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The salt gland is an organ for excreting excess
salt Salt is a mineral composed primarily of sodium chloride (NaCl), a chemical compound belonging to the larger class of salts; salt in the form of a natural crystalline mineral is known as rock salt or halite. Salt is present in vast quantitie ...
s. It is found in the cartilaginous fishes subclass
elasmobranchii Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of ...
(sharks, rays, and skates),
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s, and some
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s. Salt glands can be found in the
rectum The rectum is the final straight portion of the large intestine in humans and some other mammals, and the Gastrointestinal tract, gut in others. The adult human rectum is about long, and begins at the rectosigmoid junction (the end of the s ...
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 In biology, a tubule is a general term referring to small tube or similar type of structure. Specifically, tubule can refer to: * a small tube or fistular structure * a minute tube lined with glandular epithelium * any hollow cylindrical body stru ...
which radiate outward from the excretory canal at the center. Secretory tubules are lined with a single layer of
epithelial cells Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
. The diameter and length of these glands vary depending on the salt uptake of the species. Salt glands maintain salt balance and allow marine vertebrates to drink
seawater Seawater, or salt water, is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of about 3.5% (35 g/L, 35 ppt, 600 mM). This means that every kilogram (roughly one liter by volume) of seawater has appro ...
.
Active transport In cellular biology, ''active transport'' is the movement of molecules or ions across a cell membrane from a region of lower concentration to a region of higher concentration—against the concentration gradient. Active transport requires cellul ...
via
sodium–potassium pump The sodium–potassium pump (sodium–potassium adenosine triphosphatase, also known as Na⁺/K⁺-ATPase, Na⁺/K⁺ pump, or sodium–potassium ATPase) is an enzyme (an electrogenic transmembrane ATPase) found in the membrane of all animal c ...
, found on the
basolateral membrane Epithelial polarity is one example of the cell polarity that is a fundamental feature of many types of cells. Epithelial cells feature distinct 'apical', 'lateral' and 'basal' plasma membrane domains. Epithelial cells connect to one another via t ...
, moves salt from the
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the c ...
into the gland, where it is excreted as a concentrated
solution Solution may refer to: * Solution (chemistry), a mixture where one substance is dissolved in another * Solution (equation), in mathematics ** Numerical solution, in numerical analysis, approximate solutions within specified error bounds * Soluti ...
.


In birds

The avian salt gland has two main ducts which are a medial and a lateral. Salt gland activations occurs from increased osmolarity in the blood, stimulating the hypothalamic information processing, sending signal through the
parasympathetic nerve The parasympathetic nervous system (PSNS) is one of the three divisions of the autonomic nervous system, the others being the sympathetic nervous system and the enteric nervous system. The enteric nervous system is sometimes considered part of ...
activating
vasodilation Vasodilation is the widening of blood vessels. It results from relaxation of smooth muscle cells within the vessel walls, in particular in the large veins, large arteries, and smaller arterioles. The process is the opposite of vasoconstriction, ...
, the release of hormones (acetylcholine and vasoactive intestinal peptide).
Acetylcholine Acetylcholine (ACh) is an organic chemical that functions in the brain and body of many types of animals (including humans) as a neurotransmitter. Its name is derived from its chemical structure: it is an ester of acetic acid and choline. Part ...
binds to the receptor on the basolateral membrane of the gland. This in turn activates calcium release in the epithelial cells, opening potassium channels (flowing potassium out of the cells) on the basolateral membrane and chloride channels on the apical membrane to flow out of the cell. Ions are moved into the epithelial cells by a
Na-K-Cl cotransporter The Na-K-Cl cotransporter (NKCC) is a protein that aids in the secondary active transport of sodium, potassium, and chloride into cells. In humans there are two isoforms of this membrane transport protein, NKCC1 and NKCC2, encoded by two differ ...
, also in the basolateral membrane. Increases in sodium opens the sodium-potassium ATPase channels, removing the excess sodium back out across the basolateral membrane and allowing for potassium to come into the cell. An electrical gradient is formed from the chloride ions, allowing sodium to be passed through the
tight junctions Tight junctions, also known as occluding junctions or ''zonulae occludentes'' (singular, ''zonula occludens''), are multiprotein junctional complexes whose canonical function is to prevent leakage of solutes and water and seals between the epith ...
of the epithelial cells into the salt gland along with minimal amounts of water. As well, mitochondria rich cells are associated with changes in salt concentration, increasing with higher amounts and decreasing with lower exposure, assisting in the movement of salts. These glands excrete the
hypertonic 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- ...
sodium-chloride (with few other ions) by the stimulus of central and peripheral
osmoreceptors An osmoreceptor is a sensory receptor primarily found in the hypothalamus of most homeothermic organisms that detects changes in osmotic pressure. Osmoreceptors can be found in several structures, including two of the circumventricular organs – t ...
and volume receptors.


In reptiles

The need for salt excretion in reptiles (such as
marine iguana The marine iguana (''Amblyrhynchus cristatus''), also known as the sea iguana, saltwater iguana, or Galápagos marine iguana, is a species of iguana found only on the Galápagos Islands (Ecuador). Unique among modern lizards, it is a marine repti ...
s and
sea turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhead, ...
s) and birds (such as
petrel Petrels are tube-nosed seabirds in the bird order Procellariiformes. Description The common name does not indicate relationship beyond that point, as "petrels" occur in three of the four families within that group (all except the albatross f ...
s and
albatross Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Pacifi ...
es) reflects their having much less efficient
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
s than mammals. Unlike the
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...
of
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s, that of reptiles and birds is impermeable to salt, preventing its release. The evolution of a salt gland in early reptiles and birds allowed them to eat aquatic plants and animals with high salt concentrations. This evolutionary development does not account for the gland in
elasmobranchs Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gil ...
, suggesting
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. Some theories suggest
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
ian
tear duct The nasolacrimal duct (also called the tear duct) carries tears from the lacrimal sac of the eye into the nasal cavity. The duct begins in the eye socket between the maxillary and lacrimal bones, from where it passes downwards and backwards. The ...
s and
sweat gland Sweat glands, also known as sudoriferous or sudoriparous glands, , are small tubular structures of the skin that produce sweat. Sweat glands are a type of exocrine gland, which are glands that produce and secrete substances onto an epithelial sur ...
s may be evolutionarily related to salt glands. While human tears are high in
potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosphe ...
, most phylogeneticists disagree with the association.


See also

*
Halotolerance 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 ...
*
Lacrimal gland The lacrimal glands are paired exocrine glands, one for each eye, found in most terrestrial vertebrates and some marine mammals, that secrete the aqueous layer of the tear film. In humans, they are situated in the upper lateral region of each or ...
*
Osmoregulation 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 o ...
* Supraorbital gland


References


Further reading

*Evans, D. H. 1993. ''Osmotic and Ionic Regulation.'' pp. 315–336. In Evans, D. H. 1993. The Physiology of Fishes. CRC Press, Boca Raton, Florida. *Goldstein, D. L. 2002. ''Water and Salt Balance in Seabirds.'' pp. 467–480. In Schreiber, E. A. and J. Burger. (eds.) 2002. Biology of Marine Birds. CRC Press, Boca Raton, Florida. *Schmidt-Nielsen, K. 1959. Salt Glands. pp. 221–226. In Wessells, N. K. (comp.) 1974. Vertebrate Structures and Functions. W. H. Freeman and Company, San Francisco, CA. *Wǖrsig, B. G., T. A. Jefferson and D. J. Schmidly. 2000. The Marine Mammals of the Gulf of Mexico. Texas A&M Press, College Station, TX. {{Animalosmo Glands Vertebrate anatomy