Hypoxia (environmental)

Atmospheric hypoxia occurs naturally at high altitudes. Total atmospheric pressure 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, 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 blesmols.^[5] 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.

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Altitude

Altitude

Altitude or height 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. Although the term altitude is commonly used to mean the height above sea level of a location, in geography the term elevation is often preferred for this usage.

Atmospheric pressure

Atmospheric pressure

Atmospheric pressure, also known as barometric pressure, is the pressure within the atmosphere of Earth. The standard atmosphere is a unit of pressure defined as 101,325 Pa (1,013.25 hPa), which is equivalent to 1013.25 millibars, 760 mm Hg, 29.9212 inches Hg, or 14.696 psi. The atm unit is roughly equivalent to the mean sea-level atmospheric pressure on Earth; that is, the Earth's atmospheric pressure at sea level is approximately 1 atm.

Blesmol

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-tucos in South America, or the Spalacidae from Eurasia.

Altitude training

Altitude training

Altitude training is the practice by some endurance athletes of training for several weeks at high altitude, preferably over 2,400 metres (8,000 ft) above sea level, though more commonly at intermediate altitudes due to the shortage of suitable high-altitude locations. At intermediate altitudes, the air still contains approximately 20.9% oxygen, but the barometric pressure and thus the partial pressure of oxygen is reduced.

Hypobaric chamber

Hypobaric chamber

A hypobaric chamber, or altitude chamber, is a chamber used during aerospace or high terrestrial altitude research or training to simulate the effects of high altitude on the human body, especially hypoxia and hypobaria. Some chambers also control for temperature and relative humidity.

Oxygen depletion is a phenomenon that occurs in aquatic environments as dissolved oxygen (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; 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.^[6] 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.^[7]

Seasonal kill

Hypolimnetic oxygen depletion can lead to both summer and winter "kills". During summer stratification, inputs or organic matter and sedimentation of primary producers can increase rates of respiration in the hypolimnion. If oxygen depletion becomes extreme, aerobic organisms, like fish, may die, resulting in what is known as a "summer kill".^[8] 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".^[8]

Causes of hypoxia

Decline of oxygen saturation to anoxia, measured during the night in Kiel Fjord, Germany. Depth = 5 m

Oxygen depletion can result from a number of natural factors, but is most often a concern as a consequence of pollution and eutrophication in which plant nutrients enter a river, lake, or ocean, and phytoplankton blooms are encouraged. While phytoplankton, through photosynthesis, 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, a process that further reduces DO in the water column. If oxygen depletion progresses to hypoxia, fish kills can occur and invertebrates like worms and clams on the bottom may be killed as well.

Still frame from an underwater video of the sea floor. The floor is covered with crabs, fish, and clams apparently dead or dying from oxygen depletion.

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, 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 and areas of Puget Sound, in Washington State.^[9] The World Resources Institute 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.^[10]

Jubilee photo from Mobile Bay

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.^[11]

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.^[12]

Breakdown of lignin

Tetrapyrrol ring, the active site of Ligninperoxidase enzyme

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.^[12]

Active site of tetrapyrrol ring binding oxygen

Oxyferroheme is converted to Ferri-LiP with the addition of veratric alcohol, and gives off diatomic oxygen radical.

This is the breakdown of a confieryl alcohol by a hydrogen ion to make propanol and ortho-methoxyphenol.

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.^[12] Oxygen radicals cannot be used in the environment, and are harmful in high presence in the environment.^[13]

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.^[12]

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.^[12]

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.^[14] Breakdown of propanol and phenols occur naturally in the environment because they are both water-soluble.

Environmental factors

Drivers of hypoxia and ocean acidification intensification in upwelling shelf systems. Equatorward winds drive the upwelling of low dissolved oxygen (DO), high nutrient, and high dissolved inorganic carbon (DIC) water from above the oxygen minimum zone. Cross-shelf gradients in productivity and bottom water residence times drive the strength of DO (DIC) decrease (increase) as water transits across a productive continental shelf.^[15][16]

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:

${\displaystyle G(t)=G(0)e^{-kt}}$

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.^[17] 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.^[18]

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).^[19]

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.^[20]

Solutions

Graphs of oxygen and salinity levels at Kiel Fjord in 1998

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 reefs remove nitrogen from the water column and filter out suspended solids, subsequently reducing the likelihood or extent of harmful algal blooms or anoxic conditions.^[21] Foundational work toward the idea of improving marine water quality through shellfish cultivation was conducted by Odd Lindahl et al., using mussels in Sweden.^[22] More involved than single-species shellfish cultivation, integrated multi-trophic aquaculture mimics natural marine ecosystems, relying on polyculture 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 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, to more than 30. Spawning and growth rates of fish species such as roach and perch also increased to such an extent that they are now amongst the highest in England.^[23] For smaller-scale waters such as aquaculture ponds, pump aeration is standard.^[24]

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 seas at Kiel Fjord, Germany). New approaches of long-term monitoring of oxygen regime in the ocean observe online the behavior of fish and zooplankton, which changes drastically under reduced oxygen saturations (ecoSCOPE) and already at very low levels of water pollution.

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Ocean deoxygenation

Ocean deoxygenation

Ocean deoxygenation is the reduction of the oxygen content of the global oceans and coastal zones due to human activities as a consequence of anthropogenic emissions of carbon dioxide and eutrophication-driven excess production. It is manifest in the increasing number of coastal and estuarine hypoxic areas, or dead zones, and the expansion of oxygen minimum zones (OMZs) in the world's oceans. The decrease in oxygen content of the oceans has been fairly rapid and poses a threat to all aerobic marine life, as well as to people who depend on marine life for nutrition or livelihood.

Oxygen

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 as well as with other compounds. Oxygen is Earth's most abundant element, and after hydrogen and helium, it is the third-most abundant element in the universe. At standard temperature and pressure, two atoms of the element bind to form dioxygen, a colorless and odorless diatomic gas with the formula O2. Diatomic oxygen gas currently constitutes 20.95% of the Earth's atmosphere, though this has changed considerably over long periods of time. Oxygen makes up almost half of the Earth's crust in the form of oxides.

Oxygen saturation

Oxygen saturation

Oxygen saturation 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 can be measured with a dissolved oxygen probe such as an oxygen sensor or an optode in liquid media, usually water. The standard unit of oxygen saturation is percent (%).

Anoxic waters

Anoxic waters

Anoxic waters are areas of sea water, fresh water, or groundwater that are depleted of dissolved oxygen. The US Geological Survey defines anoxic groundwater as those with dissolved oxygen concentration of less than 0.5 milligrams per litre. Anoxic waters can be contrasted with hypoxic waters, which are low in dissolved oxygen. This condition is generally found in areas that have restricted water exchange.

Endocrine disruptor

Endocrine disruptor

Endocrine disruptors, sometimes also referred to as hormonally active agents, endocrine disrupting chemicals, or endocrine disrupting compounds are chemicals that can interfere with endocrine systems. These disruptions can cause cancerous tumors, birth defects, and other developmental disorders. Found in many household and industrial products, endocrine disruptors "interfere with the synthesis, secretion, transport, binding, action, or elimination of natural hormones in the body that are responsible for development, behavior, fertility, and maintenance of homeostasis ."

Hypolimnion

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.

Respiration (physiology)

Respiration (physiology)

In physiology, respiration is the movement of oxygen from the outside environment to the cells within tissues, and the removal of carbon dioxide in the opposite direction that's to the environment.

Attenuation

Attenuation

In physics, attenuation is the gradual loss of flux intensity through a medium. For instance, dark glasses attenuate sunlight, lead attenuates X-rays, and water and air attenuate both light and sound at variable attenuation rates.

Pollution

Pollution

Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance or energy. Pollutants, the components of pollution, can be either foreign substances/energies or naturally occurring contaminants.

Eutrophication

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 phytoplankton productivity". Water bodies with very low nutrient levels are termed oligotrophic and those with moderate nutrient levels are termed mesotrophic. Advanced eutrophication may also be referred to as dystrophic and hypertrophic conditions. Eutrophication can affect freshwater or salt water systems. In freshwater ecosystems it is almost always caused by excess phosphorus. In coastal waters on the other hand, the main contributing nutrient is more likely to be nitrogen, or nitrogen and phosphorus together. This depends on the location and other factors.

Phytoplankton

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 φυτόν, meaning 'plant', and πλαγκτός, meaning 'wanderer' or 'drifter'.

Photosynthesis

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 carbohydrate molecules, such as sugars and starches, which are synthesized from carbon dioxide and water – hence the name photosynthesis, from the Greek phōs, "light", and synthesis, "putting together". Most plants, algae, and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and supplies most of the energy necessary for life on Earth.