Annual Review of Marine Science - Volume 9, 2017

Quoting the ancient Romans: Audentes Fortuna iuvat. Being in the right place at the right time is useless if you do not grasp your Fortuna and build upon it. In this article, I expound on the milestones of my multiform research career, which over more than 40 years brought me from Venice to California to MIT; from the Venice problem to highly nonlinear, coherent structures in the ocean and atmosphere; and from the mare nostrum (the Mediterranean Sea), a laboratory for global processes, to the tropical ocean-atmosphere systems and regional coupled climate models of the Maritime Continent. The climate system, with its daunting complexity, is arguably the greatest challenge for, and the future of, the entirety of the earth sciences. Finally, living in and working for Venice has been the privilege and Fortuna of my life.

The ocean has undergone several profound biogeochemical transformations in its 4-billion-year history, and these were an integral part of the coevolution of life and the planet. This review focuses on changes in ocean redox state as controlled by changes in biological activity, nutrient concentrations, and atmospheric O2. Motivated by disparate interpretations of available geochemical data, we aim to show how quantitative modeling—spanning microbial mats, shelf seas, and the open ocean—can help constrain past ocean biogeochemical redox states and show what caused transformations between them. We outline key controls on ocean redox structure and review pertinent proxies and their interpretation. We then apply this quantitative framework to three key questions: How did the origin of oxygenic photosynthesis transform ocean biogeochemistry? How did the Great Oxidation transform ocean biogeochemistry? And how was ocean biogeochemistry transformed in the Neoproterozoic-Paleozoic?

Surface drifting buoys, or drifters, are used in oceanographic and climate research, oil spill tracking, weather forecasting, search and rescue operations, calibration and validation of velocities from high-frequency radar and from altimeters, iceberg tracking, and support of offshore drilling operations. In this review, we present a brief history of drifters, from the message in a bottle to the latest satellite-tracked, multisensor drifters. We discuss the different types of drifters currently used for research and operations as well as drifter designs in development. We conclude with a discussion of the various properties that can be observed with drifters, with heavy emphasis on a critical process that cannot adequately be observed by any other instrument: dispersion in the upper ocean, driven by turbulence at scales from waves through the submesoscale to the large-scale geostrophic eddies.

Abrupt changes in climate have occurred in many locations around the globe over the last glacial cycle, with pronounced temperature swings on timescales of decades or less in the North Atlantic. The global pattern of these changes suggests that they reflect variability in the Atlantic meridional overturning circulation (AMOC). This review examines the evidence from ocean sediments for ocean circulation change over these abrupt events. The evidence for changes in the strength and structure of the AMOC associated with the Younger Dryas and many of the Heinrich events is strong. Although it has been difficult to directly document changes in the AMOC over the relatively short Dansgaard-Oeschger events, there is recent evidence supporting AMOC changes over most of these oscillations as well. The lack of direct evidence for circulation changes over the shortest events leaves open the possibility of other driving mechanisms for millennial-scale climate variability.

The kinetic energy in ocean currents, or marine hydrokinetic (MHK) energy, is a renewable energy resource that can help meet global energy requirements. An ocean circulation model–based census shows that subtropical surface western boundary currents (WBCs) are the only nearshore, large-scale currents swift enough to drive large electricity-generating ocean turbines envisioned for future use. We review several WBCs in the context of kinetic energy extraction. The power density in the Gulf Stream off North Carolina at times reaches several thousand watts per square meter at 75 m below the surface, and the annual average power is approximately 500–1,000 W m⁻². Significant fluctuations occur with periods of 3–20 days (Gulf Stream meanders) and weeks to months (Gulf Stream path shifts). Interannual variations in annual average power occur because of year-to-year changes in these WBC motions. No large-scale turbines presently exist, and the road to establishing MHK facilities in WBCs will encounter challenges that are similar in many aspects to those associated with the development of offshore wind power.

Since preindustrial times, the ocean has removed from the atmosphere 41% of the carbon emitted by human industrial activities. Despite significant uncertainties, the balance of evidence indicates that the globally integrated rate of ocean carbon uptake is increasing in response to increasing atmospheric CO2 concentrations. The El Niño–Southern Oscillation in the equatorial Pacific dominates interannual variability of the globally integrated sink. Modes of climate variability in high latitudes are correlated with variability in regional carbon sinks, but mechanistic understanding is incomplete. Regional sink variability, combined with sparse sampling, means that the growing oceanic sink cannot yet be directly detected from available surface data. Accurate and precise shipboard observations need to be continued and increasingly complemented with autonomous observations. These data, together with a variety of mechanistic and diagnostic models, are needed for better understanding, long-term monitoring, and future projections of this critical climate regulation service.

Carbon preservation in marine sediments, supplemented by that in large lakes, is the primary mechanism that moves carbon from the active surficial carbon cycle to the slower geologic carbon cycle. Preservation rates are low relative to the rates at which carbon moves between surface pools, which has led to the preservation term largely being ignored when evaluating anthropogenic forcing of the global carbon cycle. However, a variety of anthropogenic drivers—including ocean warming, deoxygenation, and acidification, as well as human-induced changes in sediment delivery to the ocean and mixing and irrigation of continental margin sediments—all work to decrease the already small carbon preservation term. These drivers affect the cycling of both carbonate and organic carbon in the ocean. The overall effect of anthropogenic forcing in the modern ocean is to decrease delivery of carbon to sediments, increase sedimentary dissolution and remineralization, and subsequently decrease overall carbon preservation.

The events that followed the Tohoku earthquake and tsunami on March 11, 2011, included the loss of power and overheating at the Fukushima Daiichi nuclear power plants, which led to extensive releases of radioactive gases, volatiles, and liquids, particularly to the coastal ocean. The fate of these radionuclides depends in large part on their oceanic geochemistry, physical processes, and biological uptake. Whereas radioactivity on land can be resampled and its distribution mapped, releases to the marine environment are harder to characterize owing to variability in ocean currents and the general challenges of sampling at sea. Five years later, it is appropriate to review what happened in terms of the sources, transport, and fate of these radionuclides in the ocean. In addition to the oceanic behavior of these contaminants, this review considers the potential health effects and societal impacts.

Plastics contamination in the marine environment was first reported nearly 50 years ago, less than two decades after the rise of commercial plastics production, when less than 50 million metric tons were produced per year. In 2014, global plastics production surpassed 300 million metric tons per year. Plastic debris has been detected worldwide in all major marine habitats, in sizes from microns to meters. In response, concerns about risks to marine wildlife upon exposure to the varied forms of plastic debris have increased, stimulating new research into the extent and consequences of plastics contamination in the marine environment. Here, I present a framework to evaluate the current understanding of the sources, distribution, fate, and impacts of marine plastics. Despite remaining knowledge gaps in mass budgeting and challenges in investigating ecological impacts, the increasing evidence of the ubiquity of plastics contamination in the marine environment, the continued rapid growth in plastics production, and the evidence—albeit limited—of demonstrated impacts to marine wildlife support immediate implementation of source-reducing measures to decrease the potential risks of plastics in the marine ecosystem.

SAR11 is a group of small, carbon-oxidizing bacteria that reach a global estimated population size of 2.4×10²⁸ cells—approximately 25% of all plankton. They are found throughout the oceans but reach their largest numbers in stratified, oligotrophic gyres, which are an expanding habitat in the warming oceans. SAR11 likely had a Precambrian origin and, over geological time, evolved into the niche of harvesting labile, low-molecular-weight dissolved organic matter (DOM). SAR11 cells are minimal in size and complexity, a phenomenon known as streamlining that is thought to benefit them by lowering the material costs of replication and maximizing transport functions that are essential to competition at ultralow nutrient concentrations. One of the surprises in SAR11 metabolism is their ability to both oxidize and produce a variety of volatile organic compounds that can diffuse into the atmosphere. SAR11 cells divide slowly and lack many forms of regulation commonly used by bacterial cells to adjust to changing environmental conditions. As a result of genome reduction, they require an unusual range of nutrients, which leads to complex biochemical interactions with other plankton. The study of SAR11 is providing insight into the biogeochemistry of labile DOM and is affecting microbiology beyond marine science by providing a model for understanding the evolution and function of streamlined cells.

Quorum sensing (QS) is a form of chemical communication used by certain bacteria that regulates a wide range of biogeochemically important bacterial behaviors. Although QS was first observed in a marine bacterium nearly four decades ago, only in the past decade has there been a rise in interest in the role that QS plays in the ocean. It has become clear that QS, regulated by signals such as acylated homoserine lactones (AHLs) or furanosyl-borate diesters [autoinducer-2 (AI-2) molecules], is involved in important processes within the marine carbon cycle, in the health of coral reef ecosystems, and in trophic interactions between a range of eukaryotes and their bacterial associates. The most well-studied QS systems in the ocean occur in surface-attached (biofilm) communities and rely on AHL signaling. AHL-QS is highly sensitive to the chemical and biological makeup of the environment and may respond to anthropogenic change, including ocean acidification and rising sea surface temperatures.

Coccolithophores occupy a special position within the marine phytoplankton because of their production of intricate calcite scales, or coccoliths. Coccolithophores are major contributors to global ocean calcification and long-term carbon fluxes. The intracellular production of coccoliths requires modifications to cellular ultrastructure and metabolism that are surveyed here. In addition to calcification, which appears to have evolved with a diverse range of functions, several other remarkable features that likely underpin the ecological and evolutionary success of coccolithophores have recently been uncovered. These include complex and varied life cycle strategies related to abiotic and biotic interactions as well as a range of novel metabolic pathways and nutritional strategies. Together with knowledge of coccolithophore genetic and physiological variability, these findings are beginning to shed new light on species diversity, distribution, and ecological adaptation. Further advances in genetics and functional characterization at the cellular level will likely to lead to a rapid increase in this understanding.

Mixotrophs are important components of the bacterioplankton, phytoplankton, microzooplankton, and (sometimes) zooplankton in coastal and oceanic waters. Bacterivory among the phytoplankton may be important for alleviating inorganic nutrient stress and may increase primary production in oligotrophic waters. Mixotrophic phytoflagellates and dinoflagellates are often dominant components of the plankton during seasonal stratification. Many of the microzooplankton grazers, including ciliates and Rhizaria, are mixotrophic owing to their retention of functional algal organelles or maintenance of algal endosymbionts. Phototrophy among the microzooplankton may increase gross growth efficiency and carbon transfer through the microzooplankton to higher trophic levels. Characteristic assemblages of mixotrophs are associated with warm, temperate, and cold seas and with stratification, fronts, and upwelling zones. Modeling has indicated that mixotrophy has a profound impact on marine planktonic ecosystems and may enhance primary production, biomass transfer to higher trophic levels, and the functioning of the biological carbon pump.

Deep-sea fishes inhabit ∼75% of the biosphere and are a critical part of deep-sea food webs. Diet analysis and more recent trophic biomarker approaches, such as stable isotopes and fatty-acid profiles, have enabled the description of feeding guilds and an increased recognition of the vertical connectivity in food webs in a whole-water-column sense, including benthic-pelagic coupling. Ecosystem modeling requires data on feeding rates; the available estimates indicate that deep-sea fishes have lower per-individual feeding rates than coastal and epipelagic fishes, but the overall predation impact may be high. A limited number of studies have measured the vertical flux of carbon by mesopelagic fishes, which appears to be substantial. Anthropogenic activities are altering deep-sea ecosystems and their services, which are mediated by trophic interactions. We also summarize outstanding data gaps.

Baleen whales are gigantic obligate filter feeders that exploit aggregations of small-bodied prey in littoral, epipelagic, and mesopelagic ecosystems. At the extreme of maximum body size observed among mammals, baleen whales exhibit a unique combination of high overall energetic demands and low mass-specific metabolic rates. As a result, most baleen whale species have evolved filter-feeding mechanisms and foraging strategies that take advantage of seasonally abundant yet patchily and ephemerally distributed prey resources. New methodologies consisting of multi-sensor tags, active acoustic prey mapping, and hydrodynamic modeling have revolutionized our ability to study the physiology and ecology of baleen whale feeding mechanisms. Here, we review the current state of the field by exploring several hypotheses that aim to explain how baleen whales feed. Despite significant advances, major questions remain about the processes that underlie these extreme feeding mechanisms, which enabled the evolution of the largest animals of all time.

Diapause is a type of dormancy that requires preparation, typically precedes the onset of unfavorable conditions, and necessitates a period of arrest before development can proceed. Two ecologically important groups of copepods have incorporated diapausing stages into their life histories. In freshwater, estuarine, and coastal environments, species within the Centropagoidea superfamily can produce resting eggs containing embryos that may be quiescent, diapausing, or in some intermediate state. Resting eggs sink into the sediments, remain viable over months to years, and form a reservoir from which the planktonic population is reestablished. In coastal and oceanic environments, copepods within the Calanidae and Eucalanidae families can enter diapause during late juvenile (copepodid) or adult stages. These copepods accumulate large amounts of lipids before they migrate into deep water and diapause for several months. Through respiration, diapausing copepods may sequester more carbon in the deep ocean than any other biogeochemical process, and changes in diapause phenology associated with climate change (particularly reduction in diapause duration) could have a significant impact not only on regional ecosystems, but on global climate as well.

Marine zooplankton comprise a phylogenetically and functionally diverse assemblage of protistan and metazoan consumers that occupy multiple trophic levels in pelagic food webs. Within this complex network, carbon flows via alternative zooplankton pathways drive temporal and spatial variability in production-grazing coupling, nutrient cycling, export, and transfer efficiency to higher trophic levels. We explore current knowledge of the processing of zooplankton food ingestion by absorption, egestion, respiration, excretion, and growth (production) processes. On a global scale, carbon fluxes are reasonably constrained by the grazing impact of microzooplankton and the respiratory requirements of mesozooplankton but are sensitive to uncertainties in trophic structure. The relative importance, combined magnitude, and efficiency of export mechanisms (mucous feeding webs, fecal pellets, molts, carcasses, and vertical migrations) likewise reflect regional variability in community structure. Climate change is expected to broadly alter carbon cycling by zooplankton and to have direct impacts on key species.

Coral reefs provide critical services to coastal communities, and these services rely on ecosystem functions threatened by stressors. By summarizing the threats to the functioning of reefs from fishing, climate change, and decreasing water quality, we highlight that these stressors have multiple, conflicting effects on functionally similar groups of species and their interactions, and that the overall effects are often uncertain because of a lack of data or variability among taxa. The direct effects of stressors on links among functional groups, such as predator-prey interactions, are particularly uncertain. Using qualitative modeling, we demonstrate that this uncertainty of stressor impacts on functional groups (whether they are positive, negative, or neutral) can have significant effects on models of ecosystem stability, and reducing uncertainty is vital for understanding changes to reef functioning. This review also provides guidance for future models of reef functioning, which should include interactions among functional groups and the cumulative effect of stressors.

Anchovy and sardine populated productive ocean regions over hundreds of thousands of years under a naturally varying climate, and are now subject to climate change of equal or greater magnitude occurring over decades to centuries. We hypothesize that anchovy and sardine populations are limited in size by the supply of nitrogen from outside their habitats originating from upwelling, mixing, and rivers. Projections of the responses of anchovy and sardine to climate change rely on a range of model types and consideration of the effects of climate on lower trophic levels, the effects of fishing on higher trophic levels, and the traits of these two types of fish. Distribution, phenology, nutrient supply, plankton composition and production, habitat compression, fishing, and acclimation and adaptation may be affected by ocean warming, acidification, deoxygenation, and altered hydrology. Observations of populations and evaluation of model skill are essential to resolve the effects of climate change on these fish.