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Hypoxia refers to low
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...
conditions. Normally, 20.9% of the gas in the atmosphere is oxygen. The
partial pressure In a mixture of gases, each constituent gas has a partial pressure which is the notional pressure of that constituent gas as if it alone occupied the entire volume of the original mixture at the same temperature. The total pressure of an ideal g ...
of oxygen in the atmosphere is 20.9% of the total barometric pressure. In water, oxygen levels are much lower, approximately 7 ppm or 0.0007% in good quality water, and fluctuate locally depending on the presence of photosynthetic organisms and relative distance to the surface (if there is more oxygen in the air, it will diffuse across the partial pressure gradient).


Atmospheric hypoxia

Atmospheric hypoxia occurs naturally at high
altitude Altitude or height (also sometimes known as depth) is a distance measurement, usually in the vertical or "up" direction, between a reference datum and a point or object. The exact definition and reference datum varies according to the context ...
s. Total
atmospheric pressure Atmospheric pressure, also known as barometric pressure (after the barometer), is the pressure within the atmosphere of Earth. The standard atmosphere (symbol: atm) is a unit of pressure defined as , which is equivalent to 1013.25 millibars, ...
decreases as altitude increases, causing a lower partial pressure of oxygen, which is defined as hypobaric hypoxia. Oxygen remains at 20.9% of the total gas mixture, differing from
hypoxic hypoxia Generalized hypoxia is a medical condition in which the tissues of the body are deprived of the necessary levels of oxygen due to an insufficient supply of oxygen, which may be due to the composition or pressure of the breathing gas, decreased lu ...
, where the percentage of oxygen in the air (or blood) is decreased. This is common in the sealed burrows of some subterranean animals, such as
blesmol The blesmols, also known as mole-rats, or African mole-rats, are burrowing rodents of the family Bathyergidae. They represent a distinct evolution of a subterranean life among rodents much like the pocket gophers of North America, the tuco-tu ...
s. Atmospheric hypoxia is also the basis of altitude training, which is a standard part of training for elite athletes. Several companies mimic hypoxia using normobaric artificial atmosphere.


Aquatic hypoxia

Oxygen depletion is a phenomenon that occurs in aquatic environments as
dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It ca ...
(DO; molecular oxygen dissolved in the water) becomes reduced in concentration to a point where it becomes detrimental to aquatic organisms living in the system. Dissolved oxygen is typically expressed as a percentage of the oxygen that would dissolve in the water at the prevailing temperature and salinity (both of which affect the solubility of oxygen in water; see oxygen saturation and underwater). An aquatic system lacking dissolved oxygen (0% saturation) is termed anaerobic, reducing, or
anoxic The term anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of diss ...
; a system with low concentration—in the range between 1 and 30% saturation—is called hypoxic or dysoxic. Most fish cannot live below 30% saturation since they rely on oxygen to derive energy from their nutrients. Hypoxia leads to impaired reproduction of remaining fish via endocrine disruption. A "healthy" aquatic environment should seldom experience less than 80% saturation. The exaerobic zone is found at the boundary of anoxic and hypoxic zones. Hypoxia can occur throughout the water column and also at high altitudes as well as near sediments on the bottom. It usually extends throughout 20-50% of the water column, but depends on the water depth and location of pycnoclines (rapid changes in water density with depth). It can occur in 10-80% of the water column. For example, in a 10-meter water column, it can reach up to 2 meters below the surface. In a 20-meter water column, it can extend up to 8 meters below the surface.


Seasonal kill

Hypolimnetic oxygen depletion can lead to both summer and winter "kills". During summer stratification, inputs or organic matter and
sedimentation Sedimentation is the deposition of sediments. It takes place when particles in suspension settle out of the fluid in which they are entrained and come to rest against a barrier. This is due to their motion through the fluid in response to the ...
of primary producers can increase rates of respiration in the
hypolimnion The hypolimnion or under lake is the dense, bottom layer of water in a thermally- stratified lake. The word hypolimnion is derived from the Greek "limnos" meaning "lake". It is the layer that lies below the thermocline. Typically the hypolimni ...
. If oxygen depletion becomes extreme, aerobic organisms, like fish, may die, resulting in what is known as a "summer kill".Wetzel, R. G. (2001). ''Limnology: Lake and river ecosystems''. San Diego: Academic Press. The same phenomena can occur in the winter, but for different reasons. During winter, ice and snow cover can attenuate light, and therefore reduce rates of photosynthesis. The freezing over of a lake also prevents air-water interactions that allow the exchange of oxygen. This creates a lack of oxygen while respiration continues. When the oxygen becomes badly depleted, anaerobic organisms can die, resulting in a "winter kill".


Causes of hypoxia

Oxygen depletion can result from a number of natural factors, but is most often a concern as a consequence of
pollution Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, th ...
and
eutrophication Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phyt ...
in which plant nutrients enter a river, lake, or ocean, and
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. ...
blooms are encouraged. While phytoplankton, through
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
, will raise DO saturation during daylight hours, the dense population of a bloom reduces DO saturation during the night by respiration. When phytoplankton cells die, they sink towards the bottom and are decomposed by
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
, a process that further reduces DO in the water column. If oxygen depletion progresses to hypoxia,
fish kill The term fish kill, known also as fish die-off, refers to a localized die-off of fish populations which may also be associated with more generalized mortality of aquatic life.University of Florida. Gainesville, FL (2005) ''Plant Management in Fl ...
s can occur and invertebrates like
worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and no eyes (though not always). Worms vary in size from microscopic to over in length for marine polychaete wor ...
s and
clam Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shel ...
s on the bottom may be killed as well. Hypoxia may also occur in the absence of pollutants. In estuaries, for example, because freshwater flowing from a river into the sea is less dense than salt water, stratification in the water column can result. Vertical mixing between the water bodies is therefore reduced, restricting the supply of oxygen from the surface waters to the more saline bottom waters. The oxygen concentration in the bottom layer may then become low enough for hypoxia to occur. Areas particularly prone to this include shallow waters of semi-enclosed water bodies such as the Waddenzee or the
Gulf of Mexico The Gulf of Mexico ( es, Golfo de México) is an ocean basin and a marginal sea of the Atlantic Ocean, largely surrounded by the North American continent. It is bounded on the northeast, north and northwest by the Gulf Coast of the United ...
, where land run-off is substantial. In these areas a so-called " dead zone" can be created. Low dissolved oxygen conditions are often seasonal, as is the case in
Hood Canal Hood Canal is a fjord forming the western lobe, and one of the four main basins,Puget Sound Puget Sound ( ) is a sound of the Pacific Northwest, an inlet of the Pacific Ocean, and part of the Salish Sea. It is located along the northwestern coast of the U.S. state of Washington. It is a complex estuarine system of interconnected m ...
, in Washington State. The
World Resources Institute The World Resources Institute (WRI) is a global research non-profit organization established in 1982 with funding from the MacArthur Foundation under the leadership of James Gustave Speth. WRI's activities are focused on seven areas: food, for ...
has identified 375 hypoxic coastal zones around the world, concentrated in coastal areas in Western Europe, the Eastern and Southern coasts of the US, and East Asia, particularly in Japan. Hypoxia may also be the explanation for periodic phenomena such as the Mobile Bay jubilee, where aquatic life suddenly rushes to the shallows, perhaps trying to escape oxygen-depleted water. Recent widespread shellfish kills near the coasts of Oregon and Washington are also blamed on cyclic dead zone ecology.


Phytoplankton breakdown

Scientists have determined that high concentrations of minerals dumped into bodies of water causes significant growth of phytoplankton blooms. As these blooms are broken down by bacteria and other taxa, such as '' Phanerochaete chrysosporium'', oxygen is depleted by the enzymes of these organisms.


= Breakdown of lignin

= Phytoplankton are mostly made up of lignin and cellulose, which are broken down by enzymes present in organisms such as '' P. chrysosporium'', known as white-rot''.'' The breakdown of cellulose does not deplete oxygen concentration in water, but the breakdown of lignin does. This breakdown of lignin includes an oxidative mechanism, and requires the presence of dissolved oxygen to take place by enzymes like ligninperoxidase. Other fungi such as brown-rot, soft-rot, and blue stain fungi also are necessary in lignin transformation. As this oxidation takes place, CO2 is formed in its place. Ligninperoxidase (LiP) serves as the most import enzyme because it is best at breaking down lignin in these organisms. LiP disrupts C-C bonds and C-O bonds within lignin's three-dimensional structure, causing it to break down. LiP consists of ten alpha helices, two Ca2+ structural ions, as well as a heme group called a tetrapyrrol ring. Oxygen serves an important role in the catalytic cycle of LiP to form a double bond on the Fe2+ ion in the tetrapyrrol ring. Without the presence of diatomic oxygen in the water, this breakdown cannot take place because Ferrin-LiP will not be reduced into oxyferroheme. Oxygen gas is used to reduce Ferrin-LiP into oxyferroheme-LiP. Oxyferroheme and veratric alcohol combine to create oxygen radical and Ferri-LiP, which can now be used to degrade lignin. Oxygen radicals cannot be used in the environment, and are harmful in high presence in the environment. Once Ferri-LiP is present in the ligninperoxidase, it can be used to break down lignin molecules by removing one phenylpropane group at a time through either the LRET mechanism or the mediator mechanism. The LRET mechanism (long range electron transfer mechanism) transfers an electron from the tetrapyrrol ring onto a molecule of phenylpropane in a lignin. This electron moves onto a C-C or C-O bond to break one phenylpropane molecule from the lignin, breaking it down by removing one phenylpropane at a time. In the mediator mechanism, LiP enzyme is activated by the addition of hydrogen peroxide to make LiP radical, and a mediator such as veratric alcohol is added and activated creating veratric alcohol radical. Veratric alcohol radical transfers one electron to activate the phenylpropane on lignin, and the electron dismantles a C-C or C-O bond to release one phenylpropane from the lignin. As the size of a lignin molecule increases, the more difficult it is to break these C-C or C-O bonds. Three types of phenyl propane rings include coniferyl alcohol, sinapyl alcohol, and-coumaryl alcohol. LiP has a very low MolDock score, meaning there is little energy required to form this enzyme and stabilize it to carry out reactions. LiP has a MolDock score of -156.03 kcal/mol. This is energetically favorable due to its negative free energy requirements, and therefore this reaction catalyzed by LiP is likely to take place spontaneously. Breakdown of propanol and phenols occur naturally in the environment because they are both water-soluble.


= Environmental factors

= The breakdown of phytoplankton in the environment depends on the presence of oxygen, and once oxygen is no longer in the bodies of water, ligninperoxidases cannot continue to break down the lignin. When oxygen is not present in the water, the time required for breakdown of phytoplankton changes from 10.7 days to a total of 160 days. The rate of phytoplankton breakdown can be represented using this equation: G(t) = G(0)e^ In this equation, G(t) is the amount of particulate organic carbon (POC) overall at a given time, t. G(0) is the concentration of POC before breakdown takes place. k is a rate constant in year-1, and t is time in years. For most POC of phytoplankton, the k is around 12.8 years-1, or about 28 days for nearly 96% of carbon to be broken down in these systems. Whereas for anoxic systems, POC breakdown takes 125 days, over four times longer. It takes approximately 1 mg of oxygen to break down 1 mg of POC in the environment, and therefore, hypoxia takes place quickly as oxygen is used up quickly to digest POC. About 9% of POC in phytoplankton can be broken down in a single day at 18 °C. Therefore it takes about eleven days to completely break down phytoplankton. After POC is broken down, this particulate matter can be turned into other dissolved carbon, such as carbon dioxide, bicarbonate ions, and carbonate. As much as 30% of phytoplankton can be broken down into dissolved carbon. When this particulate organic carbon interacts with 350 nm ultraviolet light, dissolved inorganic carbon is formed, removing even more oxygen from the environment in the forms of carbon dioxide, bicarbonate ions, and carbonate. Dissolved inorganic carbon is made at a rate of 2.3–6.5 mg/(m3⋅day). As phytoplankton breakdown, free phosphorus and nitrogen become available in the environment, which also fosters hypoxic conditions. As the breakdown of this phytoplankton takes place, the more phosphorus turns into phosphates, and nitrogens turn into nitrates. This depletes the oxygen even more so in the environment, further creating hypoxic zones in higher quantities. As more minerals such as phosphorus and nitrogen are displaced into these aquatic systems, the growth of phytoplankton greatly increases, and after their death, hypoxic zones are formed.


Solutions

To combat hypoxia, it is essential to reduce the amount of land-derived nutrients reaching rivers in runoff. This can be done by improving sewage treatment and by reducing the amount of fertilizers leaching into the rivers. Alternately, this can be done by restoring natural environments along a river; marshes are particularly effective in reducing the amount of phosphorus and nitrogen (nutrients) in water. Other natural habitat-based solutions include restoration of shellfish populations, such as oysters.
Oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
reefs remove
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
from the water column and filter out suspended solids, subsequently reducing the likelihood or extent of
harmful algal blooms A harmful algal bloom (HAB) (or excessive algae growth) is an algal bloom that causes negative impacts to other organisms by production of natural algae-produced toxins, mechanical damage to other organisms, or by other means. HABs are sometimes ...
or anoxic conditions. Foundational work toward the idea of improving marine water quality through shellfish cultivation was conducted by Odd Lindahl et al., using mussels in Sweden. More involved than single-species shellfish cultivation, integrated multi-trophic aquaculture mimics natural marine ecosystems, relying on
polyculture In agriculture, polyculture is the practice of growing more than one crop species in the same space, at the same time. In doing this, polyculture attempts to mimic the diversity of natural ecosystems. Polyculture is the opposite of monoculture, i ...
to improve marine water quality. Technological solutions are also possible, such as that used in the redeveloped Salford Docks area of the
Manchester Ship Canal The Manchester Ship Canal is a inland waterway in the North West of England linking Manchester to the Irish Sea. Starting at the Mersey Estuary at Eastham, near Ellesmere Port, Cheshire, it generally follows the original routes of the ri ...
in England, where years of runoff from sewers and roads had accumulated in the slow running waters. In 2001 a compressed air injection system was introduced, which raised the oxygen levels in the water by up to 300%. The resulting improvement in water quality led to an increase in the number of invertebrate species, such as freshwater
shrimp Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are ref ...
, to more than 30.
Spawning Spawn is the eggs and sperm released or deposited into water by aquatic animals. As a verb, ''to spawn'' refers to the process of releasing the eggs and sperm, and the act of both sexes is called spawning. Most aquatic animals, except for aquat ...
and growth rates of fish species such as
roach Roach may refer to: Animals * Cockroach, various insect species of the order Blattodea * Common roach (''Rutilus rutilus''), a fresh and brackish water fish of the family Cyprinidae ** ''Rutilus'' or roaches, a genus of fishes * California roa ...
and
perch Perch is a common name for fish of the genus ''Perca'', freshwater gamefish belonging to the family Percidae. The perch, of which three species occur in different geographical areas, lend their name to a large order of vertebrates: the Per ...
also increased to such an extent that they are now amongst the highest in England. For smaller-scale waters such as aquaculture ponds, pump aeration is standard. In a very short time the oxygen saturation can drop to zero when offshore blowing winds drive surface water out and anoxic depth water rises up. At the same time a decline in temperature and a rise in salinity is observed (from the longterm ecological observatory in the
sea The sea, connected as the world ocean or simply the ocean, is the body of salty water that covers approximately 71% of the Earth's surface. The word sea is also used to denote second-order sections of the sea, such as the Mediterranean Sea, ...
s at Kiel Fjord, Germany). New approaches of long-term monitoring of oxygen regime in the ocean observe online the
behavior Behavior (American English) or behaviour (British English) is the range of actions and mannerisms made by individuals, organisms, systems or artificial entities in some environment. These systems can include other systems or organisms as we ...
of
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
and
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, which changes drastically under reduced oxygen saturations ( ecoSCOPE) and already at very low levels of
water pollution Water pollution (or aquatic pollution) is the contamination of water bodies, usually as a result of human activities, so that it negatively affects its uses. Water bodies include lakes, rivers, oceans, aquifers, reservoirs and groundwater. ...
.


See also

*
Algal blooms An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term ''algae'' encompasse ...
* Anoxic event * Dead zone (ecology) *
Cyanobacterial bloom Cyanotoxins are toxins produced by cyanobacteria (also known as blue-green algae). Cyanobacteria are found almost everywhere, but particularly in lakes and in the ocean where, under high concentration of phosphorus conditions, they reproduce exp ...
*
Denitrification Denitrification is a microbially facilitated process where nitrate (NO3−) is reduced and ultimately produces molecular nitrogen (N2) through a series of intermediate gaseous nitrogen oxide products. Facultative anaerobic bacteria perform denit ...
*
Eutrophication Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phyt ...
*
Hypoxia in fish Fish are exposed to large oxygen fluctuations in their aquatic environment since the inherent properties of water can result in marked spatial and temporal differences in the concentration of oxygen (see oxygenation and underwater). Fish respond ...
* Oxygen minimum zone


References


Sources

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External links


Hypoxia in the Gulf of Mexico

Scientific Assessment of Hypoxia in U.S. Coastal Waters
Council on Environmental Quality The Council on Environmental Quality (CEQ) is a division of the Executive Office of the President that coordinates federal environmental efforts in the United States and works closely with agencies and other White House offices on the developm ...

Dead zone in front of Atlantic City

Hypoxia in Oregon Waters
{{Authority control Aquatic ecology Chemical oceanography Environmental science Water quality indicators Oxygen Endocrine disruptors Limnology