Branchial arch

Gill arches supporting the gills in a pike

Branchial arches, or gill arches, are a series of bony "loops" present in fish, which support the gills. As gills are the primitive condition of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arches varies between taxa. In jawed fish, the first arch develops into the jaws, the second into the hyomandibular complex, with the posterior arches supporting gills. In amphibians and reptiles, many elements are lost including the gill arches, resulting in only the oral jaws and a hyoid apparatus remaining. In mammals and birds, the hyoid is still more simplified.

All basal vertebrates breathe with gills. The gills are carried right behind the head, bordering the posterior margins of a series of openings from the esophagus to the exterior. Each gill is supported by a cartilagenous or bony gill arch.[1] Bony fish have three pairs of arches, cartilaginous fish have five to seven pairs, and primitive jawless fish have seven. The vertebrate ancestor no doubt had more arches, as some of their chordate relatives have more than 50 pairs of gills.[2]

In amphibians and some primitive bony fish, the larvae bear external gills, branching off from the gill arches.[3] These are reduced in adulthood, their function taken over by the gills proper in fish and by lungs in most amphibians. Some amphibians retain the external larval gills in adulthood, the complex internal gill system as seen in fish apparently being irrevocably lost very early in the evolution of tetrapods.[4]

Function

The branchial system is typically used for respiration and/or feeding. Many fish have modified anterior gill arches into pharyngeal jaws, often equipped with specialized pharyngeal teeth for handling particular prey items (long, sharp teeth in carnivorous moray eels compared to broad, crushing teeth in durophagous black carp). In amphibians and reptiles, the hyoid arch is modified for similar reasons. It is often used in buccal pumping and often plays a role in tongue protusion for prey capture. In species with highly specialized ballistic tongue movements such as chameleons or some plethodontid salamanders, the hyoid system is highly modified for this purpose, while it is often hypertrophied in species which use suction feeding. Species such as snakes and monitor lizards, whose tongue has evolved into a purely sensory organ, often have very reduced hyoid systems.

Components

The primitive arrangement is 7 (possibly 8) arches, each consisting of the same series of paired (left and right) elements, in order of dorsal to ventral: Pharyngobranchial, epibranchial, ceratobranchial, hypobranchial, and basibranchial. The pharyngobranchials articulate with the neurocranium, while the left and right basibranchials connect to each other (often fusing into a single bone). When part of the hyoid system, the names of the bones are altered by replacing "branchial" with "hyal", thus "ceratobranchial" becomes "ceratohyal".

Higher vertebrates

Higher vertebrates do not have gills. The gill arches form as pharyngeal arches during embryogenesis, and lay the basis of essential structures such as jaws, the thyroid gland, the larynx, the columella (corresponding to the stapes in mammals) and in mammals the malleus and incus.[2]

References

  1. ↑ Scott, Thomas (1996). Concise encyclopedia biology. Walter de Gruyter. p. 542. ISBN 978-3-11-010661-9.
  2. 1 2 Romer, A.S. (1949): The Vertebrate Body. W.B. Saunders, Philadelphia. (2nd ed. 1955; 3rd ed. 1962; 4th ed. 1970)
  3. ↑ "The Origin of the Larva and Metamorphosis in Amphibia". The American Naturalist. Essex Institute. 91: 287. 1957. doi:10.1086/281990. JSTOR 2458911.
  4. ↑ Clack, J. A. (2002): Gaining ground: the origin and evolution of tetrapods. Indiana University Press, Bloomington, Indiana. 369 pp

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