Linkage mechanisms are especially frequent and manifold in the head of bony fishes, such as wrasses, which have evolved many specialized feeding mechanisms. Especially advanced are the linkage mechanisms of jaw protrusion. For suction feeding a system of linked four-bar linkages is responsible for the coordinated opening of the mouth and 3-D expansion of the buccal cavity. Other linkages are responsible for protrusion of the premaxilla.
Linkage systems are widely distributed in animals. The most thorough overview of the different types of linkages in animals has been provided by M. Muller, who also designed a new classification system, which is especially well suited for biological systems.
The skull of fishes is formed from a series of loosely connected bones. Lampreys and sharks only possess a cartilaginous endocranium, with both the upper and lower jaws being separate elements. Bony fishes have additional dermal bone, forming a more or less coherent skull roof in lungfish and holost fish. The lower jaw defines a chin.
The simpler structure is found in jawless fish, in which the cranium is represented by a trough-like basket of cartilaginous elements only partially enclosing the brain, and associated with the capsules for the inner ears and the single nostril. Distinctively, these fish have no jaws.
Cartilaginous fish, such as sharks, have also simple, and presumably primitive, skull structures. The cranium is a single structure forming a case around the brain, enclosing the lower surface and the sides, but always at least partially open at the top as a large fontanelle. The most anterior part of the cranium includes a forward plate of cartilage, the rostrum, and capsules to enclose the olfactory organs. Behind these are the orbits, and then an additional pair of capsules enclosing the structure of the inner ear. Finally, the skull tapers towards the rear, where the foramen magnum lies immediately above a single condyle, articulating with the first vertebra. There are, in addition, at various points throughout the cranium, smaller foramina for the cranial nerves. The jaws consist of separate hoops of cartilage, almost always distinct from the cranium proper.
In ray-finned fishes, there has also been considerable modification from the primitive pattern. The roof of the skull is generally well formed, and although the exact relationship of its bones to those of tetrapods is unclear, they are usually given similar names for convenience. Other elements of the skull, however, may be reduced; there is little cheek region behind the enlarged orbits, and little, if any bone in between them. The upper jaw is often formed largely from the premaxilla, with the maxilla itself located further back, and an additional bone, the symplectic, linking the jaw to the rest of the cranium.
Although the skulls of fossil lobe-finned fish resemble those of the early tetrapods, the same cannot be said of those of the living lungfishes. The skull roof is not fully formed, and consists of multiple, somewhat irregularly shaped bones with no direct relationship to those of tetrapods. The upper jaw is formed from the pterygoids and vomers alone, all of which bear teeth. Much of the skull is formed from cartilage, and its overall structure is reduced.
Lower jaw 
In vertebrates, the lower jaw (mandible or jawbone) is a bone forming the skull with the cranium. In lobe-finned fishes and the early fossil tetrapods, the bone homologous to the mandible of mammals is merely the largest of several bones in the lower jaw. It is referred to as the 'dentary bone, and forms the body of the outer surface of the jaw. It is bordered below by a number of splenial bones, while the angle of the jaw is formed by a lower angular bone and a suprangular bone just above it. The inner surface of the jaw is lined by a prearticular bone, while the articular bone forms the articulation with the skull proper. Finally a set of three narrow coronoid bones lie above the prearticular bone. As the name implies, the majority of the teeth are attached to the dentary, but there are commonly also teeth on the coronoid bones, and sometimes on the prearticular as well.
This complex primitive pattern has, however, been simplified to various degrees in the great majority of vertebrates, as bones have either fused or vanished entirely. In teleosts, only the dentary, articular, and angular bones remain. Cartilagenous fish, such as sharks, do not have any of the bones found in the lower jaw of other vertebrates. Instead, their lower jaw is composed of a cartilagenous structure homologous with the Meckel's cartilage of other groups. This also remains a significant element of the jaw in some primitive bony fish, such as sturgeons.
Upper jaw 
The upper jaw, or maxilla is a fusion of two bones along the palatal fissure that form the upper jaw. This is similar to the mandible (lower jaw), which is also a fusion of two halves at the mandibular symphysis. In bony fish, the maxilla is called the "upper maxilla," with the mandible being the "lower maxilla". The alveolar process of the maxilla holds the upper teeth, and is referred to as the maxillary arch. In most vertebrates, the foremost part of the upper jaw, to which the incisors are attached in mammals consists of a separate pair of bones, the premaxillae. In bony fish, both maxilla and premaxilla are relatively plate-like bones, forming only the sides of the upper jaw, and part of the face, with the premaxilla also forming the lower boundary of the nostrils. Cartilaginous fish, such as sharks and rays also lack a true maxilla. Their upper jaw is instead formed from a cartilagenous bar that is not homologous with the bone found in other vertebrates.
Pharyngeal jaws 
Pharyngeal jaws are a "second set" of jaws contained within an animal's throat, or pharynx, distinct from the primary (oral) jaws. They are believed to have originated as modified gill arches, in much the same way as oral jaws.
Although approximately 30,000 species of fishes are known to have pharyngeal jaws, in many species having their own teeth, the most notable example of animals possessing them is the moray eels of the family Muraenidae. Unlike those in other fishes known to have them, those of the moray are highly mobile. This is possibly a response to their inability to swallow as do other fishes by creating a negative pressure in the mouth, perhaps induced by their restricted environmental niche (burrows). Instead, when the moray bites prey, it first bites normally with its oral jaws, capturing the prey. Immediately thereafter, the pharyngeal jaws are brought forward and bite down on the prey to grip it; they then retract, pulling the prey down the moray eel's gullet, allowing it to be swallowed.
Wrasse jaws 
Wrasses have become a primary study species in fish-feeding biomechanics due to their jaw structure. They have protractile mouths, usually with separate jaw teeth that jut outwards. Many species can be readily recognized by their thick lips, the inside of which is sometimes curiously folded, a peculiarity which gave rise the German name of "lip-fishes" (Lippfische.)
The nasal and mandibular bones are connected at their posterior ends to the rigid neurocranium, and the superior and inferior articulations of the maxilla are joined to the anterior tips of these two bones, respectively, creating a loop of 4 rigid bones connected by moving joints. This "four-bar linkage" has the property of allowing numerous arrangements to achieve a given mechanical result (fast jaw protrusion or a forceful bite), thus decoupling morphology from function. The actual morphology of wrasses reflects this, with many lineages displaying different jaw morphology that results in the same functional output in a similar or identical ecological niche.
Shark jaws 
Jaws of sharks, like those of rays and skates, are not attached to the cranium. The jaw's surface (in comparison to the shark's vertebrae and gill arches) needs extra support due to its heavy exposure to physical stress and its need for strength. It has a layer of tiny hexagonal plates called "tesserae", which are crystal blocks of calcium salts arranged as a mosaic. This gives these areas much of the same strength found in the bony tissue found in other animals.
Generally sharks have only one layer of tesserae, but the jaws of large specimens, such as the bull shark, tiger shark, and the great white shark, have two to three layers or more, depending on body size. The jaws of a large great white shark may have up to five layers. Because sharks do not have rib cages, they can easily be crushed under their own weight on land. In the rostrum (snout), the cartilage can be spongy and flexible to absorb the power of impacts.
Shark teeth are embedded in the gums rather than directly affixed to the jaw, and are constantly replaced throughout life. Multiple rows of replacement teeth grow in a groove on the inside of the jaw and steadily move forward in comparison to a conveyor belt; some sharks lose 30,000 or more teeth in their lifetime. The rate of tooth replacement varies from once every 8 to 10 days to several months. In most species, teeth are replaced one at a time as opposed to the simultaneous replacement of an entire row, which is observed in the cookiecutter shark.
Tooth shape depends on the shark's diet: those that feed on mollusks and crustaceans have dense and flattened teeth used for crushing, those that feed on fish have needle-like teeth for gripping, and those that feed on larger prey such as mammals have pointed lower teeth for gripping and triangular upper teeth with serrated edges for cutting. The teeth of plankton-feeders such as the basking shark are small and non-functional. While the shark is moving, water passes through the mouth and over the gills in a process known as "ram ventilation". While at rest, most sharks pump water over their gills to ensure a constant supply of oxygenated water. A small number of species have lost the ability to pump water through their gills and must swim without rest. These species are obligate ram ventilators and would presumably asphyxiate if unable to move. Obligate ram ventilation is also true of some pelagic bony fish species.
Teeth on the Echinorhinus cookei jaw
Porbeagle, Lamna nasus, jaw with teeth
Dalatias licha (Kitefin shark) jaw
The teeth of extant elasmobranchs are in several series; the upper jaw is not fused to the cranium, and the lower jaw is articulated with the upper. There are several archetypal jaw suspensions: amphistyly, orbitostyly, hyostyly, and euhyostyly. In amphistyly, the palatoquadrate has a postorbital articulation with the chondrocranium from which ligaments primarily suspend it anteriorly. The hyoid articulates with the mandibular arch posteriorly, but it appears to provide little support to the upper and lower jaws. In orbitostyly, the orbital process hinges with the orbital wall and the hyoid provides the majority of suspensory support. In contrast, hyostyly involves an ethmoid articulation between the upper jaw and the cranium, while the hyoid most likely provides vastly more jaw support compared to the anterior ligaments. Finally, in euhyostyly, also known as true hyostyly, the mandibular cartilages lack a ligamentous connection to the cranium. Instead, the hyomandibular cartilages provide the only means of jaw support, while the ceratohyal and basihyal elements articulate with the lower jaw, but are disconnected from the rest of the hyoid.
The vertebrate jaw probably originally evolved in the Silurian period and appeared in the Placoderm fish which further diversified in the Devonian. Jaws are thought to derive from the pharyngeal arches that support the gills in fish. The two most anterior of these arches are thought to have become the jaw itself (see hyomandibula) and the hyoid arch, which braces the jaw against the braincase and increases mechanical efficiency. While there is no fossil evidence directly to support this theory, it makes sense in light of the numbers of pharyngeal arches that are visible in extant jawed (the Gnathostomes), which have seven arches, and primitive jawless vertebrates (the Agnatha), which have nine.
It is thought that the original selective advantage garnered by the jaw was not related to feeding, but to increased respiration efficiency. The jaws were used in the buccal pump (observable in modern fish and amphibians) that pumps water across the gills of fish or air into the lungs in the case of amphibians. Over evolutionary time the more familiar use of jaws (to humans), in feeding, was selected for and became a very important function in vertebrates.
The hyomandibula is a set of bones found in the hyoid region in most fishes. It usually plays a role in suspending the jaws and/or operculum (teleostomi only). In jawless fishes a series of gills opened behind the mouth, and these gills became supported by cartilaginous elements. The first set of these elements surrounded the mouth to form the jaw. There are ample evidences that vertebrate jaws are homologous to the gill arches of jawless fishes. The upper portion of the second embryonic arch supporting the gill became the hyomandibular bone of jawed fishes, which supports the skull and therefore links the jaw to the cranium.
Jawless fish 
See also 
- Muller, M. (1996). "A novel classification of planar four-bar linkages and its application to the mechanical analysis of animal systems". Phil. Trans. R. Soc. Lond. B 351: 689–720. doi:10.1098/rstb.1996.0065.
- Romer, Alfred Sherwood; Parsons, Thomas S. (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 173–177. ISBN 0-03-910284-X.
- Romer, Alfred Sherwood; Parsons, Thomas S. (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 244–247. ISBN 0-03-910284-X.
- The mandible is also in some sources still referred to as the inferior maxillary bone, though this is an outdated term which goes back to at least the 1858 first edition of Gray's Anatomy, if not earlier.
- OED 2nd edition, 1989.
- Entry "maxilla" in Merriam-Webster Online Dictionary.
- Mehta, Rita S.; Peter C. Wainwright (2007-09-06). "Raptorial jaws in the throat help moray eels swallow large prey". Nature 449 (7158): 79–82. doi:10.1038/nature06062. PMID 17805293. Retrieved 2007-09-06.
- Wainwright et al. (2005). "Many-to-One Mapping of Form to Function: A General Principle in Organismal Design?". Integrative & comparative biology 45: 256–262.
- Chisholm, Hugh, ed. (1911). "wrasse". Encyclopædia Britannica (11th ed.). Cambridge University Press.
- Hamlett, W. C. (1999f). Sharks, Skates and Rays: The Biology of Elasmobranch Fishes. Johns Hopkins University Press. ISBN 0-8018-6048-2. OCLC 39217534.
- Martin, R. Aidan. "Skeleton in the Corset". ReefQuest Centre for Shark Research. Retrieved 2009-08-21.
- "A Shark's Skeleton & Organs". Archived from the original on April 9, 2012. Retrieved August 14, 2009.
- Martin, R. Aidan. "Skin of the Teeth". Retrieved 2007-08-28.
- Gilbertson, Lance (1999). Zoology Laboratory Manual. New York: McGraw-Hill Companies, Inc. ISBN 0-07-237716-X.
- William J. Bennetta (1996). "Deep Breathing". Retrieved 2007-08-28.
- Wilga, C. D. 2005. Morphology and evolution of the jaw suspension in lamniform sharks. Journal of Morphology, 265, 102-119.
- Wilga, C. D., Motta, P. J. & Sanford, C. P. 2007. Evolution and ecology of feeding in elasmobranchs. Integrative and Comparative Biology, 47, 55-69.
- Wilga, C. A. D. 2008. Evolutionary divergence in the feeding mechanism of fishes. Acta Geologica Polonica, 58, 113-120.
- Clack JA (1994) "Earliest known tetrapod braincase and the evolution of the stapes and fenestra ovalis", Nature, 369, 392–394.
- For example: (1) both sets of bones are made from neural crest cells (rather than mesodermal tissue like most other bones); (2) both structures form the upper and lower bars that bend forward and are hinged in the middle; and (3) the musculature of the jaw seem homologous to the gill arches of jawless fishes. (Gilbert 2000)
- Gilbert (2000) "Evolutionary Embryology"
Other reading 
- Forey, Peter; Janvier, Philippe (2000). "Agnathans and the origin of jawed vertebrates". In Gee, Henry. Shaking the tree: readings from Nature in the history of life. USA: University of Chicago Press; Nature/Macmillan Magazines. pp. 251–266. ISBN 978-0-226-28497-2
- Gilbert, Scott F. (c2000). "The anatomical tradition: Evolutionary Embryology: Embryonic homologies". Developmental Biology. Sunderland (MA): Sinauer Associates, Inc. (NCBI). Retrieved January 2010. (3rd and 4th paras, One of the most celebrated cases...)
- Gilbert (2000). Figure 1.14. Jaw structure in the fish, reptile, and mammal. (illustration).
- Hulsey CD, GJ Fraser and JT Streelman (2005) "Evolution and development of complex biomechanical systems: 300 million years of fish jaws" Zebrafish, 2 (4): 243–257.
|Video of a slingjaw wrasse catching prey by protruding its jaw|
|Video of a red bay snook catching prey by suction feeding|