Osmoregulation

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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.

Contents

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.

Regulators and conformers

Movement of water and ions in freshwater fish Bachforelle osmoregulatoin bw en2.png
Movement of water and ions in freshwater fish
Movement of water and ions in saltwater fish Osmoseragulation Carangoides bartholomaei bw en2.png
Movement of water and ions in saltwater fish

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 environmentsmarine and fresh waterand 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.

In plants

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 waterit 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.

In animals

Humans

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]

Marine mammals

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]

Teleosts

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]

In protists

Protist Paramecium aurelia with contractile vacuoles. Paramecium contractile vacuoles.jpg
Protist Paramecium aurelia with contractile vacuoles.

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.

In bacteria

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]

Vertebrate excretory systems

Waste products of the nitrogen metabolism

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.

Achieving osmoregulation in vertebrates

Four processes occur:

See also

Related Research Articles

<span class="mw-page-title-main">Osmotic pressure</span> Measure of the tendency of a solution to take in pure solvent by osmosis

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.

Countercurrent exchange is a mechanism occurring in nature and mimicked in industry and engineering, in which there is a crossover of some property, usually heat or some chemical, between two flowing bodies flowing in opposite directions to each other. The flowing bodies can be liquids, gases, or even solid powders, or any combination of those. For example, in a distillation column, the vapors bubble up through the downward flowing liquid while exchanging both heat and mass.

<span class="mw-page-title-main">Excretion</span> Elimination by an organism of metabolic waste products

Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates, this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after leaving the cell. Excretion is an essential process in all forms of life. For example, in placental mammals, urine is expelled through the urethra, which is part of the excretory system. In unicellular organisms, waste products are discharged directly through the surface of the cell.

<span class="mw-page-title-main">Aquatic animal</span> Animal that lives in water for most or all of its lifetime

An aquatic animal is any animal, whether vertebrate or invertebrate, that lives in water for all or most of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic animals may breathe air or extract oxygen from water through specialised organs called gills, or directly through the skin. Natural environments and the animals that live in them can be categorized as aquatic (water) or terrestrial (land). This designation is polyphyletic.

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.

<span class="mw-page-title-main">Eeltail catfish</span> Family of fishes

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.

<span class="mw-page-title-main">Malpighian tubule system</span> Excretory and osmoregulatory system

The Malpighian tubule system is a type of excretory and osmoregulatory system found in some insects, myriapods, arachnids and tardigrades.

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.

<span class="mw-page-title-main">Contractile vacuole</span> Organelle used in regulating osmosis

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.

<span class="mw-page-title-main">Tonicity</span> Measure of water potential across a semi-permeable cell membrane

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.

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.

<span class="mw-page-title-main">Salt gland</span> Organ for excreting excess salt

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.

<span class="mw-page-title-main">Osmosis</span> Chemical process

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 be applied so that there is no 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 survive 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.

<span class="mw-page-title-main">Fish physiology</span> Scientific study of how the component parts of fish function together in the living fish

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 later dealing with how those components function together in the living fish. For this, at first we need to know about their intestinal morphology.

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.

The rock dove, Columbia livia, has a number of special adaptations for regulating water uptake and loss.

References

  1. "Diffusion and Osmosis". hyperphysics.phy-astr.gsu.edu. Retrieved 2019-06-20.
  2. Chen, Jiatong (Steven); Sabir, Sarah; Al Khalili, Yasir (2022), "Physiology, Osmoregulation and Excretion", StatPearls, Treasure Island (FL): StatPearls Publishing, PMID   31082152 , retrieved 2022-11-30
  3. Ortiz, Rudy M. (2001-06-01). "Osmoregulation in Marine Mammals". Journal of Experimental Biology. 204 (11): 1831–1844. doi: 10.1242/jeb.204.11.1831 . ISSN   0022-0949. PMID   11441026.
  4. Malakpour Kolbadinezhad, Salman; Coimbra, João; Wilson, Jonathan M. (2018-07-03). "Osmoregulation in the Plotosidae Catfish: Role of the Salt Secreting Dendritic Organ". Frontiers in Physiology. 9: 761. doi: 10.3389/fphys.2018.00761 . ISSN   1664-042X. PMC   6037869 . PMID   30018560.
  5. Wood, Janet M. (2011). "Bacterial Osmoregulation: A Paradigm for the Study of Cellular Homeostasis". Annual Review of Microbiology. 65 (1): 215–238. doi:10.1146/annurev-micro-090110-102815. ISSN   0066-4227. PMID   21663439.
  6. Cai, SJ; Inouye, M (5 July 2002). "EnvZ-OmpR interaction and osmoregulation in Escherichia coli". The Journal of Biological Chemistry. 277 (27): 24155–61. doi: 10.1074/jbc.m110715200 . PMID   11973328.