Showing papers in "Annual Review of Marine Science in 2016"

Multiple Stressors in a Changing World: The Need for an Improved Perspective on Physiological Responses to the Dynamic Marine Environment

Abstract: Abiotic conditions (e.g., temperature and pH) fluctuate through time in most marine environments, sometimes passing intensity thresholds that induce physiological stress. Depending on habitat and season, the peak intensity of different abiotic stressors can occur in or out of phase with one another. Thus, some organisms are exposed to multiple stressors simultaneously, whereas others experience them sequentially. Understanding these physicochemical dynamics is critical because how organisms respond to multiple stressors depends on the magnitude and relative timing of each stressor. Here, we first discuss broad patterns of covariation between stressors in marine systems at various temporal scales. We then describe how these dynamics will influence physiological responses to multi-stressor exposures. Finally, we summarize how multi-stressor effects are currently assessed. We find that multi-stressor experiments have rarely incorporated naturalistic physicochemical variation into their designs, and emphasize the importance of doing so to make ecologically relevant inferences about physiological responses to global change.

Mechanisms of Physical-Biological-Biogeochemical Interaction at the Oceanic Mesoscale

Abstract: Mesoscale phenomena are ubiquitous and highly energetic features of ocean circulation. Their influence on biological and biogeochemical processes varies widely, stemming not only from advective transport but also from the generation of variations in the environment that affect biological and chemical rates. The ephemeral nature of mesoscale features in the ocean makes it difficult to elucidate the attendant mechanisms of physical-biological-biogeochemical interaction, necessitating the use of multidisciplinary approaches involving in situ observations, remote sensing, and modeling. All three aspects are woven through this review in an attempt to synthesize current understanding of the topic, with particular emphasis on novel developments in recent years.

The Impact of Submesoscale Physics on Primary Productivity of Plankton.

Abstract: Life in the ocean relies on the photosynthetic production of phytoplankton, which is influenced by the availability of light and nutrients that are modulated by a host of physical processes. Submesoscale processes are particularly relevant to phytoplankton productivity because the timescales on which they act are similar to those of phytoplankton growth. Their dynamics are associated with strong vorticity and strain rates that occur on lateral scales of 0.1–10 km. They can support vertical velocities as large as 100 m d−1 and play a crucial role in transporting nutrients into the sunlit ocean for phytoplankton production. In regimes with deep surface mixed layers, submesoscale instabilities can cause stratification within days, thereby increasing light exposure for phytoplankton trapped close to the surface. These instabilities help to create and maintain localized environments that favor the growth of phytoplankton, contribute to productivity, and cause enormous heterogeneity in the abundance of phytopla...

Mangrove sedimentation and response to relative sea-level rise

Abstract: Mangroves occur on upper intertidal shorelines in the tropics and subtropics. Complex hydrodynamic and salinity conditions, related primarily to elevation and hydroperiod, influence mangrove distributions; this review considers how these distributions change over time. Accumulation rates of allochthonous and autochthonous sediment, both inorganic and organic, vary between and within different settings. Abundant terrigenous sediment can form dynamic mudbanks, and tides redistribute sediment, contrasting with mangrove peat in sediment-starved carbonate settings. Sediments underlying mangroves sequester carbon but also contain paleoenvironmental records of adjustments to past sea-level changes. Radiometric dating indicates long-term sedimentation, whereas measurements made using surface elevation tables and marker horizons provide shorter perspectives, indicating shallow subsurface processes of root growth and substrate autocompaction. Many tropical deltas also experience deep subsidence, which augments relative sea-level rise. The persistence of mangroves implies an ability to cope with moderately high rates of relative sea-level rise. However, many human pressures threaten mangroves, resulting in a continuing decline in their extent throughout the tropics.

Near-Inertial Internal Gravity Waves in the Ocean

Abstract: We review the physics of near-inertial waves (NIWs) in the ocean and the observations, theory, and models that have provided our present knowledge. NIWs appear nearly everywhere in the ocean as a spectral peak at and just above the local inertial period f, and the longest vertical wavelengths can propagate at least hundreds of kilometers toward the equator from their source regions; shorter vertical wavelengths do not travel as far and do not contain as much energy, but lead to turbulent mixing owing to their high shear. NIWs are generated by a variety of mechanisms, including the wind, nonlinear interactions with waves of other frequencies, lee waves over bottom topography, and geostrophic adjustment; the partition among these is not known, although the wind is likely the most important. NIWs likely interact strongly with mesoscale and submesoscale motions, in ways that are just beginning to be understood.

Ocean Research Enabled by Underwater Gliders.

Abstract: Underwater gliders are autonomous underwater vehicles that profile vertically by changing their buoyancy and use wings to move horizontally. Gliders are useful for sustained observation at relatively fine horizontal scales, especially to connect the coastal and open ocean. In this review, research topics are grouped by time and length scales. Large-scale topics addressed include the eastern and western boundary currents and the regional effects of climate variability. The accessibility of horizontal length scales of order 1 km allows investigation of mesoscale and submesoscale features such as fronts and eddies. Because the submesoscales dominate vertical fluxes in the ocean, gliders have found application in studies of biogeochemical processes. At the finest scales, gliders have been used to measure internal waves and turbulent dissipation. The review summarizes gliders' achievements to date and assesses their future in ocean observation.

Growth Rates of Microbes in the Oceans

Abstract: A microbe's growth rate helps to set its ecological success and its contribution to food web dynamics and biogeochemical processes. Growth rates at the community level are constrained by biomass and trophic interactions among bacteria, phytoplankton, and their grazers. Phytoplankton growth rates are approximately 1 d−1, whereas most heterotrophic bacteria grow slowly, close to 0.1 d−1; only a few taxa can grow ten times as fast. Data from 16S rRNA and other approaches are used to speculate about the growth rate and the life history strategy of SAR11, the most abundant clade of heterotrophic bacteria in the oceans. These strategies are also explored using genomic data. Although the methods and data are imperfect, the available data can be used to set limits on growth rates and thus on the timescale for changes in the composition and structure of microbial communities.

Changes in ocean heat, carbon content, and ventilation : a review of the first decade of GO-SHIP Global Repeat Hydrography

Abstract: Global ship-based programs, with highly accurate, full water column physical and biogeochemical observations repeated decadally since the 1970s, provide a crucial resource for documenting ocean change. The ocean, a central component of Earth's climate system, is taking up most of Earth's excess anthropogenic heat, with about 19% of this excess in the abyssal ocean beneath 2,000 m, dominated by Southern Ocean warming. The ocean also has taken up about 27% of anthropogenic carbon, resulting in acidification of the upper ocean. Increased stratification has resulted in a decline in oxygen and increase in nutrients in the Northern Hemisphere thermocline and an expansion of tropical oxygen minimum zones. Southern Hemisphere thermocline oxygen increased in the 2000s owing to stronger wind forcing and ventilation. The most recent decade of global hydrography has mapped dissolved organic carbon, a large, bioactive reservoir, for the first time and quantified its contribution to export production (∼20%) and deep-ocean oxygen utilization. Ship-based measurements also show that vertical diffusivity increases from a minimum in the thermocline to a maximum within the bottom 1,500 m, shifting our physical paradigm of the ocean's overturning circulation.

Characteristic Sizes of Life in the Oceans, from Bacteria to Whales*

Abstract: The size of an individual organism is a key trait to characterize its physiology and feeding ecology. Size-based scaling laws may have a limited size range of validity or undergo a transition from one scaling exponent to another at some characteristic size. We collate and review data on size-based scaling laws for resource acquisition, mobility, sensory range, and progeny size for all pelagic marine life, from bacteria to whales. Further, we review and develop simple theoretical arguments for observed scaling laws and the characteristic sizes of a change or breakdown of power laws. We divide life in the ocean into seven major realms based on trophic strategy, physiology, and life history strategy. Such a categorization represents a move away from a taxonomically oriented description toward a trait-based description of life in the oceans. Finally, we discuss life forms that transgress the simple size-based rules and identify unanswered questions.

Sources, Ages, and Alteration of Organic Matter in Estuaries.

Abstract: Understanding the processes influencing the sources and fate of organic matter (OM) in estuaries is important for quantifying the contributions of carbon from land and rivers to the global carbon budget of the coastal ocean. Estuaries are sites of high OM production and processing, and understanding biogeochemical processes within these regions is key to quantifying organic carbon (Corg) budgets at the land-ocean margin. These regions provide vital ecological services, including nutrient filtration and protection from floods and storm surge, and provide habitat and nursery areas for numerous commercially important species. Human activities have modified estuarine systems over time, resulting in changes in the production, respiration, burial, and export of Corg. Corg in estuaries is derived from aquatic, terrigenous, and anthropogenic sources, with each source exhibiting a spectrum of ages and lability. The complex source and age characteristics of Corg in estuaries complicate our ability to trace OM along the river-estuary-coastal ocean continuum. This review focuses on the application of organic biomarkers and compound-specific isotope analyses to estuarine environments and on how these tools have enhanced our ability to discern natural sources of OM, trace their incorporation into food webs, and enhance understanding of the fate of Corg within estuaries and their adjacent waters.

The Great Diadema antillarum Die-Off: 30 Years Later*

Abstract: In 1983–1984, the sea urchin Diadema antillarum suffered mass mortality throughout the Caribbean, Florida, and Bermuda. The demise of this herbivore contributed to a phase shift of Caribbean reefs from coral-dominated to alga-dominated communities. A compilation of published data of D. antillarum population densities shows that there has been moderate recovery since 1983, with the highest rates on islands of the eastern Caribbean. On the average the current population densities are approximately 12% of those before the die-off, apparently because of recruitment limitation, but the exact factors that are constraining the recovery are unclear. Scattered D. antillarum cohorts in some localities and aggregation of settled individuals in shallow water have created zones of higher herbivory in which juvenile coral recruitment, survivorship, and growth are higher than they are in alga-dominated areas. Unlike other stressors on Caribbean coral reefs, recent changes in D. antillarum populations progress toward aid...

Slow Microbial Life in the Seabed.

Abstract: Global microbial cell numbers in the seabed exceed those in the overlying water column, yet these organisms receive less than 1% of the energy fixed as organic matter in the ocean. The microorganisms of this marine deep biosphere subsist as stable and diverse communities with extremely low energy availability. Growth is exceedingly slow, possibly regulated by virus-induced mortality, and the mean generation times are tens to thousands of years. Intermediate substrates such as acetate are maintained at low micromolar concentrations, yet their turnover time may be several hundred years. Owing to slow growth, a cell community may go through only 10,000 generations from the time it is buried beneath the mixed surface layer until it reaches a depth of tens of meters several million years later. We discuss the efficiency of the energy-conserving machinery of subsurface microorganisms and how they may minimize energy consumption through necessary maintenance, repair, and growth.

Nitrogen and Oxygen Isotopic Studies of the Marine Nitrogen Cycle

Abstract: The marine nitrogen cycle is a complex web of microbially mediated reactions that control the inventory, distribution, and speciation of nitrogen in the marine environment. Because nitrogen is a major nutrient that is required by all life, its availability can control biological productivity and ecosystem structure in both surface and deep-ocean communities. Stable isotopes of nitrogen and oxygen in nitrate and nitrite have provided new insights into the rates and distributions of marine nitrogen cycle processes, especially when analyzed in combination with numerical simulations of ocean circulation and biogeochemistry. This review highlights the insights gained from dual-isotope studies applied at regional to global scales and their incorporation into oceanic biogeochemical models. These studies represent significant new advances in the use of isotopic measurements to understand the modern nitrogen cycle, with implications for the study of past ocean productivity, oxygenation, and nutrient status.

Cross-Shelf Exchange

Abstract: Cross-shelf exchange dominates the pathways and rates by which nutrients, biota, and materials on the continental shelf are delivered and removed. This follows because cross-shelf gradients of most properties are usually far greater than those in the alongshore direction. The resulting transports are limited by Earth's rotation, which inhibits flow from crossing isobaths. Thus, cross-shelf flows are generally weak compared with alongshore flows, and this leads to interesting observational issues. Cross-shelf flows are enabled by turbulent mixing processes, nonlinear processes (such as momentum advection), and time dependence. Thus, there is a wide range of possible effects that can allow these critical transports, and different natural settings are often governed by different combinations of processes. This review discusses examples of representative transport mechanisms and explores possible observational and theoretical paths to future progress.

Ocean Data Assimilation in Support of Climate Applications: Status and Perspectives

Abstract: Ocean data assimilation brings together observations with known dynamics encapsulated in a circulation model to describe the time-varying ocean circulation. Its applications are manifold, ranging from marine and ecosystem forecasting to climate prediction and studies of the carbon cycle. Here, we address only climate applications, which range from improving our understanding of ocean circulation to estimating initial or boundary conditions and model parameters for ocean and climate forecasts. Because of differences in underlying methodologies, data assimilation products must be used judiciously and selected according to the specific purpose, as not all related inferences would be equally reliable. Further advances are expected from improved models and methods for estimating and representing error information in data assimilation systems. Ultimately, data assimilation into coupled climate system components is needed to support ocean and climate services. However, maintaining the infrastructure and expertis...

Effects of Southern Hemisphere Wind Changes on the Meridional Overturning Circulation in Ocean Models.

Abstract: Observations show that the Southern Hemisphere zonal wind stress maximum has increased significantly over the past 30 years. Eddy-resolving ocean models show that the resulting increase in the Southern Ocean mean flow meridional overturning circulation (MOC) is partially compensated by an increase in the eddy MOC. This effect can be reproduced in the non-eddy-resolving ocean component of a climate model, providing the eddy parameterization coefficient is variable and not a constant. If the coefficient is a constant, then the Southern Ocean mean MOC change is balanced by an unrealistically large change in the Atlantic Ocean MOC. Southern Ocean eddy compensation means that Southern Hemisphere winds cannot be the dominant mechanism driving midlatitude North Atlantic MOC variability.

Ecological Insights from Pelagic Habitats Acquired Using Active Acoustic Techniques

Abstract: Marine pelagic ecosystems present fascinating opportunities for ecological investigation but pose important methodological challenges for sampling. Active acoustic techniques involve producing sound and receiving signals from organisms and other water column sources, offering the benefit of high spatial and temporal resolution and, via integration into different platforms, the ability to make measurements spanning a range of spatial and temporal scales. As a consequence, a variety of questions concerning the ecology of pelagic systems lend themselves to active acoustics, ranging from organism-level investigations and physiological responses to the environment to ecosystem-level studies and climate. As technologies and data analysis methods have matured, the use of acoustics in ecological studies has grown rapidly. We explore the continued role of active acoustics in addressing questions concerning life in the ocean, highlight creative applications to key ecological themes ranging from physiology and behavior to biogeography and climate, and discuss emerging avenues where acoustics can help determine how pelagic ecosystems function.

New Approaches to Marine Conservation Through the Scaling Up of Ecological Data

Abstract: In an era of rapid global change, conservation managers urgently need improved tools to track and counter declining ecosystem conditions. This need is particularly acute in the marine realm, where threats are out of sight, inadequately mapped, cumulative, and often poorly understood, thereby generating impacts that are inefficiently managed. Recent advances in macroecology, statistical analysis, and the compilation of global data will play a central role in improving conservation outcomes, provided that global, regional, and local data streams can be integrated to produce locally relevant and interpretable outputs. Progress will be assisted by (a) expanded rollout of systematic surveys that quantify species patterns, including some carried out with help from citizen scientists; (b) coordinated experimental research networks that utilize large-scale manipulations to identify mechanisms underlying these patterns;

Global Ocean Integrals and Means, with Trend Implications

Abstract: Understanding the ocean requires determining and explaining global integrals and equivalent average values of temperature (heat), salinity (freshwater and salt content), sea level, energy, and other properties. Attempts to determine means, integrals, and climatologies have been hindered by thinly and poorly distributed historical observations in a system in which both signals and background noise are spatially very inhomogeneous, leading to potentially large temporal bias errors that must be corrected at the 1% level or better. With the exception of the upper ocean in the current altimetric-Argo era, no clear documentation exists on the best methods for estimating means and their changes for quantities such as heat and freshwater at the levels required for anthropogenic signals. Underestimates of trends are as likely as overestimates; for example, recent inferences that multidecadal oceanic heat uptake has been greatly underestimated are plausible. For new or augmented observing systems, calculating the accuracies and precisions of global, multidecadal sampling densities for the full water column is necessary to avoid the irrecoverable loss of scientifically essential information.

The Thermodynamics of Marine Biogeochemical Cycles: Lotka Revisited.

Abstract: Nearly 100 years ago, Alfred Lotka published two short but insightful papers describing how ecosystems may organize. Principally, Lotka argued that ecosystems will grow in size and that their cycles will spin faster via predation and nutrient recycling so as to capture all available energy, and that evolution and natural selection are the mechanisms by which this occurs and progresses. Lotka's ideas have often been associated with the maximum power principle, but they are more consistent with recent developments in nonequilibrium thermodynamics, which assert that complex systems will organize toward maximum entropy production (MEP). In this review, we explore Lotka's hypothesis within the context of the MEP principle, as well as how this principle can be used to improve marine biogeochemistry models. We need to develop the equivalent of a climate model, as opposed to a weather model, to understand marine biogeochemistry on longer timescales, and adoption of the MEP principle can help create such models.

Visualizing and Quantifying Oceanic Motion

Abstract: Here I review the use of two highly complementary acoustical technologies for measuring currents in the ocean: acoustically tracked neutrally buoyant floats and vessel-mounted acoustic Doppler current profilers (ADCPs). The beauty of floats lies in their ability to efficiently and accurately visualize fluid motion in fronts and vortices and the dispersion caused by mesoscale eddy processes. Floats complement classical hydrography by articulating mechanisms and pathways by which waters spread out from their source region. Vessel-mounted ADCPs can profile the water column at O(1 km) horizontal resolution to depths greater than 1,000 m. These vessel-based scans capture in detail the cross-stream structure of fronts and eddies as well as the impact of bathymetry on currents. Sustained sampling along selected routes builds up valuable databases both for statistical studies of the submesoscale velocity field and for accurate estimates of fluid transport, as well as how these vary over time.