267:
80:
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with the North
Pacific emerging as a particular hotspot of deoxygenation due to the increased amount of time since its deep waters were last ventilated (see thermohaline circulation) and related high apparent oxygen utilization (AOU). Estimates of total oxygen loss in the global ocean range from 119 to 680 T mol decade since the 1950s. These estimates represent 2% of the global ocean oxygen inventory.
444:
quantify. Some of the processes being studied are changes in oxygen gas solubility as a result of rising ocean temperatures, as well as changes in the amount of respiration and photosynthesis occurring around OMZs. Many studies have concluded that OMZs are expanding in multiple locations, but fluctuations of modern OMZs are still not fully understood. Existing
153:
major current systems. Low oxygen water may spread (by advection) from under areas of high productivity up to these physical boundaries to create a stagnant pool of water with no direct connection to the ocean surface even though (as in the
Eastern Tropical North Pacific) there may be relatively little organic matter falling from the surface.
357:
Decreased oxygen availability results in decreases in many zooplankton species’ egg production, food intake, respiration, and metabolic rates. Temperature and salinity in areas of decreased oxygen concentrations also affect oxygen availability. Higher temperatures and salinity lower oxygen solubility
152:
Physical processes then constrain the mixing and isolate this low oxygen water from outside water. Vertical mixing is constrained due to the separation from the mixed layer by depth. Horizontal mixing is constrained by bathymetry and boundaries formed by interactions with sub-tropical gyres and other
111:
The downward flux of organic matter decreases sharply with depth, with 80–90% being consumed in the top 1,000 m (3,300 ft). The deep ocean thus has higher oxygen because rates of oxygen consumption are low compared with the supply of cold, oxygen-rich deep waters from polar regions. In the
182:
At depths in the ocean where no light can reach, aerobic respiration is the dominant process. When the oxygen in a parcel of water is consumed, the oxygen cannot be replaced without the water reaching the surface ocean. When oxygen concentrations drop to below <10 nM, microbial processes that are
443:
Research has attempted to model potential changes to OMZs as a result of rising global temperatures and human impact. This is challenging due to the many factors that could contribute to changes in OMZs. The factors used for modeling change in OMZs are numerous, and in some cases hard to measure or
219:
Life stages of organisms also have different metabolic demands. In general, younger stages tend to grow in size and advance in developmental complexity quickly. As the organism reaches maturity, metabolic demands switch from growth and development to maintenance, which requires far fewer resources.
439:
Measurement of dissolved oxygen in coastal and open ocean waters for the past 50 years has revealed a marked decline in oxygen content. This decline is associated with expanding spatial extent, expanding vertical extent, and prolonged duration of oxygen-poor conditions in all regions of the global
432:
is that substantial warming and loss of oxygen throughout the majority of the upper ocean will occur. Global warming increases ocean temperatures, especially in shallow coastal areas. When the water temperature increases, its ability to hold oxygen decreases, leading to oxygen concentrations going
259:
of the organism, but does not account for behavioral or physiological changes in organisms to compensate for reduced oxygen availability. The
Metabolic Index accounts for the supply of oxygen in terms of solubility, partial pressure, and diffusivity of oxygen in water, and the organism's metabolic
459:
The global decrease in oceanic oxygen content is statistically significant and emerging beyond the envelope of natural fluctuations. This trend of oxygen loss is accelerating, with widespread and obvious losses occurring after the 1980s. The rate and total content of oxygen loss varies by region,
378:
OMZs have changed over time due to effects from numerous global chemical and biological processes. To assess these changes, scientists utilize climate models and sediment samples to understand changes to dissolved oxygen in OMZs. Many recent studies of OMZs have focused on their fluctuations over
178:
to build organic molecules (organic matter) releasing oxygen in the process. A large fraction of the organic matter from photosynthesis becomes dissolved organic matter (DOM) that is consumed by bacteria during aerobic respiration in sunlit waters. Another fraction of organic matter sinks to the
46:
is at its lowest. This zone occurs at depths of about 200 to 1,500 m (700–4,900 ft), depending on local circumstances. OMZs are found worldwide, typically along the western coast of continents, in areas where an interplay of physical and biological processes concurrently lower the
361:
In addition to affecting their vital functions, zooplankton alter their distribution in response to hypoxic or anoxic zones. Many species actively avoid low oxygen zones, while others take advantage of their predators’ low tolerance for hypoxia and use these areas as a refuge. Zooplankton that
161:
In OMZs oxygen concentration drops to levels <10 nM at the base of the oxycline and can remain anoxic for over 700 m depth. This lack of oxygen can be reinforced or increased due to physical processes changing oxygen supply such as eddy-driven advection, sluggish ventilation, increases in
112:
surface layers, oxygen is supplied by photosynthesis of phytoplankton. Depths in between, however, have higher rates of oxygen consumption and lower rates of advective supply of oxygen-rich waters. In much of the ocean, mixing processes enable the resupply of oxygen to these waters (see
251:
equation for respiration. However, organisms have ways of altering their oxygen intake and carbon dioxide release, so the strict stoichiometric equation is not necessarily accurate. The Oxygen Supply Index accounts for oxygen solubility and partial pressure, along with the
169:
At a microscopic scale the processes causing ocean deoxygenation rely on microbial aerobic respiration. Aerobic respiration is a metabolic process that microorganisms like bacteria or archaea use to obtain energy by degrading organic matter, consuming oxygen, producing
440:
oceans. Examinations of the spatial extent of OMZs in the past through paleoceanographical methods clearly shows that the spatial extent of OMZs has expanded through time, and this expansion is coupled to ocean warming and reduced ventilation of thermocline waters.
148:
due to the influx of nutrients. Since oxygen is not being produced as a byproduct of photosynthesis below the euphotic zone, these microbes use up what oxygen is in the water as they break down the falling organic matter thus creating the lower oxygen conditions.
298:
Since bioavailability is specific to each organism and temperature, calculation of these thresholds is done experimentally by measuring activity and respiration rates under different temperature and oxygen conditions, or by collecting data from separate studies.
216:) fluctuates with the temperature of the water. As the external temperature increases, ectotherm metabolisms increase as well, increasing their demand for oxygen. Different species have different basal metabolic rates and therefore different oxygen demands.
47:
oxygen concentration (biological processes) and restrict the water from mixing with surrounding waters (physical processes), creating a "pool" of water where oxygen concentrations fall from the normal range of 4–6 mg/L to below 2 mg/L.
1935:
Vanderploeg, HA; Ludsin, SA; Ruberg, SA; Höök, TO; Pothoven, SA; Brandt, SB; Lang, GA; Liebig, JR; Cavaletto, JF (2009). "Hypoxia affects spatial distributions and overlap of pelagic fish, zooplankton, and phytoplankton in Lake Erie".
341:. The oxygen minimum zones thus play an important role in regulating the productivity and ecological community structure of the global ocean. For example, giant bacterial mats floating in the oxygen minimum zone off the west coast of
220:
Smaller organisms have higher metabolisms per unit of mass, so smaller organisms will require more oxygen per unit mass, while larger organisms generally require more total oxygen. Higher activity levels also require more oxygen.
325:) continues to live aerobically (using oxygen) in OMZs. They have highly developed gills with large surface area and thin blood-to-water diffusion distance that enables effective removal of oxygen from the water (up to 90% O
474:
that causes ocean deoxygenation is circulation changes. As the ocean warms from the surface, stratification is expected to increase, which shows a tendency for slowing down ocean circulation, which then increases ocean
119:
A distribution of the open-ocean oxygen minimum zones is controlled by the large-scale ocean circulation as well as local physical as well as biological processes. For example, wind blowing parallel to the coast causes
75:
process, lowering its concentration within the water. Therefore, the concentration of oxygen in deep water is dependent on the amount of oxygen it had when it was at the surface, minus depletion by deep sea organisms.
208:. Metabolic rates can be affected by external factors such as the temperature of the water, and internal factors such as the species, life stage, size, and activity level of the organism. The body temperature of
1896:
Vanderploeg, HA; Ludsin, SA; Cavaletto, JF; Höök, TO; Pothoven, SA; Brandt, SB; Liebig, JR; Lang, GA (2009). "Hypoxic zones as habitat for zooplankton in Lake Erie: refuges from predation or exclusion zones?".
336:
Another strategy used by some classes of bacteria in the oxygen minimum zones is to use nitrate rather than oxygen, thus drawing down the concentrations of this important nutrient. This process is called
362:
exhibit daily vertical migrations to avoid predation and low oxygen conditions also excrete ammonium near the oxycline and contribute to increased anaerobic ammonium oxidation (anammox, which produces N
395:. Throughout the history of Earth's oceans, OMZs have fluctuated on long time scales, becoming larger or smaller depending on multiple variables. The factors that change OMZs are the amount of oceanic
191:. Both processes extract elemental nitrogen from nitrogen compounds and that elemental nitrogen which does not stay in solution escapes as a gas, resulting in a net loss of nitrogen from the ocean.
235:
Several indices to measure bioavailability have been suggested: Respiration Index, Oxygen Supply Index, and the
Metabolic Index. The Respiration Index describes oxygen availability based on the
436:
Open ocean areas with no oxygen have grown more than 1.7 million square miles in the last 50 years, and coastal waters have seen a tenfold increase in low-oxygen areas in the same time.
1199:
Verberk, WCEP; Bilton, DT; Calosi, P; Spicer, JI (2011). "Oxygen supply in aquatic ectotherms: partial pressure and solubility together explain biodiversity and size patterns".
2917:
Gilly, William F.; Beman, J. Michael; Litvin, Steven Y.; Robison, Bruce H. (3 January 2013). "Oceanographic and
Biological Effects of Shoaling of the Oxygen Minimum Zone".
358:
decrease the partial pressure of oxygen. This decreased partial pressure increases organisms’ respiration rates, causing the oxygen demand of the organism to increase.
2960:
Ito, T; Nenes, A; Johnson, MS; Meskhidze, N; Deutsch, C (2016). "Acceleration of oxygen decline in the tropical
Pacific over the past decades by aerosol pollutants".
1596:
Deutsch, Curtis; Sarmiento, Jorge L.; Sigman, Daniel M.; Gruber, Nicolas; Dunne, John P. (2006). "Spatial coupling of nitrogen inputs and losses in the ocean".
2201:
Roman, MR; Pierson, JJ; Kimmel, DG; Boicourt, WC; Zhang, X (2012). "Impacts of hypoxia on zooplankton spatial distributions in the northern Gulf of Mexico".
1675:"Elliott, D.T., Pierson, J.J. and Roman, M.R., 2013. Predicting the effects of coastal hypoxia on vital rates of the planktonic copepod Acartia tonsa Dana"
1412:"Elliott, D.T., Pierson, J.J. and Roman, M.R., 2013. Predicting the effects of coastal hypoxia on vital rates of the planktonic copepod Acartia tonsa Dana"
1060:"Relationship between growth and standard metabolic rate: measurement artefacts and implications for habitat use and life-history adaptation in salmonids"
136:
nutrients (from phytodetritus, dead organisms, fecal pellets, excretions, shed shells, scales, and other parts). This "rain" of organic matter (see the
317:, special adaptations are needed to either make do with lesser amounts of oxygen or to extract oxygen from the water more efficiently. For example, the
2874:
Stramma, Lothar; Schmidtko, Sunke; Levin, Lisa A.; Johnson, Gregory C. (April 2010). "Ocean oxygen minima expansions and their biological impacts".
2273:
Stramma, Lothar; Schmidtko, Sunke; Levin, Lisa A.; Johnson, Gregory C. (April 2010). "Ocean oxygen minima expansions and their biological impacts".
179:
deep ocean forming aggregates called marine snow. These sinking aggregates are consumed via degradation of organic matter and respiration at depth.
462:
Melting of gas hydrates in bottom layers of water may result in the release of more methane from sediments and subsequent consumption of oxygen by
399:
resulting in increased respiration at greater depths, changes in the oxygen supply due to poor ventilation, and amount of oxygen supplied through
543:
227:
is important in deoxygenated systems: an oxygen quantity which is dangerously low for one species might be more than enough for another species.
994:"The effects of temperature on aerobic metabolism: towards a mechanistic understanding of the responses of ectotherms to a changing environment"
945:"The effects of temperature on aerobic metabolism: towards a mechanistic understanding of the responses of ectotherms to a changing environment"
2511:
124:
that upwells nutrients from deep water. The increased nutrients support phytoplankton blooms, zooplankton grazing, and an overall productive
1502:"Adaptations to the Deep-Sea Oxygen Minimum Layer: Oxygen Binding by the Hemocyanin of the Bathypelagic Mysid, Gnathophausia ingens Dohrn"
2321:"Sensitivity of the North Pacific oxygen minimum zone to changes in ocean circulation: A simple model calibrated by chlorofluorocarbons"
1860:"Relationship between environmental conditions and zooplankton community structure during summer hypoxia in the northern Gulf of Mexico"
174:
and obtaining energy in the form of ATP. In the ocean surface photosynthetic microorganisms called phytoplankton use solar energy and CO
294:(lethal partial pressure)- the oxygen level below which an organism cannot support the minimum respiration rate necessary for survival.
2119:"Exploring effects of hypoxia on fish and fisheries in the northern Gulf of Mexico using a dynamic spatially explicit ecosystem model"
1363:"Oxygen- And capacity-limitation of thermal tolerance: A matrix for integrating climate-related stressor effects in marine ecosystems"
2373:"Northeastern Pacific oxygen minimum zone variability over the past 70 kyr: Impact of biological production and oceanic ventilation"
19:
This article is about a natural phenomenon in the ocean at particular depths. For expansion of the OMZ due to climate change, see
329:
removal from inhaled water) and an efficient circulatory system with high capacity and high blood concentration of a protein (
2486:
1729:"Ecophysiological implications of vertical migration into oxygen minimum zones for the hyperiid amphipod Phronima sedentaria"
574:
2158:
Kraus, RT; Secor, DH; Wingate, RL (2015). "Testing the thermal-niche oxygen-squeeze hypothesis for estuarine striped bass".
308:
1816:"Ammonium excretion and oxygen respiration of tropical copepods and euphausiids exposed to oxygen minimum zone conditions"
260:
rate. The metabolic index is generally viewed as a closer approximation of oxygen bioavailability than the other indices.
471:
2030:
Stramma, L; Prince, ED; Schmidtko, S; Luo, J; Hoolihan, JP; Visbeck, M; Wallace, DWR; Brandt, P; Körtzinger, A (2012).
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523:
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gas. As hypoxic regions expand vertically and horizontally, the habitable ranges for phytoplankton, zooplankton, and
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Despite the low oxygen conditions, organisms have evolved to live in and around OMZs. For those organisms, like the
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665:
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84:
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1557:"Aerobic and anaerobic metabolism in oxygen minimum layer fishes: the role of alcohol dehydrogenase"
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project considerable reductions in oxygen and other physical-chemical variables in the ocean due to
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400:
133:
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down in the water. This compounds the effects of eutrophication in coastal zones described above.
2082:
Prince, ED; Goodyear, CP (2006). "Hypoxia-based habitat compression of tropical pelagic fishes".
429:
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While oxygen minimum zones (OMZs) occur naturally, they can be exacerbated by human impacts like
128:
at the surface. The byproducts of these blooms and the subsequent grazing sink in the form of
281:(critical partial pressure)- the oxygen level below which an organism cannot support a normal
56:
2032:"Expansion of oxygen minimum zones may reduce available habitat for tropical pelagic fishes"
782:
Lam, P; Kuypers, MM (2011). "Microbial nitrogen cycling processes in oxygen minimum zones".
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may play a key role in the region's extremely rich fisheries, as bacterial mats the size of
99:. In the upper panel a minimum in oxygen content is indicated by light blue shading between
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de
Mutsert, K; Steenbeek, J; Lewis, K; Buszowski, J; Cowan Jr., JH; Christensen, V (2016).
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1984:
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1463:"Life at stable low oxygen levels: adaptations of animals to oceanic oxygen minimum layers"
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increasingly overlap, increasing their susceptibility to predation and human exploitation.
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59:. In general, colder waters hold more oxygen than warmer waters. As water moves out of the
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Keeling, RF; Körtzinger, A; Gruber, N (2010). "Ocean
Deoxygenation in a Warming World".
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Bertagnolli, AD; Stewart, FJ (2018). "Microbial niches in marine oxygen minimum zones".
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656:
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592:"Critical role for mesoscale eddy diffusion in supplying oxygen to hypoxic ocean waters"
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Surface ocean waters generally have oxygen concentrations close to equilibrium with the
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Singer, D (2004). "Metabolic adaptation to hypoxia: cost and benefit of being small".
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Ocean deoxygenation: Everyone's problem - Causes, impacts, consequences and solutions
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488:, localized areas of dramatically reduced oxygen levels, often due to human impacts.
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1322:"The real limits to marine life : a further critique of the Respiration Index"
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1762:"Critical oxygen levels and metabolic suppression in oceanic oxygen minimum zones"
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95:(approximately the centre of the Pacific Ocean). White regions indicate section
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Deutsch, C.; Ferrel, A.; Seibel, B.; Portner, H.-O.; Huey, R. B. (4 June 2015).
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1957:
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1462:
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Makarieva, AM; Gorshkov, VG; Li, BA; Chown, SL; Reich, PB; Gavrilov, VM (2008).
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and land-based pollution from agriculture and sewage. The prediction of current
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Dynamics of Marine
Ecosystems: Biological-Physical interactions in the oceans
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2512:"Global warming, pollution supersize the oceans' oxygen-depleted dead zones"
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US Department of
Commerce, National Oceanic and Atmospheric Administration.
1997:
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970:
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Azam, F; Malfatti, F (2007). "Microbial structuring of marine ecosystems".
821:
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Cartapanis, Olivier; Tachikawa, Kazuyo; Bard, Edouard (29 October 2011).
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2320:
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Sigman, DM; Hain, MP (2012). "The biological productivity of the ocean".
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60:
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Zone in which oxygen saturation in seawater in the ocean is at its lowest
2610:
2585:
1617:
913:
2806:
Stramma, L.; Johnson, G. C.; Sprintall, J.; Mohrholz, V. (2 May 2008).
2586:"Decline in global oceanic oxygen content during the past five decades"
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1525:
1379:
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961:
944:
330:
209:
96:
79:
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van Geen, A.; Smethie, W. M.; Horneman, A.; Lee, H. (7 October 2006).
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591:
166:, and increases in ocean temperature which reduces oxygen solubility.
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453:
113:
71:
feed on this organic matter; oxygen is used as part of the bacterial
2639:
2637:
2579:
2577:
2575:
1973:"Enhancement of anammox by the excretion of diel vertical migrators"
1517:
2422:"Climate change tightens a metabolic constraint on marine habitats"
1265:"Climate change tightens a metabolic constraint on marine habitats"
125:
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2572:
2116:
1057:
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Some research has aimed to understand how OMZs have changed over
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188:
100:
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2413:
2805:
2686:
1263:
Deutsch, C; Ferrel, A; Seibel, B; Pörtner, HO; Huey, R (2015).
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2533:
2531:
2529:
2527:
2525:
1895:
91:. The data plotted show a section running north–south at the
43:
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2272:
1595:
1058:
Rosenfeld, J; Van Leeuwen, T; Richards, J; Allen, D (2015).
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1934:
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time and how they may be currently changing as a result of
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2318:
2200:
1262:
842:"Microbial respiration, the engine of ocean deoxygenation"
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There are two thresholds of oxygen required by organisms:
2869:
2867:
2801:
2799:
2646:"Declining oxygen in the global ocean and coastal waters"
2029:
1198:
1153:
Brewer, PG; Peltzer, ET (2009). "Limits to Marine Life".
589:
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991:
666:
10.1175/1520-0485(1983)013<0292:tvt>2.0.co;2
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at an average depth of ca. 1,000 m (3,300 ft).
67:, it is exposed to a rain of organic matter from above.
2916:
2876:
Deep Sea Research Part I: Oceanographic Research Papers
2808:"Expanding Oxygen-Minimum Zones in the Tropical Oceans"
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Deep Sea Research Part I: Oceanographic Research Papers
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1970:
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638:
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Gnanadesikan, A.; Bianchi, D.; Pradal, M.A. (2013).
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An organism's demand for oxygen is dependent on its
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and may lead to bacterial blooms in water below the
50:
1930:
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1938:Journal of Experimental Marine Biology and Ecology
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2157:
183:normally inhibited by oxygen can take place like
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1925:
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729:
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1319:
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83:Annual mean dissolved oxygen (upper panel) and
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1971:Bianchi, D; Babbin, AR; Galbraith, ED (2014).
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639:Luyten, J; Pedlosky, J; Stommel, H (1983).
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1109:Respiratory Physiology & Neurobiology
1083:
1019:
1009:
960:
857:
811:
664:
615:
1802:
839:
777:
775:
773:
679:
516:Biological Oceanography: An Introduction
265:
78:
2476:
1500:Sanders, N.K.; Childress, J.J. (1990).
1360:
942:
514:Lalli, Carol; Parsons, Timothy (1993).
3015:
2509:
2075:
1759:
1717:
1461:Childress, J.J.; Seibel, B.A. (1998).
1313:
1143:
1106:
1042:
2366:
2364:
2314:
2312:
2268:
2266:
1646:
770:
569:. Blackwell Scientific Publications.
2931:10.1146/annurev-marine-120710-100849
2715:10.1146/annurev.marine.010908.163855
1649:"Giant Bacteria Colonise the Oceans"
804:10.1146/annurev-marine-120709-142814
509:
507:
309:Microbiology of oxygen minimum zones
1561:The Journal of Experimental Biology
1467:The Journal of Experimental Biology
565:Mann, K.H.; Lazier, J.R.N. (1991).
452:, with potential ramifications for
13:
2644:Breitburg, D; et al. (2018).
2540:"Upper ocean O2 trends: 1958–2015"
2361:
2309:
2263:
1320:Seibel, BA; Childress, JJ (2013).
472:effect of climate change on oceans
302:
14:
3049:
2510:Gokkon, Basten (9 January 2018).
504:
51:Physical and biological processes
2104:10.1111/j.1365-2419.2005.00393.x
1647:Leahy, Stephen (20 April 2010).
412:This section is an excerpt from
199:
34:), sometimes referred to as the
2919:Annual Review of Marine Science
2729:
2695:Annual Review of Marine Science
2503:
2325:Journal of Geophysical Research
2237:
2160:Environmental Biology of Fishes
2144:10.1016/j.ecolmodel.2015.10.013
1766:Journal of Experimental Biology
1640:
1589:
1548:
1493:
1454:
1367:Journal of Experimental Biology
1100:
1051:
1036:
985:
949:Journal of Experimental Biology
784:Annual Review of Marine Science
1727:Elder, LE; Seibel, BA (2015).
1655:. Tierramérica. Archived from
673:
632:
583:
558:
532:
352:
1:
497:
333:) that readily binds oxygen.
2544:Geophysical Research Letters
2477:Manahan, Stanley E. (2005).
1864:Journal of Plankton Research
1733:Journal of Plankton Research
1692:10.1371/journal.pone.0063987
1429:10.1371/journal.pone.0063987
702:10.1126/science.248.4953.316
641:"The ventilated thermocline"
596:Geophysical Research Letters
270:Respiration- Pcrit and Pleth
7:
1958:10.1016/j.jembe.2009.07.027
1919:10.1016/j.jembe.2009.07.015
1045:Modern Engineering Dynamics
902:Nature Reviews Microbiology
846:Frontiers in Marine Science
736:Nature Reviews Microbiology
479:
156:
85:apparent oxygen utilisation
10:
3054:
1121:10.1016/j.resp.2004.02.009
883:Nature Education Knowledge
411:
373:
306:
18:
2896:10.1016/j.dsr.2010.01.005
2295:10.1016/j.dsr.2010.01.005
2223:10.1007/s12237-012-9531-x
2180:10.1007/s10641-015-0431-3
1064:Journal of Animal Ecology
813:21.11116/0000-0001-CA25-2
748:10.1038/s41579-018-0087-z
387:In geological time scales
195:Bioavailability of oxygen
859:10.3389/fmars.2018.00533
540:"World Ocean Atlas 2009"
430:climate change scenarios
401:thermohaline circulation
231:Indices and calculations
3033:Biological oceanography
2833:10.1126/science.1153847
2671:10.1126/science.aam7240
2479:Environmental chemistry
2447:10.1126/science.aaa1605
1998:10.1073/pnas.1410790111
1841:10.5194/bg-13-2241-2016
1347:10.5194/bg-10-2815-2013
1290:10.1126/science.aaa1605
1167:10.1126/science.1170756
1085:10.1111/1365-2656.12260
1011:10.1073/pnas.0802148105
407:Since industrialization
349:have been found there.
87:(lower panel) from the
38:, is the zone in which
2763:10.1073/pnas.122154899
2246:"What is a dead zone?"
2084:Fisheries Oceanography
1479:10.1242/jeb.201.8.1223
393:geological time scales
271:
108:
3038:Physical oceanography
3023:Chemical oceanography
2250:oceanservice.noaa.gov
2039:Nature Climate Change
1877:10.1093/plankt/fbs029
1746:10.1093/plankt/fbv066
307:Further information:
269:
82:
3006:. IUCN, Switzerland.
2565:10.1002/2017GL073613
2398:10.1029/2011pa002126
2346:10.1029/2005jc003192
2203:Estuaries and Coasts
2123:Ecological Modelling
2059:10.1038/nclimate1304
1361:Pörtner, HO (2010).
943:Schulte, PM (2015).
840:Robinson, C (2019).
323:Gnathophausia ingens
212:(such as fishes and
164:ocean stratification
2974:2016NatGe...9..443I
2888:2010DSRI...57..587S
2824:2008Sci...320..655S
2754:2002PNAS...99.7848K
2707:2010ARMS....2..199K
2662:2018Sci...359M7240B
2611:10.1038/nature21399
2602:2017Natur.542..335S
2556:2017GeoRL..44.4214I
2438:2015Sci...348.1132D
2432:(6239): 1132–1135.
2389:2011PalOc..26.4208C
2337:2006JGRC..11110004V
2287:2010DSRI...57..587S
2215:2012EstCo..35.1261R
2172:2015EnvBF..98.2083K
2135:2016AGUOSAH43A..07D
2096:2006FisOc..15..451P
2051:2012NatCC...2...33S
1989:2014PNAS..11115653B
1983:(44): 15653–15658.
1950:2009JEMBE.381S..92V
1911:2009JEMBE.381S.108V
1832:2016BGeo...13.2241K
1760:Seibel, BA (2011).
1653:Inter Press Service
1618:10.1038/nature05392
1610:2007Natur.445..163D
1506:Biological Bulletin
1473:(Pt 8): 1223–1232.
1338:2013BGeo...10.2815S
1281:2015Sci...348.1132D
1275:(6239): 1132–1135.
1213:2011Ecol...92.1565V
1076:2015JAnEc..84....4R
1043:Balmer, RT (2011).
1004:(44): 16994–16999.
914:10.1038/nrmicro1747
796:2011ARMS....3..317L
694:1990Sci...248..316P
657:1983JPO....13..292L
608:2013GeoRL..40.5194G
492:Ocean deoxygenation
486:Dead zone (ecology)
464:aerobic respiration
446:Earth system models
42:in seawater in the
28:oxygen minimum zone
21:ocean deoxygenation
2656:(6371): eaam7240.
1779:10.1242/jeb.049171
1574:10.1242/jeb.060236
1380:10.1242/jeb.037523
962:10.1242/jeb.118851
397:primary production
272:
109:
57:Earth's atmosphere
2962:Nature Geoscience
2818:(5876): 655–658.
2748:(12): 7848–7853.
2596:(7641): 335–339.
2488:978-1-4987-7693-6
2166:(10): 2083–2092.
1567:(11): 1905–1914.
1221:10.1890/10-2369.1
1161:(5925): 347–348.
1047:. Academic Press.
955:(12): 1856–1866.
688:(4953): 316–322.
617:10.1002/grl.50998
602:(19): 5194–5198.
576:978-1-4051-1118-8
239:available in the
89:World Ocean Atlas
40:oxygen saturation
3045:
3007:
3000:
2994:
2993:
2982:10.1038/ngeo2717
2957:
2951:
2950:
2914:
2908:
2907:
2871:
2862:
2861:
2835:
2803:
2794:
2793:
2783:
2765:
2733:
2727:
2726:
2690:
2684:
2683:
2673:
2641:
2632:
2631:
2613:
2581:
2570:
2569:
2567:
2550:(9): 4214–4223.
2535:
2520:
2519:
2507:
2501:
2500:
2474:
2468:
2467:
2449:
2417:
2411:
2410:
2400:
2377:Paleoceanography
2368:
2359:
2358:
2348:
2316:
2307:
2306:
2270:
2261:
2260:
2258:
2256:
2241:
2235:
2234:
2209:(5): 1261–1269.
2198:
2192:
2191:
2155:
2149:
2148:
2146:
2114:
2108:
2107:
2079:
2073:
2072:
2070:
2036:
2027:
2021:
2020:
2010:
2000:
1968:
1962:
1961:
1932:
1923:
1922:
1893:
1882:
1881:
1879:
1855:
1846:
1845:
1843:
1826:(8): 2241–2255.
1811:
1800:
1799:
1781:
1757:
1751:
1750:
1748:
1724:
1715:
1714:
1704:
1694:
1670:
1661:
1660:
1659:on 24 June 2010.
1644:
1638:
1637:
1593:
1587:
1586:
1576:
1552:
1546:
1545:
1497:
1491:
1490:
1458:
1452:
1451:
1441:
1431:
1407:
1401:
1400:
1382:
1358:
1352:
1351:
1349:
1317:
1311:
1310:
1292:
1260:
1251:
1250:
1232:
1207:(8): 1565–1572.
1196:
1187:
1186:
1150:
1141:
1140:
1104:
1098:
1097:
1087:
1055:
1049:
1048:
1040:
1034:
1033:
1023:
1013:
989:
983:
982:
964:
940:
934:
933:
897:
891:
890:
878:
872:
871:
861:
837:
826:
825:
815:
779:
768:
767:
731:
722:
721:
677:
671:
670:
668:
636:
630:
629:
619:
587:
581:
580:
562:
556:
555:
553:
551:
536:
530:
529:
511:
69:Aerobic bacteria
3053:
3052:
3048:
3047:
3046:
3044:
3043:
3042:
3028:Aquatic ecology
3013:
3012:
3011:
3010:
3001:
2997:
2958:
2954:
2915:
2911:
2872:
2865:
2804:
2797:
2734:
2730:
2691:
2687:
2642:
2635:
2582:
2573:
2536:
2523:
2508:
2504:
2489:
2475:
2471:
2418:
2414:
2369:
2362:
2331:(C10): C10004.
2317:
2310:
2271:
2264:
2254:
2252:
2242:
2238:
2199:
2195:
2156:
2152:
2115:
2111:
2080:
2076:
2034:
2028:
2024:
1969:
1965:
1933:
1926:
1894:
1885:
1856:
1849:
1812:
1803:
1758:
1754:
1725:
1718:
1671:
1664:
1645:
1641:
1604:(7124): 163–7.
1594:
1590:
1553:
1549:
1518:10.2307/1541830
1498:
1494:
1459:
1455:
1408:
1404:
1359:
1355:
1318:
1314:
1261:
1254:
1197:
1190:
1151:
1144:
1105:
1101:
1056:
1052:
1041:
1037:
990:
986:
941:
937:
908:(10): 782–791.
898:
894:
879:
875:
838:
829:
780:
771:
742:(12): 723–729.
732:
725:
678:
674:
645:J Phys Oceanogr
637:
633:
588:
584:
577:
563:
559:
549:
547:
538:
537:
533:
526:
512:
505:
500:
482:
477:
476:
417:
409:
389:
376:
365:
355:
339:denitrification
328:
319:giant red mysid
311:
305:
303:Life in the OMZ
292:
279:
257:
233:
225:bioavailability
202:
197:
185:denitrification
177:
173:
159:
138:biological pump
122:Ekman transport
53:
24:
17:
12:
11:
5:
3051:
3041:
3040:
3035:
3030:
3025:
3009:
3008:
2995:
2968:(6): 443–447.
2952:
2925:(1): 393–420.
2909:
2882:(4): 587–595.
2863:
2795:
2728:
2701:(1): 199–229.
2685:
2633:
2571:
2521:
2502:
2487:
2469:
2412:
2360:
2308:
2281:(4): 587–595.
2262:
2236:
2193:
2150:
2109:
2090:(6): 451–464.
2074:
2022:
1963:
1924:
1883:
1870:(7): 602–613.
1847:
1820:Biogeosciences
1801:
1772:(2): 326–336.
1752:
1739:(5): 897–911.
1716:
1662:
1639:
1588:
1547:
1512:(3): 286–294.
1492:
1453:
1402:
1373:(6): 881–893.
1353:
1326:Biogeosciences
1312:
1252:
1188:
1142:
1115:(3): 215–228.
1099:
1050:
1035:
984:
935:
892:
873:
827:
769:
723:
672:
651:(2): 292–309.
631:
582:
575:
557:
531:
524:
502:
501:
499:
496:
495:
494:
489:
481:
478:
475:deoxygenation.
468:carbon dioxide
466:of methane to
450:climate change
426:climate models
422:climate change
418:
410:
408:
405:
388:
385:
381:climate change
375:
372:
363:
354:
351:
326:
304:
301:
296:
295:
290:
286:
277:
255:
249:stoichiometric
232:
229:
206:metabolic rate
201:
198:
196:
193:
175:
171:
158:
155:
142:microbial loop
93:180th meridian
52:
49:
15:
9:
6:
4:
3:
2:
3050:
3039:
3036:
3034:
3031:
3029:
3026:
3024:
3021:
3020:
3018:
3005:
2999:
2991:
2987:
2983:
2979:
2975:
2971:
2967:
2963:
2956:
2948:
2944:
2940:
2936:
2932:
2928:
2924:
2920:
2913:
2905:
2901:
2897:
2893:
2889:
2885:
2881:
2877:
2870:
2868:
2859:
2855:
2851:
2847:
2843:
2839:
2834:
2829:
2825:
2821:
2817:
2813:
2809:
2802:
2800:
2791:
2787:
2782:
2777:
2773:
2769:
2764:
2759:
2755:
2751:
2747:
2743:
2739:
2732:
2724:
2720:
2716:
2712:
2708:
2704:
2700:
2696:
2689:
2681:
2677:
2672:
2667:
2663:
2659:
2655:
2651:
2647:
2640:
2638:
2629:
2625:
2621:
2617:
2612:
2607:
2603:
2599:
2595:
2591:
2587:
2580:
2578:
2576:
2566:
2561:
2557:
2553:
2549:
2545:
2541:
2534:
2532:
2530:
2528:
2526:
2517:
2516:Mongabay News
2513:
2506:
2498:
2494:
2490:
2484:
2481:. CRC Press.
2480:
2473:
2465:
2461:
2457:
2453:
2448:
2443:
2439:
2435:
2431:
2427:
2423:
2416:
2408:
2404:
2399:
2394:
2390:
2386:
2383:(4): PA4208.
2382:
2378:
2374:
2367:
2365:
2356:
2352:
2347:
2342:
2338:
2334:
2330:
2326:
2322:
2315:
2313:
2304:
2300:
2296:
2292:
2288:
2284:
2280:
2276:
2269:
2267:
2251:
2247:
2240:
2232:
2228:
2224:
2220:
2216:
2212:
2208:
2204:
2197:
2189:
2185:
2181:
2177:
2173:
2169:
2165:
2161:
2154:
2145:
2140:
2136:
2132:
2128:
2124:
2120:
2113:
2105:
2101:
2097:
2093:
2089:
2085:
2078:
2069:
2064:
2060:
2056:
2052:
2048:
2044:
2040:
2033:
2026:
2018:
2014:
2009:
2004:
1999:
1994:
1990:
1986:
1982:
1978:
1974:
1967:
1959:
1955:
1951:
1947:
1943:
1939:
1931:
1929:
1920:
1916:
1912:
1908:
1905:: S108–S120.
1904:
1900:
1892:
1890:
1888:
1878:
1873:
1869:
1865:
1861:
1854:
1852:
1842:
1837:
1833:
1829:
1825:
1821:
1817:
1810:
1808:
1806:
1797:
1793:
1789:
1785:
1780:
1775:
1771:
1767:
1763:
1756:
1747:
1742:
1738:
1734:
1730:
1723:
1721:
1712:
1708:
1703:
1698:
1693:
1688:
1685:(5): e63987.
1684:
1680:
1676:
1669:
1667:
1658:
1654:
1650:
1643:
1635:
1631:
1627:
1623:
1619:
1615:
1611:
1607:
1603:
1599:
1592:
1584:
1580:
1575:
1570:
1566:
1562:
1558:
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1539:
1535:
1531:
1527:
1523:
1519:
1515:
1511:
1507:
1503:
1496:
1488:
1484:
1480:
1476:
1472:
1468:
1464:
1457:
1449:
1445:
1440:
1435:
1430:
1425:
1422:(5): e63987.
1421:
1417:
1413:
1406:
1398:
1394:
1390:
1386:
1381:
1376:
1372:
1368:
1364:
1357:
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1339:
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1327:
1323:
1316:
1308:
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1300:
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1218:
1214:
1210:
1206:
1202:
1195:
1193:
1184:
1180:
1176:
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1168:
1164:
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1126:
1122:
1118:
1114:
1110:
1103:
1095:
1091:
1086:
1081:
1077:
1073:
1069:
1065:
1061:
1054:
1046:
1039:
1031:
1027:
1022:
1017:
1012:
1007:
1003:
999:
995:
988:
980:
976:
972:
968:
963:
958:
954:
950:
946:
939:
931:
927:
923:
919:
915:
911:
907:
903:
896:
888:
884:
877:
869:
865:
860:
855:
851:
847:
843:
836:
834:
832:
823:
819:
814:
809:
805:
801:
797:
793:
789:
785:
778:
776:
774:
765:
761:
757:
753:
749:
745:
741:
737:
730:
728:
719:
715:
711:
707:
703:
699:
695:
691:
687:
683:
676:
667:
662:
658:
654:
650:
646:
642:
635:
627:
623:
618:
613:
609:
605:
601:
597:
593:
586:
578:
572:
568:
561:
545:
541:
535:
527:
525:0-7506-2742-5
521:
517:
510:
508:
503:
493:
490:
487:
484:
483:
473:
469:
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461:
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451:
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427:
423:
415:
404:
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398:
394:
384:
382:
371:
369:
359:
350:
348:
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