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Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the
mouth In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...
. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can be extracted),
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...
(such as
lipase Lipase ( ) is a family of enzymes that catalyzes the hydrolysis of fats. Some lipases display broad substrate scope including esters of cholesterol, phospholipids, and of lipid-soluble vitamins and sphingomyelinases; however, these are usually tr ...
and amylase), antimicrobial agents (such as secretory
IgA Iga may refer to: Arts and entertainment * Ambush at Iga Pass, a 1958 Japanese film * Iga no Kagemaru, Japanese manga series * Iga, a set of characters from the Japanese novel '' The Kouga Ninja Scrolls'' Biology * ''Iga'' (beetle), a gen ...
, and
lysozymes Lysozyme (EC 3.2.1.17, muramidase, ''N''-acetylmuramide glycanhydrolase; systematic name peptidoglycan ''N''-acetylmuramoylhydrolase) is an antimicrobial enzyme produced by animals that forms part of the innate immune system. It is a glycosid ...
). The enzymes found in saliva are essential in beginning the process of digestion of dietary starches and fats. These enzymes also play a role in breaking down food particles entrapped within dental crevices, thus protecting teeth from bacterial decay. Saliva also performs a lubricating function, wetting
food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is inge ...
and permitting the initiation of
swallowing Swallowing, sometimes called deglutition in scientific contexts, is the process in the human or animal body that allows for a substance to pass from the mouth, to the pharynx, and into the esophagus, while shutting the epiglottis. Swallowing i ...
, and protecting the oral mucosa from drying out. Various animal species have special uses for saliva that go beyond predigestion. Some swifts use their gummy saliva to build nests. '' Aerodramus'' nests form the basis of bird's nest soup. Cobras, vipers, and certain other members of the venom clade hunt with venomous saliva injected by
fang A fang is a long, pointed tooth. In mammals, a fang is a modified maxillary tooth, used for biting and tearing flesh. In snakes, it is a specialized tooth that is associated with a venom gland (see snake venom). Spiders also have external fang ...
s. Some
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s produce silk fiber from silk proteins stored in modified salivary glands (which are unrelated to the vertebrate ones).


Composition

Produced in salivary glands, human saliva comprises 99.5% water, but also contains many important substances, including
electrolyte An electrolyte is a medium containing ions that is electrically conducting through the movement of those ions, but not conducting electrons. This includes most soluble salts, acids, and bases dissolved in a polar solvent, such as water. Upon dis ...
s, mucus, antibacterial compounds and various enzymes. Medically, constituents of saliva can noninvasively provide important diagnostic information related to oral and systemic diseases. * Water: 99.5% *
Electrolyte An electrolyte is a medium containing ions that is electrically conducting through the movement of those ions, but not conducting electrons. This includes most soluble salts, acids, and bases dissolved in a polar solvent, such as water. Upon dis ...
s: ** 2–21 mmol/L sodium (lower than blood plasma) ** 10–36 mmol/L potassium (higher than plasma) ** 1.2–2.8 mmol/L calcium (similar to plasma) ** 0.08–0.5 mmol/L magnesium ** 5–40 mmol/L chloride (lower than plasma) ** 25 mmol/L bicarbonate (higher than plasma) ** 1.4–39 mmol/L phosphate **
Iodine Iodine is a chemical element with the symbol I and atomic number 53. The heaviest of the stable halogens, it exists as a semi-lustrous, non-metallic solid at standard conditions that melts to form a deep violet liquid at , and boils to a vi ...
(mmol/L concentration is usually higher than plasma, but dependent variable according to dietary iodine intake) * Mucus (mucus in saliva mainly consists of mucopolysaccharides and
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glycos ...
s) * Antibacterial compounds ( thiocyanate, hydrogen peroxide, and secretory immunoglobulin A) *
Epidermal growth factor Epidermal growth factor (EGF) is a protein that stimulates cell growth and differentiation by binding to its receptor, EGFR. Human EGF is 6-k Da and has 53 amino acid residues and three intramolecular disulfide bonds. EGF was originally descr ...
(EGF) * Various enzymes; most notably: ** α- amylase (EC3.2.1.1), or ptyalin, secreted by the acinar cells of the parotid and submandibular glands, starts the digestion of starch before the food is even swallowed; it has a pH optimum of 7.4 ** Lingual lipase, which is secreted by the acinar cells of the sublingual gland; has a pH optimum around 4.0 so it is not activated until entering the acidic environment of the stomach ** Kallikrein, an enzyme that proteolytically cleaves high-molecular-weight kininogen to produce bradykinin, which is a vasodilator; it is secreted by the acinar cells of all three major salivary glands ** Antimicrobial enzymes that kill bacteria: *** Lysozyme ***Salivary
lactoperoxidase Lactoperoxidase is a peroxidase enzyme secreted from mammary, salivary and other mucosal glands including the lungs, bronchii and nose that functions as a natural and the first line of defense against bacteria and viruses. Lactoperoxidase is a m ...
*** Lactoferrin *** Immunoglobulin A **
Proline Proline (symbol Pro or P) is an organic acid classed as a proteinogenic amino acid (used in the biosynthesis of proteins), although it does not contain the amino group but is rather a secondary amine. The secondary amine nitrogen is in the prot ...
-rich proteins (function in enamel formation, Ca2+-binding, microbe killing and lubrication) ** Minor enzymes including: salivary
acid phosphatase Acid phosphatase (EC 3.1.3.2, acid phosphomonoesterase', phosphomonoesterase, glycerophosphatase, acid monophosphatase, acid phosphohydrolase, acid phosphomonoester hydrolase, uteroferrin, acid nucleoside diphosphate phosphatase, orthophosphoric-m ...
s A+B, N-acetylmuramoyl-L-alanine amidase, NAD(P)H dehydrogenase (quinone),
superoxide dismutase Superoxide dismutase (SOD, ) is an enzyme that alternately catalyzes the dismutation (or partitioning) of the superoxide () radical into ordinary molecular oxygen (O2) and hydrogen peroxide (). Superoxide is produced as a by-product of oxygen me ...
,
glutathione transferase Glutathione ''S''-transferases (GSTs), previously known as ligandins, are a family of eukaryotic and prokaryotic phase II metabolic isozymes best known for their ability to catalyze the conjugation of the reduced form of glutathione (GSH) to x ...
, class 3 aldehyde dehydrogenase,
glucose-6-phosphate isomerase Glucose-6-phosphate isomerase (GPI), alternatively known as phosphoglucose isomerase/phosphoglucoisomerase (PGI) or phosphohexose isomerase (PHI), is an enzyme ( ) that in humans is encoded by the ''GPI'' gene on chromosome 19. This gene enc ...
, and tissue kallikrein (function unknown) * Cells: possibly as many as 8 million human and 500 million bacterial cells per mL. The presence of bacterial products (small organic acids, amines, and thiols) causes saliva to sometimes exhibit a foul odor. *
Opiorphin Opiorphin is an endogenous chemical compound first isolated from human saliva. Initial research with mice shows the compound has a painkilling effect greater than that of morphine. It works by stopping the normal breakup of enkephalins, natural ...
, a pain-killing substance found in human saliva *
Haptocorrin Haptocorrin also known as transcobalamin-1 (TC-1) or cobalophilin is a transcobalamin protein that in humans is encoded by the gene. The essential function of haptocorrin is protection of the acid-sensitive vitamin B12, vitamin B12 while it moves ...
, a protein which binds to vitamin B12 to protect it against degradation in the stomach, before it binds to intrinsic factor.


Daily salivary output

Experts debate the amount of saliva that a healthy person produces. Production is estimated at 1500ml per day and researchers generally accept that during sleep the amount drops significantly. In humans, the
submandibular gland The paired submandibular glands (historically known as submaxillary glands) are major salivary glands located beneath the floor of the mouth. They each weigh about 15 grams and contribute some 60–67% of unstimulated saliva secretion; on stimula ...
contributes around 70 to 75% of secretions, while the parotid gland secretes about 20 to 25%; small amounts are secreted from the other salivary glands.


Functions

Saliva contributes to the digestion of food and to the maintenance of oral hygiene. Without normal salivary function the frequency of dental caries, gum disease ( gingivitis and periodontitis), and other oral problems increases significantly. Saliva limits the growth of bacterial pathogens and is a major factor in sustaining systemic and oral health through the prevention of tooth decay and the removal of sugars and other food sources for microbes.


Lubricant

Saliva coats the oral mucosa mechanically protecting it from trauma during eating, swallowing, and speaking. Mouth soreness is very common in people with reduced saliva ( xerostomia) and food (especially dry food) sticks to the inside of the mouth.


Digestion

The digestive functions of saliva include moistening food and helping to create a food
bolus Bolus may refer to: Geography * Bolus, Iran, a village in Ardabil Province, Iran * Bolus, or Baulus, an Anatolian village on the site of ancient Berissa Medicine * Bolus (digestion), a ball-shaped mass moving through the digestive tract * Bolus ...
. The lubricative function of saliva allows the food bolus to be passed easily from the mouth into the esophagus. Saliva contains the enzyme amylase, also called ptyalin, which is capable of breaking down
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
into simpler sugars such as maltose and dextrin that can be further broken down in the small intestine. About 30% of starch digestion takes place in the mouth cavity. Salivary glands also secrete salivary lipase (a more potent form of lipase) to begin fat digestion. Salivary
lipase Lipase ( ) is a family of enzymes that catalyzes the hydrolysis of fats. Some lipases display broad substrate scope including esters of cholesterol, phospholipids, and of lipid-soluble vitamins and sphingomyelinases; however, these are usually tr ...
plays a large role in fat digestion in newborn infants as their pancreatic lipase still needs some time to develop.


Role in taste

Saliva is very important in the
sense of taste The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
. It is the liquid medium in which chemicals are carried to taste receptor cells (mostly associated with lingual papillae). People with little saliva often complain of dysgeusia (i.e. disordered taste, e.g. reduced ability to taste, or having a bad, metallic taste at all times). A rare condition identified to affect taste is that of '' 'Saliva Hypernatrium''', or excessive amounts of sodium in saliva that is ''not'' caused by any other condition (e.g., ''
Sjögren syndrome Sjögren syndrome or Sjögren's syndrome (SjS, SS) is a Chronic disease, long-term autoimmune disease that affects the body's moisture-producing (lacrimal and salivary) glands, and often seriously affects other organ systems, such as the lungs, k ...
''), causing everything to taste 'salty'.


Other

* Saliva maintains the pH of the mouth. Saliva is supersaturated with various ions. Certain salivary proteins prevent precipitation, which would form salts. These ions act as a buffer, keeping the acidity of the mouth within a certain range, typically pH 6.2–7.4. This prevents minerals in the dental hard tissues from dissolving. * Saliva secretes
carbonic anhydrase The carbonic anhydrases (or carbonate dehydratases) () form a family of enzymes that catalyze the interconversion between carbon dioxide and water and the dissociated ions of carbonic acid (i.e. bicarbonate and hydrogen ions). The active site ...
(gustin), which is thought to play a role in the development of taste buds. * Saliva contains EGF. EGF results in cellular proliferation, differentiation, and survival. EGF is a low-molecular-weight polypeptide first purified from the mouse submandibular gland, but since then found in many human tissues including submandibular gland, parotid gland. Salivary EGF, which seems also regulated by dietary inorganic
iodine Iodine is a chemical element with the symbol I and atomic number 53. The heaviest of the stable halogens, it exists as a semi-lustrous, non-metallic solid at standard conditions that melts to form a deep violet liquid at , and boils to a vi ...
, also plays an important physiological role in the maintenance of oro-esophageal and gastric tissue integrity. The biological effects of salivary EGF include healing of oral and gastroesophageal ulcers, inhibition of gastric acid secretion, stimulation of DNA synthesis as well as mucosal protection from intraluminal injurious factors such as gastric acid, bile acids, pepsin, and trypsin and to physical, chemical and bacterial agents.


Production

The production of saliva is stimulated both by the
sympathetic nervous system The sympathetic nervous system (SNS) is one of the three divisions of the autonomic nervous system, the others being the parasympathetic nervous system and the enteric nervous system. The enteric nervous system is sometimes considered part of th ...
and the
parasympathetic 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 t ...
. The saliva stimulated by sympathetic innervation is thicker, and saliva stimulated parasympathetically is more fluid-like. Sympathetic stimulation of saliva is to facilitate respiration, whereas parasympathetic stimulation is to facilitate digestion. Parasympathetic stimulation leads to acetylcholine (ACh) release onto the salivary acinar cells. ACh binds to muscarinic receptors, specifically M3, and causes an increased intracellular calcium ion concentration (through the IP3/DAG second messenger system). Increased calcium causes vesicles within the cells to fuse with the apical cell membrane leading to secretion. ACh also causes the salivary gland to release kallikrein, an enzyme that converts kininogen to lysyl-bradykinin. Lysyl-bradykinin acts upon blood vessels and capillaries of the salivary gland to generate vasodilation and increased
capillary permeability Vascular permeability, often in the form of capillary permeability or microvascular permeability, characterizes the capacity of a blood vessel wall to allow for the flow of small molecules (drugs, nutrients, water, ions) or even whole cells (lymph ...
, respectively. The resulting increased blood flow to the
acini An acinus (; plural, acini; adjective, acinar or acinous) refers to any cluster of cells that resembles a many-lobed "berry," such as a raspberry ('' acinus'' is Latin for "berry"). The berry-shaped termination of an exocrine gland, where the ...
allows the production of more saliva. In addition, Substance P can bind to
Tachykinin Tachykinin peptides are one of the largest families of neuropeptides, found from amphibians to mammals. They were so named due to their ability to rapidly induce contraction of gut tissue. The tachykinin family is characterized by a common C-termi ...
NK-1 receptors leading to increased intracellular calcium concentrations and subsequently increased saliva secretion. Lastly, both parasympathetic and sympathetic nervous stimulation can lead to myoepithelium contraction which causes the expulsion of secretions from the secretory acinus into the ducts and eventually to the oral cavity. Sympathetic stimulation results in the release of norepinephrine. Norepinephrine binding to α-adrenergic receptors will cause an increase in intracellular calcium levels leading to more fluid vs. protein secretion. If norepinephrine binds β-adrenergic receptors, it will result in more protein or enzyme secretion vs. fluid secretion. Stimulation by norepinephrine initially decreases blood flow to the salivary glands due to constriction of blood vessels but this effect is overtaken by vasodilation caused by various local vasodilators. Saliva production may also be pharmacologically stimulated by the so-called
sialagogue __NOTOC__ A sialogogue (also spelled sialagogue, ptysmagogue or ptyalagogue) is a drug or substance that increases the flow rate of saliva. Sialogogues can be used in the treatment of xerostomia (the subjective feeling of having a dry mouth), to st ...
s. It can also be suppressed by the so-called
antisialagogue Antisialagogues are drugs or substances that decrease the flow rate of saliva and their effect is opposite to that of sialagogues. Their origin may be both natural and synthetic. Anticholinergics generally have antisialagogue effects, and most pro ...
s.


Behaviour


Spitting

Spitting Spitting is the act of forcibly ejecting saliva or other substances from the mouth. The act is often done to get rid of unwanted or foul-tasting substances in the mouth, or to get rid of a large buildup of mucus. Spitting of small saliva drop ...
is the act of forcibly ejecting saliva or other substances from the mouth. In many parts of the world, it is considered rude and a social taboo, and has even been outlawed in many countries. In Western countries, for example, it has often been outlawed for reasons of public decency and attempting to reduce the spread of disease; however, these laws are often not strictly enforced. In Singapore, the fine for spitting may be as high as SGD$2,000 for multiple offenses, and one can even be arrested. In some other parts of the world, such as in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, expectoration is more socially acceptable (even if officially disapproved of or illegal), and spittoons are still a common appearance in some cultures. Some animals, even humans in some cases, use spitting as an automatic defensive maneuver.
Camel A camel (from: la, camelus and grc-gre, κάμηλος (''kamēlos'') from Hebrew or Phoenician: גָמָל ''gāmāl''.) is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. C ...
s are well known for doing this, though most domestic camels are trained not to. Because saliva can contain large amounts of virus copies in infected individuals (for example, in people infected with SARS-CoV-2), spitting in public places can pose a health hazard to the public.


Glue to construct bird nests

Many birds in the swift family, Apodidae, produce a viscous saliva during nesting season to glue together materials to construct a nest. Two species of swifts in the genus Aerodramus build their nests using only their saliva, the base for bird's nest soup.


Wound licking

A common belief is that saliva contained in the mouth has natural disinfectants, which leads people to believe it is beneficial to " lick their wounds". Researchers at the University of Florida at Gainesville have discovered a protein called
nerve growth factor Nerve growth factor (NGF) is a neurotrophic factor and neuropeptide primarily involved in the regulation of growth, maintenance, proliferation, and survival of certain target neurons. It is perhaps the prototypical growth factor, in that it was on ...
(NGF) in the saliva of
mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
. Wounds doused with NGF healed twice as fast as untreated and unlicked wounds; therefore, saliva can help to heal wounds in some species. NGF has not been found in human saliva; however, researchers find human saliva contains such antibacterial agents as secretory mucin,
IgA Iga may refer to: Arts and entertainment * Ambush at Iga Pass, a 1958 Japanese film * Iga no Kagemaru, Japanese manga series * Iga, a set of characters from the Japanese novel '' The Kouga Ninja Scrolls'' Biology * ''Iga'' (beetle), a gen ...
, lactoferrin, lysozyme and peroxidase. It has not been shown that human licking of wounds disinfects them, but licking is likely to help clean the wound by removing larger contaminants such as dirt and may help to directly remove infective bodies by brushing them away. Therefore, licking would be a way of wiping off pathogens, useful if clean water is not available to the animal or person.


Classical conditioning

In Pavlov's experiment, dogs were conditioned to salivate in response to a ringing bell, this stimulus is associated with a meal or hunger. Salivary secretion is also associated with nausea. Saliva is usually formed in the mouth through an act called
gleeking Spitting is the act of forcibly ejecting saliva or other substances from the mouth. The act is often done to get rid of unwanted or foul-tasting substances in the mouth, or to get rid of a large buildup of mucus. Spitting of small saliva drop ...
, which can be voluntary or involuntary.


Making alcoholic beverages

Some old cultures chewed grains to produce alcoholic beverages, such as
chicha ''Chicha'' is a fermented (alcoholic) or non-fermented beverage of Latin America, emerging from the Andes and Amazonia regions. In both the pre- and post-Spanish conquest periods, corn beer (''chicha de jora'') made from a variety of maize land ...
,
kasiri Kasiri, also known as kaschiri and cassava beer, is an alcoholic drink made from cassava by Amerindians in Suriname and Guyana. The roots of the cassava plant are grated, diluted in water, and pressed in a cylindrical basketwork press to extract t ...
or sake.


Substitutes

A number of commercially available saliva substitutes exist.


See also

* Basic reproduction number *
Spittle cures Fasting spittle – saliva produced first thing in the morning, before breakfast – has been used to treat a wide variety of diseases for many hundreds of years. Spittle cures are usually considered to be more effective if fasting spitt ...


References


Further reading

* * * * * *


External links

* {{Authority control Body fluids Excretion