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Hypoxia (hypo: "below", oxia: "oxygenated") refers to low oxygen conditions. Hypoxia is problematic for air-breathing organisms, yet it is essential for many anaerobic organisms. Hypoxia applies to many situations, but usually refers to the atmosphere and natural waters. [3]
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. [4] 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.
An aquatic system lacking dissolved oxygen (0% saturation) is termed anaerobic, reducing, or anoxic .
In water, oxygen levels are approximately 7 ppm or 0.0007% in good quality water, but fluctuate. [5] Many organisms require hypoxic conditions. Oxygen is poisonous to anaerobic bacteria for example. [3]
Oxygen depletion is typically expressed as a percentage of the oxygen that would dissolve in the water at the prevailing temperature and salinity. 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]
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]
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.
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]
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 are mostly made up of lignin and cellulose, which are broken down by oxidative mechanism, which consume oxygen. [12]
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:
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. [15] 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. [16]
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). [17]
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. [18]
An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in fresh water or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term algae encompasses many types of aquatic photosynthetic organisms, both macroscopic multicellular organisms like seaweed and microscopic unicellular organisms like cyanobacteria. Algal bloom commonly refers to the rapid growth of microscopic unicellular algae, not macroscopic algae. An example of a macroscopic algal bloom is a kelp forest.
Eutrophication is a general term describing a process in which nutrients accumulate in a body of water, resulting in an increased growth of microorganisms that may deplete the oxygen of water. Eutrophication may occur naturally or as a result of human actions. Manmade, or cultural, eutrophication occurs when sewage, industrial wastewater, fertilizer runoff, and other nutrient sources are released into the environment. Such nutrient pollution usually causes algal blooms and bacterial growth, resulting in the depletion of dissolved oxygen in water and causing substantial environmental degradation.
The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmospheric, terrestrial, and marine ecosystems. The conversion of nitrogen can be carried out through both biological and physical processes. Important processes in the nitrogen cycle include fixation, ammonification, nitrification, and denitrification. The majority of Earth's atmosphere (78%) is atmospheric nitrogen, making it the largest source of nitrogen. However, atmospheric nitrogen has limited availability for biological use, leading to a scarcity of usable nitrogen in many types of ecosystems.
An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environments and are an example of an ecotone. Estuaries are subject both to marine influences such as tides, waves, and the influx of saline water, and to fluvial influences such as flows of freshwater and sediment. The mixing of seawater and freshwater provides high levels of nutrients both in the water column and in sediment, making estuaries among the most productive natural habitats in the world.
Limnology is the study of inland aquatic ecosystems. The study of limnology includes aspects of the biological, chemical, physical, and geological characteristics of fresh and saline, natural and man-made bodies of water. This includes the study of lakes, reservoirs, ponds, rivers, springs, streams, wetlands, and groundwater. Water systems are often categorized as either running (lotic) or standing (lentic).
Dead zones are hypoxic (low-oxygen) areas in the world's oceans and large lakes. Hypoxia occurs when dissolved oxygen (DO) concentration falls to or below 2 ml of O2/liter. When a body of water experiences hypoxic conditions, aquatic flora and fauna begin to change behavior in order to reach sections of water with higher oxygen levels. Once DO declines below 0.5 ml O2/liter in a body of water, mass mortality occurs. With such a low concentration of DO, these bodies of water fail to support the aquatic life living there. Historically, many of these sites were naturally occurring. However, in the 1970s, oceanographers began noting increased instances and expanses of dead zones. These occur near inhabited coastlines, where aquatic life is most concentrated.
The oxygen minimum zone (OMZ), sometimes referred to as the shadow zone, is the zone in which oxygen saturation in seawater in the ocean 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 oxygen concentration and restrict the water from mixing with surrounding waters, creating a "pool" of water where oxygen concentrations fall from the normal range of 4–6 mg/L to below 2 mg/L.
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 (%).
In biogeochemistry, remineralisation refers to the breakdown or transformation of organic matter into its simplest inorganic forms. These transformations form a crucial link within ecosystems as they are responsible for liberating the energy stored in organic molecules and recycling matter within the system to be reused as nutrients by other organisms.
A lake ecosystem or lacustrine ecosystem includes biotic (living) plants, animals and micro-organisms, as well as abiotic (non-living) physical and chemical interactions. Lake ecosystems are a prime example of lentic ecosystems, which include ponds, lakes and wetlands, and much of this article applies to lentic ecosystems in general. Lentic ecosystems can be compared with lotic ecosystems, which involve flowing terrestrial waters such as rivers and streams. Together, these two ecosystems are examples of freshwater ecosystems.
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.
The phosphorus cycle is the biogeochemical cycle that involves the movement of phosphorus through the lithosphere, hydrosphere, and biosphere. Unlike many other biogeochemical cycles, the atmosphere does not play a significant role in the movement of phosphorus, because phosphorus and phosphorus-based materials do not enter the gaseous phase readily, as the main source of gaseous phosphorus, phosphine, is only produced in isolated and specific conditions. Therefore, the phosphorus cycle is primarily examined studying the movement of orthophosphate (PO4)3-, the form of phosphorus that is most commonly seen in the environment, through terrestrial and aquatic ecosystems.
Monomictic lakes are holomictic lakes that mix from top to bottom during one mixing period each year. Monomictic lakes may be subdivided into cold and warm types.
The Trophic State Index (TSI) is a classification system designed to rate water bodies based on the amount of biological productivity they sustain. Although the term "trophic index" is commonly applied to lakes, any surface water body may be indexed.
Ocean deoxygenation is the reduction of the oxygen content in different parts of the ocean due to human activities. There are two areas where this occurs. Firstly, it occurs in coastal zones where eutrophication has driven some quite rapid declines in oxygen to very low levels. This type of ocean deoxygenation is also called dead zones. Secondly, ocean deoxygenation occurs also in the open ocean. In that part of the ocean, there is nowadays an ongoing reduction in oxygen levels. As a result, the naturally occurring low oxygen areas are now expanding slowly. This expansion is happening as a consequence of human caused climate change. The resulting decrease in oxygen content of the oceans poses a threat to marine life, as well as to people who depend on marine life for nutrition or livelihood. A decrease in ocean oxygen levels affects how productive the ocean is, how nutrients and carbon move around, and how marine habitats function.
Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word πλαγκτός (planktós), meaning "wandering" or "drifting", and bacterium, a Latin term coined in the 19th century by Christian Gottfried Ehrenberg. They are found in both seawater and fresh water.
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. Fish respond to hypoxia with varied behavioral, physiological, and cellular responses to maintain homeostasis and organism function in an oxygen-depleted environment. The biggest challenge fish face when exposed to low oxygen conditions is maintaining metabolic energy balance, as 95% of the oxygen consumed by fish is used for ATP production releasing the chemical energy of nutrients through the mitochondrial electron transport chain. Therefore, hypoxia survival requires a coordinated response to secure more oxygen from the depleted environment and counteract the metabolic consequences of decreased ATP production at the mitochondria.
Nutrient cycling in the Columbia River Basin involves the transport of nutrients through the system, as well as transformations from among dissolved, solid, and gaseous phases, depending on the element. The elements that constitute important nutrient cycles include macronutrients such as nitrogen, silicate, phosphorus, and micronutrients, which are found in trace amounts, such as iron. Their cycling within a system is controlled by many biological, chemical, and physical processes.
An oxygen minimum zone (OMZ) is characterized as an oxygen-deficient layer in the world's oceans. Typically found between 200 m to 1500 m deep below regions of high productivity, such as the western coasts of continents. OMZs can be seasonal following the spring-summer upwelling season. Upwelling of nutrient-rich water leads to high productivity and labile organic matter, that is respired by heterotrophs as it sinks down the water column. High respiration rates deplete the oxygen in the water column to concentrations of 2 mg/L or less forming the OMZ. OMZs are expanding, with increasing ocean deoxygenation. Under these oxygen-starved conditions, energy is diverted from higher trophic levels to microbial communities that have evolved to use other biogeochemical species instead of oxygen, these species include nitrate, nitrite, sulphate etc. Several Bacteria and Archea have adapted to live in these environments by using these alternate chemical species and thrive. The most abundant phyla in OMZs are Pseudomonadota, Bacteroidota, Actinomycetota, and Planctomycetota.
Lake metabolism represents a lake's balance between carbon fixation and biological carbon oxidation. Whole-lake metabolism includes the carbon fixation and oxidation from all organism within the lake, from bacteria to fishes, and is typically estimated by measuring changes in dissolved oxygen or carbon dioxide throughout the day.
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