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Embryology

Oral Biology

Alex Forrest
Associate Professor of Forensic Odontology Forensic Science Research & Innovation Centre, Griffith University Consultant Forensic Odontologist, Queensland Health Forensic and Scientific Services, 39 Kessels Rd, Coopers Plains, Queensland, Australia 4108

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Learning Objectives

1. You should be able to understand and explain the basic embryological formation of the face, including the times at which crucial events occur. 2. You should understand and be able to identify the major possible causes of interruption to the formation of the face, and to explain the clinical consequences that result. 3. You should be able to describe and explain the fates of the derivatives of branchial arches I,II,III,IV and VI in man.

Learning Objectives

4. You should be able to relate your embryological knowledge to recognition of patterns in the head and neck, such as the innervation of the palate, and the innervation of major muscle groups in the head.

Early Development

Early Development

As you will remember, the embryo develops from the original fertilized cell, and as cell numbers increase, they become specialized in different regions to form tissues and organs. Developing neural and blood systems provide communications between the different cells and groups of cells.

From: http://www.luc.edu/depts/biol ogy/devf.htm

Early Development

Recall that the embryo develops into a threelayered plate shape that folds into a tube to create the primitive gut, and curves at the head and tail to end up with a C-shaped form that curves around the developing heart.

Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 14

What about the Germ Layers?

Initially, however, the embryo forms a twolayered (or bilaminar) embryonic disk. Mesoderm develops in the third week.

Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 14

Formation of Mesoderm

Modified from http://connection.lww.com/products/sadler/images/figurelarge4-1.jpg

At week three, a structure called the primitive streak develops.

Formation of Mesoderm

Modified from http://connection.lww.com/products/sadler/images/figurelarge4-1.jpg

Ectodermal cells migrate towards this primitive streak, and then invaginate beneath it and spread between the ectoderm and endoderm to form the third layer.

Formation of Mesoderm

Modified from http://connection.lww.com/products/sadler/images/figurelarge4-1.jpg

This process continues until mesoderm separates the ectoderm and endoderm, except at the head end and down at the far tail end.

Formation of Mesoderm

Modified from http://connection.lww.com/products/sadler/images/figurelarge4-1.jpg

Clearly, however, in the head of a developed organism we still have mesoderm. But to understand where it comes from, we need to examine the formation of the nervous system.

Development of Nervous System

The nervous system begins its development towards the head end of the embryo. It begins as a thickening in the ectoderm in this area called the neural plate.

Modified from http://connection.lww.com/products/sadler/images/figurelarge11-1.jpg

Development of Nervous System

Modified from http://publish.uwo.ca/~jkiernan/nrltub.gif

The edges of the neural plate now begin to thicken, producing raised margins to the structure. Between the raised edges lies the neural groove.

Development of Nervous System

Modified from http://publish.uwo.ca/~jkiernan/nrltub.gif

The neural folds continue to develop until they meet on top of the neural fold to produce a neural tube. This will become the ventricles and central canal of the nervous system.

Development of Nervous System

Modified from http://publish.uwo.ca/~jkiernan/nrltub.gif

Notice however the group of cells that develop alongside the growing neural folds called the neural crest, and shown in the diagram above.

Development of Nervous System

Modified from http://publish.uwo.ca/~jkiernan/nrltub.gif

As you can see, the neural crest cells separate from the remainder of the neural tube to form a group of cells in their own right.

Development of Ectomesenchyme

Modified from http://publish.uwo.ca/~jkiernan/nrltub.gif

There is ongoing debate in the literature as to whether the neural crest cells really are a derivative of the developing nervous cell population or are a distinct group themselves.
e.g: Weston JA, Yoshida H, Robinson V, Nishikawa S and Fraser ST (2004). Neural crest and the origin of ectomesenchyme: neural fold heterogeneity suggests an alternative hypothesis. Dev Dyn. 229: 118-30.

Development of Ectomesenchyme

Either way, these cells are most important to the continued development of the embryo. They migrate extensively and give rise to components of the peripheral nervous system including sensory ganglia, sympathetic neurons, Schwann cells, meninges and pigment cells.

Development of Ectomesenchyme

They also give rise to the embryonic connective tissue of the facial region and to the cartilages of the branchial arches. This embryonic connective tissue is known as ectomesenchyme, to differentiate it from the mesenchyme produced at the primitive streak.

Development of Ectomesenchyme

It has been regarded as a fourth germ layer by some authors, and there are cogent reasons for thinking of it in this way.

Development of Ectomesenchyme

It has properties slightly different to those of mesenchyme elsewhere in the body. For instance, the bones of the face and cranium (but not of the base of the skull, which derive from conventional mesoderm) form by intramembranous ossification rather than the endochondral method common in other bones.

Treacher Collins Syndrome

http://www.dental.mu.edu/oralpath/lesions/treachercollins/treachercollins.htm

You should look up Treacher Collins Syndrome, which results from interference in the migration of neural crest cells in the facial area.

Origin of Muscles

Note that the skeletal muscles of the head and neck do not derive from the ectomesenchyme. They develop from somites or somitomeres which migrate into the branchial arches or the frontonasal process. (Somitomeres are similar to somites but derived from paraxial mesoderm. They arise as 7 pairs and give rise to facial muscles, muscles of mastication, extraocular muscles and some muscles of the pharynx.)

Facial Development

Facial Development

The face develops from five embryonic tissue masses, also known as processes. They all arise by rapid multiplication of neural crest cells which originate from ectoderm:
Frontonasal Process Mandibular Processes (paired) Maxillary Processes (paired)

Modified from Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 24

Mouth

The developing mouth is given a special name: the stomodeum or stomatodeum.

From http://www.llk.ie/cleftpalate/embryology.shtml

Mouth

It appears as a depression in the embryonic surface at about the fourth week of development, so it is an early feature.
From http://www.llk.ie/cleftpalate/embryology.shtml

Mouth

Eventually, the stomatodeum will give rise to the oral cavity. At first, it is lined with oral ectoderm, which will give rise to the teeth and finally become the oral epithelium.

Mouth

The floor of this depression pushes against the developing gut, and the wall that separates them is termed the buccopharyngeal membrane. It represents the meeting of ectoderm and endoderm.

Modified from http://www.bartleby.com/107/241.html

Mouth

Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 29

This soon breaks down so that the stomatodeum can open directly into the primitive pharynx of the foregut.

Mouth

At the sides, the stomatodeum becomes limited by the formation of the first pair of branchial arches, which give rise to the lower part of the face.

Modified from Berkovitz, BKB et al. Color Atlas of Oral Anatomy, Histology & Embryology, London, Mosby-Wolfe, 2nd Ed 1992, p. 231

Neck

Development of the neck begins at the same time as development of the face, in the fourth week of embryonic life. It is completed before birth. It arises from the branchial arches and the primitive pharynx, which is part of the digestive tube.

Neck

The caudal part of the primitive pharynx forms the oesophagus, and a ventral outgrowth leads to formation of the larynx, and trachea, which end at the developing lungs.

Arey, Developmental Anatomy, Philadelphia, WB Saunders, 4th Ed 1941, p 218

Branchial Arches

The branchial arches are bulges on the embryo visible immediately beneath the developing brain mass. The first one will form the lower part of the face.
Modified from: Arey, Developmental Anatomy, Philadelphia, WB Saunders, 4th Ed 1941, p. 145

Branchial Arches

Of the six branchial arches, only five survive in man.

From: http://anatome.ncl.ac.uk/tutorials/arch/images/branchial.jpg

Branchial Arches

Each branchial arch contains a portion of primitive striated muscle tissue, some nervous tissue from the neural crest, some vascular tissue, and a bar of cartilage in its mesodermal core. Each arch therefore contains an artery and a nerve.

Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 20

Branchial Arches

The branchial arches are covered on the external surface by ectoderm, and on the internal surface by endoderm, and support the lateral wall of the primitive pharynx.

Modified from: http://anatome.ncl.ac.uk/tutorials/arch/images/branchial.jpg

Branchial Arches

The first two arches are mainly involved in the formation of the face, and the remaining arches are more involved in the formation of structures within the neck.

Modified from Arey, Developmental Anatomy, Philadelphia, WB Saunders, 4th Ed 1941, p 146

1st Branchial Arch

Called the mandibular arch. Cartilage is termed Meckel's cartilage, after the anatomist who first described it. It forms the mandible and the maxillae, the muscles of mastication and the mandibular division of the trigeminal nerve.
Modified from: Grays Anatomy, London, Longman, 35th Ed, 1973, p.117

1st Branchial Arch

The cartilage of the mandibular arch gives rise to the incus and malleus bones, the anterior ligament of the malleus and the sphenomandibular ligament, but the bones of the mandible and maxilla do not form from it. Instead they form by intramembranous ossification following degeneration of the cartilage.

1st Branchial Arch

Modified from: Grays Anatomy, London, Longman, 35th Ed, 1973, p.132

The nerve becomes mandibular division of the trigeminal nerve, and migrates with the muscles.

1st Branchial Arch

The muscle tissue becomes the muscles of mastication, tensor tympani and tensor veli palatini as well as the mylohyoid muscle and the anterior belly of the digastric. All of these muscles take their motor nerve supplies from branches of the mandibular division of the trigeminal nerve. The artery of the first arch does not survive.

2nd Branchial Arch

The second arch is called the hyoid arch. Its cartilage is termed Reichert's cartilage, again after the anatomist who first described it.

Modified from: Grays Anatomy, London, Longman, 35th Ed, 1973, p.117

2nd Branchial Arch

Modified from: Grays Anatomy, London, Longman, 35th Ed, 1973, p.132

Its muscle tissue forms the muscles of facial expression, and its nerve is the facial nerve (VII).

2nd Branchial Arch

The Muscles of Facial Expression therefore all share a common motor nerve supply the facial nerve. Its muscle tissue also forms the stapedius muscle, the stylohyoid and the posterior belly of digastric.

2nd Branchial Arch

Its cartilage contributes to the stapes, styloid process of the temporal bone, and some of the hyoid bone including the upper part of its body and the lesser cornu (to which the stylohyoid ligament attaches), as well as the stylohyoid ligament. The artery of the second arch also degenerates and cannot be recognized in the adult.

The cartilages of the remaining arches are not named.

3rd Branchial Arch

Modified from: Grays Anatomy, London, Longman, 35th Ed, 1973, p.132

The cartilage of the third arch contributes to the hyoid bone. Its muscle becomes the stylopharyngeus muscle, and its nerve becomes the glossopharyngeal nerve (IX).

3rd Branchial Arch

Its artery becomes the common carotid artery and it also contributes to the proximal part of the internal carotid artery.

4th - 6th Branchial Arch

The fourth and sixth arches fuse. Their cartilages contribute to most of the laryngeal cartilages (thyroid, cricoid, arytenoid, corniculate and cuneiform cartilages).

4th - 6th Branchial Arch

The muscles of the fourth arch are innervated by its nerve, the superior laryngeal branch of the vagus nerve (X) The nerve of the sixth arch becomes the recurrent laryngeal branch of vagus (X), which supplies the intrinsic muscles of the larynx.

4th - 6th Branchial Arch

The artery of the fourth arch becomes the arch of the aorta on the left side, and contributes to the right subclavian and brachiocephalic arteries. The artery of the sixth arch contributes to the pulmonary arteries.

5th Branchial Arch

The fifth arch degenerates and becomes unrecognizable in man.

Branchial Pouches & Clefts

A branchial pouch is the fold seen on the inside aspect of the branchial complex between adjacent arches. Branchial cleft is the name given to the fold seen on the outside.

Branchial Pouches & Clefts

The first pharyngeal pouch becomes the auditory tube and middle ear. The first pharyngeal cleft becomes the external auditory meatus, and the tympanic membrane remains as the structure resulting from their meeting.

Branchial Pouches & Clefts

The second pharyngeal pouch becomes the tonsillar fossa between the palataglossal and palatopharyngeal folds. As development of the embryo continues, this develops a mass of lymphoid tissue called the palatine tonsil.

Formation of the Face

The face develops between the 24th and 38th day of gestation. In the early stages, the development of the face is dominated by the changes that create the primitive nasal cavities.
Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 26

Formation of the Face

At about 24 days, the first branchial arch starts to bud off a process called the maxillary process. The stomatodeum is now limited by the developing brain rostrally, by the first Modified from Ten Cate, AR, Oral Histology, Development, branchial arch caudally, Structure & Function, St. Louis, CV Mosby, 1980, p. 24 and by the newly formed maxillary process laterally. The first arch also contains the mandibular process, and contains Meckel's cartilage.

Formation of the Face

The frontonasal process develops two nasal pits, and tissue builds up around them in a horseshoe shape to form the lateral and medial nasal processes.

Modified from Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 24

Formation of the Face

The maxillary process grows medially and approaches the lateral and medial nasal processes. At this stage, it remains separated from them by a groove.
Modified from Berkovitz, BKB et al. Color Atlas of Oral Anatomy, Histology & Embryology, London, Mosby-Wolfe, 2nd Ed 1992, p. 232

Formation of the Face

From Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 27

The continued central growth of the maxillary process pushes the medial nasal process towards the midline.

Formation of the Face

Modified from: Arey, Developmental Anatomy, Philadelphia, WB Saunders, 4th Ed 1941, p. 199

Here they fuse to create the middle part of the nose, the middle part of the upper lip, the anterior part of the maxilla and the primary palate.

Formation of the Face

Modified from: Arey, Developmental Anatomy, Philadelphia, WB Saunders, 4th Ed 1941, p. 199

This is known as the premaxilla, and it carries the odontogenic epithelium for the maxillary incisors.

Formation of the Palate

Modified from: Arey, Developmental Anatomy, Philadelphia, WB Saunders, 4th Ed 1941, p. 199

It is, however, only after the formation of the secondary palate that the distinction between the oral and nasal cavities can be made clearly.

Formation of the Palate

The formation of the secondary palate takes place between the 7th and 8th weeks of development. It results from the fusion of shelves which arise from the maxillary processes.
Modified from: Berkovitz, BKB et al. Color Atlas of Oral Anatomy, Histology & Embryology, London, Mosby-Wolfe, 2nd Edition 1992, p. 234

Formation of the Palate

The nerve of the primary palate is the incisive branch of the long nasopalatine branch of the maxillary division of the trigeminal nerve (V).

From Jamieson, EB, Illustrations of Regional Anatomy, Section II Head and Neck, Edinburgh, Livingstone Ltd, 8th Edition, p 35.

Formation of the Palate

The nerve of the secondary palate is the greater palatine branch of the maxillary division of V.

From Jamieson, EB, Illustrations of Regional Anatomy, Section II Head and Neck, Edinburgh, Livingstone Ltd, 8th Edition, p 35.

Formation of the Palate

This photograph shows the portions of the adult palate derived from the primary and secondary palates. Correlate this with the palatal nerve supply.

Formation of the Tongue


A mesenchymal swelling arises in the midline in the mandibular process of the first branchial arch. It is called the tuberculum impar. Two other swellings appear, one on either side of the tuberculum impar, and these enlarge rapidly and merge with each other and with the tuberculum to form a large mass.
Modified from: Berkovitz, BKB et al. Color Atlas of Oral Anatomy, Histology & Embryology, London, Mosby-Wolfe, 2nd Edition 1992, p. 247

Formation of the Tongue

The mucous membrane of the anterior two-thirds of the tongue is formed from this mass. This is why the anterior two-thirds of the tongue have the mandibular division of the trigeminal nerve (via the lingual nerve) as their sensory nerve supply.
Modified from: Berkovitz, BKB et al. Color Atlas of Oral Anatomy, Histology & Embryology, London, Mosby-Wolfe, 2nd Edition 1992, p. 247

Formation of the Tongue

The nerve of the second arch contributes taste fibres to the anterior two-thirds of the tongue, via the chorda tympani.

Formation of the Tongue


The posterior part of the tongue arises from the hypobranchial eminence, a large midline swelling from the third branchial arch, the nerve of which becomes the glossopharyngeal. Therefore, the posterior third of the tongue comes to be supplied by IX.
Modified from: Berkovitz, BKB et al. Color Atlas of Oral Anatomy, Histology & Embryology, London, Mosby-Wolfe, 2nd Edition 1992, p. 247

Formation of the Mandible

From Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 36

The mandible develops in the mandibular process, which we have already mentioned. The cartilage of the first arch is called Meckel's cartilage.

Formation of the Mandible

It is important to realize that, while this cartilage is in close proximity to the developing mandible, it does not make a large contribution to it.
Modified from Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 38

Formation of the Mandible

The two mandibular processes fuse in the midline to form a mandibular arch. Now, the bone of the mandible begins to form in the mesenchymal tissue that condenses laterally to the cartilage, and the cartilage begins to disappear.

From Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 36

Formation of the Mandible

From: Grays Anatomy, London, Longman, 35th Ed, 1973, p.283

The bone forms a trough which contains the inferior dental nerve.

Formation of the Mandible

From http://www.usc.edu/hsc/dental/ohisto/Cards/odev/17_bb.html

The bone above this forms a series of compartments for the individual teeth, and finally closes over the tooth germs to form a roof over the trough.

Formation of the Mandible

By 10 weeks of age, the rudimentary mandible is formed almost entirely from intramembranous ossification with little direct involvement by Meckel's cartilage.

Formation of the Mandible

From: Grays Anatomy, London, Longman, 35th Ed, 1973, p.283

The cartilage does, however, contribute to the development of the malleus of the middle ear and its ligament, and to the sphenomandibular ligament.

Formation of the Mandible

From: Grays Anatomy, London, Longman, 35th Ed, 1973, p.283

There is some evidence that Meckel's cartilage may contribute to a small extent to the formation of the mandible forwards of the position of the mental foramen.

Formation of the Maxilla

The maxilla also develops from a condensation of mesenchyme from the first branchial arch. There is no branchial arch cartilage in the maxillary process, so the bone formation is entirely by intramembranous ossification. The maxillary sinus forms in the 16th week.

Origin Unknown

Congenital Defects

You will by now be aware that the development of a person is a very complicated and finely-balanced process. Malfunctions of this process at different points can produce congenital defects. Some of these will be genetic in origin, but environmental factors including drugs can also be responsible.

Congenital Defects

The types of environmental factors that can affect the embryo can be classified into five groups:
1. Infectious agents 2. Ionizing radiation 3. Drugs 4. Hormones 5. Nutritional deficiencies

Congenital Defects

Among the most common of the defects that occur are the orofacial clefts. These constitute approximately 13% of the reported anomalies and are the second most common reported malformation.

Modified from Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 45

Modified from Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 43

Origin Unknown

Modified from Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 45

Modified from Ten Cate, AR, Oral Histology, Development, Structure & Function, St. Louis, CV Mosby, 1980, p. 43

Origin Unknown

Origin Unknown

The End

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