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Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration of electrolytes (salts in solution which in this case is represented by body fluid) to keep the body fluids from becoming too diluted or concentrated. Osmotic pressure is a measure of the tendency of water to move into one solution from another by osmosis. [1] The higher the osmotic pressure of a solution, the more water tends to move into it. Pressure must be exerted on the hypertonic side of a selectively permeable membrane to prevent diffusion of water by osmosis from the side containing pure water.
Although there may be hourly and daily variations in osmotic balance, an animal is generally in an osmotic steady state over the long term. Organisms in aquatic and terrestrial environments must maintain the right concentration of solutes and amount of water in their body fluids; this involves excretion (getting rid of metabolic nitrogen wastes and other substances such as hormones that would be toxic if allowed to accumulate in the blood) through organs such as the skin and the kidneys.
Two major types of osmoregulation are osmoconformers and osmoregulators. Osmoconformers match their body osmolarity to their environment actively or passively. Most marine invertebrates are osmoconformers, although their ionic composition may be different from that of seawater. In a strictly osmoregulating animal, the amounts of internal salt and water are held relatively constant in the face of environmental changes. It requires that intake and outflow of water and salts be equal over an extended period of time.
Organisms that maintain an internal osmolarity different from the medium in which they are immersed have been termed osmoregulators. They tightly regulate their body osmolarity, maintaining constant internal conditions. They are more common in the animal kingdom. Osmoregulators actively control salt concentrations despite the salt concentrations in the environment. An example is freshwater fish. The gills actively uptake salt from the environment by the use of mitochondria-rich cells. Water will diffuse into the fish, so it excretes a very hypotonic (dilute) urine to expel all the excess water. A marine fish has an internal osmotic concentration lower than that of the surrounding seawater, so it tends to lose water and gain salt. It actively excretes salt out from the gills. Most fish are stenohaline, which means they are restricted to either salt or fresh water and cannot survive in water with a different salt concentration than they are adapted to. However, some fish show an ability to effectively osmoregulate across a broad range of salinities; fish with this ability are known as euryhaline species, e.g., flounder. Flounder have been observed to inhabit two disparate environments—marine and fresh water—and it is inherent to adapt to both by bringing in behavioral and physiological modifications.
Some marine fish, like sharks, have adopted a different, efficient mechanism to conserve water, i.e., osmoregulation. They retain urea in their blood in relatively higher concentration. Urea damages living tissues so, to cope with this problem, some fish retain trimethylamine oxide, which helps to counteract urea's destabilizing effects on cells. Sharks, having slightly higher solute concentration (i.e., above 1000 mOsm which is sea solute concentration), do not drink water like fresh water fish.
While there are no specific osmoregulatory organs in higher plants, the stomata are important in regulating water loss through evapotranspiration, and on the cellular level the vacuole is crucial in regulating the concentration of solutes in the cytoplasm. Strong winds, low humidity and high temperatures all increase evapotranspiration from leaves. Abscisic acid is an important hormone in helping plants to conserve water—it causes stomata to close and stimulates root growth so that more water can be absorbed.
Plants share with animals the problems of obtaining water but, unlike in animals, the loss of water in plants is crucial to create a driving force to move nutrients from the soil to tissues. Certain plants have evolved methods of water conservation.
Xerophytes are plants that can survive in dry habitats, such as deserts, and are able to withstand prolonged periods of water shortage. Succulent plants such as the cacti store water in the vacuoles of large parenchyma tissues. Other plants have leaf modifications to reduce water loss, such as needle-shaped leaves, sunken stomata, and thick, waxy cuticles as in the pine. The sand-dune marram grass has rolled leaves with stomata on the inner surface.
Hydrophytes are plants that grow in aquatic habitats; they may be floating, submerged, or emergent, and may grow in seasonal (rather than permanent) wetlands. In these plants the water absorption may occur through the whole surface of the plant, e.g., the water lily, or solely through the roots, as in sedges. These plants do not face major osmoregulatory challenges from water scarcity, but aside from species adapted for seasonal wetlands, have few defenses against desiccation.
Halophytes are plants living in soils with high salt concentrations, such as salt marshes or alkaline soils in desert basins. They have to absorb water from such a soil which has higher salt concentration and therefore lower water potential(higher osmotic pressure). Halophytes cope with this situation by activating salts in their roots. As a consequence, the cells of the roots develop lower water potential which brings in water by osmosis. The excess salt can be stored in cells or excreted out from salt glands on leaves. The salt thus secreted by some species help them to trap water vapours from the air, which is absorbed in liquid by leaf cells. Therefore, this is another way of obtaining additional water from air, e.g., glasswort and cord-grass.
Mesophytes are plants living in lands of temperate zone, which grow in well-watered soil. They can easily compensate the water lost by transpiration through absorbing water from the soil. To prevent excessive transpiration they have developed a waterproof external covering called cuticle.
Kidneys play a very large role in human osmoregulation by regulating the amount of water reabsorbed from glomerular filtrate in kidney tubules, which is controlled by hormones such as antidiuretic hormone (ADH), aldosterone, and angiotensin II. For example, a decrease in water potential is detected by osmoreceptors in the hypothalamus, which stimulates ADH release from the pituitary gland to increase the permeability of the walls of the collecting ducts in the kidneys. Therefore, a large proportion of water is reabsorbed from fluid in the kidneys to prevent too much water from being excreted. [2]
Drinking is not common behavior in pinnipeds and cetaceans. Water balance is maintained in marine mammals by metabolic and dietary water, while accidental ingestion and dietary salt may help maintain homeostasis of electrolytes. The kidneys of pinnipeds and cetaceans are lobed in structure, unlike those of non-bears among terrestrial mammals, but this specific adaptation does not confer any greater concentrating ability. Unlike most other aquatic mammals, manatees frequently drink fresh water and sea otters frequently drink saltwater. [3]
In teleost (advanced ray-finned) fishes, the gills, kidney and digestive tract are involved in maintenance of body fluid balance, as the main osmoregulatory organs. Gills in particular are considered the primary organ by which ionic concentration is controlled in marine teleosts.
Unusually, the catfishes in the eeltail family Plotosidae have an extra-branchial salt-secreting dendritic organ. The dendritic organ is likely a product of convergent evolution with other vertebrate salt-secreting organs. The role of this organ was discovered by its high NKA and NKCC activity in response to increasing salinity. However, the Plotosidae dendritic organ may be of limited use under extreme salinity conditions, compared to more typical gill-based ionoregulation. [4]
Amoeba makes use of contractile vacuoles to collect excretory wastes, such as ammonia, from the intracellular fluid by diffusion and active transport. As osmotic action pushes water from the environment into the cytoplasm, the vacuole moves to the surface and pumps the contents into the environment.
Bacteria respond to osmotic stress by rapidly accumulating electrolytes or small organic solutes via transporters whose activities are stimulated by increases in osmolarity. The bacteria may also turn on genes encoding transporters of osmolytes and enzymes that synthesize osmoprotectants. [5] The EnvZ/OmpR two-component system, which regulates the expression of porins, is well characterized in the model organism E. coli . [6]
Ammonia is a toxic by-product of protein metabolism and is generally converted to less toxic substances after it is produced then excreted; mammals convert ammonia to urea, whereas birds and reptiles form uric acid to be excreted with other wastes via their cloacas.
Four processes occur:
Osmotic pressure is the minimum pressure which needs to be applied to a solution to prevent the inward flow of its pure solvent across a semipermeable membrane. It is also defined as the measure of the tendency of a solution to take in its pure solvent by osmosis. Potential osmotic pressure is the maximum osmotic pressure that could develop in a solution if it were separated from its pure solvent by a semipermeable membrane.
Halotolerance is the adaptation of living organisms to conditions of high salinity. Halotolerant species tend to live in areas such as hypersaline lakes, coastal dunes, saline deserts, salt marshes, and inland salt seas and springs. Halophiles are organisms that live in highly saline environments, and require the salinity to survive, while halotolerant organisms can grow under saline conditions, but do not require elevated concentrations of salt for growth. Halophytes are salt-tolerant higher plants. Halotolerant microorganisms are of considerable biotechnological interest.
In the kidney, the loop of Henle is the portion of a nephron that leads from the proximal convoluted tubule to the distal convoluted tubule. Named after its discoverer, the German anatomist Friedrich Gustav Jakob Henle, the loop of Henle's main function is to create a concentration gradient in the medulla of the kidney.
The syndrome of inappropriate antidiuretic hormone secretion (SIADH), also known as the syndrome of inappropriate antidiuresis (SIAD), is characterized by a physiologically inappropriate release of antidiuretic hormone (ADH) either from the posterior pituitary gland, or an abnormal non-pituitary source. Unsuppressed ADH causes a physiologically inappropriate increase in solute-free water being reabsorbed by the tubules of the kidney to the venous circulation leading to hypotonic hyponatremia.
The eeltail catfish are a family (Plotosidae) of catfish whose tails are elongated in an eel-like fashion. These catfishes are native to the Indian Ocean and western Pacific from Japan to Australia and Fiji. The family includes about 41 species in 10 genera. About half of the species are freshwater, occurring in Australia and New Guinea.
Osmoconformers are marine organisms that maintain an internal environment which is isotonic to their external environment. This means that the osmotic pressure of the organism's cells is equal to the osmotic pressure of their surrounding environment. By minimizing the osmotic gradient, this subsequently minimizes the net influx and efflux of water into and out of cells. Even though osmoconformers have an internal environment that is isosmotic to their external environment, the types of ions in the two environments differ greatly in order to allow critical biological functions to occur.
A contractile vacuole (CV) is a sub-cellular structure (organelle) involved in osmoregulation. It is found predominantly in protists and in unicellular algae. It was previously known as pulsatile or pulsating vacuole.
In chemical biology, tonicity is a measure of the effective osmotic pressure gradient; the water potential of two solutions separated by a partially-permeable cell membrane. Tonicity depends on the relative concentration of selective membrane-impermeable solutes across a cell membrane which determine the direction and extent of osmotic flux. It is commonly used when describing the swelling-versus-shrinking response of cells immersed in an external solution.
Osmotic concentration, formerly known as osmolarity, is the measure of solute concentration, defined as the number of osmoles (Osm) of solute per litre (L) of solution. The osmolarity of a solution is usually expressed as Osm/L, in the same way that the molarity of a solution is expressed as "M". Whereas molarity measures the number of moles of solute per unit volume of solution, osmolarity measures the number of osmoles of solute particles per unit volume of solution. This value allows the measurement of the osmotic pressure of a solution and the determination of how the solvent will diffuse across a semipermeable membrane (osmosis) separating two solutions of different osmotic concentration.
Plasma osmolality measures the body's electrolyte–water balance. There are several methods for arriving at this quantity through measurement or calculation.
Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall.
Euryhaline organisms are able to adapt to a wide range of salinities. An example of a euryhaline fish is the short-finned molly, Poecilia sphenops, which can live in fresh water, brackish water, or salt water.
Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds, water, CO2, phosphates, sulphates, etc. Animals treat these compounds as excretes. Plants have metabolic pathways which transforms some of them (primarily the oxygen compounds) into useful substances.
The salt gland is an organ for excreting excess salts. It is found in the cartilaginous fishes subclass elasmobranchii, seabirds, and some reptiles. Salt glands can be found in the rectum of sharks. Birds and reptiles have salt glands located in or on the skull, usually in the eyes, nose, or mouth. These glands are lobed containing many secretory tubules which radiate outward from the excretory canal at the center. Secretory tubules are lined with a single layer of epithelial cells. The diameter and length of these glands vary depending on the salt uptake of the species.
Osmosis is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential to a region of low water potential, in the direction that tends to equalize the solute concentrations on the two sides. It may also be used to describe a physical process in which any solvent moves across a selectively permeable membrane separating two solutions of different concentrations. Osmosis can be made to do work. Osmotic pressure is defined as the external pressure required to prevent net movement of solvent across the membrane. Osmotic pressure is a colligative property, meaning that the osmotic pressure depends on the molar concentration of the solute but not on its identity.
Osmoprotectants or compatible solutes are small organic molecules with neutral charge and low toxicity at high concentrations that act as osmolytes and help organisms to survive in extreme osmotic stress. Osmoprotectants can be placed in three chemical classes: betaines and associated molecules, sugars and polyols, and amino acids. These molecules accumulate in cells and balance the osmotic difference between the cell's surroundings and the cytosol. In plants, their accumulation can increase survival during stresses such as drought. In extreme cases, such as in bdelloid rotifers, tardigrades, brine shrimp, and nematodes, these molecules can allow cells to survive being completely dried out and let them enter a state of suspended animation called cryptobiosis.
Fish physiology is the scientific study of how the component parts of fish function together in the living fish. It can be contrasted with fish anatomy, which is the study of the form or morphology of fishes. In practice, fish anatomy and physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the latter dealing with how those components function together in the living fish.
The common raven, also known as the northern raven, is a large, all-black passerine bird. Found across the Northern Hemisphere, it is the most widely distributed of all corvids. Their Northern range encompasses Arctic and temperate regions of Eurasia and North America, and they reach as far South as Northern Africa and Central America. The common raven is an incredibly versatile passerine to account for this distribution, and their physiology varies with this versatility. This article discusses its physiology, including its homeostasis, respiration, circulatory system, and osmoregulation.
An ionocyte (formerly called a chloride cell) is a mitochondrion-rich cell within ionoregulatory organs of animals, such as teleost fish gill, insect Malpighian tubules, crustacean gills, antennal glands and maxillary glands, and copepod Crusalis organs. These cells contribute to the maintenance of optimal osmotic, ionic, and acid-base levels within metazoans. In aquatic invertebrates, ionocytes perform the functions of both ion uptake and ion excretion. In marine teleost fish, by expending energy to power the enzyme Na+/K+-ATPase and in coordination with other protein transporters, ionocytes pump excessive sodium and chloride ions against the concentration gradient into the ocean. Conversely, freshwater teleost ionocytes use this low intracellular environment to attain sodium and chloride ions into the organism, and also against the concentration gradient. In larval fishes with underdeveloped / developing gills, ionocytes can be found on the skin and fins.
The rock dove, Columbia livia, has a number of special adaptations for regulating water uptake and loss.