Tooth
A tooth (pl.: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, 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 outermost embryonic germ layer, the ectoderm.
This article is about the structure found in many animals. For the teeth of humans, see Human tooth. For other uses, see Tooth (disambiguation).
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 crocodilians. In most teleost fish, however, the teeth are attached to the outer surface of the bone, while in lizards they are attached to the inner surface of the jaw by one side. In cartilaginous fish, such as sharks, the teeth are attached by tough ligaments to the hoops of cartilage that form the jaw.[1]
Monophyodonts are animals that develop only one set of teeth, while diphyodonts grow an early set of deciduous teeth and a later set of permanent or "adult" teeth. Polyphyodonts grow many sets of teeth. For example, sharks, 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 is due in part to this qualification. Some rodents, such as voles and guinea pigs (but not mice), as well as lagomorpha (rabbits, hares and pikas), have continuously growing molars in addition to incisors.[2][3] Also, tusks (in tusked mammals) grow almost throughout life.[4]
Teeth are not always attached to the jaw, as they are in mammals. In many reptiles 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 denticles 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.[1] Though "modern" teeth-like structures with dentine and enamel have been found in late conodonts, they are now supposed to have evolved independently of later vertebrates' teeth.[5][6]
Living amphibians 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 fangs 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 use a standardised dental formula to describe the precise pattern in any given group.[1]
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Amphibians
All amphibians have pedicellate teeth, which are modified to be flexible due to connective tissue and uncalcified dentine that separates the crown from the base of the tooth.[38]
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.[39] 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.[40]
Etymology
The word tooth comes from Proto-Germanic *tanþs, derived from the Proto-Indo-European *h₁dent-, which was composed of the root *h₁ed- 'to eat' plus the active participle suffix *-nt, therefore literally meaning 'that which eats'.[7]
The irregular plural form teeth is the result of Germanic umlaut whereby vowels immediately preceding a high vocalic in the following syllable were raised. As the nominative plural ending of the Proto-Germanic consonant stems (to which *tanþs belonged) was *-iz, the root vowel in the plural form *tanþiz (changed by this point to *tą̄þi via unrelated phonological processes) was raised to /œː/, and later unrounded to /eː/, resulting in the tōþ/tēþ alternation attested from Old English. Cf. also Old English bōc/bēċ 'book/books' and 'mūs/mȳs' 'mouse/mice', from Proto-Germanic *bōks/bōkiz and *mūs/mūsiz respectively.
Cognate with Latin dēns, Greek ὀδούς (odous), and Sanskrit dát.
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Origin
Teeth are assumed to have evolved either from ectoderm 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 pharyngeal teeth (primarily formed in the pharynx of jawless vertebrates) (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, either ectoderm or endoderm).[4][8]
The genes governing tooth development in mammals are homologous to those involved in the development of fish scales.[9] Study of a tooth plate of a fossil of the extinct fish Romundina stellina 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.[10]
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Reptiles
The teeth of reptiles are replaced constantly throughout their lives. Crocodilian 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.[41]
Fossilization and taphonomy
Because teeth are very resistant, often preserved when bones are not,[52] and reflect the diet of the host organism, they are very valuable to archaeologists and palaeontologists.[53] 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 had dentine in their exoskeletons, which may have functioned in defense or for sensing their environments.[54] Dentine can be as hard as the rest of teeth and is composed of collagen fibres, reinforced with hydroxyapatite.[54]
Though teeth are very resistant, they also can be brittle and highly susceptible to cracking.[55] 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.[56] Enamel is quickly decalcified in acids,[57] perhaps by dissolution by plant acids or via diagenetic solutions, or in the stomachs of vertebrate predators.[56] Enamel can be lost by abrasion or spalling,[56] and is lost before dentine or bone are destroyed by the fossilisation process.[57] In such a case, the 'skeleton' of the teeth would consist of the dentine, with a hollow pulp cavity.[56]
The organic part of dentine, conversely, is destroyed by alkalis.[57]
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