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Annual Review of Marine Science - Volume 13, 2021
Volume 13, 2021
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Right Place, Right Time: An Informal Memoir
Vol. 13 (2021), pp. 1–21More LessMy career spanned the revolution in understanding of the large-scale fluid ocean, as modern electronics produced vast new capabilities. I started in the days of almost purely mechanical instruments operated by seagoing scientists, ones not so different from those used more than a century earlier. Elegant theories existed of hypothetical steady-state oceans. Today, we understand that the ocean is a highly turbulent fluid, interacting over global scales, and it is now studied by large teams using spacecraft and diverse sets of self-contained in situ instrumentation. Mine was an accidental career: I was lucky to be in the right place at the right time.
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Natural and Anthropogenic Drivers of Acidification in Large Estuaries
Vol. 13 (2021), pp. 23–55More LessOceanic 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 impacts 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 coastal upwelling, river–ocean mixing, air–water gas exchange, biological production and subsequent aerobic and 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.
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The Dissolution Rate of CaCO3 in the Ocean
Vol. 13 (2021), pp. 57–80More LessThe 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.
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The Biogeochemistry of Marine Polysaccharides: Sources, Inventories, and Bacterial Drivers of the Carbohydrate Cycle
Vol. 13 (2021), pp. 81–108More LessPolysaccharides are major components of macroalgal and phytoplankton biomass and constitute a large fraction of the organic matter produced and degraded in the ocean. Until recently, however, our knowledge of marine polysaccharides was limited due to their great structural complexity, the correspondingly complicated enzymatic machinery used by microbial communities to degrade them, and a lack of readily applied means to isolate andcharacterize polysaccharides in detail. Advances in carbohydrate chemistry, bioinformatics, molecular ecology, and microbiology have led to new insights into the structures of polysaccharides, the means by which they are degraded by bacteria, and the ecology of polysaccharide production and decomposition. Here, we survey current knowledge, discuss recent advances, and present a new conceptual model linking polysaccharide structural complexity and abundance to microbially driven mechanisms of polysaccharide processing. We conclude by highlighting specific future research foci that will shed light on this central but poorly characterized component of the marine carbon cycle.
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The Complexity of Spills: The Fate of the Deepwater Horizon Oil
Vol. 13 (2021), pp. 109–136More LessThe 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.
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Physiological Responses of Fish to Oil Spills
Vol. 13 (2021), pp. 137–160More LessMillions of tons of oil are spilled in aquatic environments every decade, and this oil has the potential to greatly impact fish populations. Here, we review available information on the physiological effects of oil and polycyclic aromatic hydrocarbons on fish. Oil toxicity affects multiple biological systems, including cardiac function, cholesterol biosynthesis, peripheral and central nervous system function, the stress response, and osmoregulatory and acid–base balance processes. We propose that cholesterol depletion may be a significant contributor to impacts on cardiac, neuronal, and synaptic function as well as reduced cortisol production and release. Furthermore, it is possible that intracellular calcium homeostasis—a part of cardiotoxic and neuronal function that is affected by oil exposure—may be related to cholesterol depletion. A detailed understanding of oil impacts and affected physiological processes is emerging, but knowledge of their combined effects on fish in natural habitats is largely lacking. We identify key areas deserving attention in future research.
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New Microbial Biodiversity in Marine Sediments
Vol. 13 (2021), pp. 161–175More LessMicrobes in marine sediments represent a large portion of the biosphere, and resolving their ecology is crucial for understanding global ocean processes. Single-gene diversity surveys have revealed several uncultured lineages that are widespread in ocean sediments and whose ecological roles are unknown, and advancements in the computational analysis of increasingly large genomic data sets have made it possible to reconstruct individual genomes from complex microbial communities. Using these metagenomic approaches to characterize sediments is transforming our view of microbial communities on the ocean floor and the biodiversity of the planet. In recent years, marine sediments have been a prominent source of new lineages in the tree of life. The incorporation of these lineages into existing phylogenies has revealed that many belong to distinct phyla, including archaeal phyla that are advancing our understanding of the origins of cellular complexity and eukaryotes. Detailed comparisons of the metabolic potentials of these new lineages have made it clear that uncultured bacteria and archaea are capable of mediating key previously undescribed steps in carbon and nutrient cycling.
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Production of Extracellular Reactive Oxygen Species by Marine Biota
Vol. 13 (2021), pp. 177–200More LessReactive 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.
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Variations in Ocean Mixing from Seconds to Years
Vol. 13 (2021), pp. 201–226More LessOver the past several decades, there has developed a community-wide appreciation for the importance of mixing at the smallest scales to geophysical fluid dynamics on all scales. This appreciation has spawned greater participation in the investigation of ocean mixing and new ways to measure it. These are welcome developments given the tremendous separation in scales between the basins, ) m, and the turbulence, ) m, and the fact that turbulence that leads to thermodynamically irreversible mixing in high-Reynolds-number geophysical flows varies by at least eight orders of magnitude in both space and time. In many cases, it is difficult to separate the dependencies because measurements are sparse, also in both space and time. Comprehensive shipboard turbulence profiling experiments supplemented by Doppler sonar current measurements provide detailed observations of the evolution of the vertical structure of upper-ocean turbulence on timescales of minutes to weeks. Recent technical developments now permit measurements of turbulence in the ocean, at least at a few locations, for extended periods. This review summarizes recent and classic results in the context of our expanding knowledge of the temporal variability of ocean mixing, beginning with a discussion of the timescales of the turbulence itself (seconds to minutes) and how turbulence-enhanced mixing varies over hours, days, tidal cycles, monsoons, seasons, and El Niño–Southern Oscillation timescales (years).
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Oceanic Frontogenesis
Vol. 13 (2021), pp. 227–253More LessFrontogenesis is the fluid-dynamical processes that rapidly sharpen horizontal density gradients and their associated horizontal velocity shears. It is a positive feedback process where the ageostrophic, overturning secondary circulation in the cross-front plane accelerates the frontal sharpening until an arrest occurs through frontal instability and other forms of turbulent mixing. Several well-known types of oceanic frontal phenomena are surveyed, their impacts on oceanic system functioning are assessed, and future research is envisioned.
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Combining Modern and Paleoceanographic Perspectives on Ocean Heat Uptake
Vol. 13 (2021), pp. 255–281More LessMonitoring Earth's energy imbalance requires monitoring changes in the heat content of the ocean. Recent observational estimates indicate that ocean heat uptake is accelerating in the twenty-first century. Examination of estimates of ocean heat uptake over the industrial era, the Common Era of the last 2,000 years, and the period since the Last Glacial Maximum, 20,000 years ago, permits a wide perspective on modern-day warming rates. In addition, this longer-term focus illustrates how the dynamics of the deep ocean and the cryosphere were active in the past and are still active today. The large climatic shifts that started with the melting of the great ice sheets have involved significant ocean heat uptake that was sustained over centuries and millennia, and modern-ocean heat content changes are small by comparison.
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Historical Estimates of Surface Marine Temperatures
Vol. 13 (2021), pp. 283–311More LessSurface temperature documents our changing climate, and the marine record represents one of the longest widely distributed, observation-based estimates. Measurements of near-surface marine air temperature and sea-surface temperature have been recorded on platforms ranging from sailing ships to autonomous drifting buoys. The raw observations show an imprint of differing measurement methods and are sparse in certain periods and regions. This review describes how the real signal of global climate change can be determined from these sparse and noisy observations, including the quantification of measurement method–dependent biases and the reduction of spurious signals. Recent progress has come from analysis of the observations at increasing levels of granularity and from accounting for artifacts in the data that depend on platform types, measurement methods, and environmental conditions. Cutting across these effects are others caused by how the data were recorded, transcribed, and archived. These insights will be integrated into the next generation of global products quantified with validated estimates of uncertainty and the dependencies of its correlation structure. Further analysis of these records using improved data, metadata, and methods will certainly uncover more idiosyncrasies and new ways to improve the record.
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Marine Heatwaves
Vol. 13 (2021), pp. 313–342More LessOcean 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 thechoice of climatological baseline periods in defining extremes and how to communicate findings in the context of societal needs.
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Turbulence and Coral Reefs
Vol. 13 (2021), pp. 343–373More LessThe interaction of coral reefs, both chemically and physically, with the surrounding seawater is governed, at the smallest scales, by turbulence. Here, we review recent progress in understanding turbulence in the unique setting of coral reefs—how it influences flow and the exchange of mass and momentum both above and within the complex geometry of coral reef canopies. Flow above reefs diverges from canonical rough boundary layers due to their large and highly heterogeneous roughness and the influence of surface waves. Within coral canopies, turbulence is dominated by large coherent structures that transport momentum both into and away from the canopy, but it is also generated at smaller scales as flow is forced to move around branches or blades, creating wakes. Future work interpreting reef-related observations or numerical models should carefully consider the influence that spatial variation has on momentum and scalar flux.
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The Hydrodynamics of Jellyfish Swimming
Vol. 13 (2021), pp. 375–396More LessJellyfish have provided insight into important components of animal propulsion, such as suction thrust, passive energy recapture, vortex wall effects, and the rotational mechanics of turning. These traits are critically important to jellyfish because they must propel themselves despite severe limitations on force production imposed by rudimentary cnidarian muscular structures. Consequently, jellyfish swimming can occur only by careful orchestration of fluid interactions. Yet these mechanics may be more broadly instructive because they also characterize processes shared with other animal swimmers, whose structural and neurological complexity can obscure these interactions. In comparison with other animal models, the structural simplicity, comparative energetic efficiency, and ease of use in laboratory experimentation allow jellyfish to serve as favorable test subjects for exploration of the hydrodynamic bases of animal propulsion. These same attributes also make jellyfish valuable models for insight into biomimetic or bioinspired engineeringof swimming vehicles. Here, we review advances in understanding of propulsive mechanics derived from jellyfish models as a pathway toward the application of animal mechanics to vehicle designs.
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Marine Parasites and Disease in the Era of Global Climate Change
Vol. 13 (2021), pp. 397–420More LessClimate change affects ecological processes and interactions, including parasitism. Because parasites are natural components of ecological systems, as well as agents of outbreak and disease-induced mortality, it is important to summarize current knowledge of the sensitivity of parasites to climate and identify how to better predict their responses to it. This need is particularly great in marine systems, where the responses of parasites to climate variables are less well studied than those in other biomes. As examples of climate's influence on parasitism increase, they enable generalizations of expected responses as well as insight into useful study approaches, such as thermal performance curves that compare the vital rates of hosts and parasites when exposed to several temperatures across a gradient. For parasites not killed by rising temperatures, some simple physiological rules, including the tendency of temperature to increase the metabolism of ectotherms and increase oxygen stress on hosts, suggest that parasites’ intensity and pathologies might increase. In addition to temperature, climate-induced changes in dissolved oxygen, ocean acidity, salinity, and host and parasite distributions also affect parasitism and disease, but these factors are much less studied. Finally, because parasites are constituents of ecological communities, we must consider indirect and secondary effects stemming from climate-induced changes in host–parasite interactions, which may not be evident if these interactions are studied in isolation.
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Incorporating Biological Traits into Conservation Strategies
Vol. 13 (2021), pp. 421–443More LessImplementation of marine conservation strategies, such as increasing the numbers, extent, and effectiveness of protected areas (PAs), can help achieve conservation and restoration of ocean health and associated goods and services. Despite increasing recognition of the importance of including aspects of ecological functioning in PA design, the physical characteristics of habitats and simple measures of species diversity inform most PA designations. Marine and terrestrial ecologists have recently been using biological traits to assess community dynamics, functioning, and vulnerability to anthropogenic impacts. Here, we explore potential trait-based marine applications to advance PA design. We recommend strategies to integrate biological traits into (a) conservation objectives (e.g., by assessing and predicting impacts and vulnerability), (b) PA spatial planning (e.g., mapping ecosystem functions and functional diversity hot spots), and (c) time series monitoring protocols (e.g., using functional traits to detect recoveries). We conclude by emphasizing the need for pragmatic tools to improve the efficacy of spatial planning and monitoring efforts.
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Emerging Solutions to Return Nature to the Urban Ocean
Vol. 13 (2021), pp. 445–477More LessUrban 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 grayinfrastructure, 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.
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Ocean Optimism: Moving Beyond the Obituaries in Marine Conservation
Vol. 13 (2021), pp. 479–499More LessWhile the ocean has suffered many losses, there is increasing evidence that important progress is being made in marine conservation. Examples include striking recoveries of once-threatened species, increasing rates of protection of marine habitats, more sustainably managed fisheries and aquaculture, reductions in some forms of pollution, accelerating restoration of degraded habitats, and use of the ocean and its habitats to sequester carbon and provide clean energy. Many of these achievements have multiple benefits, including improved human well-being. Moreover, better understanding of how to implement conservation strategies effectively, new technologies and databases, increased integration of the natural and social sciences, and use of indigenous knowledge promise continued progress. Enormous challenges remain, and there is no single solution; successful efforts typically are neither quick nor cheap and require trust and collaboration. Nevertheless, a greater focus on solutions and successes will help them to become the norm rather than the exception.
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Amazon Sediment Transport and Accumulation Along the Continuum of Mixed Fluvial and Marine Processes
Vol. 13 (2021), pp. 501–536More LessSediment transfer from land to ocean begins in coastal settings and, for large rivers such as the Amazon, has dramatic impacts over thousands of kilometers covering diverse environmental conditions. In the relatively natural Amazon tidal river, combinations of fluvial and marine processes transition toward the ocean, affecting the transport and accumulation of sediment in floodplains and tributary mouths. The enormous discharge of Amazon fresh water causes estuarine processes to occur on the continental shelf, where much sediment accumulation creates a large clinoform structure and where additional sediment accumulates along its shoreward boundary in tidal flats and mangrove forests. Some remaining Amazon sediment is transported beyond the region near the river mouth, and fluvial forces on it diminish. Numerous perturbations to Amazon sediment transport and accumulation occur naturally, but human actions will likely dominate future change, and now is the time to document, understand, and mitigate their impacts.
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The Origin of Modern Atolls: Challenging Darwin's Deeply Ingrained Theory
Vol. 13 (2021), pp. 537–573More LessIn 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.
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