ALBERT

Description: Ocean temperature variability is a fundamental component of the Earth's climate system, and extremes in this variability affect the health of marine ecosystems around the world. The study of marine heatwaves has emerged as a rapidly growing field of research, given notable extreme warm-water events that have occurred against a background trend of global ocean warming. This review summarizes the latest physical and statistical understanding of marine heatwaves based on how they are identified, defined, characterized, and monitored through remotely sensed and in situ data sets. We describe the physical mechanisms that cause marine heatwaves, along with their global distribution, variability, and trends. Finally, we discuss current issues in this developing research area, including considerations related to the choice of climatological baseline periods in defining extremes and how to communicate findings in the context of societal needs. Expected final online publication date for the Annual Review of Marine Science, Volume 13 is January 4, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: In 1842, Darwin identified three types of reefs: fringing reefs, which are directly attached to volcanic islands; barrier reefs, which are separated from volcanic islands by lagoons; and ring reefs, which enclose only a lagoon and are defined as atolls. Moreover, he linked these reef types through an evolutionary model in which an atoll is the logical end point of a subsiding volcanic edifice, as he was unaware of Quaternary glaciations. As an alternative, starting in the 1930s, several authors proposed the antecedent karst model; in this model, atolls formed as a direct interaction between subsidence and karst dissolution that occurred preferentially in the bank interiors rather than on their margins through exposure during glacial lowstands of sea level. Atolls then developed during deglacial reflooding of the glacial karstic morphologies by preferential stacked coral-reef growth along their margins. Here, a comprehensive new model is proposed, based on the antecedent karst model and well-established sea-level fluctuations during the last 5 million years, by demonstrating that most modern atolls from the Maldives Archipelago and from the tropical Pacific and southwest Indian Oceans are rooted on top of late Pliocene flat-topped banks. The volcanic basement, therefore, has had no influence on the late Quaternary development of these flat-topped banks into modern atolls. During the multiple glacial sea-level lowstands that intensified throughout the Quaternary, the tops of these banks were karstified; then, during each of the five mid-to-late Brunhes deglaciations, coral reoccupied their raised margins and grew vertically, keeping up with sea-level rise and creating the modern atolls. Expected final online publication date for the Annual Review of Marine Science, Volume 13 is January 4, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: The large-scale dynamics of ocean oxygenation have changed dramatically throughout Earth's history, in step with major changes in the abundance of O2 in the atmosphere and changes to marine nutrient availability. A comprehensive mechanistic understanding of this history requires insights from oceanography, marine geology, geochemistry, geomicrobiology, evolutionary ecology, and Earth system modeling. Here, we attempt to synthesize the major features of evolving ocean oxygenation on Earth through more than 3 billion years of planetary history. We review the fundamental first-order controls on ocean oxygen distribution and summarize the current understanding of the history of ocean oxygenation on Earth from empirical and theoretical perspectives—integrating geochemical reconstructions of oceanic and atmospheric chemistry, genomic constraints on evolving microbial metabolism, and mechanistic biogeochemical models. These changes are used to illustrate primary regimes of large-scale ocean oxygenation and to highlight feedbacks that can act to stabilize and destabilize the ocean–atmosphere system in anoxic, low-oxygen, and high-oxygen states. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: The SUP05 clade of gammaproteobacteria (Thioglobaceae) comprises both primary producers and primary consumers of organic carbon in the oceans. Host-associated autotrophs are a principal source of carbon and other nutrients for deep-sea eukaryotes at hydrothermal vents, and their free-living relatives are a primary source of organic matter in seawater at vents and in marine oxygen minimum zones. Similar to other abundant marine heterotrophs, such as SAR11 and Roseobacter, heterotrophic Thioglobaceae use the dilute pool of osmolytes produced by phytoplankton for growth, including methylated amines and sulfonates. Heterotrophic members are common throughout the ocean, and autotrophic members are abundant at hydrothermal vents and in anoxic waters; combined, they can account for more than 50% of the total bacterial community. Studies of both cultured and uncultured representatives from this diverse family are providing novel insights into the shifting biogeochemical roles of autotrophic and heterotrophic bacteria that cross oxic–anoxic boundary layers in the ocean. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: A key Earth system science question is the role of atmospheric deposition in supplying vital nutrients to the phytoplankton that form the base of marine food webs. Industrial and vehicular pollution, wildfires, volcanoes, biogenic debris, and desert dust all carry nutrients within their plumes throughout the globe. In remote ocean ecosystems, aerosol deposition represents an essential new source of nutrients for primary production. The large spatiotemporal variability in aerosols from myriad sources combined with the differential responses of marine biota to changing fluxes makes it crucially important to understand where, when, and how much nutrients from the atmosphere enter marine ecosystems. This review brings together existing literature, experimental evidence of impacts, and new atmospheric nutrient observations that can be compared with atmospheric and ocean biogeochemistry modeling. We evaluate the contribution and spatiotemporal variability of nutrient-bearing aerosols from desert dust, wildfire, volcanic, and anthropogenic sources, including the organic component, deposition fluxes, and oceanic impacts. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: Marine ecosystems are increasingly impacted by global environmental changes, including warming temperatures, deoxygenation, and ocean acidification. Marine scientists recognize intuitively that these environmental changes are translated into community changes via organismal physiology. However, physiology remains a black box in many ecological studies, and coexisting species in a community are often assumed to respond similarly to environmental stressors. Here, we emphasize how greater attention to physiology can improve our ability to predict the emergent effects of ocean change. In particular, understanding shifts in the intensity and outcome of species interactions such as competition and predation requires a sharpened focus on physiological variation among community members and the energetic demands and trophic mismatches generated by environmental changes. Our review also highlights how key species interactions that are sensitive to environmental change can operate as ecological leverage points through which small changes in abiotic conditions are amplified into large changes in marine ecosystems. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: A small subset of marine microbial enzymes and surface transporters have a disproportionately important influence on the cycling of carbon and nutrients in the global ocean. As a result, they largely determine marine biological productivity and have been the focus of considerable research attention from microbial oceanographers. Like all biological catalysts, the activity of these keystone biomolecules is subject to control by temperature and pH, leaving the crucial ecosystem functions they support potentially vulnerable to anthropogenic environmental change. We summarize and discuss both consensus and conflicting evidence on the effects of sea surface warming and ocean acidification for five of these critical enzymes [carbonic anhydrase, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO), nitrogenase, nitrate reductase, and ammonia monooxygenase] and one important transporter (proteorhodopsin). Finally, we forecast how the responses of these few but essential biocatalysts to ongoing global change processes may ultimately help to shape the microbial communities and biogeochemical cycles of the future greenhouse ocean. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: Many large marine predators make excursions from surface waters to the deep ocean below 200 m. Moreover, the ability to access meso- and bathypelagic habitats has evolved independently across marine mammals, reptiles, birds, teleost fishes, and elasmobranchs. Theoretical and empirical evidence suggests a number of plausible functional hypotheses for deep-diving behavior. Developing ways to test among these hypotheses will, however, require new ways to quantify animal behavior and biophysical oceanographic processes at coherent spatiotemporal scales. Current knowledge gaps include quantifying ecological links between surface waters and mesopelagic habitats and the value of ecosystem services provided by biomass in the ocean twilight zone. Growing pressure for ocean twilight zone fisheries creates an urgent need to understand the importance of the deep pelagic ocean to large marine predators. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: The water mass transformation (WMT) framework weaves together circulation, thermodynamics, and biogeochemistry into a description of the ocean that complements traditional Eulerian and Lagrangian methods. In so doing, a WMT analysis renders novel insights and predictive capabilities for studies of ocean physics and biogeochemistry. In this review, we describe fundamentals of the WMT framework and illustrate its practical analysis capabilities. We show how it provides a robust methodology to characterize and quantify the impact of physical processes on buoyancy and other thermodynamic fields. We also detail how to extend WMT to insightful analysis of biogeochemical cycles.

Description: Urban and periurban ocean developments impact 1.5% of the global exclusive economic zones, and the demand for ocean space and resources is increasing. As we strive for a more sustainable future, it is imperative that we better design, manage, and conserve urban ocean spaces for both humans and nature. We identify three key objectives for more sustainable urban oceans: reduction of urban pressures, protection and restoration of ocean ecosystems, and support of critical ecosystem services. We describe an array of emerging evidence-based approaches, including greening gray infrastructure, restoring habitats, and developing biotechnologies. We then explore new economic instruments and incentives for supporting these new approaches and evaluate their feasibility in delivering these objectives. Several of these tools have the potential to help bring nature back to the urban ocean while also addressing some of the critical needs of urban societies, such as climate adaptation, seafood production, clean water, and recreation, providing both human and environmental benefits in some of our most impacted ocean spaces. Expected final online publication date for the Annual Review of Marine Science, Volume 13 is January 3, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: The CARIACO (Carbon Retention in a Colored Ocean) Ocean Time-Series Program station, located at 10.50°N, 64.66°W, observed biogeochemical and ecological processes in the Cariaco Basin of the southwestern Caribbean Sea from November 1995 to January 2017. The program completed 232 monthly core cruises, 40 sediment trap deployment cruises, and 40 microbiogeochemical process cruises. Upwelling along the southern Caribbean Sea occurs from approximately November to August. High biological productivity (320–628 g C m−2y−1) leads to large vertical fluxes of particulate organic matter, but only approximately 9–10 g C m−2y−1fall to the bottom sediments (∼1–3% of primary production). A diverse community of heterotrophic and chemoautotrophic microorganisms, viruses, and protozoa thrives within the oxic–anoxic interface. A decrease in upwelling intensity from approximately 2003 to 2013 and the simultaneous overfishing of sardines in the region led to diminished phytoplankton bloom intensities, increased phytoplankton diversity, and increased zooplankton densities. The deepest waters of the Cariaco Basin exhibited long-term positive trends in temperature, salinity, hydrogen sulfide, ammonia, phosphate, methane, and silica. Earthquakes and coastal flooding also resulted in the delivery of sediment to the seafloor. The program's legacy includes climate-quality data from suboxic and anoxic habitats and lasting relationships between international researchers.

Description: The use of unoccupied aircraft systems (UASs, also known as drones) in science is growing rapidly. Recent advances in microelectronics and battery technology have resulted in the rapid development of low-cost UASs that are transforming many industries. Drones are poised to revolutionize marine science and conservation, as they provide essentially on-demand remote sensing capabilities at low cost and with reduced human risk. A variety of multirotor, fixed-wing, and transitional UAS platforms are capable of carrying various optical and physical sampling payloads and are being employed in almost every subdiscipline of marine science and conservation. This article provides an overview of the UAS platforms and sensors used in marine science and conservation missions along with example physical, biological, and natural resource management applications and typical analytical workflows. It concludes with details on potential effects of UASs on marine wildlife and a look to the future of UASs in marine science and conservation.

Description: In this article, I use the Estimating the Circulation and Climate of the Ocean version 4 (ECCO4) reanalysis to estimate the residual meridional overturning circulation, zonally averaged, over the separate Atlantic and Indo-Pacific sectors. The abyssal component of this estimate differs quantitatively from previously published estimates that use comparable observations, indicating that this component is still undersampled. I also review recent conceptual models of the oceanic meridional overturning circulation and of the mid-depth and abyssal stratification. These theories show that dynamics in the Antarctic circumpolar region are essential in determining the deep and abyssal stratification. In addition, they show that a mid-depth cell consistent with observational estimates is powered by the wind stress in the Antarctic circumpolar region, while the abyssal cell relies on interior diapycnal mixing, which is bottom intensified.

Description: We have known for more than 45 years that microplastics in the ocean are carriers of microbially dominated assemblages. However, only recently has the role of microbial interactions with microplastics in marine ecosystems been investigated in detail. Research in this field has focused on three main areas: ( a) the establishment of plastic-specific biofilms (the so-called plastisphere); ( b) enrichment of pathogenic bacteria, particularly members of the genus Vibrio, coupled to a vector function of microplastics; and ( c) the microbial degradation of microplastics in the marine environment. Nevertheless, the relationships between marine microorganisms and microplastics remain unclear. In this review, we deduce from the current literature, new comparative analyses, and considerations of microbial adaptation concerning plastic degradation that interactions between microorganisms and microplastic particles should have rather limited effects on the ocean ecosystems. The majority of microorganisms growing on microplastics seem to belong to opportunistic colonists that do not distinguish between natural and artificial surfaces. Thus, microplastics do not pose a higher risk than natural particles to higher life forms by potentially harboring pathogenic bacteria. On the other hand, microplastics in the ocean represent recalcitrant substances for microorganisms that are insufficient to support prokaryotic metabolism and will probably not be microbially degraded in any period of time relevant to human society. Because we cannot remove microplastics from the ocean, proactive action regarding research on plastic alternatives and strategies to prevent plastic entering the environment should be taken promptly.

Description: Reactive oxygen species (ROS) are produced ubiquitously across the tree of life. Far from being synonymous with toxicity and harm, biological ROS production is increasingly recognized for its essential functions in signaling, growth, biological interactions, and physiochemical defense systems in a diversity of organisms, spanning microbes to mammals. Part of this shift in thinking can be attributed to the wide phylogenetic distribution of specialized mechanisms for ROS production, such as NADPH oxidases, which decouple intracellular and extracellular ROS pools by directly catalyzing the reduction of oxygen in the surrounding aqueous environment. Furthermore, biological ROS production contributes substantially to natural fluxes of ROS in the ocean, thereby influencing the fate of carbon, metals, oxygen, and climate-relevant gases. Here, we review the taxonomic diversity, mechanisms, and roles of extracellular ROS production in marine bacteria, phytoplankton, seaweeds, and corals, highlighting the ecological and biogeochemical influences of this fundamental and remarkably widespread process. Expected final online publication date for the Annual Review of Marine Science, Volume 13 is January 4, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: Oceanic uptake of anthropogenic carbon dioxide (CO2) from the atmosphere has changed ocean biogeochemistry and threatened the health of organisms through a process known as ocean acidification (OA). Such large-scale changes affect ecosystem functions and can have effects on societal uses, fisheries resources, and economies. In many large estuaries, anthropogenic CO2-induced acidification is enhanced by strong stratification, long water residence times, eutrophication, and a weak acid–base buffer capacity. In this article, we review how a variety of processes influence aquatic acid–base properties in estuarine waters, including river–ocean mixing, upwelling, air–water gas exchange, biological production and subsequent respiration, anaerobic respiration, calcium carbonate (CaCO3) dissolution, and benthic inputs. We emphasize the spatial and temporal dynamics of partial pressure of CO2 ( pCO2), pH, and calcium carbonate mineral saturation states. Examples from three large estuaries—Chesapeake Bay, the Salish Sea, and Prince William Sound—are used to illustrate how natural and anthropogenic processes and climate change may manifest differently across estuaries, as well as the biological implications of OA on coastal calcifiers. Expected final online publication date for the Annual Review of Marine Science, Volume 13 is January 4, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: The Deepwater Horizon oil spill was the largest, longest-lasting, and deepest oil accident to date in US waters. As oil and natural gas jetted from release points at 1,500-m depth in the northern Gulf of Mexico, entrainment of the surrounding ocean water into a buoyant plume, rich in soluble hydrocarbons and dispersed microdroplets of oil, created a deep (1,000-m) intrusion layer. Larger droplets of liquid oil rose to the surface, forming a slick of mostly insoluble, hydrocarbon-type compounds. A variety of physical, chemical, and biological mechanisms helped to transform, remove, and redisperse the oil and gas that was released. Biodegradation removed up to 60% of the oil in the intrusion layer but was less efficient in the surface slick, due to nutrient limitation. Photochemical processes altered up to 50% (by mass) of the floating oil. The surface oil expression changed daily due to wind and currents, whereas the intrusion layer flowed southwestward. A portion of the weathered surface oil stranded along shorelines. Oil from both surface and intrusion layers were deposited onto the seafloor via sinking marine oil snow. The biodegradation rates of stranded or sedimented oil were low, with resuspension and redistribution transiently increasing biodegradation. The subsequent research efforts increased our understanding of the fate of spilled oil immensely, with novel insights focusing on the importance of photooxidation, the microbial communities driving biodegradation, and the formation of marine oil snow that transports oil to the seafloor. Expected final online publication date for the Annual Review of Marine Science, Volume 13 is January 4, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: The dissolution of CaCO3 minerals in the ocean is a fundamental part of the marine alkalinity and carbon cycles. While there have been decades of work aimed at deriving the relationship between dissolution rate and mineral saturation state (a so-called rate law), no real consensus has been reached. There are disagreements between laboratory- and field-based studies and differences in rates for inorganic and biogenic materials. Rates based on measurements on suspended particles do not always agree with rates inferred from measurements made near the sediment–water interface of the actual ocean. By contrast, the freshwater dissolution rate of calcite has been well described by bulk rate measurements from a number of different laboratories, fit by basic kinetic theory, and well studied by atomic force microscopy and vertical scanning interferometry to document the processes at the atomic scale. In this review, we try to better unify our understanding of carbonate dissolution in the ocean via a relatively new, highly sensitive method we have developed combined with a theoretical framework guided by the success of the freshwater studies. We show that empirical curve fits of seawater data as a function of saturation state do not agree, largely because the curvature is itself a function of the thermodynamics. Instead, we show that models that consider both surface energetic theory and the complicated speciation of seawater and calcite surfaces in seawater are able to explain most of the most recent data. This new framework can also explain features of the historical data that have not been previously explained. The existence of a kink in the relationship between rate and saturation state, reflecting a change in dissolution mechanism, may be playing an important role in accelerating CaCO3 dissolution in key sedimentary environments. Expected final online publication date for the Annual Review of Marine Science, Volume 13 is January 3, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Description: Archaea are ubiquitous and abundant members of the marine plankton. Once thought of as rare organisms found in exotic extremes of temperature, pressure, or salinity, archaea are now known in nearly every marine environment. Though frequently referred to collectively, the planktonic archaea actually comprise four major phylogenetic groups, each with its own distinct physiology and ecology. Only one group—the marine Thaumarchaeota—has cultivated representatives, making marine archaea an attractive focus point for the latest developments in cultivation-independent molecular methods. Here, we review the ecology, physiology, and biogeochemical impact of the four archaeal groups using recent insights from cultures and large-scale environmental sequencing studies. We highlight key gaps in our knowledge about the ecological roles of marine archaea in carbon flow and food web interactions. We emphasize the incredible uncultivated diversity within each of the four groups, suggesting there is much more to be done.