Molar (tooth)

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Molar
Lower wisdom tooth.jpg
A lower wisdom tooth after extraction.
Gray997.png
Permanent teeth of right half of lower dental arch, seen from above: In this diagram, a healthy wisdom tooth (third, rearmost molar) is included
Details
Artery Posterior superior alveolar artery
Identifiers
Latin dentes molaers
MeSH D008963
TA98 A05.1.03.007
TA2 910
FMA 55638
Anatomical terminology
Image showing molar teeth and their arrangement in the mouth of an adult human being 3D Medical Animation Still Showing Types of Teeth.jpg
Image showing molar teeth and their arrangement in the mouth of an adult human being

The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name molar derives from Latin, molaris dens, meaning "millstone tooth", from mola, millstone and dens, tooth. Molars show a great deal of diversity in size and shape across mammal groups. The third molar of humans is a vestigial organ, as it has lost its original function.

Contents

Human anatomy

In humans, the molar teeth have either four or five cusps. Adult humans have 12 molars, in four groups of three at the back of the mouth. The third, rearmost molar in each group is called a wisdom tooth. It is the last tooth to appear, breaking through the front of the gum at about the age of 20, although this varies from individual to individual. Race can also affect the age at which this occurs, with statistical variations between groups. [1] In some cases, it may not even erupt at all.

The human mouth contains upper (maxillary) and lower (mandibular) molars. They are: maxillary first molar, maxillary second molar, maxillary third molar, mandibular first molar, mandibular second molar, and mandibular third molar.

Mammal evolution

In mammals, the crown of the molars and premolars is folded into a wide range of complex shapes. The basic elements of the crown are the more or less conical projections called cusps and the valleys that separate them. The cusps contain both dentine and enamel, whereas minor projections on the crown, called crenulations, are the result of different enamel thickness. Cusps are occasionally joined to form ridges and expanded to form crests. Cingula are often incomplete ridges that pass around the base of the crown. [2]

Mammalian, multicusped cheek teeth probably evolved from single-cusped teeth in synapsids, although the diversity of therapsid molar patterns and the complexity in the molars of the earliest mammals make determining how this happened impossible. According to the widely accepted "differentiation theory", additional cusps have arisen by budding or outgrowth from the crown, while the rivalling "concrescence theory" instead proposes that complex teeth evolved by the clustering of originally separate conical teeth. Therian mammals (placentals and marsupials) are generally agreed to have evolved from an ancestor with tribosphenic cheek teeth, with three main cusps arranged in a triangle. [2]

Comparison of cheek teeth in various taxa: 1, pelicosaur; 2, Dromatherium (a Triassic cynodont); 3, Microconodon (a Triassic eucynodont); 4, Spalacotherium (a Cretaceous symmetrodont); 5, Amphitherium (a Jurassic mammal) Cambridge Natural History Mammalia Fig 039.png
Comparison of cheek teeth in various taxa: 1, pelicosaur; 2, Dromatherium (a Triassic cynodont); 3, Microconodon (a Triassic eucynodont); 4, Spalacotherium (a Cretaceous symmetrodont); 5, Amphitherium (a Jurassic mammal)

Morphology

Generalized cusp of a mammalian molar: ant, anterior; pos, posterior; ci, lingual cingulum; pa, paraconid; pr, protoconid; me, metaconid; hy, hypoconid; hl, hypoconulid; ec, entocristid; tb, talonid basin Australosphenidan molar labeled.png
Generalized cusp of a mammalian molar: ant, anterior; pos, posterior; ci, lingual cingulum; pa, paraconid; pr, protoconid; me, metaconid; hy, hypoconid; hl, hypoconulid; ec, entocristid; tb, talonid basin

Each major cusp on an upper molar is called a cone and is identified by a prefix dependent on its relative location on the tooth: proto-, para-, meta-, hypo-, and ento-. Suffixes are added to these names: -id is added to cusps on a lower molar (e.g., protoconid); -ule to a minor cusp (e.g., protoconulid). A shelf-like ridge on the lower part of the crown (on an upper molar) is called a cingulum; the same feature on the lower molar a cingulid, and a minor cusp on these, for example, a cingular cuspule or conulid. [3]

Tribosphenic

Generalized tribosphenic molar: The protocone is on the lingual (tongue) side, while the anterior paracone and posterior metacone are on the buccal (cheek) side of the jaw). Tribosphenic molar.svg
Generalized tribosphenic molar: The protocone is on the lingual (tongue) side, while the anterior paracone and posterior metacone are on the buccal (cheek) side of the jaw).

The design that is considered one of the most important characteristics of mammals is a three-cusped shape called a tribosphenic molar. This molar design has two important features: the trigonid, or shearing end, and the talonid, or crushing heel. In modern tribosphenic molars, the trigonid is towards the front of the jaw and the talonid is towards the rear.

The tribosphenic tooth is found in insectivores and young platypuses (adults have no teeth). Upper molars look like three-pointed mountain ranges; lowers look like two peaks and a third off to the side.

The tribosphenic design appears primitively in all groups of mammals. Some paleontologists believe that it developed independently in monotremes (or australosphenidans), rather than being inherited from an ancestor that they share with marsupials and placentals (or boreosphenidans); but this idea has critics and the debate is still going on. [4] For example, the dentition of the Early Cretaceous monotreme Steropodon is similar to those of Peramus and dryolestoids, which suggests that monotremes are related to some pre-tribosphenic therian mammals, [5] but, on the other hand, the status of neither of these two groups is well-established.

Some Jurassic mammals, such as Shuotherium and Pseudotribos , have "reversed tribosphenic" molars, in which the talonid is towards the front. This variant is regarded as an example of convergent evolution. [6]

From the primitive tribosphenic tooth, molars have diversified into several unique morphologies. In many groups, a fourth cusp, the hypocone (hypoconid), subsequently evolved (see below).

Quadrate

Pig tooth Pig tooth.jpg
Pig tooth

Quadrate (also called quadritubercular or euthemorphic) molars have an additional fourth cusp on the lingual (tongue) side called the hypocone, located posterior to the protocone. Quadrate molars appeared early in mammal evolution and are present in many species, including hedgehogs, raccoons, and many primates, including humans. [7] There may be a fifth cusp.

In many mammals, additional smaller cusps called conules appear between the larger cusps. They are named after their locations, e.g. a paraconule is located between a paracone and a metacone, a hypoconulid is located between a hypoconid and an entoconid. [7]

Bunodont

Upper and lower dentition of a chimpanzee Chimpjaw.png
Upper and lower dentition of a chimpanzee

In bunodont molars, the cusps are low and rounded hills rather than sharp peaks. They are most common among omnivores such as pigs, bears, and humans. [7] Bunodont molars are effective crushing devices and often basically quadrate in shape. [8]

Hypsodont

Hypsodont dentition is characterized by high-crowned teeth and enamel that extends far past the gum line, which provides extra material for wear and tear. [9] Some examples of animals with hypsodont dentition are cattle and horses, all animals that feed on gritty, fibrous material. Hypsodont molars can continue to grow throughout life, for example in some species of Arvicolinae (herbivorous rodents). [7]

Hypsodont molars lack both a crown and a neck. The occlusal surface is rough and mostly flat, adapted for crushing and grinding plant material. The body is covered with cementum both above and below the gingival line, below which is a layer of enamel covering the entire length of the body. The cementum and the enamel invaginate into the thick layer of dentin. [10]

Brachydont

The opposite condition to hypsodont is called brachydont or brachyodont (from brachys, "short"). It is a type of dentition characterized by low-crowned teeth. Human teeth are brachydont. [7]

A brachydont tooth has a crown above the gingival line and a neck just below it, and at least one root. A cap of enamel covers the crown and extends down to the neck. Cementum is only found below the gingival line. The occlusal surfaces tend to be pointed, well-suited for holding prey and tearing and shredding. [10]

Zalambdodont

Zalambdodont molars have three cusps, one larger on the lingual side and two smaller on the labial side, joined by two crests that form a V- or λ-shape. The larger inner cusp might be homologous with the paracone in a tribosphenic molar, but can also be fused with the metacone. The protocone is typically missing. The two smaller labial cusps are located on an expanded shelf called the stylar shelf. Zalambdodont molars are found in, for example, golden moles and solenodons. [7]

Dilambdodont

Like zalambdodont molars, dilambdodont molars have a distinct ectoloph, but are shaped like two lambdas or a W. On the lingual side, at the bottom of the W, are the metacone and paracone, and the stylar shelf is on the labial side. A protocone is present lingual to the ectoloph. Dilambdodont molars are present in shrews, moles, and some insectivorous bats. [7]

Lophodont

Lophodont molars of Elephas (left) and Loxodonta (center), compared to the nonlophodont mastodon (right) Elefantenzaehne-drawing.jpg
Lophodont molars of Elephas (left) and Loxodonta (center), compared to the nonlophodont mastodon (right)

Lophodont teeth are easily identified by the differentiating patterns of ridges or lophs of enamel interconnecting the cusps on the crowns. Present in most herbivores, these patterns of lophs can be a simple, ring-like edge, as in mole rats, or a complex arrangement of series of ridges and cross-ridges, as those in odd-toed ungulates, such as equids. [8]

Lophodont molars have hard and elongated enamel ridges called lophs oriented either along or perpendicular to the dental row. Lophodont molars are common in herbivores that grind their food thoroughly. Examples include tapirs, manatees, and many rodents. [7]

When two lophs form transverse, often ring-shaped, ridges on a tooth, the arrangement is called bilophodont. This pattern is common in primates, but can also be found in lagomorphs (hares, rabbits, and pikas) and some rodents. [7] [8]

Extreme forms of lophodonty in elephants and some rodents (such as Otomys ) is known as loxodonty. [7] The African elephant belongs to a genus called Loxodonta because of this feature.

Selenodont

In selenodont molars (so-named after moon goddess Selene), the major cusp is elongated into crescent-shaped ridge. Examples include most even-toed ungulates, such as cattle and deer. [7] [8]

Secodont

Carnassials of a Eurasian wolf Carnassials.jpg
Carnassials of a Eurasian wolf

Many carnivorous mammals have enlarged and blade-like teeth especially adapted for slicing and chopping called carnassials. A general term for such blade-like teeth is secodont or plagiaulacoid. [7]

See also

Notes

  1. Rozkovcová, E; Marková, M; Dolejsí, J (1999). "Studies on agenesis of third molars amongst populations of different origin". Sbornik Lekarsky. 100 (2): 71–84. PMID   11220165.
  2. 1 2 Zhao, Weiss & Stock 2000 , Acquisition of multi-cusped cheek teeth in mammals, p. 154
  3. Myers et al. 2013b
  4. Stokstad 2001
  5. Luo, Cifelli & Kielan-Jaworowska 2001
  6. Luo, Ji & Yuan 2007
  7. 1 2 3 4 5 6 7 8 9 10 11 12 Myers et al. 2013a
  8. 1 2 3 4 Lawlor 1979 , pp. 13–4
  9. Flynn, Wyss & Charrier 2007
  10. 1 2 Kwan, Paul W.L. (2007). "Digestive system I" (PDF). Tufts University. Archived from the original (PDF) on 2012-09-13. Retrieved 2013-05-18.

Related Research Articles

Canine tooth tooth

In mammalian oral anatomy, the canine teeth, also called dog teeth, fangs, cuspids or eye teeth, are the relatively long, pointed teeth. They can appear more flattened however, causing them to resemble incisors and leading them to be called incisiform. They developed and are used primarily for firmly holding food in order to tear it apart, and occasionally as weapons. They are often the largest teeth in a mammal's mouth. Individuals of most species that develop them normally have four, two in the upper jaw and two in the lower, separated within each jaw by incisors; humans and dogs are examples. In most species, canines are the anterior-most teeth in the maxillary bone.

Ferugliotherium is a genus of fossil mammals in the family Ferugliotheriidae from the Campanian and/or Maastrichtian period of Argentina. It contains a single species, Ferugliotherium windhauseni, which was first described in 1986. Although originally interpreted on the basis of a single brachydont (low-crowned) molar as a member of Multituberculata, an extinct group of small, rodent-like mammals, it was recognized as related to the hypsodont (high-crowned) Sudamericidae following the discovery of additional material in the early 1990s. After a jaw of the sudamericid Sudamerica was described in 1999, these animals were no longer considered to be multituberculates and a few fossils that were previously considered to be Ferugliotherium were assigned to unspecified multituberculates instead. Since 2005, a relationship between gondwanatheres and multituberculates has again received support. A closely related animal, Trapalcotherium, was described in 2009 on the basis of a single tooth.

Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on gritty, fibrous material. The opposite condition is called brachydont.

Maxillary central incisor tooth

The maxillary central incisor is a human tooth in the front upper jaw, or maxilla, and is usually the most visible of all teeth in the mouth. It is located mesial to the maxillary lateral incisor. As with all incisors, their function is for shearing or cutting food during mastication (chewing). There is typically a single cusp on each tooth, called an incisal ridge or incisal edge. Formation of these teeth begins at 14 weeks in utero for the deciduous (baby) set and 3–4 months of age for the permanent set.

Maxillary lateral incisor

The maxillary lateral incisors are a pair of upper (maxillary) teeth that are located laterally from both maxillary central incisors of the mouth and medially from both maxillary canines. As with all incisors, their function is for shearing or cutting food during mastication, commonly known as chewing. There are generally no cusps on the teeth, but the rare condition known as talon cusps are most prevalent on the maxillary lateral incisors. The surface area of the tooth used in eating is called an incisal ridge or incisal edge. Though relatively the same, there are some minor differences between the deciduous (baby) maxillary lateral incisor and that of the permanent maxillary lateral incisor. The maxillary lateral incisors occlude in opposition to the mandibular lateral incisors.

Mandibular first molar

The mandibular first molar or six-year molar is the tooth located distally from both the mandibular second premolars of the mouth but mesial from both mandibular second molars. It is located on the mandibular (lower) arch of the mouth, and generally opposes the maxillary (upper) first molars and the maxillary 2nd premolar in normal class I occlusion. The function of this molar is similar to that of all molars in regard to grinding being the principal action during mastication, commonly known as chewing. There are usually five well-developed cusps on mandibular first molars: two on the buccal, two lingual, and one distal. The shape of the developmental and supplementary grooves, on the occlusal surface, are describes as being 'M' shaped. There are great differences between the deciduous (baby) mandibular molars and those of the permanent mandibular molars, even though their function are similar. The permanent mandibular molars are not considered to have any teeth that precede it. Despite being named molars, the deciduous molars are followed by permanent premolars.

Dental anatomy

Dental anatomy is a field of anatomy dedicated to the study of human tooth structures. The development, appearance, and classification of teeth fall within its purview. Tooth formation begins before birth, and the teeth's eventual morphology is dictated during this time. Dental anatomy is also a taxonomical science: it is concerned with the naming of teeth and the structures of which they are made, this information serving a practical purpose in dental treatment.

Cusp (anatomy)

A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth. The concept is also used with regard to the valve between the right atrium and the right ventricle in the human heart. This valve is closed during ventricular contraction by the tricuspid valve, so named because it usually consists of three cusps or leaflets.

Dental pertains to the teeth, including dentistry. Topics related to the dentistry, the human mouth and teeth include:

Dryolestoidea is an extinct clade of Mesozoic mammals that only contains two orders. It has been suggested that this group is closely related to modern therian mammals.

<i>Ambondro mahabo</i> Species of small mammal from the middle Jurassic of Madagascar

Ambondro mahabo is a mammal from the Middle Jurassic (Bathonian) Isalo III Formation of Madagascar. The only described species of the genus Ambondro, it is known from a fragmentary lower jaw with three teeth, interpreted as the last premolar and the first two molars. The premolar consists of a central cusp with one or two smaller cusps and a cingulum (shelf) on the inner, or lingual, side of the tooth. The molars also have such a lingual cingulum. They consist of two groups of cusps: a trigonid of three cusps at the front and a talonid with a main cusp, a smaller cusp, and a crest at the back. Features of the talonid suggest that Ambondro had tribosphenic molars, the basic arrangement of molar features also present in marsupial and placental mammals. It is the oldest known mammal with putatively tribosphenic teeth; at the time of its discovery it antedated the second oldest example by about 25 million years.

<i>Carletonomys</i> species of mammal

Carletonomys cailoi is an extinct rodent from the Pleistocene (Ensenadan) of Buenos Aires Province, Argentina. Although known only from a single maxilla with the first molar, its features are so distinctive that it is placed in its own genus, Carletonomys. Discovered in 1998 and formally described in 2008, it is part of a well-defined group of oryzomyine rodents that also includes Holochilus, Noronhomys, Lundomys, and Pseudoryzomys. This group is characterized by progressive semiaquatic specializations and a reduction in the complexity of molar morphology.

Many different terms have been proposed for features of the tooth crown in mammals.

<i>Brachytarsomys mahajambaensis</i> species of mammal

Brachytarsomys mahajambaensis is an extinct rodent from northwestern Madagascar. It is known from nine isolated molars found in several sites during fieldwork that started in 2001. First described in 2010, it is placed in the genus Brachytarsomys together with two larger living species, which may differ in some details of molar morphology. The presence of B. mahajambaensis, a rare element in the local rodent fauna, suggests that the region was previously more humid.

<i>Agathaeromys</i> genus of mammals

Agathaeromys is an extinct genus of oryzomyine rodents from the Pleistocene of Bonaire, Netherlands Antilles. Two species are known, which differ in size and some details of tooth morphology. The larger A. donovani, the type species, is known from hundreds of teeth, found in four localities that are probably 900,000 to 540,000 years old. A. praeuniversitatis, the smaller species, is known from 35 teeth found in a single fossil site, which is probably 540,000 to 230,000 years old.

Nycticebus linglom is a fossil strepsirrhine primate from the Miocene of Thailand. Known only from a single tooth, an upper third molar, it is thought to be related to the living slow lorises, but the material is not sufficient to assign the species to Nycticebus with certainty, and the species name therefore uses open nomenclature. With a width of 1.82 mm, this tooth is very small for a primate. It is triangular in shape, supported by a single root, and shows three main cusps, in addition to various crests. The absence of a fourth cusp, the hypocone, distinguishes it from various other prosimian primates.

Dermotherium is a genus of fossil mammals closely related to the living colugos, a small group of gliding mammals from Southeast Asia. Two species are recognized: D. major from the Late Eocene of Thailand, based on a single fragment of the lower jaw, and D. chimaera from the Late Oligocene of Thailand, known from three fragments of the lower jaw and two isolated upper molars. In addition, a single isolated upper molar from the Early Oligocene of Pakistan has been tentatively assigned to D. chimaera. All sites where fossils of Dermotherium have been found probably developed in forested environments and the fossil species probably were forest dwellers like living colugos, but whether they already had the gliding adaptations of the living species is unknown.

Indraloris is a fossil primate from the Miocene of India and Pakistan in the family Sivaladapidae. Two species are now recognized: I. himalayensis from Haritalyangar, India and I. kamlialensis from the Pothohar Plateau, Pakistan. Other material from the Potwar Plateau may represent an additional, unnamed species. Body mass estimates range from about 2 kg (4.4 lb) for the smaller I. kamlialensis to over 4 kg (8.8 lb) for the larger I. himalayensis.

<i>Apeomyoides</i> genus of mammals

Apeomyoides savagei is a fossil rodent from the Miocene of the United States, the only species in the genus Apeomyoides. It is known from fragmentary jaws and isolated teeth from a site in the early Barstovian, around 15–16 million years ago, of Nevada. Together with other species from scattered localities in the United States, Japan, and Europe, Apeomyoides is classified in the subfamily Apeomyinae of the extinct rodent family Eomyidae. Apeomyines are a rare but widespread group that may have been adapted to a relatively dry habitat.

The Grit, not grass hypothesis is an evolutionary hypothesis that explains the evolution of high-crowned teeth, particularly in New World mammals. The hypothesis is that the ingestion of gritty soil is the primary driver of hypsodont tooth development, not the silica-rich composition of grass, as was previously thought.

References