BACKGROUND Oxygen concentrations in both the open ocean and coastal waters have been declining since at least the middle of the 20th century. This oxygen loss, or deoxygenation, is one of the most important changes occurring in an ocean increasingly modified by human activities that have raised temperatures, CO 2 levels, and nutrient inputs and have altered the abundances and distributions of marine species. Oxygen is fundamental to biological and biogeochemical processes in the ocean. Its decline can cause major changes in ocean productivity, biodiversity, and biogeochemical cycles. Analyses of direct measurements at sites around the world indicate that oxygen-minimum zones in the open ocean have expanded by several million square kilometers and that hundreds of coastal sites now have oxygen concentrations low enough to limit the distribution and abundance of animal populations and alter the cycling of important nutrients. ADVANCES In the open ocean, global warming, which is primarily caused by increased greenhouse gas emissions, is considered the primary cause of ongoing deoxygenation. Numerical models project further oxygen declines during the 21st century, even with ambitious emission reductions. Rising global temperatures decrease oxygen solubility in water, increase the rate of oxygen consumption via respiration, and are predicted to reduce the introduction of oxygen from the atmosphere and surface waters into the ocean interior by increasing stratification and weakening ocean overturning circulation. In estuaries and other coastal systems strongly influenced by their watershed, oxygen declines have been caused by increased loadings of nutrients (nitrogen and phosphorus) and organic matter, primarily from agriculture; sewage; and the combustion of fossil fuels. In many regions, further increases in nitrogen discharges to coastal waters are projected as human populations and agricultural production rise. Climate change exacerbates oxygen decline in coastal systems through similar mechanisms as those in the open ocean, as well as by increasing nutrient delivery from watersheds that will experience increased precipitation. Expansion of low-oxygen zones can increase production of N 2 O, a potent greenhouse gas; reduce eukaryote biodiversity; alter the structure of food webs; and negatively affect food security and livelihoods. Both acidification and increasing temperature are mechanistically linked with the process of deoxygenation and combine with low-oxygen conditions to affect biogeochemical, physiological, and ecological processes. However, an important paradox to consider in predicting large-scale effects of future deoxygenation is that high levels of productivity in nutrient-enriched coastal systems and upwelling areas associated with oxygen-minimum zones also support some of the world’s most prolific fisheries. OUTLOOK Major advances have been made toward understanding patterns, drivers, and consequences of ocean deoxygenation, but there is a need to improve predictions at large spatial and temporal scales important to ecosystem services provided by the ocean. Improved numerical models of oceanographic processes that control oxygen depletion and the large-scale influence of altered biogeochemical cycles are needed to better predict the magnitude and spatial patterns of deoxygenation in the open ocean, as well as feedbacks to climate. Developing and verifying the next generation of these models will require increased in situ observations and improved mechanistic understanding on a variety of scales. Models useful for managing nutrient loads can simulate oxygen loss in coastal waters with some skill, but their ability to project future oxygen loss is often hampered by insufficient data and climate model projections on drivers at appropriate temporal and spatial scales. Predicting deoxygenation-induced changes in ecosystem services and human welfare requires scaling effects that are measured on individual organisms to populations, food webs, and fisheries stocks; considering combined effects of deoxygenation and other ocean stressors; and placing an increased research emphasis on developing nations. Reducing the impacts of other stressors may provide some protection to species negatively affected by low-oxygen conditions. Ultimately, though, limiting deoxygenation and its negative effects will necessitate a substantial global decrease in greenhouse gas emissions, as well as reductions in nutrient discharges to coastal waters.

Declining oxygen in the global ocean and coastal waters.

A comparison of Eastern Boundary Upwelling Ecosystems

Marine photosynthetic plankton are responsible for approximately 50 petagrams (10(15)) of carbon per year of net primary production, an amount equivalent to that on land. This primary production supports essentially all life in the oceans and profoundly affects global biogeochemical cycles and climate. This review discusses the general distribution of primary production in the sea, the processes that regulate this distribution, and how marine primary production is sensitive to climate variability and change. Statistical modes of ocean variability and their characteristic interannual to multi-decadal timescales over the last century are described. Recent in situ and satellite time-series of primary production can be clearly linked to interannual ocean variability. Global marine primary production appears to have increased over the past several decades in association with multi-decadal variations. A paleoclimate record extends discussion to the centennial scale, providing contrasting insights into how marine primary production might vary in the future.

Marine primary production in relation to climate variability and change.

Coastal ocean upwelling ecosystems generally represent the most productive large marine ecosystems of the world's oceans, in terms of both primary production rates and tonnages of exploitable fish produced. The Peruvian upwelling system, in particular, stands out as a major factor in world fish production. The Pacific trade winds have traditionally been considered to be the primary driving force for the upwelling system off Peru, but are projected to weaken as climate change proceeds. This leads to concern that the upwelling process in the Peru system, to which its productivity is linked, may likewise weaken. However, other mechanisms involving greenhouse-associated intensification of thermal low-pressure cells over the coastal landmasses of upwelling regions suggest general intensification of wind-driven ocean upwelling in coastal upwelling regions of the world's oceans. But although certain empirical results have supported this expectation, it has not been consistently corroborated in climate model simulations, possibly because the scale of the coastal intensification may be small relative to the scales that are appropriately reflected in the standard models. Here we summarize available evidence for the intensification mechanism and present a proxy test that uses variations in water vapor, the dominant natural greenhouse gas, to offer multiple-realization empirical evidence for action of the proposed mechanism in the real world situation. While many potential consequences to the future of marine ecosystems would codepend on climate change-related changes in the thermocline and nutricline structures, an important subset, involving potential increased propensities for hypoxia, noxious gas eruptions, toxic red tide blooms, and/or jellyfish outbreaks, may depend more directly on changes in the upwelling-favorable wind itself. A prospective role of fisheries in either mitigating or reinforcing this particular class of effects is suggested.

Greenhouse gas, upwelling-favorable winds, and the future of coastal ocean upwelling ecosystems.

Early diagenesis of redox-sensitive trace metals in the Peru upwelling area – response to ENSO-related oxygen fluctuations in the water column

. Climate and ocean ecosystem variability has been well recognized during the twentieth century but it is unclear if modern ocean biogeochemistry is susceptible to the large, abrupt shifts that characterized the Late Quaternary. Time series from marine sediments off Peru show an abrupt centennial-scale biogeochemical regime shift in the early nineteenth century, of much greater magnitude and duration than present day multi-decadal variability. A rapid expansion of the subsurface nutrient-rich, oxygen-depleted waters resulted in the present-day higher biological productivity, including pelagic fish. The shift was likely driven by a northward migration of the Intertropical Convergence Zone and the South Pacific Subtropical High to their present day locations, coupled with a strengthening of Walker circulation, towards the end of the Little Ice Age. These findings reveal the potential for large reorganizations in tropical Pacific climate with immediate effects on ocean biogeochemical cycling and ecosystem structure.

/pdf/rapid-reorganization-in-ocean-biogeochemistry-off-peru-57317fx1j3.pdf

Rapid reorganization in ocean biogeochemistry off Peru towards the end of the Little Ice Age

Variations in foraminifera over the last 460 years from laminated sediments off the coast of Peru

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Rapid reorganization in ocean biogeochemistry off Peru towards the end of the Little Ice Age

Variations in foraminifera over the last 460 years from laminated sediments off the coast of Peru