timonen, and Cécile Guieu
Atmos. Chem. Phys., 21, 10625–10641, https://doi.org/10.5194/acp-21-10625-2021, https://doi.org/10.5194/acp-21-10625-2021, 2021
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In this work, we present observations of the organic aerosol content in primary sea spray aerosols (SSAs) continuously generated along a 5-week cruise in the Mediterranean. This information is combined with seawater biogeochemical properties also measured continuously along the ship track to develop a number of parametrizations that can be used in models to determine SSA organic content in oligotrophic waters that represent 60 % of the oceans from commonly measured seawater variables.
Alexander Braun, Marina Spona-Friedl, Maria Avramov, Martin Elsner, Federico Baltar, Thomas Reinthaler, Gerhard J. Herndl, and Christian Griebler
Biogeosciences, 18, 3689–3700, https://doi.org/10.5194/bg-18-3689-2021, https://doi.org/10.5194/bg-18-3689-2021, 2021
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It is known that CO2 fixation by photoautotrophic organisms is the major sink from the atmosphere. While biologists are aware that CO2 fixation also occurs in heterotrophic organisms, this route of inorganic carbon, and its quantitative role, is hardly recognized in biogeochemistry. We demonstrate that a considerable amount of CO2 is fixed annually through anaplerotic reactions in heterotrophic organisms, and a significant quantity of inorganic carbon is temporally sequestered in biomass.
Gerd Krahmann, Damian L. Arévalo-Martínez, Andrew W. Dale, Marcus Dengler, Anja Engel, Nicolaas Glock, Patricia Grasse, Johannes Hahn, Helena Hauss, Mark Hopwood, Rainer Kiko, Alexandra Loginova, Carolin R. Löscher, Marie Maßmig, Alexandra-Sophie Roy, Renato Salvatteci, Stefan Sommer, Toste Tanhua, and Hela Mehrtens
Earth Syst. Sci. Data Discuss., https://doi.org/10.5194/essd-2020-308, https://doi.org/10.5194/essd-2020-308, 2021
Preprint withdrawn
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The project "Climate-Biogeochemistry Interactions in the Tropical Ocean" (SFB 754) was a multidisciplinary research project active from 2008 to 2019 aimed at a better understanding of the coupling between the tropical climate and ocean circulation and the ocean's oxygen and nutrient balance. On 34 research cruises, mainly in the Southeast Tropical Pacific and the Northeast Tropical Atlantic, 1071 physical, chemical and biological data sets were collected.
France Van Wambeke, Elvira Pulido, Philippe Catala, Julie Dinasquet, Kahina Djaoudi, Anja Engel, Marc Garel, Sophie Guasco, Barbara Marie, Sandra Nunige, Vincent Taillandier, Birthe Zäncker, and Christian Tamburini
Biogeosciences, 18, 2301–2323, https://doi.org/10.5194/bg-18-2301-2021, https://doi.org/10.5194/bg-18-2301-2021, 2021
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Michaelis–Menten kinetics were determined for alkaline phosphatase, aminopeptidase and β-glucosidase in the Mediterranean Sea. Although the ectoenzymatic-hydrolysis contribution to heterotrophic prokaryotic needs was high in terms of N, it was low in terms of C. This study points out the biases in interpretation of the relative differences in activities among the three tested enzymes in regard to the choice of added concentrations of fluorogenic substrates.
Jonathan V. Trueblood, Alessia Nicosia, Anja Engel, Birthe Zäncker, Matteo Rinaldi, Evelyn Freney, Melilotus Thyssen, Ingrid Obernosterer, Julie Dinasquet, Franco Belosi, Antonio Tovar-Sánchez, Araceli Rodriguez-Romero, Gianni Santachiara, Cécile Guieu, and Karine Sellegri
Atmos. Chem. Phys., 21, 4659–4676, https://doi.org/10.5194/acp-21-4659-2021, https://doi.org/10.5194/acp-21-4659-2021, 2021
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Sea spray aerosols (SSAs) can be an important source of ice-nucleating particles (INPs) that impact cloud properties over the oceans. In the Mediterranean Sea, we found that the INPs in the seawater surface microlayer increased by an order of magnitude after a rain dust event that impacted iron and bacterial abundances. The INP properties of SSA (INPSSA) increased after a 3 d delay. Outside this event, INPSSA could be parameterized as a function of the seawater biogeochemistry.
Birthe Zäncker, Michael Cunliffe, and Anja Engel
Biogeosciences, 18, 2107–2118, https://doi.org/10.5194/bg-18-2107-2021, https://doi.org/10.5194/bg-18-2107-2021, 2021
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Fungi are found in numerous marine environments. Our study found an increased importance of fungi in the Ionian Sea, where bacterial and phytoplankton counts were reduced, but organic matter was still available, suggesting fungi might benefit from the reduced competition from bacteria in low-nutrient, low-chlorophyll (LNLC) regions.
Emilio Marañón, France Van Wambeke, Julia Uitz, Emmanuel S. Boss, Céline Dimier, Julie Dinasquet, Anja Engel, Nils Haëntjens, María Pérez-Lorenzo, Vincent Taillandier, and Birthe Zäncker
Biogeosciences, 18, 1749–1767, https://doi.org/10.5194/bg-18-1749-2021, https://doi.org/10.5194/bg-18-1749-2021, 2021
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The concentration of chlorophyll is commonly used as an indicator of the abundance of photosynthetic plankton (phytoplankton) in lakes and oceans. Our study investigates why a deep chlorophyll maximum, located near the bottom of the upper, illuminated layer develops in the Mediterranean Sea. We find that the acclimation of cells to low light is the main mechanism involved and that this deep maximum represents also a maximum in the biomass and carbon fixation activity of phytoplankton.
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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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