Branchial arch

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Gill arches supporting the gills in a pike Gills (esox).jpg
Gill arches supporting the gills in a pike

Branchial arches or gill arches are a series of paired bony/cartilaginous "loops" behind the throat (pharyngeal cavity) of fish, which support the fish gills. As chordates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arches varies between taxa. In all jawed fish (gnathostomes), the first arch pair (mandibular arches) develops into the jaw, the second gill arches (the hyoid arches) develop into the hyomandibular complex (which supports the back of the jaw and the front of the gill series), and the remaining posterior arches (simply called branchial arches) support the gills. In tetrapods, a mostly terrestrial clade evolved from lobe-finned fish, many pharyngeal arch elements are lost, including the gill arches. In amphibians and reptiles, only the oral jaws and a hyoid apparatus remains, and in mammals and birds the hyoid is simplified further to support the tongue and floor of the mouth. In mammals, the first and second pharyngeal arches also give rise to the auditory ossicles.

Contents

Most vertebrates are aquatic and breathe with gills, where water comes in contact for exchanging dissolved oxygen before flowing out through a series of openings (gill slits) to the outside. Each gill is supported by a cartilaginous or bony gill arch, [1] which helps to maintain the gill's surface area. Bony fish (osteichthyans, mostly teleost ray-finned fish) have four pairs of arches, cartilaginous fish (chondrichthyans) have five to seven pairs, and the more basal jawless fish ("agnathans") have up to seven. The Cambrian ancestors of vertebrates no doubt had more gill 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 out from the gill arches. [3] These regress upon adulthood, their function taken over by the gills proper in fish, or by lungs (which are homologous to swim bladders) and cutaneous respiration in most amphibians. Some neotenic amphibians (such as the axolotl) 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 posterior 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 protrusion 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. order from dorsal-most (highest) to ventral-most (lowest), these elements are the pharyngobranchial, epibranchial, ceratobranchial, hypobranchial, and basibranchial. The pharyngobranchials may 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 arch, the names of the bones are altered by replacing "-branchial" with "-hyal", thus "ceratobranchial" becomes "ceratohyal". [5]

Amniotes

Amniotes 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] Studies on placoderms also show that the shoulder girdle also originated from gill arches. [6]

Related Research Articles

<span class="mw-page-title-main">Chondrichthyes</span> Class of jawed cartilaginous fishes

Chondrichthyes is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or bony fish, which have skeletons primarily composed of bone tissue. Chondrichthyes are aquatic vertebrates with paired fins, paired nares, placoid scales, conus arteriosus in the heart, and a lack of opercula and swim bladders. Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates.

<span class="mw-page-title-main">Osteichthyes</span> Diverse group of vertebrates with skeletons of bone rather than cartilage

Osteichthyes, also known as osteichthyans or commonly referred to as the bony fish, is a diverse superclass of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes and the extinct placoderms and acanthodians, which have endoskeletons primarily composed of cartilage. The vast majority of extant fish are members of Osteichthyes, being an extremely diverse and abundant group consisting of 45 orders, over 435 families and 28,000 species. It is the largest class of vertebrates in existence today, encompassing most aquatic vertebrates, as well as all semi-aquatic and terrestrial vertebrates.

<span class="mw-page-title-main">Vertebrate</span> Subphylum of chordates

Vertebrates are animals with a backbone or spine, consisting of vertebrae and intervertebral discs. The vertebrae are irregular bones, and the intervertebral discs are of fibrocartilage. The vertebral column surrounds and protects the spinal cord. The other feature unique to vertebrates is the presence of a cranium, or skull.

<span class="mw-page-title-main">Tongue</span> Muscular organ in the mouth of most vertebrates

The tongue is a muscular organ in the mouth of a typical tetrapod. It manipulates food for chewing and swallowing as part of the digestive process, and is the primary organ of taste. The tongue's upper surface (dorsum) is covered by taste buds housed in numerous lingual papillae. It is sensitive and kept moist by saliva and is richly supplied with nerves and blood vessels. The tongue also serves as a natural means of cleaning the teeth. A major function of the tongue is the enabling of speech in humans and vocalization in other animals.

<span class="mw-page-title-main">Gnathostomata</span> Infraphylum of vertebrates

Gnathostomata are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. Most gnathostomes have retained ancestral traits like true teeth, a stomach, and paired appendages. Other traits are elastin, a horizontal semicircular canal of the inner ear, myelin sheaths of neurons, and an adaptive immune system which has discrete lymphoid organs, and uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene.

<span class="mw-page-title-main">Jaw</span> Opposable articulated structure at the entrance of the mouth

The jaws are a pair of opposable articulated structures at the entrance of the mouth, typically used for grasping and manipulating food. The term jaws is also broadly applied to the whole of the structures constituting the vault of the mouth and serving to open and close it and is part of the body plan of humans and most animals.

<span class="mw-page-title-main">Hyoid bone</span> Bone situated in the neck between the chin and the thyroid cartilage

The hyoid bone is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical vertebra.

<span class="mw-page-title-main">Fish anatomy</span> Study of the form or morphology of fishes

Fish anatomy is the study of the form or morphology of fish. It can be contrasted with fish physiology, which is the study of how the component parts of fish function together in the living fish. In practice, fish anatomy and fish physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the latter dealing with how those components function together in living fish.

<span class="mw-page-title-main">Acanthodii</span> Paraphyletic group of cartilaginous fishes

Acanthodii or acanthodians is an extinct class of gnathostomes. They are currently considered to represent a paraphyletic grade of various fish lineages basal to extant Chondrichthyes, which includes living sharks, rays, and chimaeras. Acanthodians possess a mosaic of features shared with both osteichthyans and chondrichthyans. In general body shape, they were similar to modern sharks, but their epidermis was covered with tiny rhomboid platelets like the scales of holosteians.

<span class="mw-page-title-main">Teleostomi</span> Clade of jawed vertebrates

Teleostomi is an obsolete taxon of jawed vertebrates that supposedly includes the tetrapods, bony fish, and the wholly extinct acanthodian fish. Key characters of this group include an operculum and a single pair of respiratory openings, features which were lost or modified in some later representatives. The teleostomes include all jawed vertebrates except the chondrichthyans and the extinct class Placodermi.

<span class="mw-page-title-main">Adelospondyli</span> Extinct order of amphibians

Adelospondyli is an order of elongated, presumably aquatic, Carboniferous amphibians. They have a robust skull roofed with solid bone, and orbits located towards the front of the skull. The limbs were almost certainly absent, although some historical sources reported them to be present. Despite the likely absence of limbs, adelospondyls retained a large part of the bony shoulder girdle. Adelospondyls have been assigned to a variety of groups in the past. They have traditionally been seen as members of the subclass Lepospondyli, related to other unusual early tetrapods such as "microsaurs", "nectrideans", and aïstopods. Analyses such as Ruta & Coates (2007) have offered an alternate classification scheme, arguing that adelospondyls were actually far removed from other lepospondyls, instead being stem-tetrapod stegocephalians closely related to the family Colosteidae.

<span class="mw-page-title-main">Pharyngeal arch</span> Embryonic precursor structures in vertebrates

The pharyngeal arches, also known as visceral arches, are transient structures seen in the embryonic development of humans and other vertebrates, that are recognisable precursors for many structures. In fish, the arches support the gills and are known as the branchial arches, or gill arches.

<span class="mw-page-title-main">Gill slit</span> Individual opening to a gill

Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of cartilaginous fish such as sharks and rays, as well as deep-branching vertebrates such as lampreys. In contrast, bony fishes have a single outer bony gill covering called an operculum.

<span class="mw-page-title-main">Meckel's cartilage</span>

In humans, the cartilaginous bar of the mandibular arch is formed by what are known as Meckel's cartilages also known as Meckelian cartilages; above this the incus and malleus are developed. Meckel's cartilage arises from the first pharyngeal arch.

<i>Gerrothorax</i> Extinct genus of amphibians

Gerrothorax is an extinct genus of temnospondyl amphibian from the Triassic period of Greenland, Germany, Poland, Sweden, and possibly Thailand. It is known from a single species, G. pulcherrimus, although several other species such as G. pustuloglomeratus have been named in the past.

<span class="mw-page-title-main">Skull roof</span> Roofing bones of the skull

The skull roof or the roofing bones of the skull are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium.

<span class="mw-page-title-main">Fish jaw</span>

Most bony fishes have two sets of jaws made mainly of bone. The primary oral jaws open and close the mouth, and a second set of pharyngeal jaws are positioned at the back of the throat. The oral jaws are used to capture and manipulate prey by biting and crushing. The pharyngeal jaws, so-called because they are positioned within the pharynx, are used to further process the food and move it from the mouth to the stomach.

<span class="mw-page-title-main">Evolution of fish</span> Origin and diversification of fish through geologic time

The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include Haikouichthys. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans.

<span class="mw-page-title-main">Fish gill</span> Organ that allows fish to breathe underwater

Fish gills are organs that allow fish to breathe underwater. Most fish exchange gases like oxygen and carbon dioxide using gills that are protected under gill covers (operculum) on both sides of the pharynx (throat). Gills are tissues that are like short threads, protein structures called filaments. These filaments have many functions including the transfer of ions and water, as well as the exchange of oxygen, carbon dioxide, acids and ammonia. Each filament contains a capillary network that provides a large surface area for exchanging oxygen and carbon dioxide.

Ozarcus is an extinct genus of symmoriiform cartilaginous fish from the Carboniferous period of Arkansas. The type species, Ozarcus mapesae, was named in 2014 based on cartilaginous skulls from the Serpukhovian-age Fayetteville Formation. The genus is named after the Ozark Mountains while the species was named after its discoverer, G. K. Mapes.

References

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  2. 1 2 Romer, A.S. (1949): The Vertebrate Body. W.B. Saunders, Philadelphia. (2nd ed. 1955; 3rd ed. 1962; 4th ed. 1970)
  3. Szarski, Henryk (1957). "The Origin of the Larva and Metamorphosis in Amphibia". The American Naturalist. 91 (860). Essex Institute: 287. doi:10.1086/281990. JSTOR   2458911. S2CID   85231736.
  4. Clack, J. A. (2002): Gaining ground: the origin and evolution of tetrapods. Indiana University Press, Bloomington, Indiana. 369 pp
  5. Pradel, Alan; Maisey, John G.; Tafforeau, Paul; Mapes, Royal H.; Mallatt, Jon (16 April 2014). "A Palaeozoic shark with osteichthyan-like branchial arches". Nature. 509 (7502): 608–611. Bibcode:2014Natur.509..608P. doi:10.1038/nature13195. ISSN   1476-4687. PMID   24739974. S2CID   3504437.
  6. Brazeau et al, Fossil evidence for a pharyngeal origin of the vertebrate pectoral girdle, Nature volume 623, pages550–554 (2023)