The pharyngeal arches, also known as visceral arches'','' are structures seen in the

embryonic development An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...

vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...

s that are recognisable precursors for many structures. In

fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...

, the arches are known as the

branchial arches 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 ar ...

, or gill arches. In the

human embryo Human embryonic development, or human embryogenesis, is the development and formation of the human embryo. It is characterised by the processes of cell division and cellular differentiation of the embryo that occurs during the early stages of d ...

, the arches are first seen during the fourth week of development. They appear as a series of outpouchings of

mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical Emb ...

on both sides of the developing

pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its ...

. The vasculature of the pharyngeal arches is known as the

aortic arches The aortic arches or pharyngeal arch arteries (previously referred to as branchial arches in human embryos) are a series of six paired embryological vascular structures which give rise to the great arteries of the neck and head. They are ventral ...

. In fish, the branchial arches support the

gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they ar ...

Structure

In vertebrates, the pharyngeal arches are derived from all three germ layers (the primary layers of cells that form during embryogenesis). Neural crest cells enter these arches where they contribute to features of the

skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, t ...

and

facial skeleton The facial skeleton comprises the ''facial bones'' that may attach to build a portion of the skull. The remainder of the skull is the braincase. In human anatomy and development, the facial skeleton is sometimes called the ''membranous visceroc ...

such as bone and cartilage. However, the existence of pharyngeal structures before neural crest cells evolved is indicated by the existence of neural crest-independent mechanisms of pharyngeal arch development. The first, most anterior pharyngeal arch gives rise to the oral jaw. The second arch becomes the

hyoid The hyoid bone (lingual bone or tongue-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. ...

and jaw support. In fish, the other posterior arches contribute to the branchial skeleton, which support the gills; in tetrapods the anterior arches develop into components of the ear, tonsils, and thymus. The genetic and developmental basis of pharyngeal arch development is well characterized. It has been shown that Hox genes and other developmental genes such as ''

DLX The DLX (pronounced "Deluxe") is a RISC processor architecture designed by John L. Hennessy and David A. Patterson, the principal designers of the Stanford MIPS and the Berkeley RISC designs (respectively), the two benchmark examples of RISC desi ...

'' are important for patterning the anterior/posterior and dorsal/ventral axes of the branchial arches. Some fish species have a second set of jaws in their throat, known as pharyngeal jaws, which develop using the same genetic pathways involved in oral jaw formation. During human and all

development, a series of pharyngeal arch pairs form in the developing

embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...

. These project forward from the back of the embryo toward the front of the face and neck. Each arch develops its own artery, nerve that controls a distinct muscle group, and skeletal tissue. The arches are numbered from 1 to 6, with 1 being the arch closest to the head of the embryo, and arch 5 existing only transiently. These grow and join in the ventral midline. The first arch, as the first to form, separates the mouth pit or stomodeum from the

pericardium The pericardium, also called pericardial sac, is a double-walled sac containing the heart and the roots of the great vessels. It has two layers, an outer layer made of strong connective tissue (fibrous pericardium), and an inner layer made of ...

. By differential growth the neck elongates and new arches form, so the pharynx has six arches ultimately. Each pharyngeal arch has a

cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck a ...

stick, a

muscle Skeletal muscles (commonly referred to as muscles) are Organ (biology), organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other ...

component that differentiates from the cartilaginous tissue, an artery, and a

cranial nerve Cranial nerves are the nerves that emerge directly from the brain (including the brainstem), of which there are conventionally considered twelve pairs. Cranial nerves relay information between the brain and parts of the body, primarily to and f ...

. Each of these is surrounded by

mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every ...

. Arches do not develop simultaneously but instead possess a "staggered" development. Pharyngeal pouches form on the endodermal side between the arches, and pharyngeal grooves (or clefts) form from the lateral

ectodermal The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from the o ...

surface of the

neck The neck is the part of the body on many vertebrates that connects the head with the torso. The neck supports the weight of the head and protects the nerves that carry sensory and motor information from the brain down to the rest of the body. In ...

region to separate the arches. In fish the pouches line up with the clefts, and these thin segments become

s. In mammals the

endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the gast ...

and

ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from the o ...

not only remain intact but also continue to be separated by a

layer. The development of the pharyngeal arches provides a useful landmark with which to establish the precise stage of embryonic development. Their formation and development corresponds to Carnegie stages 10 to 16 in

mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fu ...

, and Hamburger–Hamilton stages 14 to 28 in the

chicken The chicken (''Gallus gallus domesticus'') is a domestication, domesticated junglefowl species, with attributes of wild species such as the grey junglefowl, grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster ...

. Although there are six pharyngeal arches, in humans the fifth arch exists only transiently during

embryogenesis An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...

First arch

The first pharyngeal arch also mandibular arch (corresponding to the first branchial arch or gill arch of

), is the first of six pharyngeal arches that develops during the fourth week of development.William J. Larsen (2001). Human embryology. Edinburgh: Churchill Livingstone. It is located between the stomodeum and the first pharyngeal groove.

Processes

This arch divides into a

maxillary process Maxillary process is commonly an alternate name for the maxillary prominence. Maxillary process may also refer to the maxillary process of inferior nasal concha, which curves downward and laterally; it articulates with the maxilla and forms a par ...

and a

mandibular process The mandibular prominence is an embryological structure which gives rise to the lower portion of the face. The mandible and lower lip derive from it. The mesenchymal cells within the mandibular prominence condense to form Meckel's cartilage. It i ...

, giving rise to structures including the

bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, an ...

s of the lower two-thirds of the face and the jaw. The maxillary process becomes the

maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...

(or upper jaw, although there are large differences among animals), and

palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly s ...

while the mandibular process becomes the

mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bon ...

or lower jaw. This arch also gives rise to the muscles of mastication.

Meckel's cartilage

Meckel's cartilage forms in the

of the mandibular process and eventually regresses to form the incus and

malleus The malleus, or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibrations ...

of the

middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in ...

, the anterior ligament of the malleus and the

sphenomandibular ligament The sphenomandibular ligament (internal lateral ligament) is one of the three ligaments of the temporomandibular joint. It is situated medially to - and generally separate from - the articular capsule of the joint. Superiorly, it is attached to th ...

. The

or lower jaw forms by perichondral

ossification Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in ...

using Meckel's cartilage as a 'template', but the maxillary does ''not'' arise from direct ossification of Meckel's cartilage.

Derivatives

The skeletal elements and muscles are derived from mesoderm of the pharyngeal arches. Skeletal *

and incus of the

and

* spine of sphenoid bone *

palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the throat. Together with the maxillae, they comprise the hard palate. (''Palate'' is derived from the Latin ...

* squamous part of temporal bone * anterior ligament of malleus Muscles * muscles of mastication (chewing) ** masseter **

medial Medial may refer to: Mathematics * Medial magma, a mathematical identity in algebra Geometry * Medial axis, in geometry the set of all points having more than one closest point on an object's boundary * Medial graph, another graph that re ...

and

lateral pterygoid muscle The lateral pterygoid muscle (or external pterygoid muscle) is a muscle of mastication. It has two heads. It lies superior to the medial pterygoid muscle. It is supplied by pterygoid branches of the maxillary artery, and the lateral pterygoid ne ...

s ** temporalis * mylohyoid muscle * digastric muscle, anterior belly * tensor veli palatini muscle * tensor tympani muscle Other

Mucous membrane A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It i ...

and glands of the anterior two thirds of the tongue are derived from

and

of the arch.

Nerve supply

The mandibular and maxillary branches of the

trigeminal nerve In neuroanatomy, the trigeminal nerve (literal translation, lit. ''triplet'' nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for Sense, sensation in the face and motor functions ...

(CN V) innervate the structures derived from the corresponding processes of the first arch. In some lower animals, each arch is supplied by two cranial nerves. The nerve of the arch itself runs along the cranial side of the arch and is called post-trematic nerve of the arch. Each arch also receives a branch from the nerve of the succeeding arch called the pre-trematic nerve which runs along the caudal border of the arch. In human embryo, a double innervation is seen only in the first pharyngeal arch. The mandibular nerve is the post-trematic nerve of the first arch and

chorda tympani The chorda tympani is a branch of the facial nerve that originates from the taste buds in the front of the tongue, runs through the middle ear, and carries taste messages to the brain. It joins the facial nerve (cranial nerve VII) inside the facia ...

(branch of facial nerve) is the pre-trematic nerve. This double innervation is reflected in the nerve supply of anterior two-thirds of

tongue The tongue is a muscular organ in the mouth of a typical tetrapod. It manipulates food for mastication 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 bu ...

which is derived from the first arch.

Blood supply

The artery of the first arch is the first

aortic arch The aortic arch, arch of the aorta, or transverse aortic arch () is the part of the aorta between the ascending and descending aorta. The arch travels backward, so that it ultimately runs to the left of the trachea. Structure The aorta begins ...

,McMinn, R., 1994. ''Last's anatomy: Regional and applied (9th ed).'' which partially persists as the

maxillary artery The maxillary artery supplies deep structures of the face. It branches from the external carotid artery just deep to the neck of the mandible. Structure The maxillary artery, the larger of the two terminal branches of the external carotid artery, ...

Second arch

The second pharyngeal arch or hyoid arch, is the second of fifth pharyngeal arches that develops in fetal life during the fourth week of development and assists in forming the side and front of the

Reichert's cartilage

Cartilage in the second pharyngeal arch is referred to as Reichert's cartilage and contributes to many structures in the fully developed adult. In contrast to the Meckel's cartilage of the

first pharyngeal arch The pharyngeal arches, also known as visceral arches'','' are structures seen in the embryonic development of vertebrates that are recognisable precursors for many structures. In fish, the arches are known as the branchial arches, or gill arch ...

it does not constitute a continuous element, and instead is composed of two distinct cartilaginous segments joined by a faint layer of

. Dorsal ends of Reichert's cartilage

ossify Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...

during development to form the stapes of the

before being incorporated into the middle ear cavity, while the ventral portion ossifies to form the lesser cornu and upper part of the body of the

hyoid bone The hyoid bone (lingual bone or tongue-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 verteb ...

. Caudal to what will eventually become the stapes, Reichert's cartilage also forms the styloid process of the

temporal bone The temporal bones are situated at the sides and base of the skull, and lateral to the temporal lobes of the cerebral cortex. The temporal bones are overlaid by the sides of the head known as the temples, and house the structures of the ears ...

. The cartilage between the

and styloid process will not remain as development continues, but its

perichondrium The perichondrium (from Greek el, περί, peri, around, label=none and el, χόνδρος, chondros, cartilage, label=none) is a layer of dense irregular connective tissue that surrounds the cartilage of developing bone. It consists of two se ...

will eventually form the

stylohyoid ligament The stylohyoid ligament is a ligament that connects the hyoid bone to the temporal styloid process (of the temporal bone of the skull). Structure The stylohyoid ligament connects the lesser horn of hyoid bone to the styloid process of the tem ...

Derivatives

Skeletal From the cartilage of the second arch arises * Stapes, *

Temporal styloid process The temporal styloid process is a slender bony process of the temporal bone extending downward and forward from the undersurface of the temporal bone just below the ear. The styloid process gives attachments to several muscles, and ligaments. Str ...

, *

Stylohyoid ligament The stylohyoid ligament is a ligament that connects the hyoid bone to the temporal styloid process (of the temporal bone of the skull). Structure The stylohyoid ligament connects the lesser horn of hyoid bone to the styloid process of the tem ...

, and *

Lesser cornu The hyoid bone (lingual bone or tongue-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 ...

of the

. Muscles *

Facial muscles The facial muscles are a group of striated skeletal muscles supplied by the facial nerve (cranial nerve VII) that, among other things, control facial expression. These muscles are also called mimetic muscles. They are only found in mammals, al ...

* Occipitofrontalis muscle *

Platysma The platysma muscle is a superficial muscle of the human neck that overlaps the sternocleidomastoid. It covers the anterior surface of the neck superficially. When it contracts, it produces a slight wrinkling of the neck, and a "bowstring" eff ...

* Stylohyoid muscle *Posterior belly of digastric muscle * Stapedius muscle *Auricular muscles

Nerve supply

Facial nerve The facial nerve, also known as the seventh cranial nerve, cranial nerve VII, or simply CN VII, is a cranial nerve that emerges from the pons of the brainstem, controls the muscles of facial expression, and functions in the conveyance of ta ...

(CN VII)

Blood supply

The artery of the second arch is the second

, which gives origin to the

stapedial artery In human anatomy, the stapedial branch of posterior auricular artery, or stapedial artery for short, is a small artery supplying the stapedius muscle in the inner ear. Structure In humans In humans, the stapedial artery is normally present in the ...

in some mammals but atrophies in humans.

Muscles derived from the pharyngeal arches

Pharyngeal muscles or Branchial muscles are

striated muscle Striations means a series of ridges, furrows or linear marks, and is used in several ways: * Glacial striation * Striation (fatigue), in material * Striation (geology), a ''striation'' as a result of a geological fault * Striation Valley, in Anta ...

s of the head and neck. Unlike skeletal muscles that developmentally come from somites, pharyngeal muscles are developmentally formed from the pharyngeal arches. Most of the skeletal musculature supplied by the cranial nerves ( special visceral efferent) is pharyngeal. Exceptions include, but are not limited to, the extraocular muscles and some of the muscles of the tongue. These exceptions receive general somatic efferent innervation.

First arch

All of the

pharyngeal muscles The pharyngeal muscles are a group of muscles that form the pharynx, which is posterior to the oral cavity, determining the shape of its lumen, and affecting its sound properties as the primary resonating cavity. The pharyngeal muscles (involunta ...

that come from the first pharyngeal arch are innervated by the mandibular divisions of the

. These muscles include all the muscles of mastication, the anterior belly of the digastric, the mylohyoid, tensor tympani, and

tensor veli palatini The tensor veli palatini muscle (tensor palati or tensor muscle of the velum palatinum) is a broad, thin, ribbon-like muscle in the head that tenses the soft palate. Structure The tensor veli palatini is found anterior-lateral to the levator ve ...

Second arch

All of the pharyngeal muscles of the second pharyngeal arch are innervated by the

facial nerve The facial nerve, also known as the seventh cranial nerve, cranial nerve VII, or simply CN VII, is a cranial nerve that emerges from the pons of the brainstem, controls the muscles of facial expression, and functions in the conveyance of ta ...

. These muscles include the muscles of facial expression, the posterior belly of the digastric, the stylohyoid muscle, the auricular muscle and the stapedius muscle of the middle ear.

Third arch

There is only one muscle of the third pharyngeal arch, the

stylopharyngeus The stylopharyngeus is a muscle in the head that stretches between the temporal styloid process and the pharynx. Structure The stylopharyngeus is a long, slender muscle, cylindrical above, flattened below. It arises from the medial side of the ...

. The stylopharyngeus and other structures from the third pharyngeal arch are all innervated by the

glossopharyngeal nerve The glossopharyngeal nerve (), also known as the ninth cranial nerve, cranial nerve IX, or simply CN IX, is a cranial nerve that exits the brainstem from the sides of the upper medulla, just anterior (closer to the nose) to the vagus nerve. Be ...

Fourth and sixth arches

All the pharyngeal muscles of the fourth and sixth arches are innervated by the superior laryngeal and the recurrent laryngeal branches of the

vagus nerve The vagus nerve, also known as the tenth cranial nerve, cranial nerve X, or simply CN X, is a cranial nerve that interfaces with the parasympathetic control of the heart, lungs, and digestive tract. It comprises two nerves—the left and rig ...

. These muscles include all the muscles of the palate (exception of the

which is innervated by the

), all the muscles of the pharynx (except

which is innervated by the

), and all the muscles of the larynx.

In humans

Amniotes have five arches, numbered 1 to 5. Older literature reports the fifth arch as the sixth arch, the fifth being absent. More is known about the fate of the first arch than the remaining four. The first three contribute to structures above the larynx, whereas the last two contribute to the larynx and

trachea The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends from th ...

. The recurrent laryngeal nerves are produced from the nerve of arch 5, and the laryngeal cartilages from arches 4 and 5. The superior laryngeal branch of the vagus nerve arises from arch 4. Its arteries, which project between the nerves of the fourth and fifth arches, become the left-side arch of the aorta and the right subclavian artery. On the right side, the artery of arch 5 is obliterated while, on the left side, the artery persists as the

ductus arteriosus The ''ductus arteriosus'', also called the ''ductus Botalli'', named after the Italian physiologist Leonardo Botallo, is a blood vessel in the developing fetus connecting the trunk of the pulmonary artery to the proximal descending aorta. It all ...

; circulatory changes immediately following birth cause the vessel to close down, leaving a remnant, the

ligamentum arteriosum The ligamentum arteriosum (arterial ligament), also known as the Ligament of Botallo or Harvey's ligament, is a small ligament attaching the aorta to the pulmonary artery. It serves no function in adults but is the remnant of the ductus arteriosu ...

. During growth, these arteries descend into their ultimate positions in the chest, creating the elongated recurrent paths.

References

External links

* {{Authority control Vertebrate anatomy Pharyngeal arches Animal developmental biology

Structure

First arch

Processes

Meckel's cartilage

Derivatives

Nerve supply

Blood supply

Second arch

Reichert's cartilage

Derivatives

Nerve supply

Blood supply

Muscles derived from the pharyngeal arches

First arch

Second arch

Third arch

Fourth and sixth arches

In humans

See also

References

External links