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A tooth ( : teeth) is a hard, calcified structure found in the jaws (or
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 ...
s) of many vertebrates and used to break down
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 ...
. Some animals, particularly carnivores and
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s, also use teeth to help with capturing or wounding prey, tearing food, for defensive purposes, to intimidate other animals often including their own, or to carry prey or their young. The roots of teeth are covered by gums. Teeth are not made of bone, but rather of multiple tissues of varying density and hardness that originate from the embryonic
germ layer A germ layer is a primary layer of cells that forms during embryonic development. The three germ layers in vertebrates are particularly pronounced; however, all eumetazoans (animals that are sister taxa to the sponges) produce two or three pr ...
, the
ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from t ...
. The general structure of teeth is similar across the vertebrates, although there is considerable variation in their form and position. The teeth of mammals have deep roots, and this pattern is also found in some fish, and in
crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
s. In most teleost fish, however, the teeth are attached to the outer surface of the bone, while in
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s they are attached to the inner surface of the jaw by one side. In
cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue ...
, such as sharks, the teeth are attached by tough
ligament A ligament is the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the: * Peritoneal li ...
s to the hoops of
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck an ...
that form the jaw. Some animals develop only one set of teeth (monophyodonts) while others are
diphyodont A diphyodont is any animal with two ss of tooth (animal), teeth, initially the ''deciduous teeth, deciduous'' set and consecutively the ''permanent teeth, permanent'' set. Most mammals are diphyodonts—as to chew their food they need a strong, dura ...
s, i.e. they have an early set of
deciduous teeth Deciduous teeth or primary teeth, also informally known as baby teeth, milk teeth, or temporary teeth,Illustrated Dental Embryology, Histology, and Anatomy, Bath-Balogh and Fehrenbach, Elsevier, 2011, page 255 are the first set of teeth in the ...
and a later set of permanent or "adult" teeth. Still others develop many sets ( polyphyodonts). Sharks, for example, grow a new set of teeth every two weeks to replace worn teeth. Most extant mammals including humans are diphyodonts, but there are exceptions including elephants, kangaroos, and manatees, all of which are polyphyodonts. Rodent incisors grow and wear away continually through gnawing, which helps maintain relatively constant length. The industry of the
beaver Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers ar ...
is due in part to this qualification. Many rodents such as voles and guinea pigs, but not
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' ...
, as well as leporidae like
rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s, have continuously growing molars in addition to incisors. Also, tusks (in tusked mammals) grow almost throughout life. Teeth are not always attached to the jaw, as they are in mammals. In many
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 and fish, teeth are attached to the palate or to the floor of the mouth, forming additional rows inside those on the jaws proper. Some teleosts even have teeth in the pharynx. While not true teeth in the usual sense, the
dermal denticle A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as we ...
s of sharks are almost identical in structure and are likely to have the same evolutionary origin. Indeed, teeth appear to have first evolved in sharks, and are not found in the more primitive jawless fish – while lampreys do have tooth-like structures on the tongue, these are in fact, composed of keratin, not of dentine or enamel, and bear no relationship to true teeth. Though "modern" teeth-like structures with dentine and enamel have been found in late
conodont Conodonts (Greek ''kōnos'', "cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, which ...
s, they are now supposed to have evolved independently of later vertebrates' teeth.nature.com, ''Fossil scans reveal origins of teeth'', 16 October 2013
/ref> Living
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 typically have small teeth, or none at all, since they commonly feed only on soft foods. In reptiles, teeth are generally simple and conical in shape, although there is some variation between species, most notably the venom-injecting
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 of snakes. The pattern of incisors, canines, premolars and molars is found only in mammals, and to varying extents, in their evolutionary ancestors. The numbers of these types of teeth vary greatly between species;
zoologists This is a list of notable zoologists who have published names of new taxa under the International Code of Zoological Nomenclature. A * Abe – Tokiharu Abe (1911–1996) * Abeille de Perrin, Ab. – Elzéar Abeille de Perrin (1843–1910) * ...
use a standardised dental formula to describe the precise pattern in any given group.


Etymology

The word ''tooth'' comes from Proto-Germanic *''tanthu''- which in turn comes from Proto-Indo-European *''h₁dent''-, which was composed of the root *''h₁ed''- ("to eat") plus the active participle suffix -''nt'', therefore it literally meant "that which eats". Cognate with Latin ''dēns'', Greek ὀδούς ''odous'', and Sanskrit ''dát''.


Origin

Teeth are assumed to have evolved either from
ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from t ...
denticles (scales, much like those on the skin of sharks) that folded and integrated into the mouth (called the "outside–in" theory), or from
endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the gast ...
pharyngeal teeth (primarily formed in the pharynx of
jawless vertebrates Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct ( conodonts and ostracoderms) species. Among recent animals, cyclosto ...
) (the "inside–out" theory). In addition, there is another theory stating that neural crest gene regulatory network, and neural crest-derived ectomesenchyme are the key to generate teeth (with any
epithelium 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 intercellul ...
, either ectoderm or endoderm). The genes governing tooth development in mammals are
homologous Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor *Sequence homology, biological homology between DNA, RNA, or protein sequences * Homologous chrom ...
to those involved in the development of fish scales. Study of a tooth plate of a fossil of the extinct fish ''
Romundina stellina ''Romundina'' is a small, heavily armored extinct genus of acanthothoracid placoderms which lived in shallow marine environments in the early Devonian (Lochkovian). The name ''Romundina'' honors Canadian geologist and paleontologist Dr. Róm ...
'' showed that the teeth and scales were made of the same tissues, also found in mammal teeth, lending support to the theory that teeth evolved as a modification of scales.


Mammals

Teeth are among the most distinctive (and long-lasting) features of
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 ...
species. Paleontologists use teeth to identify fossil species and determine their relationships. The shape of the animal's teeth are related to its diet. For example, plant matter is hard to digest, so
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s have many molars for chewing and grinding. Carnivores, on the other hand, have canine teeth to kill prey and to tear meat. Mammals, in general, are
diphyodont A diphyodont is any animal with two ss of tooth (animal), teeth, initially the ''deciduous teeth, deciduous'' set and consecutively the ''permanent teeth, permanent'' set. Most mammals are diphyodonts—as to chew their food they need a strong, dura ...
, meaning that they develop two sets of teeth. In humans, the first set (the "baby," "milk," "primary" or " deciduous" set) normally starts to appear at about six months of age, although some babies are born with one or more visible teeth, known as
neonatal teeth Natal teeth are teeth that are present above the gumline (have already erupted) at birth, and neonatal teeth are teeth that emerge through the gingiva during the first month of life (the neonatal period). The incidence of neonatal teeth varies c ...
. Normal tooth eruption at about six months is known as teething and can be painful. Kangaroos, elephants, and manatees are unusual among mammals because they are polyphyodonts.


Aardvark

In aardvarks, teeth lack enamel and have many pulp tubules, hence the name of the order Tubulidentata.


Canines

In dogs, the teeth are less likely than humans to form dental cavities because of the very high pH of dog saliva, which prevents enamel from demineralizing. Sometimes called cuspids, these teeth are shaped like points (cusps) and are used for tearing and grasping food.


Cetaceans

Like human teeth, whale teeth have polyp-like protrusions located on the root surface of the tooth. These polyps are made of cementum in both species, but in human teeth, the protrusions are located on the outside of the root, while in whales the nodule is located on the inside of the pulp chamber. While the roots of human teeth are made of cementum on the outer surface, whales have cementum on the entire surface of the tooth with a very small layer of enamel at the tip. This small enamel layer is only seen in older whales where the cementum has been worn away to show the underlying enamel. The toothed whale is a
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
of the
cetacean Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
s characterized by having teeth. The teeth differ considerably among the species. They may be numerous, with some dolphins bearing over 100 teeth in their jaws. On the other hand, the
narwhal The narwhal, also known as a narwhale (''Monodon monoceros''), is a medium-sized toothed whale that possesses a large "tusk" from a protruding canine tooth. It lives year-round in the Arctic waters around Greenland, Canada and Russia. It is o ...
s have a giant unicorn-like tusk, which is a tooth containing millions of sensory pathways and used for sensing during feeding, navigation, and mating. It is the most neurologically complex tooth known. Beaked whales are almost toothless, with only bizarre teeth found in males. These teeth may be used for feeding but also for demonstrating aggression and showmanship.


Primates

In humans (and most other primates) there are usually 20 primary (also "baby" or "milk") teeth, and later up to 32 permanent teeth. Four of these 32 may be third molars or wisdom teeth, although these are not present in all adults, and may be removed surgically later in life. Among primary teeth, 10 of them are usually found in the maxilla (i.e. upper jaw) and the other 10 in the mandible (i.e. lower jaw). Among permanent teeth, 16 are found in the maxilla and the other 16 in the mandible. Most of the teeth have uniquely distinguishing features.


Horse

An adult horse has between 36 and 44 teeth. The enamel and dentin layers of horse teeth are intertwined. All horses have 12 premolars, 12 molars, and 12 incisors. Generally, all male equines also have four canine teeth (called tushes) between the molars and incisors. However, few female horses (less than 28%) have canines, and those that do usually have only one or two, which many times are only partially erupted. A few horses have one to four
wolf teeth Wolf teeth are small, peg-like horse teeth, which sit just in front of (or rostral to) the first cheek teeth of horses and other equids. They are vestigial first premolars, and the first cheek tooth is referred to as the second premolar even wh ...
, which are vestigial premolars, with most of those having only one or two. They are equally common in male and female horses and much more likely to be on the upper jaw. If present these can cause problems as they can interfere with the horse's bit contact. Therefore, wolf teeth are commonly removed. Horse teeth can be used to estimate the animal's age. Between birth and five years, age can be closely estimated by observing the eruption pattern on milk teeth and then permanent teeth. By age five, all permanent teeth have usually erupted. The horse is then said to have a "full" mouth. After the age of five, age can only be conjectured by studying the wear patterns on the incisors, shape, the angle at which the incisors meet, and other factors. The wear of teeth may also be affected by diet, natural abnormalities, and
cribbing Crib may refer to: *Bach (New Zealand), a type of modest beach house, called a crib in the southern half of the South Island e.g. Otago and Southland *Box crib, a wooden frame used to stabilise a heavy object during a rescue, jacking, construction ...
. Two horses of the same age may have different wear patterns. A horse's incisors, premolars, and molars, once fully developed, continue to erupt as the grinding surface is worn down through chewing. A young adult horse will have teeth which are long, with the majority of the crown remaining below the gumline in the dental socket. The rest of the tooth will slowly emerge from the jaw, erupting about each year, as the horse ages. When the animal reaches old age, the crowns of the teeth are very short and the teeth are often lost altogether. Very old horses, if lacking molars, may need to have their
fodder Fodder (), also called provender (), is any agriculture, agricultural foodstuff used specifically to feed domesticated livestock, such as cattle, domestic rabbit, rabbits, sheep, horses, chickens and pigs. "Fodder" refers particularly to food g ...
ground up and soaked in water to create a soft mush for them to eat in order to obtain adequate nutrition.


Proboscideans

Elephants' tusks are specialized incisors for digging food up and fighting. Some elephant teeth are similar to those in manatees, and it is notable that elephants are believed to have undergone an aquatic phase in their evolution. At birth, elephants have a total of 28 molar plate-like grinding teeth not including the tusks. These are organized into four sets of seven successively larger teeth which the elephant will slowly wear through during its lifetime of chewing rough plant material. Only four teeth are used for chewing at a given time, and as each tooth wears out, another tooth moves forward to take its place in a process similar to a conveyor belt. The last and largest of these teeth usually becomes exposed when the animal is around 40 years of age, and will often last for an additional 20 years. When the last of these teeth has fallen out, regardless of the elephant's age, the animal will no longer be able to chew food and will die of starvation.


Rabbit

Rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s and other lagomorphs usually shed their deciduous teeth before (or very shortly after) their birth, and are usually born with their permanent teeth. The teeth of rabbits complement their diet, which consists of a wide range of vegetation. Since many of the foods are abrasive enough to cause attrition, rabbit teeth grow continuously throughout life. Rabbits have a total of 6 incisors, three upper premolars, three upper molars, two lower premolars, and two lower molars on each side. There are no canines. Three to four millimeters of the tooth is worn away by incisors every week, whereas the posterior teeth require a month to wear away the same amount. The incisors and cheek teeth of rabbits are called aradicular hypsodont teeth. This is sometimes referred to as an elodent dentition. These teeth grow or erupt continuously. The growth or eruption is held in balance by dental abrasion from chewing a diet high in fiber.


Rodents

Rodents have upper and lower hypselodont incisors that can continuously grow enamel throughout its life without having properly formed roots. These teeth are also known as aradicular teeth, and unlike humans whose ameloblasts die after tooth development, rodents continually produce enamel, they must wear down their teeth by gnawing on various materials. Enamel and
dentin Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
are produced by the enamel organ, and growth is dependent on the presence of stem cells, cellular amplification, and cellular maturation structures in the odontogenic region. Rodent incisors are used for cutting wood, biting through the skin of fruit, or for defense. This allows for the rate of wear and tooth growth to be at equilibrium. The microstructure of rodent incisor enamel has shown to be useful in studying the phylogeny and systematics of rodents because of its independent evolution from the other dental traits. The enamel on rodent incisors are composed of two layers: the inner portio interna (PI) with Hunter-Schreger bands (HSB) and an outer portio externa (PE) with radial enamel (RE). It usually involves the differential regulation of the epithelial
stem cell In multicellular organisms, stem cells are undifferentiated or partially differentiated cells that can differentiate into various types of cells and proliferate indefinitely to produce more of the same stem cell. They are the earliest type o ...
niche in the tooth of two rodent species, such as guinea pigs.Tummers M and Thesleff I. Root or crown: a developmental choice orchestrated by the differential regulation of the epithelial stem cell niche in the tooth of two rodent species. Development (2003). 130(6):1049-57.AM Hunt. A description of the molar teeth and investing tissues of normal guinea pigs. J Dent Res. (1959) 38(2):216-31. The teeth have enamel on the outside and exposed
dentin Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
on the inside, so they self-sharpen during gnawing. On the other hand, continually growing molars are found in some rodent species, such as the
sibling vole The East European vole (''Microtus mystacinus'') is a species of vole (rodent) in the family Cricetidae. Distribution and habitat It is found in Albania, Bulgaria, Finland, Greece, Iran, Svalbard (accidentally introduced), North Macedonia, Roma ...
and the guinea pig. There is variation in the dentition of the rodents, but generally, rodents lack canines and premolars, and have a space between their incisors and molars, called the diastema region.


Manatee

Manatees are polyphyodont with mandibular molars developing separately from the jaw and are encased in a bony shell separated by soft tissue.


Walrus

Walrus tusks are canine teeth that grow continuously throughout life.


Fish

Fish, such as sharks, may go through many teeth in their lifetime. The replacement of multiple teeth is known as polyphyodontia. A class of prehistoric shark are called cladodonts for their strange forked teeth. Unlike the continuous shedding of functional teeth seen in modern sharks, the majority of
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
chondrichthyan lineages retained all tooth generations developed throughout the life of the animal. This replacement mechanism is exemplified by the tooth whorl-based dentitions of
acanthodians Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, which ...
, which include the oldest known toothed vertebrate, ''Qianodus'' ''duplicis'.''


Amphibians

All amphibians have
pedicellate teeth Pedicellate teeth are a tooth morphology today unique to modern amphibians, but also seen in a variety of extinct labyrinthodonts. Pedicellate teeth consist of a tooth crown and a base (both composed of dentine) separated by a layer of uncalcifie ...
which are modified to be flexible due to connective tissue and uncalcified
dentin Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
e that separates the crown from the base of the tooth. Most amphibians exhibit teeth that have a slight attachment to the jaw or acrodont teeth. Acrodont teeth exhibit limited connection to the dentary and have little enervation. This is ideal for organisms who mostly use their teeth for grasping, but not for crushing and allows for rapid regeneration of teeth at a low energy cost. Teeth are usually lost in the course of feeding if the prey is struggling. Additionally, amphibians that undergo a metamorphosis develop bicuspid shaped teeth.


Reptiles

The teeth of
reptiles Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...
are replaced constantly throughout their lives.
Crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
juveniles replace teeth with larger ones at a rate as high as one new tooth per socket every month. Once mature, tooth replacement rates can slow to two years and even longer. Overall, crocodilians may use 3,000 teeth from birth to death. New teeth are created within old teeth.


Birds

A skull of Ichthyornis discovered in 2014 suggests that the beak of birds may have evolved from teeth to allow chicks to escape their shells earlier, and thus avoid predators and also to penetrate protective covers such as hard earth to access underlying food.


Invertebrates

True teeth are unique to vertebrates, although many invertebrates have analogous structures often referred to as teeth. The organisms with the simplest genome bearing such tooth-like structures are perhaps the parasitic worms of the family
Ancylostomatidae The Ancylostomatidae are a family of worms that includes the hookworms. Genera of Ancylostomatidae * '' Agriostomum'' * ''Ancylostoma'' * ''Bunostomum ''Bunostomum'' is a genus of nematodes of the small intestine of ruminants and camelids. ...
. For example, the hookworm '' Necator americanus'' has two dorsal and two ventral cutting plates or teeth around the anterior margin of the buccal capsule. It also has a pair of subdorsal and a pair of subventral teeth located close to the rear. Historically the European medicinal leech, another invertebrate parasite, has been used in medicine to remove blood from patients. They have three jaws (tripartite) that resemble saws in both appearance and function, and on them are about 100 sharp teeth used to incise the host. The incision leaves a mark that is an inverted Y inside of a circle. After piercing the skin and injecting
anticoagulant Anticoagulants, commonly known as blood thinners, are chemical substances that prevent or reduce coagulation of blood, prolonging the clotting time. Some of them occur naturally in blood-eating animals such as leeches and mosquitoes, where the ...
s ( hirudin) and anaesthetics, they suck out blood, consuming up to ten times their body weight in a single meal. In some species of Bryozoa, the first part of the stomach forms a muscular gizzard lined with
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
ous teeth that crush armoured prey such as
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s. Wave-like
peristaltic Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, which ...
contractions then move the food through the stomach for digestion.
Mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s have a structure called a
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
which bears a ribbon of
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
ous teeth. However, these teeth are histologically and developmentally different from vertebrate teeth and are unlikely to be
homologous Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor *Sequence homology, biological homology between DNA, RNA, or protein sequences * Homologous chrom ...
. For example, vertebrate teeth develop from a neural crest
mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every o ...
-derived dental papilla, and the neural crest is specific to vertebrates, as are tissues such as enamel. The
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
is used by molluscs for feeding and is sometimes compared rather inaccurately to a tongue. It is a minutely toothed,
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
ous ribbon, typically used for scraping or cutting food before the food enters the oesophagus. The radula is unique to molluscs, and is found in every class of mollusc apart from
bivalves Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
. Within the gastropods, the
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
is used in feeding by both herbivorous and
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
snails and
slug Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word ''slug'' is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a smal ...
s. The arrangement of teeth (also known as denticles) on the radula ribbon varies considerably from one group to another as shown in the diagram on the left. Predatory marine snails such as the Naticidae use the radula plus an acidic secretion to bore through the shell of other molluscs. Other predatory marine snails, such as the
Conidae Conidae, with the current common name of "cone snails", is a taxonomic family (previously subfamily) of predatory sea snails, marine gastropod molluscs in the superfamily Conoidea. The 2014 classification of the superfamily Conoidea, groups onl ...
, use a specialized radula tooth as a poisoned harpoon. Predatory
pulmonate Pulmonata or pulmonates, is an informal group (previously an order, and before that a subclass) of snails and slugs characterized by the ability to breathe air, by virtue of having a pallial lung instead of a gill, or gills. The group includ ...
land
slug Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word ''slug'' is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a smal ...
s, such as the ghost slug, use elongated razor-sharp teeth on the radula to seize and devour
earthworms An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. Th ...
. Predatory cephalopods, such as
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
, use the radula for cutting prey. In most of the more ancient lineages of gastropods, the radula is used to graze by scraping
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s and other microscopic algae off rock surfaces and other substrates.
Limpet Limpets are a group of aquatic snails that exhibit a conical shell shape (patelliform) and a strong, muscular foot. Limpets are members of the class Gastropoda, but are polyphyletic, meaning the various groups called "limpets" descended indep ...
s scrape
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
from rocks using radula equipped with exceptionally hard rasping teeth. These teeth have the strongest known tensile strength of any biological material, outperforming spider silk. The mineral protein of the limpet teeth can withstand a tensile stress of 4.9  GPa, compared to 4 GPa of spider silk and 0.5 GPa of human teeth.


Fossilization and taphonomy

Because teeth are very resistant, often preserved when bones are not, and reflect the diet of the host organism, they are very valuable to archaeologists and palaeontologists. Early fish such as the thelodonts had scales composed of dentine and an enamel-like compound, suggesting that the origin of teeth was from scales which were retained in the mouth. Fish as early as the late
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
had dentine in their exoskeletons, which may have functioned in defense or for sensing their environments.Teaford, Mark F and Smith, Moya Meredith, 2007. ''Development, Function and Evolution of Teeth'', Cambridge University Press. , Chapter 5. Dentine can be as hard as the rest of teeth and is composed of collagen fibres, reinforced with
hydroxyapatite Hydroxyapatite, also called hydroxylapatite (HA), is a naturally occurring mineral form of calcium apatite with the formula Ca5(PO4)3(OH), but it is usually written Ca10(PO4)6(OH)2 to denote that the crystal unit cell comprises two entities. ...
. Though teeth are very resistant, they also can be brittle and highly susceptible to cracking. However, cracking of the tooth can be used as a diagnostic tool for predicting bite force. Additionally, enamel fractures can also give valuable insight into the diet and behaviour of archaeological and fossil samples. Decalcification removes the enamel from teeth and leaves only the organic interior intact, which comprises dentine and cementine. Enamel is quickly decalcified in acids, perhaps by dissolution by plant acids or via diagenetic solutions, or in the stomachs of vertebrate predators. Enamel can be lost by abrasion or spalling, and is lost before dentine or bone are destroyed by the fossilisation process. In such a case, the 'skeleton' of the teeth would consist of the dentine, with a hollow pulp cavity. The organic part of dentine, conversely, is destroyed by alkalis.


See also

* Animal tooth development * Dragon's teeth (mythology)


References


Sources

*


External links

* {{Authority control Animal anatomy Speech organs