Under Ocean Acidification, Embryos of a Key Forage Fish Struggle to Hatch

A potential ripple effect from carbon in the atmosphere could have severe impacts throughout the ocean ecosystem

This photo shows sand lance embryos that have and have not hatched. Sand lance have trouble hatching at future ocean CO2 levels (photo courtesy of Emma Cross).

By Elaina Hancock. Reposted from UConn Today, 7 April 2022

When carbon is emitted into the atmosphere, about a quarter of it is absorbed by the earth’s oceans. As the oceans serve as a massive ‘sink’ for carbon, there are changes to the water’s pH – a measure of how acidic or basic water is. As oceans absorb carbon, their water becomes more acidic, a process called ocean acidification (OA). For years, researchers have worked to understand what effect this could have on marine life.

While most research so far shows that fish are fairly resilient to OA, new research from UConn, the University of Washington, the National Oceanic and Atmospheric Administration (NOAA), and Southern Connecticut State University, shows that an important forage fish for the Northwest Atlantic called sand lance is very sensitive to OA, and that this could have considerable ecosystem impacts by 2100. The team’s findings have just been published in Marine Ecology Progress Series 687.

Sand lance spawn in the winter months in offshore environments that tend to have stable, low levels of CO2, explains UConn Department of Marine Sciences researcher and lead author Hannes Baumann.

“Marine organisms are not living in a uniform ocean,” Baumann says. “In near shore environments, large CO2 fluctuations between day and night and between seasons are the norm, and the fish and other organisms are adapted to this variability. When we stumbled upon sand lances we suspected they are different. We thought that a fish that lives in a more open-ocean offshore environment might be more sensitive than the near-shore fish because there’s just much less variability.”

The project was a collaboration with physical oceanographers, including Assistant Professor of Marine Sciences Samantha Siedlecki and Michael Alexander from NOAA’s Physical Sciences Laboratory in Boulder, Colorado, who modeled CO2 levels in 2050 and 2100 for a specific part of the Gulf of Maine where sand lance spawn. Then Baumann and his team reared sand lance embryos in the lab under experimentally higher CO2 levels matching the projected levels.

There are instances of direct fish mortality as result of elevated CO2, but they are rare, says Baumann. However, sand lance embryos proved to be exceptionally sensitive, and fewer embryos hatched under future oceanic CO2 conditions. The researchers repeated the experiments three more times to avoid jumping to conclusions but each time they observed the same result.

“We found that embryo survival-to-hatch decreased sharply with increasing CO2 levels in the water, concluding that this is one of the most CO2-sensitive fish species studied thus far,” Baumann says.

Sand lances are surely one of the most important forage fish here on the Northwest Atlantic shelf… The humpback whales, sharks, tuna, cod, shearwaters, terns — you name it — they are all relying on sand lance.

With this interdisciplinary approach combining model forecasts and serial experimentation the researchers arrived at a picture that is much more specific.

“We consequently applied principles of serial experimentation, which is a most timely and important topic in ocean acidification research right now,” Baumann says. “Because our findings are backed up by repeated independent evidence, they are more robust than many published ocean acidification studies to date.”

In addition to preventing many sand lance embryos from developing normally, the researchers document a second negative, and novel, response to elevated CO2. Higher CO2 levels appear to make it harder for embryos to hatch.

Baumann explains the lowered pH likely renders enzymes needed for successful hatching less effective, leaving the embryos unable to break through their eggshell (chorion) to hatch.

The results show that by 2100, due to acidification, sand lance hatching success could be reduced to 71% of today’s levels. Since sand lance are such a critical component of the food web of the Northwest Atlantic, this marked decrease in sand lance would have profound impacts throughout the ecosystem.

“Sand lances are surely one of the most important forage fish here on the Northwest Atlantic shelf,” Baumann says. “Their range spans from the Mid Atlantic Bight all the way to Greenland. Where we studied them, on Stellwagen Bank, they are called the backbone of the ecosystem. The humpback whales, sharks, tuna, cod, shearwaters, terns — you name it — they are all relying on sand lance, and if sand lance productivity goes down, we will see ripple effects to all these higher trophic animals. Even though we humans don’t fish for sand lance, we need to take care of the species because it has such a huge effect on everything else.”

Baumann says this study supports the hypothesis that offshore, high latitude marine organisms like the sand lance may be among the most vulnerable to OA. As a result, these organisms and food webs will likely be impacted first and soon, and we must act now.

Previous research has focused on opportunistically chosen species when testing their sensitivity for ocean acidification, says Baumann, but this should change.

“We need strategic thinking about what species we are testing next, because we cannot test every marine fish species, that’s an impossible task. We should concentrate on fish species that are likely the most vulnerable, and therefore the ones that are probably being affected first and this research makes a compelling argument that those are the fish species at higher latitudes and in more offshore than nearshore environments.”

DMS researchers contribute to study on copepod climate adaptation

One of the most difficult challenges facing scientists is predicting how organisms will respond to rapid global change. A collaboration between oceanographers at the University of Connecticut and evolutionary biologists at the University of Vermont is looking into how copepods (tiny crustaceans that rival insects as the most abundant animals on the planet) adapt to ocean warming and acidification. This requires understanding the underlying genomic mechanisms that allow these animals to adapt, and the constraints to adaptation. This study by Reid Brennan and collaborators is a lucid example of this approach, identifying sets of genes that are linked to copepod adaptation to stressful new environments, and showing that the ability of these animals to respond to changing conditions is challenged after prolonged adaptation. Therefore, there are limits to adaptation that can constrain the resilience of animal populations to environmental stress.


DMS faculty contributes textbook chapter on Fish Ecology

3rd March 2022. DMS faculty Hannes Baumann contributed a chapter to the new textbook Marine Biology: a functional approach to the oceans & their organisms (Taylor & Francis), which has just been published. The chapter is based on Baumann's long-running class "Ecology of Fishes" (MARN4018/5018), touching on a large variety topics including fish evolution, zoogeography, metabolism, growth, reproduction & basic concepts of fisheries science. The book is geared towards advanced undergraduate and graduate students, stimulating interest while encouraging readers to seek out further in-depth sources.

"With about 28,000 known species, fishes make up more than half of all known vertebrates (Helfman et al. 2009). Over the course of their long evolutionary history they radiated in every conceivable aquatic habitat, from the open ocean and deep-sea trenches to shelf seas, estuaries and lakes, to rivers and the smallest streams and ponds. They are found in subzero Antarctic waters, altitudes of over 4,000 m and even acidic desert springs of > 40°C (Moyle and Cech 2004). The fascinating adaptations to these habitats have produced a mind-bending diversity of form and function, a difference in size that spans more than three magnitudes (0.01 – 18 m), and a profusion of reproductive strategies. Apart from their diversity and unique evolutionary history, fishes are of intense scientific interest for economic reasons, because they comprise the nutritional foundation for a large part of humanity (Costanza et al. 1997) and their exploitation over time has led to thriving – and warring – civilizations. Today, the impetus of sustainable fish management at a time of rapid ecological re-organization due to man-made climate change has made the study of fish ecology and fish stock productivity as urgent and important as ever."

Fig.1: Origin, evolution, and systematics of fishes. A – Origin hypothesis. Early during chordate evolution, sessile arm feeders (pterobranchs) gave rise to gill feeders. In one line, free-swimming filter-feeding larvae lost their sessile stage and evolved into the first, gill-feeding vertebrates (redrawn after Romer and Parsons 1977). B – Evolution and relative abundance of major fish lines through time. Most of today’s fish groups originated in the Devonian; ray-finned fishes became the dominant fish group during the Meso- and Cenozoic (numbers refer to million year ago, Mya). C – Abridged overview of Actinopterygii systematics showing select major orders (-formes) and Perciform families (-idae) sorted top to bottom from ancestral to most derived groups. Most fishes are Teleosts, and within those, most belong to the Euteleosts. Acanthopterygii evolved fin spines; the most species-rich vertebrate order are the Perciformes (after Moyle and Cech 2004).

Marine Sciences selected to partner with AGU Bridge Program

The Department of Marine Sciences (DMS) was recently selected to participate in AGU Bridge Program (https://www.agu.org/bridge-program), which matches under-represented minority students with graduate Earth Science programs in the U.S.  As a Bridge Partner, DMS will work to increase the diversity of the marine sciences and create a more welcoming environment for people from a variety of backgrounds.

The official announcement is available on the AGU website:  https://fromtheprow.agu.org/announcing-the-2021-bridge-program-partners/.

DMS professor George McManus chairs new SCOR working group

The Department of Marine Sciences is proud to announce that Prof. George McManus will co-chair a new SCOR working group titled

Mixotrophy in the Oceans – Novel Experimental designs and Tools for a new trophic paradigm (MixONET) [link to: https://scor-int.org/group/mixotrophy-in-the-oceans-novel-experimental-designs-and-tools-for-a-new-trophic-paradigm-mixonet]

The new working group has 4 broad objectives:

1. Biological oceanography databases and the mixoplankton paradigm: Advocate for the realignment of existing plankton-facing databases in light of the mixoplankton paradigm. Identify connections between mixoplankton communities and essential ocean variables.

2. Biological oceanography research methods under the mixoplankton paradigm: Re-evaluate extant standard biological oceanographic research methods and practices for application under the mixoplankton paradigm. For example, conventional fixatives often destroy the delicate mixoplankton, while more gentle ones obscure the presence of chloroplasts. Also, pico- and nano-planktonic organisms are routinely counted using flow cytometry or epifluorescence microscopy; while standard protocols can discriminate between pigmented and colourless plankton, they are not geared for identification and quantification of mixoplankton.

3. Development of new biological oceanography methods accounting for primary and secondary productions by mixoplankton: Evaluate development of (a) routine new methods and simple protocols that could be incorporated routinely in ongoing monitoring programmes to better quantify mixoplankton and interpret their activities; and (b) new experimental and observing methods (including autonomous technologies) for quantifying and monitoring mixoplanktonic abundance and activity.

4. Ocean literacy: Development of multi-lingual training material for Early Career Researchers (ECRs), ecosystem managers, teachers and students, to enhance ocean literacy. The mixoplankton paradigm needs to be brought to the attention of students through to policy makers. A Decision Support Tool (DST) will be developed to aid configuration of mixoplankton-centric experiments to determine contributions to primary versus secondary production by these organisms.

The other co-Chair is Dr. Aditee Mitra of Cardiff University in Wales.

SCOR (Scientific Committee on Ocean Research) is an international non-governmental non-profit organization. It’s activities focus on promoting international cooperation in planning and conducting oceanographic research, and solving methodological and conceptual problems that hinder research. SCOR covers all areas of ocean science and cooperates with other organizations with common interests to conduct many SCOR activities. SCOR also conducts several different activities to build the capacity for ocean science in developing countries and every SCOR activity includes members from developing countries. Scientists from thirty-three nations have formed national SCOR committees as a foundation for international SCOR. Approximately 550 scientists from 57 countries currently participate in SCOR activities.

Ann Bucklin organizes special issue in the ICES Journal of Marine Science

Patterns of Biodiversity of Marine Zooplankton Based on Molecular Analysis is the latest themed set of articles from​ ICES Journal of Marine Science. (See https://academic.oup.com/icesjms/issue/78/9#1302581-6403476 ). This collection showcases the ongoing refinement of molecular approaches for analysis of zooplankton diversity.

ICES (International Council for the Exploration of the Sea) commissioned a cartoon by Bas Köhler and announced the publication (see: https://www.ices.dk/news-and-events/news-archive/news/Pages/TSZooplankton.aspx).

The motivators for the special issue are members of the SCOR WG157 MetaZooGene (see: https://metazoogene.org/ ), chaired by Ann Bucklin, who also authored the introductory paper, New insights into biodiversity, biogeography, ecology, and evolution of marine zooplankton based on molecular approaches (see https://doi.org/10.1093/icesjms/fsab198) with co-authors, Katja T.C.A. Peijnenburg (NL), Ksenia Kosobokova (RU), and Ryuji J. Machida (TW).

Celebrate Black in Marine Science week (Nov. 28-Dec.4)

Black in Marine Science (BIMS) is a premier organization aimed to celebrate Black marine scientists, spread environmental awareness and inspire the next generation of scientific thought leaders.  Please join us in celebrating and amplifying the work of Black marine scientists by tuning into their many panels, workshops, or keynotes during the week. All of the events are free and will be streamed to the BIMTV YouTube channel (YouTube channel). More information about BIMS can be found here: https://www.blackinmarinescience.org/

Eos highlights DMS graduate student research

Graduate student Halle Berger’s recent publication in AGU Advances has been selected for an Editor’s Highlight in Eos (fewer than 2% of papers are selected for this).


You can read more about it and hear from Halle in UConn Today’s highlight of the paper here: https://today.uconn.edu/2021/10/understanding-how-a-crabs-complex-life-cycle-will-respond-to-climate-change/

Berger, H.M., S.A. Siedlecki, C.M. Matassa, S.R. Alin, I.C. Kaplan, E.E. Hodgson, D. Pilcher, E.L. Norton, and J.A. Newton (2021): Seasonality and life history complexity determine vulnerability of Dungeness crab to multiple climate stressors. AGU Advances, 2(4), e2021AV000456, doi: 10.1029/2021AV000456, Published online (open access).

The interaction of exposure and the degree of negative response (consequence) to reduced pH, low oxygen and warming temperature on the vulnerability of Dungeness crab eggs, larvae, juvenile and adult life stages assessed from life stage distribution simulations (distribution map) and Lagrangian particle tracking simulations (transport model). Credit: Berger et al., 2021, Figure 3

Jim O’Donnell Attends COP26

UConn Professor of Marine Sciences and CIRCA’s Executive Director Jim O’Donnell traveled across the Atlantic this week to attend the COP 26 conference, where leaders from around the globe are meeting in Glasgow, Scotland to confront the climate crisis. So in addition to resources and news articles we’re collecting about the conference, we are fortunate to have someone on the inside! Read on for Jim’s daily perspective along with photos, helpful links, and other insights he shares from what we hope will be an earth-changing climate conference:  https://circa.uconn.edu/news-announcements/.

See the UConn Today article:  https://today.uconn.edu/2021/11/cop26-a-uconn-perspective-on-the-climate-change-summit/

Fall 2021 Department Achievements

Fall 2021 Department Achievements


Prof. Hans Dam

Received the 2021 UConn Faculty Excellence Award for Graduate Teaching. This Award recognizes a faculty member with a distinguished record of sustained teaching excellence through outstanding instruction, engaging students thoroughly in the process of learning, and contributing significantly to the intellectual life of the University.  


CIRCA – Prof. James O’Donnell

The Connecticut legislature’s 2021-23 budget provided an additional $5 million to CIRCA to expand Resilient Connecticut activities and advance fundable projects. CIRCA will continue to support development of innovative adaptation approaches for flood and heat vulnerability along with expert advice on climate issues to communities in Hartford, New London, and Middlesex Counties. 

Prof. Hannes Baumann

Connecticut SeaGrant: PI Baumann together with collaborators from CTDEEP received funding to investigate the causes and ecosystem consequences of the recent, steep increase in Black Sea Bass in Long Island Sound. 

Jo-Marie Kasinak (graduate student, Prof. Vaudrey) & Prof. Vaudrey

Connecticut SeaGrant: Toward a deeper understanding of human connections with ocean environments: Ocean Identity (OI) as a novel construct, research instrument, and assessment tool. (2022-2024), $143,309, PIs Kelly, Kasinak, McKinley, Vaudrey, & Mattei. 

Prof. Robert Mason

NSF Chemical Oceanography: Methylated mercury sources and cycling in the high latitude North Atlantic. (2021-2023), $283,534, PI Mason. 

Prof. Samantha Siedlecki

NSF: Regional climate change projections to enable equitable ocean planning for the blue economy (2021-2022), PIs. Pinsky, Hice-Dunton, Siedlecki, & St. Martin. This project aims to enable climate-ready, coordinated, and inclusive decision making throughout the blue economy and spark a new generation of durable blue development.

Prof. Vlahos

NSF: Arctic Marginal Ice Zone Alkalinity (AMIZA). PI Vlahos. This project is studying the components of carbonate alkalinity in the changing Arctic with a focus on the transient ice melt zones.

NIH: Chronic Kidney Disease. PI Vlahos. Lead PI Shuchi Anand, Stanford University. This project is a continuation of our efforts with colleagues in Sri Lanka and at Stanford to expand our 300 person cohort to a 900 person longitudinal study on the progression of kidney disease and water quality.


Prof. Peter Auster

Prof. Auster presents a chapter as part of an international effort to inform delegations to the United Nations about the status and effects of human activities on the global ocean. (Levin, L. A., Auster, P., Clark, M. R., Hall-Spencer, J. M., Hopcroft, R., Ingels, J., Metaxas, A., Narayanaswamy, B., Tuhumwire, J. T., Yasuhara, M. (2021).  Continental slopes and submarine canyons.  Chapter 7J, p. 395-420, in: The Second World Ocean Assessment, World Ocean Assessment II. United Nations, New York.)

Prof. Auster addresses the confusion in ecological terminology used in international agreements to manage fisheries impacts on the high seas. (Watling, L., Auster, P. J. (2021). Vulnerable marine ecosystems, communities, and indicator species: confusing concepts for conservation of seamounts. Frontiers in Marine Science 8:622586.)

Prof. Auster and colleagues demonstrate that simple GoPro cameras can be used to quantify the role of oyster aquaculture cages as fish habitat. (Mercaldo-Allen, R., Clark, P., Liu, Y., Phillips, G., Redman, D., Auster, P. J., Estela, E., Milke, L., Verkade, A., Rose, J. M.  (2021). Exploring video and eDNA metabarcoding methods to assess oyster aquaculture cages as fish habitat.  Aquaculture Environment Interactions 13:277-294.)

Prof. Paola Batta-Lona

Prof. Batta-Lona and colleagues examined how environmental conditions affect the distribution of zooplankton in the Gulf of Mexico. (Cicala, F., Arteaga, M., Herzka, S., Martinez, M., Hereu, C., Jimenez Rosenberg, S. P. A., Saavedra, A., Robles, J., Gomez, R., Batta-Lona, P. G., Galindo Sanchez, C. E. (2021). Environmental conditions drive zooplankton community structure in the deep-water region of the southern Gulf of Mexico: a molecular approach. Molecular Ecology.) 

Prof. Batta-Lona and colleagues used DNA (metabarcoding) to look at the diversity of zooplankton in the Gulf of Mexico. (Martinez, M., Hereu, C., Galindo Sanchez, C. E., Arteaga, M., Batta-Lona, P. G., Saavedra, A., Robles, J., Jimenez Rosenberg, S. P. A., Herzka, S. (2021). Epipelagic zooplankton diversity in the deep water region of the Gulf of Mexico: A metabarcoding survey.  ICES Journal of Marine Science.)

Prof. Zofia Baumann

Prof. Baumann and colleagues present a study that demonstrates the liver detoxifies previously-bioaccumulated methylmercury based on analyses of tissues from three waterbird species. (Poulin, B.A., Janssen, S.E., Rosera, T.J., Krabbenhoft, D.P., Eagles-Smith, C.A., Ackerman, J.T., Stewart, A.R., Kim, E., Baumann, Z., Kim, J.H. and Manceau, A., 2021. Isotope fractionation from in vivo methylmercury detoxification in waterbirds. ACS Earth and Space Chemistry, 5(5), pp.990-997.)

Prof. Ann Bucklin

Ann Bucklin, chair of the Scientific Committee for Ocean Research (SCOR) Working Group WG157 presents with other members of WG157 a review paper examining global patterns of biodiversity of marine zooplankton using DNA barcodes or short sequences of cytochrome oxidase I (COI) that discriminate and identify species and announce a reference database for identification of species from DNA barcoding and metabarcoding of pelagic biodiversity, with advanced search functions by ocean region and taxonomic group. (Bucklin A., Peijnenburg, K. T. C. A., Kosobokova, K. N., O’Brien, T. D., Blanco-Bercial, L., Cornils, A., Falkenhaug, T., Hopcroft, R. R., Hosia, A., Laakmann, S., Li, C., Martell, L., Questel, J. M., Wall-Palmer, D., Wang, M., Wiebe, P. H., Weydmann-Zwolicka, A. (2021). Toward a global reference database of COI barcodes for marine zooplankton. Marine Biology.)

Photo by R.R. Hopcroft and C. Clarke (UAF) and L.P. Madin (WHOI); see: http://www.cmarz.org/galleries.html 

Profs. Hans Dam, Michael Finiguerra, Hannes Baumann

Profs. Dam, Finiguerra, and Baumann show that zooplankton adapt quickly, but with limited capacity, to ocean warming and acidification, which is both encouraging and sobering news for the response of animal populations to rapid climate change. (Dam, H. G., deMayo, J. A., Park, G., Norton, L., He, X., Finiguerra, M. B., Baumann, H., Brennan, R. S., Pespeni, M. H. (2021). Rapid, but limited, zooplankton adaptation to simultaneous warming and acidification. Nature Climate Change, 11, 780-786.)

Prof. Leonel Romero 

Prof. Romero and colleagues propose a new approach to realistically model wave effects on currents, overcoming several limitations of state-of-the-art coupled wave-ocean models. (Romero, L., Hypolite, D., McWilliams, J. C. (2021). Representing Wave Effects on Currents. Ocean Modelling, 167, 101873.)

Prof. Sandy Shumway

Prof. Shumway edited a book titled “Molluscan Shellfish Aquaculture: A Practical Guide” as a usable manual for those interested in an up-to-date introduction to the field. The book covers each of the major cultured species of cultural importance. (Shumway, Sandy, Ed. (2021) Molluscan Shellfish Aquaculture: A Practical Guide. 5M Publishing.)

Prof. Samantha Siedlecki

Prof. Siedlecki and colleagues present work showing that the  projected changes for carbon variables like pCO2 and pH in the California Current System (CCS) using a high resolution model are modified by coastal processes resolved in the downscaled projections relative to the projected global simulation, suggesting downscaled projections are necessary to more accurately project future conditions of these variables. (Siedlecki, S. A., Pilcher, D., Howard, E. M., Deutsch, C., MacCready, P., Norton, E. L., Frenzel, H., Newton, J., Feely, R. A., Alin, S. R., Klinger, T. (2021). Coastal processes modify projections of some climate-driven stressors in the California Current System, Biogeosciences, 18, 2871–2890.)

Prof. Siedlecki, graduate student Kelly McGarry, and colleagues present a combination of regional high-resolution simulations that project ocean acidification (OA) conditions for the Gulf of Maine into 2050, the results of which indicate that the aragonite saturation state (one measure of OA) declines and the entire GOM will experiences biologically critical conditions for most of the year. (Siedlecki, S. A., Salisbury, J., Gledhill, D. K., Bastidas, C., Meseck, S., McGarry, K., … & Morrison, R. (2021). Projecting ocean acidification impacts for the Gulf of Maine to 2050: New tools and expectations. Elementa: Science of the Anthropocene, 9(1):00062.) 

Prof. Penny Vlahos

Prof. Vlahos, graduate student Emma Shipley, and colleagues present an interdisciplinary study that connects drinking water quality to the progression of kidney disease in rice farmers in the Sri Lankan dry zone. (Vlahos, P., Schensul, S., Anand, S., Shipley, E., Diyabalanage, S., Hu, C., Ha, T., Staniec, A., Haider, L., Schensul, J., Hewavitarne, P., Silva, T., Chandrajith, R., Nanayakkara, N. (Accepted). Water Sources and Kidney Function: Investigating Chronic Kidney Disease of Unknown Etiology in a Prospective Study. NPJ Clean Water.)

Halle Berger (graduate student, Profs. Samantha Siedlecki and Catherine Matassa)

Berger and colleagues present a vulnerability assessment for Dungeness crab to climate change which revealed that population-level vulnerability to future hypoxia is most severe overall due to increased exposure of the critical adult stage during the upwelling season. (Berger, H. M., Siedlecki, S. A., Matassa, C. M., Alin, S. R., Kaplan, I. C., Hodgson, E. E., Pilcher, D. J., Norton, E. L., Newton, J. A. (2021). Seasonality and life history complexity determine vulnerability of Dungeness crab to multiple climate stressors. AGU Advances, 2, e2021AV000456.) 

Tyler Griffin (graduate student, Prof. Evan Ward)

Griffin and colleagues demonstrate that antibiotics can be used as effective tools to experimentally diminish the gut microbiomes of suspension-feeding animals, like oysters and mussels. (Griffin, T. W., Pierce, M. L., Nigro, L. M., Holohan, B., & Ward, J. E. (2021). An examination of the use of antibiotics as a method to experimentally perturb the microbiota of suspension-feeding bivalves. Invertebrate Biology.  e12352.)

Allison Staniec (PhD graduate, Prof. Penny Vlahos)

Staniec and colleagues present a study identifying the role of sea spray in gas exchange in an article that was also featured in Nature Highlights: Big Potential for Tiny Droplets. (Staniec, A., Vlahos, P., Monahan, E. C. (2021) The role of sea spray in atmosphere-ocean gas exchange. Nature Geoscience, 14, 593-598.)