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Annual Review of Marine Science - Volume 1, 2009
Volume 1, 2009
- Preface
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Wally's Quest to Understand the Ocean's CaCO3 Cycle
Vol. 1 (2009), pp. 1–18More LessAspects of the production and dissolution of CaCO3 hard parts dominate the literature regarding contemporary marine chemistry and paleoceanography. During my long career I have contributed more than 200 papers related to this subject. In this prefatory article in the first volume of the Annual Review of Marine Science, I recount what I consider to be the highlights of my attempts to understand the cycle of CaCO3 in today's ocean and in oceans of the past. These studies began in the Bahamas in the early 1960s and then quickly graduated to the world ocean. Although much of my research has involved stable and radioisotopes contained in shells and coral directed toward reconstruction of the late Quaternary operation of the earth system, in this review I concentrate on carbonate chemistry and, in particular, the compensation in the deep sea for the overproduction of CaCO3 by marine organisms.
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A Decade of Satellite Ocean Color Observations*
Vol. 1 (2009), pp. 19–42More LessAfter the successful Coastal Zone Color Scanner (CZCS, 1978–1986) demonstration that quantitative estimations of geophysical variables such as chlorophyll a and diffuse attenuation coefficient could be derived from top of the atmosphere radiances, a number of international missions with ocean color capabilities were launched beginning in the late 1990s. Most notable were those with global data acquisition capabilities, i.e., the Ocean Color and Temperature Sensor (OCTS, Japan, 1996–1997), the Sea-viewing Wide Field-of-view Sensor (SeaWiFS, United States, 1997-present), two Moderate Resolution Imaging Spectroradiometers (MODIS, United States, Terra/2000-present and Aqua/2002-present), the Global Imager (GLI, Japan, 2002–2003), and the Medium Resolution Imaging Spectrometer (MERIS, European Space Agency, 2002-present). These missions have provided data of exceptional quality and continuity, allowing for scientific inquiries into a wide variety of marine research topics not possible with the CZCS. This review focuses on the scientific advances made over the past decade using these data sets.
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Chemistry of Marine Ligands and Siderophores
Vol. 1 (2009), pp. 43–63More LessMarine microorganisms are presented with unique challenges to obtain essential metal ions required to survive and thrive in the ocean. The production of organic ligands to complex transition metal ions is one strategy to both facilitate uptake of specific metals, such as iron, and to mitigate the potential toxic effects of other metal ions, such as copper. A number of important trace metal ions are complexed by organic ligands in seawater, including iron, cobalt, nickel, copper, zinc, and cadmium, thus defining the speciation of these metal ions in the ocean. In the case of iron, siderophores have been identified and structurally characterized. Siderophores are low molecular weight iron-binding ligands produced by marine bacteria. Although progress has been made toward the identity of in situ iron-binding ligands, few compounds have been identified that coordinate the other trace metals. Deciphering the chemical structures and production stimuli of naturally produced organic ligands and the organisms they come from is fundamental to understanding metal speciation and bioavailability. The current evidence for marine ligands, with an emphasis on siderophores, and discussion of the importance and implications of metal-binding ligands in controlling metal speciation and cycling within the world's oceans are presented.
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Particle Aggregation
Vol. 1 (2009), pp. 65–90More LessA basic problem in marine biogeochemistry is understanding material and elemental distributions and fluxes in the oceans, and a key part of this problem is understanding the processes that affect particulate material in the ocean. Aggregation of particulate material is a primary process because it alters the transport properties of particulate material and provides a mechanism for transferring material from the dissolved into the particulate pools. Aggregation theory not only provides a framework for understanding these processes, but it also provides a means for making predictions and has been successfully used to predict maximum particle concentrations in the oceans and the fate of diatom blooms (including those from iron fertilization), the size spectra of particles in the oceans, and the size distributions of trace metals. Here we review the basic theory involved, summarize recent developments, and explore unresolved issues.
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Marine Chemical Technology and Sensors for Marine Waters: Potentials and Limits
Vol. 1 (2009), pp. 91–115More LessA significant need exists for in situ sensors that can measure chemical species involved in the major processes of primary production (photosynthesis and chemosynthesis) and respiration. Some key chemical species are O2, nutrients (N and P), micronutrients (metals), pCO2, dissolved inorganic carbon (DIC), pH, and sulfide. Sensors need to have excellent detection limits, precision, selectivity, response time, a large dynamic concentration range, low power consumption, robustness, and less variation of instrument response with temperature and pressure, as well as be free from fouling problems (biological, physical, and chemical). Here we review the principles of operation of most sensors used in marine waters. We also show that some sensors can be used in several different oceanic environments to detect the target chemical species, whereas others are useful in only one environment because of various limitations. Several sensors can be used truly in situ, whereas many others involve water brought into a flow cell via tubing to the analyzer in the environment or aboard ship. Multi-element sensors that measure many chemical species in the same water mass should be targeted for further development.
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Centuries of Human-Driven Change in Salt Marsh Ecosystems
Vol. 1 (2009), pp. 117–141More LessSalt marshes are among the most abundant, fertile, and accessible coastal habitats on earth, and they provide more ecosystem services to coastal populations than any other environment. Since the Middle Ages, humans have manipulated salt marshes at a grand scale, altering species composition, distribution, and ecosystem function. Here, we review historic and contemporary human activities in marsh ecosystems—exploitation of plant products; conversion to farmland, salt works, and urban land; introduction of non-native species; alteration of coastal hydrology; and metal and nutrient pollution. Unexpectedly, diverse types of impacts can have a similar consequence, turning salt marsh food webs upside down, dramatically increasing top down control. Of the various impacts, invasive species, runaway consumer effects, and sea level rise represent the greatest threats to salt marsh ecosystems. We conclude that the best way to protect salt marshes and the services they provide is through the integrated approach of ecosystem-based management.
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Macro-Ecology of Gulf of Mexico Cold Seeps
Vol. 1 (2009), pp. 143–168More LessShortly after the discovery of chemosynthetic ecosystems at deep-sea hydrothermal vents, similar ecosystems were found at cold seeps in the Gulf of Mexico. Over the past two decades, these sites have become model systems for understanding the physiology of the symbiont-containing megafauna and the ecology of seep communities worldwide. Symbiont-containing bivalves and siboglinid polychaetes dominate the communities, including five bathymodiolin mussel species and six vestimentiferan (siboglinid polychaete) species in the Gulf of Mexico. The mussels include the first described examples of methanotrophic symbiosis and dual methanotrophic/thiotrophic symbiosis. Studies with the vestimentiferans have demonstrated their potential for extreme longevity and their ability to use posterior structures for subsurface exchange of dissolved metabolites. Ecological investigations have demonstrated that the vestimentiferans function as ecosystem engineers and identified a community succession sequence from a specialized high-biomass endemic community to a low-biomass community of background fauna over the life of a hydrocarbon seep site.
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Ocean Acidification: The Other CO2 Problem
Vol. 1 (2009), pp. 169–192More LessRising atmospheric carbon dioxide (CO2), primarily from human fossil fuel combustion, reduces ocean pH and causes wholesale shifts in seawater carbonate chemistry. The process of ocean acidification is well documented in field data, and the rate will accelerate over this century unless future CO2 emissions are curbed dramatically. Acidification alters seawater chemical speciation and biogeochemical cycles of many elements and compounds. One well-known effect is the lowering of calcium carbonate saturation states, which impacts shell-forming marine organisms from plankton to benthic molluscs, echinoderms, and corals. Many calcifying species exhibit reduced calcification and growth rates in laboratory experiments under high-CO2 conditions. Ocean acidification also causes an increase in carbon fixation rates in some photosynthetic organisms (both calcifying and noncalcifying). The potential for marine organisms to adapt to increasing CO2 and broader implications for ocean ecosystems are not well known; both are high priorities for future research. Although ocean pH has varied in the geological past, paleo-events may be only imperfect analogs to current conditions.
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Marine Chemical Ecology: Chemical Signals and Cues Structure Marine Populations, Communities, and Ecosystems
Vol. 1 (2009), pp. 193–212More LessChemical cues constitute much of the language of life in the sea. Our understanding of biotic interactions and their effects on marine ecosystems will advance more rapidly if this language is studied and understood. Here, I review how chemical cues regulate critical aspects of the behavior of marine organisms from bacteria to phytoplankton to benthic invertebrates and water column fishes. These chemically mediated interactions strongly affect population structure, community organization, and ecosystem function. Chemical cues determine foraging strategies, feeding choices, commensal associations, selection of mates and habitats, competitive interactions, and transfer of energy and nutrients within and among ecosystems. In numerous cases, the indirect effects of chemical signals on behavior have as much or more effect on community structure and function as the direct effects of consumers and pathogens. Chemical cues are critical for understanding marine systems, but their omnipresence and impact are inadequately recognized.
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Advances in Quantifying Air-Sea Gas Exchange and Environmental Forcing*
Vol. 1 (2009), pp. 213–244More LessThe past decade has seen a substantial amount of research on air-sea gas exchange and its environmental controls. These studies have significantly advanced the understanding of processes that control gas transfer, led to higher quality field measurements, and improved estimates of the flux of climate-relevant gases between the ocean and atmosphere. This review discusses the fundamental principles of air-sea gas transfer and recent developments in gas transfer theory, parameterizations, and measurement techniques in the context of the exchange of carbon dioxide. However, much of this discussion is applicable to any sparingly soluble, non-reactive gas. We show how the use of global variables of environmental forcing that have recently become available and gas exchange relationships that incorporate the main forcing factors will lead to improved estimates of global and regional air-sea gas fluxes based on better fundamental physical, chemical, and biological foundations.
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Atmospheric Iron Deposition: Global Distribution, Variability, and Human Perturbations*
Natalie M. Mahowald, Sebastian Engelstaedter, Chao Luo, Andrea Sealy, Paulo Artaxo, Claudia Benitez-Nelson, Sophie Bonnet, Ying Chen, Patrick Y. Chuang, David D. Cohen, Francois Dulac, Barak Herut, Anne M. Johansen, Nilgun Kubilay, Remi Losno, Willy Maenhaut, Adina Paytan, Joseph M. Prospero, Lindsey M. Shank, and Ronald L. SiefertVol. 1 (2009), pp. 245–278More LessAtmospheric inputs of iron to the open ocean are hypothesized to modulate ocean biogeochemistry. This review presents an integration of available observations of atmospheric iron and iron deposition, and also covers bioavailable iron distributions. Methods for estimating temporal variability in ocean deposition over the recent past are reviewed. Desert dust iron is estimated to represent 95% of the global atmospheric iron cycle, and combustion sources of iron are responsible for the remaining 5%. Humans may be significantly perturbing desert dust (up to 50%). The sources of bioavailable iron are less well understood than those of iron, partly because we do not know what speciation of the iron is bioavailable. Bioavailable iron can derive from atmospheric processing of relatively insoluble desert dust iron or from direct emissions of soluble iron from combustion sources. These results imply that humans could be substantially impacting iron and bioavailable iron deposition to ocean regions, but there are large uncertainties in our understanding.
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Contributions of Long-Term Research and Time-Series Observations to Marine Ecology and Biogeochemistry
Vol. 1 (2009), pp. 279–302More LessTime-series observations form a critical element of oceanography. New interdisciplinary efforts launched in the past two decades complement the few earlier, longer-running time series to build a better, though still poorly resolved, picture of lower-frequency ocean variability, the climate processes that drive variability, and the implications for food web dynamics, carbon storage, and climate feedbacks. Time series also enlarge our understanding of ecological processes and are integral for improving models of physical-biogeochemical-ecological ocean dynamics. The major time-series observatories go well beyond simple monitoring of core ocean properties, although that important activity forms the critical center of all time-series efforts. Modern ocean time series have major process and experimental components, entrain ancillary programs, and have integrated modeling programs for deriving a better understanding of the observations and the changing, three-dimensional ocean in which the observatories are embedded.
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Clathrate Hydrates in Nature
Vol. 1 (2009), pp. 303–327More LessScientific knowledge of natural clathrate hydrates has grown enormously over the past decade, with spectacular new findings of large exposures of complex hydrates on the sea floor, the development of new tools for examining the solid phase in situ, significant progress in modeling natural hydrate systems, and the discovery of exotic hydrates associated with sea floor venting of liquid CO2. Major unresolved questions remain about the role of hydrates in response to climate change today, and correlations between the hydrate reservoir of Earth and the stable isotopic evidence of massive hydrate dissociation in the geologic past. The examination of hydrates as a possible energy resource is proceeding apace for the subpermafrost accumulations in the Arctic, but serious questions remain about the viability of marine hydrates as an economic resource. New and energetic explorations by nations such as India and China are quickly uncovering large hydrate findings on their continental shelves.
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Hypoxia, Nitrogen, and Fisheries: Integrating Effects Across Local and Global Landscapes
Vol. 1 (2009), pp. 329–349More LessAnthropogenic nutrient enrichment and physical characteristics result in low dissolved oxygen concentrations (hypoxia) in estuaries and semienclosed seas throughout the world. Published research indicates that within and near oxygen-depleted waters, finfish and mobile macroinvertebrates experience negative effects that range from mortality to altered trophic interactions. Chronic exposure to hypoxia and fluctuating oxygen concentrations impair reproduction, immune responses, and growth. We present an analysis of hypoxia, nitrogen loadings, and fisheries landings in 30 estuaries and semienclosed seas worldwide. Our results suggest that hypoxia does not typically reduce systemwide fisheries landings below what would be predicted from nitrogen loadings, except where raw sewage is released or particularly sensitive species lose critical habitat. A number of compensatory mechanisms limit the translation of local-scale effects of hypoxia to the scale of the whole system. Hypoxia is, however, a serious environmental challenge that should be considered in fisheries management strategies and be a direct target of environmental restoration.
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The Oceanic Vertical Pump Induced by Mesoscale and Submesoscale Turbulence
Vol. 1 (2009), pp. 351–375More LessThe motivation to study the vertical exchanges of tracers associated with mesoscale eddies is that the mean concentration of most oceanic tracers changes rapidly with depth. Because mesoscale processes may transport these tracers vertically, biogeochemists hypothesized that these vertical exchanges may strongly affect global tracer budgets. This hypothesis has motivated a large number of biogeochemical studies that we review here by focusing on the significant advances that have been achieved and the remaining issues and uncertainties. The main question that emerges concerns the importance of the submesoscales (10 km in the horizontal) in these vertical exchanges. Independently, in the past decade, fluid dynamicists examined the three-dimensional properties of submesoscales generated by a mesoscale (100 km in the horizontal) turbulent eddy field. We review their results and discuss how the vertical exchanges associated with these submesoscales may answer the issues raised by biogeochemical studies and inspire future directions.
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An Inconvenient Sea Truth: Spread, Steepness, and Skewness of Surface Slopes
Vol. 1 (2009), pp. 377–415More LessBréon and Henriot (BH) have collected eight million globally distributed satellite images of sunglitter, which yield a few simple, robust rules about the statistics of surface slopes: 1) constant angular spread, 2) linear steepness, and 3) sigmoid (near stepwise) skewness (all with respect to wind speed). Yet the information is sparse because it says nothing about time and space scales. The BH rules are an inconvenient sea truth, too fundamental to be ignored, too incomplete to be understood. With regard to BH rule 1 (BH:1), I suggest that the constant spread is associated with a wake-like geometry of the short gravities. Steepness linearity (BH:2) remains an enigma. Skewness (BH:3) appears to be correlated with a rather sudden onset of breaking for winds above 4 m s−1. I do not think that skewness comes from parasitic capillaries. These are tentative conclusions; I look forward to intensive sea-going experiments over the next few years demolishing the proposed interpretations.
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Loss of Sea Ice in the Arctic*
Vol. 1 (2009), pp. 417–441More LessThe Arctic sea ice cover is in decline. The areal extent of the ice cover has been decreasing for the past few decades at an accelerating rate. Evidence also points to a decrease in sea ice thickness and a reduction in the amount of thicker perennial sea ice. A general global warming trend has made the ice cover more vulnerable to natural fluctuations in atmospheric and oceanic forcing. The observed reduction in Arctic sea ice is a consequence of both thermodynamic and dynamic processes, including such factors as preconditioning of the ice cover, overall warming trends, changes in cloud coverage, shifts in atmospheric circulation patterns, increased export of older ice out of the Arctic, advection of ocean heat from the Pacific and North Atlantic, enhanced solar heating of the ocean, and the ice-albedo feedback. The diminishing Arctic sea ice is creating social, political, economic, and ecological challenges.
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Larval Dispersal and Marine Population Connectivity
Vol. 1 (2009), pp. 443–466More LessConnectivity, or the exchange of individuals among marine populations, is a central topic in marine ecology. For most benthic marine species with complex life cycles, this exchange occurs primarily during the pelagic larval stage. The small size of larvae coupled with the vast and complex fluid environment they occupy hamper our ability to quantify dispersal and connectivity. Evidence from direct and indirect approaches using geochemical and genetic techniques suggests that populations range from fully open to fully closed. Understanding the biophysical processes that contribute to observed dispersal patterns requires integrated interdisciplinary approaches that incorporate high-resolution biophysical modeling and empirical data. Further, differential postsettlement survival of larvae may add complexity to measurements of connectivity. The degree to which populations self recruit or receive subsidy from other populations has consequences for a number of fundamental ecological processes that affect population regulation and persistence. Finally, a full understanding of population connectivity has important applications for management and conservation.
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