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: 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: 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: 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: Tides are changing worldwide at rates not explained by astronomical forcing. Rather, the observed evolution of tides and other long waves, such as storm surges, is influenced by shelf processes and changes to the roughness, depth, width, and length of embayments, estuaries, and tidal rivers. In this review, we focus on processes in estuaries and tidal rivers, because that is where the largest changes to tidal properties are occurring. Recent literature shows that changes in tidal amplitude have been ubiquitous worldwide over the past century, often in response to wetland reclamation, channel dredging, and other environmental changes. While tidal amplitude changes are sometimes slight (

Description: Monitoring 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. 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: In this article, we analyze the impacts of climate change on Antarctic marine ecosystems. Observations demonstrate large-scale changes in the physical variables and circulation of the Southern Ocean driven by warming, stratospheric ozone depletion, and a positive Southern Annular Mode. Alterations in the physical environment are driving change through all levels of Antarctic marine food webs, which differ regionally. The distributions of key species, such as Antarctic krill, are also changing. Differential responses among predators reflect differences in species ecology. The impacts of climate change on Antarctic biodiversity will likely vary for different communities and depend on species range. Coastal communities and those of sub-Antarctic islands, especially range-restricted endemic communities, will likely suffer the greatest negative consequences of climate change. Simultaneously, ecosystem services in the Southern Ocean will likely increase. Such decoupling of ecosystem services and endemic species will require consideration in the management of human activities such as fishing in Antarctic marine ecosystems.

Description: In the last few decades, numerous studies have investigated the impacts of simulated ocean acidification on marine species and communities, particularly those inhabiting dynamic coastal systems. Despite these research efforts, there are many gaps in our understanding, particularly with respect to physiological mechanisms that lead to pathologies. In this review, we trace how carbonate system disturbances propagate from the coastal environment into marine invertebrates and highlight mechanistic links between these disturbances and organism function. We also point toward several processes related to basic invertebrate biology that are severely understudied and prevent an accurate understanding of how carbonate system dynamics influence organismic homeostasis and fitness-related traits. We recommend that significant research effort be directed to studying cellular phenotypes of invertebrates acclimated or adapted to elevated seawater pCO2 using biochemical and physiological methods.

Description: Photosynthesis evolved in the ocean more than 2 billion years ago and is now performed by a wide range of evolutionarily distinct organisms, including both prokaryotes and eukaryotes. Our appreciation of their abundance, distributions, and contributions to primary production in the ocean has been increasing since they were first discovered in the seventeenth century and has now been enhanced by data emerging from the Tara Oceans project, which performed a comprehensive worldwide sampling of plankton in the upper layers of the ocean between 2009 and 2013. Largely using recent data from Tara Oceans, here we review the geographic distributions of phytoplankton in the global ocean and their diversity, abundance, and standing stock biomass. We also discuss how omics-based information can be incorporated into studies of photosynthesis in the ocean and show the likely importance of mixotrophs and photosymbionts.

Description: This narrative is a personal account of my evolution as a student of phytoplankton and the ocean. Initially I focused on phytoplankton nutrient physiology and uptake, later switching to photosynthetic physiology. Better models of photosynthesis naturally require a better understanding of spectral underwater light fields and absorption coefficients, which precipitated my involvement in the nascent field of bio-optical oceanography. Establishment of the now 34-year-old summer graduate course in ocean optics, which continues to attract students from around the globe, is a legacy of my jumping into optics. The importance of social interactions in advancing science cannot be underestimated; a prime example is how a TGIF gathering led to my immersion in the world of autonomous underwater vehicles for the past two decades of my career. Working with people who you like and respect is also critical; I believe collegial friendship played a major role in the great success of the 2008 North Atlantic Bloom Experiment.

Description: Implementation 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 used 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. 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: With the decline of reef-building corals on tropical reefs, sponges have emerged as an important component of changing coral reef ecosystems. Seemingly simple, sponges are highly diverse taxonomically, morphologically, and in terms of their relationships with symbiotic microbes, and they are one of nature's richest sources of novel secondary metabolites. Unlike most other benthic organisms, sponges have the capacity to disrupt boundary flow as they pump large volumes of seawater into the water column. This seawater is chemically transformed as it passes through the sponge body as a consequence of sponge feeding, excretion, and the activities of microbial symbionts, with important effects on carbon and nutrient cycling and on the organisms in the water column and on the adjacent reef. In this review, we critically evaluate developments in the recently dynamic research area of sponge ecology on tropical reefs and provide a perspective for future studies.

Description: Bays in coastal upwelling regions are physically driven and biochemically fueled by their interaction with open coastal waters. Wind-driven flow over the shelf imposes a circulation in the bay, which is also influenced by local wind stress and thermal bay–ocean density differences. Three types of bays are recognized based on the degree of exposure to coastal currents and winds (wide-open bays, square bays, and elongated bays), and the characteristic circulation and stratification patterns of each type are described. Retention of upwelled waters in bays allows for dense phytoplankton blooms that support productive bay ecosystems. Retention is also important for the accumulation of larvae, which accounts for high recruitment in bays. In addition, bays are coupled to the shelf ecosystem through export of plankton-rich waters during relaxation events. Ocean acidification and deoxygenation are a concern in bays because local extrema can develop beneath strong stratification.

Description: Much of the global cooling during ice ages arose from changes in ocean carbon storage that lowered atmospheric CO2. A slew of mechanisms, both physical and biological, have been proposed as key drivers of these changes. Here we discuss the current understanding of these mechanisms with a focus on how they altered the theoretically defined soft-tissue and biological disequilibrium carbon storage at the peak of the last ice age. Observations and models indicate a role for Antarctic sea ice through its influence on ocean circulation patterns, but other mechanisms, including changes in biological processes, must have been important as well, and may have been coordinated through links with global air temperature. Further research is required to better quantify the contributions of the various mechanisms, and there remains great potential to use the Last Glacial Maximum and the ensuing global warming as natural experiments from which to learn about climate-driven changes in the marine ecosystem.

Description: More than two-thirds of global biomass consists of vascular plants. A portion of the detritus they generate is carried into the oceans from land and highly productive blue carbon ecosystems—salt marshes, mangrove forests, and seagrass meadows. This large detrital input receives scant attention in current models of the global carbon cycle, though for blue carbon ecosystems, increasingly well-constrained estimates of biomass, productivity, and carbon fluxes, reviewed in this article, are now available. We show that the fate of this detritus differs markedly from that of strictly marine origin, because the former contains lignocellulose—an energy-rich polymer complex of cellulose, hemicelluloses, and lignin that is resistant to enzymatic breakdown. This complex can be depolymerized for nutritional purposes by specialized marine prokaryotes, fungi, protists, and invertebrates using enzymes such as glycoside hydrolases and lytic polysaccharide monooxygenases to release sugar monomers. The lignin component, however, is less readily depolymerized, and detritus therefore becomes lignin enriched, particularly in anoxic sediments, and forms a major carbon sink in blue carbon ecosystems. Eventual lignin breakdown releases a wide variety of small molecules that may contribute significantly to the oceanic pool of recalcitrant dissolved organic carbon. Marine carbon fluxes and sinks dependent on lignocellulosic detritus are important ecosystem services that are vulnerable to human interventions. These services must be considered when protecting blue carbon ecosystems and planning initiatives aimed at mitigating anthropogenic carbon emissions.

Description: Jellyfish 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 engineering of 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. 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: Over 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, ?(107) m, and the turbulence, ? (10−2) 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). 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: Apex predators play pivotal roles in marine ecosystems, mediated principally through diet and nutrition. Yet, compared with terrestrial animals, the nutritional ecology of marine predators is poorly understood. One reason is that the field has adhered to an approach that evaluates diet principally in terms of energy gain. Studies in terrestrial systems, by contrast, increasingly adopt a multidimensional approach, the nutritional geometry framework, that distinguishes specific nutrients and calories. We provide evidence that a nutritional approach is likewise relevant to marine apex predators, then demonstrate how nutritional geometry can characterize the nutrient and energy content of marine prey. Next, we show how this framework can be used to reconceptualize ecological interactions via the ecological niche concept, and close with a consideration of its application to problems in marine predator research.

Description: My 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. 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: While 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. 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: Advances in technologies for molecular observation are leading to novel types of data, including gene, transcript, protein, and metabolite levels, which are fundamentally different from the types traditionally compared with microbial ecosystem models, such as biomass (e.g., chlorophyll a) and nutrient concentrations. A grand challenge is to use these data to improve predictive models and use models to explain observed patterns. This article presents a framework that aligns observations and models along the dimension of abstraction or biological organization—from raw sequences to ecosystem patterns for observations, and from sequence simulators to ecological theory for models. It then reviews 16 studies that compared model results with molecular observations. Molecular data can and are being combined with microbial ecosystem models, but to keep up with and take advantage of the full scope of observations, models need to become more mechanistically detailed and complex, which is a technical and cultural challenge for the ecological modeling community.

Description: Climate 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. 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 biogeochemical cycles of trace elements and their isotopes (TEIs) constitute an active area of oceanographic research due to their role as essential nutrients for marine organisms and their use as tracers of oceanographic processes. Selected TEIs also provide diagnostic information about the physical, geological, and chemical processes that supply or remove solutes in the ocean. Many of these same TEIs provide information about ocean conditions in the past, as their imprint on marine sediments can be interpreted to reflect changes in ocean circulation, biological productivity, the ocean carbon cycle, and more. Other TEIs have been introduced as the result of human activities and are considered contaminants. The development and implementation of contamination-free methods for collecting and analyzing samples for TEIs revolutionized marine chemistry, revealing trace element distributions with oceanographically consistent features and new insights about the processes regulating them. Despite these advances, the volume and geographic coverage of high-quality TEI data by the end of the twentieth century were insufficient to constrain their global biogeochemical cycles. To accelerate progress in this field of research, marine geochemists developed a coordinated international effort to systematically study the marine biogeochemical cycles of TEIs—the GEOTRACES program. Following a decade of planning and implementation, GEOTRACES launched its main field effort in 2010. This review, roughly midway through the field program, summarizes the steps involved in designing the program, its management structure, and selected findings.

Description: Compared with terrestrial ecosystems, marine ecosystems have a higher proportion of heterotrophic biomass. Building from this observation, we define the North Atlantic biome as the region where the large, lipid-rich copepod Calanus finmarchicus is the dominant mesozooplankton species. This species is superbly adapted to take advantage of the intense pulse of productivity associated with the North Atlantic spring bloom. Most of the characteristic North Atlantic species, including cod, herring, and right whales, rely on C. finmarchicus either directly or indirectly. The notion of a biome rests inherently on an assumption of stability, yet conditions in the North Atlantic are anything but stable. Humans have reduced the abundance of many fish and whales (though some recovery is underway). Humans are also introducing physical and chemical trends associated with global climate change. Thus, the future of the North Atlantic depends on the biome's newest species, Homo sapiens.

Description: Glacial–interglacial cycles have constituted a primary mode of climate variability over the last 2.6 million years of Earth's history. While glacial periods cannot be seen simply as a reverse analogue of future warming, they offer an opportunity to test our understanding of the response of precipitation patterns to a much wider range of conditions than we have been able to directly observe. This review explores key features of precipitation patterns associated with glacial climates, which include drying in large regions of the tropics and wetter conditions in substantial parts of the subtropics and midlatitudes. I describe the evidence for these changes and examine the potential causes of hydrological changes during glacial periods. Central themes that emerge include the importance of atmospheric circulation changes in determining glacial–interglacial precipitation changes at the regional scale, the need to take into account climatic factors beyond local precipitation amount when interpreting proxy data, and the role of glacial conditions in suppressing the strength of Northern Hemisphere monsoon systems.

Description: One major objective of aquatic microbial ecology is to understand the distribution of microbial populations over space and time and in response to environmental factors. Perhaps more importantly, it is crucial to quantify how those microbial cells affect biogeochemical processes of interest, such as primary production, nitrogen cycling, or the breakdown of pollutants. One valuable approach to link microbial identity to activity is to carry out incubations with stable-isotope-labeled substrates and then quantify the isotope incorporation by individual microbial cells using nanoscale secondary ion mass spectrometry (NanoSIMS). This review summarizes recent efforts in this field, highlights novel methods, describes studies investigating rare metabolisms as well as widespread microbial activity, and hopes to provide a framework to increase the use and capabilities of NanoSIMS for microbial biogeochemical studies in the future.

Description: Biogeochemical-Argo (BGC-Argo) is a network of profiling floats carrying sensors that enable observation of as many as six essential biogeochemical and bio-optical variables: oxygen, nitrate, pH, chlorophyll a, suspended particles, and downwelling irradiance. This sensor network represents today's most promising strategy for collecting temporally and vertically resolved observations of biogeochemical properties throughout the ocean. All data are freely available within 24 hours of transmission. These data fill large gaps in ocean-observing systems and support three ambitions: gaining a better understanding of biogeochemical processes (e.g., the biological carbon pump and air–sea CO2 exchanges) and evaluating ongoing changes resulting from increasing anthropogenic pressure (e.g., acidification and deoxygenation); managing the ocean (e.g., improving the global carbon budget and developing sustainable fisheries); and carrying out exploration for potential discoveries. The BGC-Argo network has already delivered extensive high-quality global data sets that have resulted in unique scientific outcomes from regional to global scales. With the proposed expansion of BGC-Argo in the near future, this network has the potential to become a pivotal observation system that links satellite and ship-based observations in a transformative manner.

Description: Experimental evolution and the associated theory are underutilized in marine microbial studies; the two fields have developed largely in isolation. Here, we review evolutionary tools for addressing four key areas of ocean global change biology: linking plastic and evolutionary trait changes, the contribution of environmental variability to determining trait values, the role of multiple environmental drivers in trait change, and the fate of populations near their tolerance limits. Wherever possible, we highlight which data from marine studies could use evolutionary approaches and where marine model systems can advance our understanding of evolution. Finally, we discuss the emerging field of marine microbial experimental evolution. We propose a framework linking changes in environmental quality (defined as the cumulative effect on population growth rate) with population traits affecting evolutionary potential, in order to understand which evolutionary processes are likely to be most important across a range of locations for different types of marine microbes.

Description: The geographic distributions of marine species are changing rapidly, with leading range edges following climate poleward, deeper, and in other directions and trailing range edges often contracting in similar directions. These shifts have their roots in fine-scale interactions between organisms and their environment—including mosaics and gradients of temperature and oxygen—mediated by physiology, behavior, evolution, dispersal, and species interactions. These shifts reassemble food webs and can have dramatic consequences. Compared with species on land, marine species are more sensitive to changing climate but have a greater capacity for colonization. These differences suggest that species cope with climate change at different spatial scales in the two realms and that range shifts across wide spatial scales are a key mechanism at sea. Additional research is needed to understand how processes interact to promote or constrain range shifts, how the dominant responses vary among species, and how the emergent communities of the future ocean will function.

Description: Microbes 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. 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: Polysaccharides 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 and characterize 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. 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: Salt marshes are recognized as valuable resources that are threatened by climate change and human activities. Better management and planning for these ecosystems will depend on understanding which marshes are most vulnerable, what is driving their change, and what their future trajectory is likely to be. Both observations and models have provided inconsistent answers to these questions, likely in part because of comparisons among sites and/or models that differ significantly in their characteristics and processes. Some of these differences almost certainly arise from processes that are not fully accounted for in marsh morphodynamic models. Here, we review distinguishing properties of marshes, important processes missing from many morphodynamic models, and key measurements missing from many observational studies. We then suggest some comparisons between models and observations that will provide critical tests and insights to improve our ability to forecast future change in these coastal landscapes.

Description: Ocean ecosystems are experiencing unprecedented rates of climate and anthropogenic change, which can often initiate stress in marine organisms. Symbioses, or associations between different organisms, are plentiful in the ocean and could play a significant role in facilitating organismal adaptations to stressful ocean conditions. This article reviews current knowledge about the role of symbiosis in marine organismal acclimation and adaptation. It discusses stress and adaptations in symbioses from coral reef ecosystems, which are among the most affected environments in the ocean, including the relationships between corals and microalgae, corals and bacteria, anemones and clownfish, and cleaner fish and client fish. Despite the importance of this subject, knowledge of how marine organisms adapt to stress is still limited, and there are vast opportunities for research and technological development in this area. Attention to this subject will enhance our understanding of the capacity of symbioses to alleviate organismal stress in the oceans.

Description: The 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. 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: Millions 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. 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.