Talk:BGD Lecture - Face and Ear Development: Difference between revisions

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==2019==


== Introduction ==
ILI redevelopment.


[[File:Stage16-18 face.jpg|thumb]]
The face is the anatomical feature which is truly unique to each human, though the basis of its general development is identical for all humans and similar to that seem for other species. The face has a complex origin arising from a number of head structures and sensitive to a number of teratogens during critical periods of its development. The related structures of upper lip and palate significantly contribute to the majority of face abnormalities.


The head and neck structures are more than just the face, and are derived from pharyngeal arches 1 - 6 with the face forming from arch 1 and 2 and the frontonasal prominence.  Each arch contains similar Arch components derived from endoderm, mesoderm, neural crest and ectoderm. These components though will form different structures depending on their arch origin. Because the head contains many different structures also review notes on [[Sensory System Development|Special Senses]]), [[Respiratory System Development|Respiratory]], Integumentary (Teeth), [[Endocrine System Development|Endocrine]] (thyroid, parathyroid, pituitary, thymus) and [[Ultrasound]]- Cleft lip/palate.


==Lecture Objectives==
==2016==
* List the main structures derived from the pharyngeal arches, pouches and clefts. 
Removed these images
* Know the stages and structures involved in the development of the face.  
<gallery>
* Predict the results of abnormal development of the face and palate.
File:Skull anterior.gif|Anterior (anterior fontenelle, sutures, mandible)
* Briefly summarise the development of the tongue.
File:Skull_superior.gif|Superior (anterior fontenelle, sutures)
File:Skull lateral view.gif|Lateral view ( suture, mandible)
</gallery>


[[File:Pharyngeal arch structure cartoon.gif]][[File:Stage13 pharyngeal arch excerpts.gif|300px]]


== Textbook References ==
Original movie media player format
[[File:Pharyngeal_arch_cartilages.jpg|thumb]]
* '''The Developing Human: Clinically Oriented Embryology''' (8th Edition) by Keith L. Moore and T.V.N Persaud - Moore & Persaud Chapter Chapter 10 The Pharyngeal Apparatus pp201 - 240.
* '''Larsen’s Human Embryology''' by GC. Schoenwolf, SB. Bleyl, PR. Brauer and PH. Francis-West -  Chapter 12 Development of the Head, the Neck, the Eyes, and the Ears pp349 - 418.
* [[Head Development]]


<mediaplayer width='410' height='340' image="http://php.med.unsw.edu.au/embryology/images/7/7d/Chicken-neural-crest-migration-01.jpg">File:Chicken-neural crest migration 01.mp4</mediaplayer>


==Movies==
==2013==


{| border='0px'
* [http://embryology.med.unsw.edu.au/embryology/index.php?title=BGD_Lecture_-_Face_and_Ear_Development&oldid=122946 2013]
|-
| [[File:Stage16-18_face_01_icon.jpg|120px|link=Movie_-_Stage_16_to_18_Face]]
| [[File:Stage15-22 head 01 icon.jpg|120px|link=Movie_-_Stage_15_to_22_Head]]
| [[File:Face 001 icon.jpg|120px|link=Development_Animation_-_Face]]
| [[File:Palate_001 icon.jpg|120px|link=Development_Animation_-_Palate 1]]
| [[File:Palate_002 icon.jpg|120px|link=Development_Animation_-_Palate 2]]
| [[File:Tongue_001 icon.jpg|120px|link=Development_Animation_-_Tongue]]
|-
| [[Movie_-_Stage_16_to_18_Face|Face]]


Stage 16 to 18
==2012 ==
| [[Movie_-_Stage_15_to_22_Head|Head]]


Stage 15 to 22
[[File:Stage16-18 face.jpg|thumb]]
| [[Development_Animation_-_Face|Face]]
| [[Development_Animation_-_Palate 1|Palate 1]]
| [[Development_Animation_-_Palate 2|Palate 2]]
| [[Development_Animation_-_Tongue|Tongue]]
|-
|}


==Animation of Face Development==
{|
| <Flowplayer width="420" height="500" autoplay="true">Face_001.flv</Flowplayer>


[[Media:Face_001.mov|Quicktime version]]
[[File:Pharyngeal arch structure cartoon.gif]][[File:Stage13 pharyngeal arch excerpts.gif|300px]]
|
'''Development of the Face'''


===Textbooks===
* '''Before We Are Born''' (5th ed.) Moore and Persaud Chapter 20: p460-479
* '''Essentials of Human Embryology''', Larson Chapter 12: p252-272


This animation shows a ventral view of development of the human face from approximately week 5 through to neonate.
===Online Textbooks===
* '''Developmental Biology''' (6th ed.)  Gilbert, Scott F. Sunderland (MA): Sinauer Associates, Inc.; c2000. [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=dbio.figgrp.5455%20 Evolution of the mammalian middle ear bones from the reptilian jaw] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=dbio.figgrp.5460 Chick embryo rhombomere neural crest cells] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=dbio.table.3135 Some derivatives of the pharyngeal arches] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowSection&rid=dbio.section.2871 Formation of the Neural Tube] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowSection&rid=dbio.section.2884 Differentiation of the Neural Tube] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowSection&rid=dbio.section.2894 Tissue Architecture of the Central Nervous System] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowSection&rid=dbio.section.2908 Neuronal Types] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowSection&rid=dbio.section.2937 Snapshot Summary: Central Nervous System and Epidermis]


The separate embryonic components that contribute to the face have been colour coded.
* '''Neuroscience''' Purves, Dale; Augustine, George J.; Fitzpatrick, David; Katz, Lawrence C.; LaMantia, Anthony-Samuel; McNamara, James O.; Williams, S. Mark. Sunderland (MA): Sinauer Associates, Inc. ; c2001 [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.chapter.879 The Auditory System] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.section.894 The Inner Ear] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.section.893 The Middle Ear] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.section.891 The External Ear] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.chapter.1447 Early Brain Development] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.chapter.1546 Construction of Neural Circuits] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.chapter.1640 Modification of Brain Circuits as a Result of Experience]


* '''Frontonasal Prominence (white)'''
* '''Molecular Biology of the Cell''' (4th Edn) Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter. New York: Garland Publishing; 2002. [http://www.ncbi.nlm.nih.gov:80/books/bv.fcgi?db=Books&rid=mboc4.section.3963 Neural Development] | [http://www.ncbi.nlm.nih.gov:80/books/bv.fcgi?db=Books&rid=mboc4.figgrp.3966 The three phases of neural development]
* <font color=blueviolet>'''Frontonasal Prominence - Lateral nasal'''</font> (purple)
* <font color=yellowgreen>'''Frontonasal Prominence - Medial nasal'''</font> (green)
* <font color=gold>'''Pharyngeal Arch 1 - Maxillary prominence'''</font> (yellow)
* <font color=darkorange>'''Pharyngeal Arch 1 - Mandibular prominence'''</font> (orange)
* '''Stomodeum (black)'''


The [[S#stomodeum|stomodeum]] is the primordial mouth region and a surface central depression lying between the forebrain bulge and the heart bulge. At the floor of the stomodeum indentation is the [[B#buccopharyngeal membrane|buccopharyngeal membrane]] (oral membrane).
* '''Clinical Methods''' [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=cm.chapter.1949 63. Cranial Nerves IX and X: The Glossopharyngeal and Vagus Nerves] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=cm.chapter.3847 The Tongue] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=cm.chapter.3777 126. The Ear and Auditory System] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=cm.chapter.3627#3654 An Overview of the Head and Neck - Ears and Hearing] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=cm.chapter.3897 Audiometry]  


Note the complex origin of the maxillary region (upper jaw) requiring the fusion of several embryonic elements, abnormalities of this process lead to [[C#cleft lip|cleft lip]] and [[C#cleft palate|cleft palate]].
* '''Health Services/Technology Assessment Text (HSTAT)''' Bethesda (MD): National Library of Medicine (US), 2003 Oct. [http://www.ncbi.nlm.nih.gov:80/books/bv.fcgi?db=Books&rid=hstat1a.section.25014#25029 Developmental Disorders Associated with Failure to Thrive]  


See also the movie ([[Quicktime_Movie_-_Stage_16_to_18_Face|Quicktime]] | [[Quicktime_Movie_-_Stage_16_to_18_Face|Flash]]) showing a similar view of human embryo faces between [[Carnegie stage 16|Carnegie stage 16]] to [[Carnegie stage 18|18]].
* '''Eurekah Bioscience Collection'''[http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=eurekah.chapter.53006 Cranial Neural Crest and Development of the Head Skeleton]


===Search ===


* '''Bookshelf'''  [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=hearing+development hearing development]


* '''Pubmed''' [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=hearing+development hearing development]


'''Links:''' [[Quicktime Development Animation - Face]] | [[Media:Face_001.mov|Quicktime version]] | [[Head Development]]
==Links==
|}
* NIDCD - [http://www.nidcd.nih.gov/health/balance/balance_disorders.asp Balance Disorders]


==Buccopharyngeal Membrane==
== Terms ==
'''altricial animal''' - Term used to describe an animal born in a helpless state, with incomplete development of sensory systems at birth. For example rats and mice are born with incomplete development of visual and auditory systems.


These images of the Stage 11 embryo show the breakdown of the buccopharyngeal membrane.
'''ampulla''' - Term used to describe an anatomical dilation of a tube or canal lumen. Anatomical description of the opening end of the uterine tube lying above the ovary and the enlarged initial segmeny of the semicircular canals of the inner ear vestibular system. (More? [ear6.htm Inner Ear] | [genitalXXuterus.htm Genital System - Female Uterus])
<gallery>
File:Stage11 sem4c.jpg|Low power ventral view of the Buccopharyngeal Membrane
File:Stage11 sem3c.jpg|Higher power ventrolateral view of the Buccopharyngeal Membrane
File:Stage11 sem2c.jpg|Close up view of the degenerating Buccopharyngeal Membrane
</gallery>


'''aneurism''' - (Greek, ''aneurysma'' = a widening, aneurysm) A term used to describe an abnormal widening of a vessel or anatomical tubal structure.


==The Pharynx==
'''aquaeductus vestibuli '''- see vestibular aqueduct
[[File:Head arches cartoon.jpg|300px]]
[[File:Stage13 B2 excerpt.gif]]
[[File:Pharynx cartoon.jpg|300px]]


The cavity within the pharyngeal arches forms the pharynx.  
'''auditory neuropathy''' - (AN) abnormality of transmission of sound information to the brain.


* begins at the buccopharyngeal membrane (oral membrane), apposition of ectoderm with endoderm (no mesoderm between)
'''auditory tube '''- (eustachian tube) between the middle ear and oral cavity, has a bony (tympanic 1/3) and cartilaginous (pharyngeal 2/3) portion. The main role is equalization of pressure and fluid drainage in the middle ear.
* expands behind pharyngeal arches
* narrows at glottis and bifurcation of gastrointestinal (oesophagus) and respiratory (trachea) systems
* regions on roof, walls and floor have important contributions to endocrine in oral and neck regions
* also contributes to tongue development


== Pharyngeal Arch Components ==
'''auricular hillock '''- see hillock
[[File:Pharyngeal arch structure cartoon.gif]]


Major features to identify for each: '''arch''', '''pouch''', '''groove''' and '''membrane'''. Contribute to the formation of head and neck and in the human appear at the 4th week. The first arch contributes the majority of upper and lower jaw structures.
'''atresia''' - narrowing, usually of an anatomical tube or cavity.


==Pharyngeal Arch Development==
'''autophagocytosis''' - (Greek, auto = self, phagy = eating, also called autophagy) a cell death mechanism that uses the cell's own lysosomes to self digest.


* branchial arch (Greek. ''branchia'' = gill)
'''border cells''' - columnar cells within the organ of Corti on the medial portion of the basilar membrane.
* arch consists of all 3 trilaminar embryo layers
* ectoderm- outside
* mesoderm- core of mesenchyme
* endoderm- inside


===Neural Crest ===
'''canalis reuniens''' - (ductus reuniens, canaliculus reuniens, canalis reuniens, Hensen's canal, Hensen's duct, uniting canal, canalis reuniens of Hensen) short narrow canal connecting the cochlea duct to the saccule. (Victor Hensen, 1835-1924)
[[File:Head arches cartoon.jpg|thumb|neural crest migration]]
* Mesenchyme invaded by neural crest generating connective tissue components
* cartilage, bone, ligaments
* arises from midbrain and hindbrain region


===Arch Features===
'''cerumen''' - (ear wax) produced by glands in the skin of the outer portion of the ear canal.


Each arch contains: artery, cartilage, nerve, muscular component
'''chondrified''' - the developmental differentiation of cartilage from mesenchye, an embryonic connective tissue.


Arches and Phanynx Form the face, tongue, lips, jaws, palate, pharynx and neck cranial nerves, sense organ components, glands
'''cristae ampullaris''' - located in the ampulla of the membranous semicircular canals a region with both supporting and hair cells. The hair cell cilia are embedded in the gelatinous cupula.


* Humans have 5 arches - 1, 2, 3, 4, 6 (Arch 5 does not form or regresses  rapidly)
'''claudius cells '''- (cells of Claudius) columnar cells with microvilli overlying the basilar membrane and extend from Hensen's cells to the spiral prominence. Barrier cells that lie external to the organ of corti in endolymph.
* from in rostro-caudal sequence, Arch 1 to 6  from week 4 onwards
* arch 1 and 2 appear at time of closure of cranial neuropore


* Face - mainly arch 1 and 2
'''cochlear sac '''- embryonic structure, which will form the coiled cochlear duct and contribute to the saccule.
* Neck components - arch 3 and 4 (arch 4 and 6 fuse)


Arch Features
'''cochlear aqueduct''' - a bony channel containing the fibrous periotic duct. It connects the basal turn of the cochlea perilymphatic space with the subarachnoid space of the posterior cranial cavity.
[[File:Stage13_B2_excerpt.gif]]
** arch
** groove
*** externally separates each arch
**** also called a cleft
*** only first pair persist as external auditory meatus
** pouch
*** internally separates each arch
*** pockets from the pharynx
** membrane
*** ectoderm and endoderm contact regions
*** only first pair persist as tympanic membrane
* Pharyngeal Arch 1 (Mandibular Arch) has 2 prominences
** smaller upper- maxillary forms maxilla, zygomatic bone and squamous part of temporal
** larger lower- mandibular, forms mandible
* Pharyngeal Arch 2  (Hyoid Arch)
** forms most of hyoid bone
* Arch 3 and 4
** neck structures


==Arch Arteries==
'''cochlin''' - major constituent of the inner ear extracellular matrix.


* Arch 1 - mainly lost, form part of maxillary artery
'''collagen type II''' - major constituent of the inner ear extracellular matrix.
* Arch 2 - stapedial arteries
* Arch 3 - common carotid arteries, internal carotid arteries
* Arch 4 - left forms part of aortic arch, right forms part right subclavian artery
* Arch 6 - left forms part of left pulmonary artery , right forms part of right pulmonary artery


placental vein -> liver -> heart -> truncus arteriosus -> aortic sac -> '''arch arteries''' -> dorsal aorta -> placental artery
'''conductive loss''' - term used to describe one of the two major classes of hearing loss involving external and middle ear abnormalities (other form is Sensorineural loss).


==Arch Cartilage==
'''connexins '''- channel proteins of the gap junctions that allow rapid communication between adjacent cells. The two connexins Cx26 and Cx30 are the major proteins of cochlear gap junctions.
[[File:Meckel.jpg|thumb|Meckel's cartilage, first pharyngeal arch]]
[[File:Pharyngeal_arch_cartilages.jpg|thumb|Pharyngeal arch cartilages]]
* Arch 1 - Meckel's cartilage, horseshoe shaped 
** dorsal ends form malleus and incus
** midpart forms ligaments (ant. malleus, sphenomandibular)
** ventral part forms mandible template


* Arch 2 - Reichert's cartilage
'''connexin 26''' - A strikingly high proportion (50%) of congenital bilateral nonsyndromic sensorineural deafness cases have been linked to mutations in the GJB2 coding for the connexin26
** dorsal ends form stapes and Temporal bone styloid process
** ventral part ossifies to form hyoid bone components
** lesser cornu and superior body


* Arch 3- forms greater cornu and inferior part of hyoid
'''cupular deposits''' - basophilic material on the cupulae of the semicircular ducts, an postnatal ageing phenomenon seen in some vestibular labyrinth.


* Arch 4&6- form laryngeal cartilages, except epiglottis (from hypobranchial eminence)
'''clinical weeks''' - taken from last menstrual period (LMP) and therefore approximately two weeks before fertilization occurs.


==Arch Muscle==
Deiters' cells
* Arch 1 - muscles of mastication, mylohyoid, tensor tympanic, ant. belly digastric
* Arch 2 - muscles of facial expression, stapedius, stylohyoid, post. belly digastric
* Arch 3 - stylopharyngeus
* Arch 4&6 - crycothyroid, pharynx constrictors, larynx muscles, oesophagus (st. muscle)


==Arch Nerve==
'''discoidin domain receptor 1''' - (DDR1) a tyrosine kinase receptor activated by native collagen, expressed in the basement membrane and with fibrillar collagens. Found in basal cells of the stria vascularis, type III fibrocytes, and cells lining the basilar membrane of the organ of Corti. {Meyer zum Gottesberge, 2008 #1877}
* Arch 1 - CN V trigeminal, caudal 2/3 maxillary and mandibular, cranial 1/3 sensory nerve of heaad and neck, mastication motor
* Arch 2 - CN VII facial
* Arch 3 - CN IX glossopharyngeal
* Arch 4&6  - CN X vagus, arch 4- superior laryngeal, arch 6- recurrent laryngeal


==Arch Pouches==
ductus utriculosaccularis -  
* Arch 1 - elongates to form tubotympanic recess, tympanic cavity, mastoid antrum, eustachian tube
* Arch 2 - forms tonsillar sinus, mostly oblierated by palatine tonsil
* Arch 3  - forms inferior parathyroid and thymus
* Arch 4 - forms superior parathyroid, parafollicular cells of Thyroid


==Thyroid Gland ==
'''endochondral ossification''' - the process of bone formation from a pre-existing cartilage template.


* not a pouch structure
endoderm -
* first endocrine organ to develop day 24
* from floor of pharynx
* descends thyroglossal duct (which closes)
* upper end at foramen cecum


==Anterior Pituitary==
endolymphatic fluid -
[[File:Historic-pituitary.jpg|thumb|Pituitary]]
* not a pouch structure
* boundary epitheilal ectoderm in the roof of the pharynx
* forms a pocket (Rathke's pouch) that comes into contact with the ectoderm of developing brain.
** Rathke's pouch is named after German embryologist and anatomist Martin Heinrich Rathke (1793 — 1860).


==Face Development==
'''endolymphatic sac''' - inner ear structure that has anatomically both an intraosseous and extraosseous component. Th e sac has functions regulating endolymph that are both secretory and absorptive. Also the site of endolymphatic sac tumors either sporadical occurring or associated with the autosomal-dominant von Hippel-Lindau (VHL) disease, due to a germ line mutation.
[[File:Stage16-18 face animation.gif|right]]
Begins week 4 centered around stomodeum, external depression at oral membrane


5 initial primordia from neural crest mesenchyme
'''embryological weeks''' - taken from the time of fertilization which typically occurs around the middle (day 14), or just after, of the typical 28 day menstrual cycle.
* single frontonasal prominence (FNP) - forms forehead, nose dorsum and apex
* nasal placodes develop later bilateral,  pushed medially
* paired maxillary prominences - form upper cheek and upper lip
* paired mandibular prominences - lower cheek, chin and lower lip


[[:File:Face animation.gif]]
'''Emx2''' - homeobox gene affecting middle ear and inner ear development.


==Head/Skull==
'''eustachian tube''' - (auditory tube) A cavity linking the pharynx to the middle ear, which develops from the first pharyngeal pouch. Named after Bartolomeo Eustachi (1500 - 1574) an Italian anatomist.


* chondrocranium forms base of skull
'''external auditory meatus''' - (ear canal) develops from the first pharyngeal cleft.
* in lower vertebrates encases brain
* cranial vault
* calveria
* facial skeleton
* pharyngeal arches


==Sensory Placodes ==
'''ear wax '''- see cerumen.


'''MH -''' will cover sensory placodes in Senses Lecture.
epithelia -


* During week 4 a series of thickened surface ectodermal patches form in pairs rostro-caudally in the head region.
'''espins''' - calcium-resistant actin-bundling proteins enriched in hair cell stereocilia and sensory cell microvilli and spiral ganglion neurons (SGNs)
* Recent research suggests that all sensory placodes may arise from common panplacodal primordium origin around the neural plate, and then differentiate to eventually have different developmental fates.
* These sensory placodes will later contribute key components of each of our special senses (vision, hearing and smell). Other species have a number of additional placodes which form other sensory structures (fish, lateral line receptor). Note that their initial postion on the developing head is significantly different to their final position in the future sensory system


===Otic placode===
'''eustachian tube''' - (auditory tube) between the middle ear and oral cavity, equalization of pressure in the middle ear.
in the stage 13/14 embryo (shown below) the otic placode has sunk from the surface ectoderm to form a hollow epithelial ball, the otocyst, which now lies beneath the surface surrounded by mesenchyme (mesoderm). The epithelia of this ball varies in thickness and has begun to distort, it will eventually form the inner ear membranous labyrinth.


===Lens placode===
external auditory canal -
lies on the surface, adjacent to the outpocketing of the nervous system (which will for the retina) and will form the lens.


===Nasal placode===
'''fenestra ovalis''' - (oval window) separates the tympanic cavity from the vestibule of the osseous labyrinth.
has 2 components (medial and lateral) and will form the nose olefactory epithelium.


==Head Growth==
'''fenestra rotunda''' - (round window) separates the tympanic cavity from the scala tympani of the cochlea.
[[File:Fetal_head_lateral.jpg|thumb|Fetal Head (12 weeks) showing bone and cartilage]]
* continues postnatally - fontanelle allow head distortion on birth and early growth
* bone plates remain unfused to allow growth, puberty growth of face


===Skull Overview===
'''fetus''' - (foetus) term used to describe human development after the 8th week (10th clinical week, LPM) and covers the developmental periods of second and third trimester.


Chondrocranium - formed from paraxial mesoderm
'''fibroblast growth factor 1''' - (Fgf-1) a growth factor released from cochlea sensory epithelium which stimulates spiral ganglion neurite branching.
* cranial end of vertebral column
* modified vertebral elements
* occipital and cervical sclerotome
* bone preformed in cartilage (endochondrial ossification)


Cranial Vault and Facial Skeleton - formed from neural crest
'''fibroblast growth factor 8''' - (Fgf-8) a growth factor released by inner hair cells which regulates pillar cell number, position and rate of development.
* muscle is paraxial mesoderm
* somitomeres and occipital somites


Calveria - bone has no cartilage (direct ossification of mesenchyme)  
'''fibroblast growth factor receptor 3''' - (Fgfr-3) a tyrosine kinase receptor with a role in the commitment, differentiation and position of pillar cells in the organ of corti
* bones do not fuse, fibrous sutures 1. allow distortion to pass through birth canal 2. allow growth of the brain
* 6 fontanelles,  posterior closes at 3 months, anterior closes at 18 months
{| class="prettytable"
| [[File:Skull anterior.gif]]
| [[File:Skull_superior.gif]]
| [[File:Skull lateral view.gif]]


|-
'''fundamental frequency''' - (natural frequency) the lowest frequency in a harmonic series, for the female voice this is about 225 Hz.
| anterior view
| superior view
| lateral view


|-
'''helicotrema''' - term used to describe the cochlear apex.
| showing anterior fontenelle, sutures, mandible
| showing anterior fontenelle, sutures
| showing suture, mandible


|}
Hes - (hairy and enhancer of split) family of factors, which has been shown to be a general negative regulator of neurogenesis {Zheng, 2000 #1936}.


==Palate==
'''hillock''' - a small hill, used to describe the six surface elevations on pharyngeal arch one and two.
The palate has two key stages of development during embryonic and an early fetal involving the fusion of structures (epithelia to mesenchymal).


=== Embryonic===
Hindbrain - Invaginate -
Primary palate, fusion in the human embryo between stage 17 and 18, from an epithelial seam to the mesenchymal bridge.


[[File:Stage17-18 Primary palate.gif]]
'''Incus''' - (anvil) auditory ossicle


===Fetal===
inner phalangeal cells
[[File:Stage_22_image_061.jpg|thumb]]
Secondary palate, fusion in the human embryo in week 9. This requires the early palatal shelves growth, elevation and fusion during the early embryonic period. The fusion event is to both each other and the primary palate. [[:File:Palatal shelves animation.gif|palatal shelf elevation]] | [[:File:palate.gif|secondary palate]]


==Ear Auricles==
'''inner pillar cells''' - organ of Corti cells arranged in rows and form a boundary between the single row of inner hair cells and three rows of outer hair cells. These cells have surface-associated microtubule bundles.


* form from 6 hillocks (week 5)
inner sulcus - area of the cochlear duct
* 3 on each of arch 1 and 2


==Tongue Development==
interdental region -
[[File:Tongue1.png|200px]] [[File:Tongue2.png|200px]] [[File:Tongue3.png|200px]]


* Ectoderm of the first arch surrounding the stomodeum forms the epithelium lining the buccal cavity.  
'''internal auditory meatus''' - (internal acoustic meatus, IAM) Anatomical canal in which CN VII and CN VIII ganglia reside and pass through to the brainstem. This bony canal lies between the posterior surface of the petrous pyramid and the bony labyrinth within the dense petrous bone. Also associated clinically with the site where acoustic neuromas may occur.  
* Also the salivary glands, enamel of the teeth, epithelium of the body of the tongue.  
** As the tongue develops "inside" the floor of the oral cavity, it is not readily visible in the external views of the embryonic (Carnegie) stages of development.


* Contributions from all arches, which changes with time
'''Kolliker's organ''' - (Kollicker's organ, greater epithelial ridge) Developing cochlear structure consisting of columnar-shaped supporting cells filling the inner sulcus and lying directly under the tectorial membrane. This transient organ regresses and generates the space of the inner sulcus. Rudolph Albert von Kolliker (1817-1905)??
* begins as swelling rostral to foramen cecum, '''median tongue bud'''  


** Arch 1 - oral part of tongue (ant 3/2)
lateral semicircular duct - Limbus -  
** Arch 2 - initial contribution to surface is lost
** Arch 3 - pharyngeal part of tongue (post 1/3)
** Arch 4 - epiglottis and adjacent regions


[[:File:tongue.gif|tongue development animation]] | [http://embryology.med.unsw.edu.au/Movies/larsen/tongue.mov Development of the Tongue]
'''LMP''' - acronym for last menstrual period, used to clinically measure gestation.


===Tongue muscle===
'''malleus''' - (hammer) auditory ossicle
[[File:Tongue-muscle.jpg|thumb|tongue muscle]]
* Tongue muscles originate from the somites. Tongue muscles develop before masticatory muscles and is completed by birth.
* Masticatory muscles originate from the somitomeres. These muscles develop late and are not complete even at birth.


===Salivary Glands===
'''mastoid process''' - of temporal bone


* epithelial buds in oral cavity (wk 6-7) extend into mesenchyme
'''Math1''' - homolog of the Drosophila proneural gene atonal, necessary and sufficient for the production of hair cells in the mouse inner ear. {Chen, 2002 #1932}Negatively regulated by Hes1 and Hes5
* parotid, submandibular, sublingual


== Abnormalities ==
'''meatal plug''' - temporary blockage of the external auditory meatus which forms at the end of the embryonic period and remains present until the seventh month.
[[File:Stage18-19 Cleft Primary Palate.jpg|thumb|300px|Stage 18-19 Cleft Primary Palate]]
===Cleft Lip and Palate===
* 300+ different abnormalities, different cleft forms and extent, upper lip and ant. maxilla, hard and soft palate


====Cleft Palate====
'''meatus''' - anatomical opening, cavity or space (external acoustic meatus,internal auditory meatus)
* Cleft palate has the International Classification of Diseases code 749.0.
* In Australia the national rate (1982-1992) for this abnormalitity in births was 4.8 - 6/10,000 births, which represented 1,530 infants 5.5% were stillborn and 11.5% liveborn died during neonatal period and slightly more common in twin births than singleton.


====Cleft Lip====
'''Meckel's cartilage''' - first pharyngeal ach cartilage, located within the mandibular prominence. This cartilage first appears at stage 16, stage 20 the beginning of membranous ossification. Named after Johann Friedrich Meckel, (1781 - 1833) a German anatomist. (http://www.whonamedit.com/doctor.cfm/1840.html)
* The International Classification of Diseases code 749.1 for isolated cleft lip and 749.2 for cleft lip with cleft palate.
* In Australia the national rate (1982-1992) for this abnormalitity was 8.1 - 9.9 /10,000 births. Of 2,465 infants 6.2% were stillborn and 7.8% liveborn died during neonatal period and the rate was similar in singleton and twin births.


===First Arch Syndrome ===
membranous labyrinth - Mesenchyme - Mesoderm - Microtia - Modiolus -
* There are 2 major types of associated first arch syndromes, Treacher Collins (Mandibulofacial dysostosis) and Pierre Robin (Pierre Robin complex or sequence), both result in extensive facial abnormalites.


==== Treacher Collins Syndrome====
'''mucopolysaccharidosis''' - (MPS IIIB, Sanfilippo Syndrome type B) abnormality caused by a deficiency in the lysosomal enzyme N-acetyl-glucosaminidase (Naglu). Children with MPS IIIB develop abnormal hearing, and mental functioning culminating in early death.


====Pierre Robin Syndrome ====
'''netrin-1''' - secreted growth factor, expressed in the organ of Corti and spiral ganglion cells, role in process outgrowth.
* Hypoplasia of the mandible, cleft palate, eye and ear defects.
* Initial defect is small mandible (micrognathia) resulting in posterior displacement of tongue and a bilateral cleft palate.


===DiGeorge Syndrome===
neural tube -
* absence of thymus and parathyroid glands, 3rd and 4th pouch do not form
* disturbance of cervical neural crest migration


===Cysts===
'''olivocochlear''' - brainstem cholinergic and GABAergic efferent system that innervates sensory cells and sensory neurons of the inner ear.
* Many different types


=== Facial Clefts===
organ of Corti - organ of Corti protein II - (OCP-II) cytosolic protein or transcription factor?
** extremely rare
* Holoprosencephaly
** shh abnormality


===Maternal Effects ===
'''otolithic membrane''' - extracellular matrix that cover the sensory epithelia of the inner ear.
* Retinoic Acid - present in skin ointments
*1988 associated with facial developmental abnormalities


===Fetal Alcohol Syndrome===
'''ossicle''' - (small bone) the individual bone of the three middle ear bones (auditory ossicles), which reduce vibrational amplitude but increase force to drive fluid-filled inner ear.
[[File:FASface.jpg|left]]
Due to alcohol in early development (week 3+) leading to both facial and neurological abnormalities
* lowered ears, small face, mild+ retardation
* Microcephaly - leads to small head circumference
* Short Palpebral fissure - opening of eye
* Epicanthal folds - fold of skin at inside of corner of eye
* Flat midface
* Low nasal bridge
* Indistinct Philtrum - vertical grooves between nose and mouth
* Thin upper lip
* Micrognathia - small jaw


Exposure of embryos in vitro to ethanol simulates premature differentiation of prechondrogenic mesenchyme of the facial primordia (1999)
ossify -


[http://embryology.med.unsw.edu.au/Defect/page5a.htm Fetal Alcohol Syndrome]
otic capsule -


otic cup


== Table - Structures derived from Arches ==
otic placode -  


{| class="prettytable" border=1
otic vesicle -
| '''Arch'''
| '''Nerve'''
| '''Skeletal Structures'''
| '''Muscles'''
| '''Ligaments'''


|-
'''otoconin''' - inner ear biominerals required for vestibular apparatus function.
| 1 (maxillary/mandibular)
| trigeminal (V)
| mandible, maxilla, malleus, incus
| &nbsp;
| ant lig of malleus, sphenomandibular ligament


|-
'''otogelin''' - (Otog) an inner ear specific glycoprotein expressed in cochlea cells at different developmental times.
| 2 (hyoid)  
| facial (VII)
| stapes, styloid process, lesser cornu of hyoid, upper part of body of hyoid bone
| &nbsp;
| stylohyoid ligament


|-
'''otolithic membrane''' - a membrane within the utricle and saccule containing embedded hair cell cilia and small crystalline bodies of calcium carbonate (otoliths). Functions to detect head motion.
| 3
| glossopharyngeal (IX)
| greater cornu of hyoid, lower part of body of hyoid bone
| &nbsp;
|


|-
'''otoliths''' - small crystalline bodies of calcium carbonate found within the otolitic membrane of the utricle and saccule.
| 4 & 6
| superior laryngeal and recurrent laryngeal branch of vagus (X)
| thyroid, cricoid, arytenoid, corniculate and cuneform cartilages
| &nbsp;
|


|}
'''ototoxic''' - compound or drug causing temporary or permanent hearing loss.


=== Structures derived from Arches ===
'''outer hair cells''' - (OHCs) three rows of hair cells that function to increase basilar membrane motion through a local mechanical feedback process within the cochlea, the "cochlear amplifier".


{| class="prettytable" border=1
'''outer pillar cells''' - arranged in rows and form a boundary between the single row of inner hair cells and three rows of outer hair cells.
| '''Arch'''
| '''Nerve'''
| '''Skeletal Structures'''
| '''Muscles'''
| '''Ligaments'''


|-
'''paratubal musculature''' - muscles lying beside the auditory (Eustachian) tube. The tensor veli, palatini (TVP) and tensor tympani muscles.
| 1 (maxillary/mandibular)
| trigeminal (V)
| mandible, maxilla, malleus, incus
| &nbsp;
| ant lig of malleus, sphenomandibular ligament


|-
perilymph - perilymphatic space - Periotic Capsule - petrous portion - of temporal bone
| 2 (hyoid)
| facial (VII)
| stapes, styloid process, lesser cornu of hyoid, upper part of body of hyoid bone
| &nbsp;
| stylohyoid ligament


|-
'''pejvakin gene''' - in humans, two missense mutations in this gene cause nonsyndromic recessive deafness (DFNB59) by affecting the function of auditory neurons.
| 3
| glossopharyngeal (IX)
| greater cornu of hyoid, lower part of body of hyoid bone
| &nbsp;
|


|-
pharyngeal archpharyngeal pouchpharyngeal membranePharynx
| 4 & 6
| superior laryngeal and recurrent laryngeal branch of vagus (X)
| thyroid, cricoid, arytenoid, corniculate and cuneform cartilages
| &nbsp;
|


|}
'''pillar cells''' - (PC) form an inner and outer row of support cells that form a boundary between inner and outer hair cells.
'''Skeletal Structures'''


Placode


{| class="prettytable" border=1
'''preyer reflex''' - ear flick in mouse in response to sound.
| [[Image:mandible_sm.jpg]]
| mandible


|-
presbyacusis
| [[Image:hyoid_bone_sm.jpg]]
| hyoid


|-
'''prestin''' - a motor protein structurally similar to the anion transporter family expressed in cochlear outer hair cells.
| [[Image:larynx_cartilage_sm.jpg]]
| &nbsp;


|}
'''preauricular tag''' - skin tags located in front of the external ear opening, are common in neonates and in most cases are normal, though in some cases are indicative of other associated abnormalities.


=== Structures derived from Pouches ===
primordium-
Each pouch is lined with endoderm and generates specific structures.


'''protocadherin 15''' - (Pcdh15) required for initial formation of stereocilia bundles and changes in the actin meshwork within hair cells. The Ames waltzer (av) mouse mutant has both auditory and vestibular abnormalities from a mutation in this gene.


{| class="prettytable" border=1
'''Reichert's cartilage''' - pharyngeal ach 2 cartilage, named after Karl Bogislaus Reichert (1811 - 1883) a German anatomist.
| <center>'''POUCH'''</center>
| '''Overall Structure'''
| '''Specific Structures'''


|-
'''Reissner's membrane''' - (vestibular membrane, vestibular wall) is a membrane located inside the cochlea separating the scala media from scala vestibuli. Named after Ernst Reissner (1824-1878) a German anatomist. “It primarily functions as a diffusion barrier, allowing nutrients to travel from the perilymph to the endolymph of the membranous labyrinth.
| <center>1</center>
| tubotympanic recess
| tympanic membrane, tympanic cavity, mastoid antrum, auditory tube


|-
rhombomere -
| <center>2</center>
| intratonsillar cleft
| crypts of palatine tonsil, lymphatic nodules of palatine tonsil


|-
Saccular macula -  
| <center>3</center>
| inferior parathyroid gland, thymus gland
|


|-
Saccule - (Latin, sacculus = a small pouch)
| <center>4</center>
| superior parathyroid gland, ultimobranchial body
|


|-
sacculocollic reflex -  
| <center>5</center>
| becomes part of 4th pouch
|


|}
scala tympani - one of the three Cochlea cavities, it is filled with perilymph.


=== Structures derived from Grooves ===
'''Scarpa's ganglion''' - (vestibular ganglion) primary afferent vestibular neuron ganglion of the vestibular nerve. Located within the internal auditory meatus.
Only the '''first''' '''groove''' differentiates into an adult structure and forms part of the external acoustic meatus.
 
=== Structures derived from Membranes ===
At the bottom of each groove lies the membrane which is formed from the contact region of ectodermal groove and endodermal pouch. Only the '''first''' '''membrane''' differentiates into an adult structure and forms the tympanic membrane.
 
== References ==
===Textbooks===
* '''The Developing Human: Clinically Oriented Embryology''' (8th Edition) by Keith L. Moore and T.V.N Persaud - Moore & Persaud Chapter Chapter 10 The Pharyngeal Apparatus pp201 - 240.
* '''Larsen’s Human Embryology''' by GC. Schoenwolf, SB. Bleyl, PR. Brauer and PH. Francis-West -  Chapter 12 Development of the Head, the Neck, the Eyes, and the Ears pp349 - 418.
 
===Online Textbooks===
* '''Developmental Biology''' by Gilbert, Scott F. Sunderland (MA): Sinauer Associates, Inc.; c2000 [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=dbio.figgrp.43 Figure 1.3. Pharyngeal arches] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=dbio.table.3135 Table 13.2. Some derivatives of the pharyngeal arches] | [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=dbio.section.3109#3133 The Cranial Neural Crest]
 
* '''Madame Curie Bioscience Database''' Chapters taken from the Madame Curie Bioscience Database (formerly, Eurekah Bioscience Database) [http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=eurekah&part=A53006 Cranial Neural Crest and Development of the Head Skeleton] | [http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=eurekah&part=ch2957 Neural Crest Cells and the Community of Plan for Craniofacial Development: Historical Debates and Current Perspectives]
 
===Search ===


* '''Bookshelf''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=pharyngeal_arch pharyngeal arch] | [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=head_development head development] |  [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=face_development face development]
'''semicircular canals''' - series of fluid-filled loops of the inner ear required for balance and sensing acceleration.


* '''Pubmed''' [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=pharyngeal_arch pharyngeal arch] | [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=head_development head development] | [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=face_development face development] |
sensorineural - term used to describe one of the two major classes of hearing loss involving the central pathway from the cochlear (other form is conductive loss).


==Movies==
'''space of Nuel''' - within the cochlea, an organ of Corti space between the outer pillar cells and the phalangeal and hair cells. Named after Jean-Pierre Nuel (1847-1920) a Belgian ophthalmologist.
{| border='0px'
|-
| [[File:Face 001 icon.jpg|90px|link=Development_Animation_-_Face]]
| [[File:Palate_001 icon.jpg|90px|link=Development_Animation_-_Palate 1]]
| [[File:Palate_002 icon.jpg|90px|link=Development_Animation_-_Palate 2]]
| [[File:Tongue_001 icon.jpg|90px|link=Development_Animation_-_Tongue]]
|-
| [[Development_Animation_-_Face|Face]]
| [[Development_Animation_-_Palate 1|Palate 1]]
| [[Development_Animation_-_Palate 2|Palate 2]]
| [[Development_Animation_-_Tongue|Tongue]]
|-
|}


== External Links ==
'''spiral ganglion neurons''' - (SGN) innervate the inner (Type I) and outer (Type II) hair cells of the cochlea.
'''Embryo Images Unit:''' [http://www.med.unc.edu/embryo_images/ Embryo Images Online] | [http://www.med.unc.edu/embryo_images/unit-hednk/hednk_htms/hednktoc.htm Craniofacial Development] | [http://www.med.unc.edu/embryo_images/unit-hednk/hednk_htms/hednk001.htm Cell Populations] | [http://www.med.unc.edu/embryo_images/unit-hednk/hednk_htms/hednk007.htm Pharyngeal Arches] | [http://www.med.unc.edu/embryo_images/unit-hednk/hednk_htms/hednk024.htm Tongue] | [http://www.med.unc.edu/embryo_images/unit-hednk/hednk_htms/hednk026.htm Nose and Upper Lip] | [http://www.med.unc.edu/embryo_images/unit-hednk/hednk_htms/hednk033.htm Palate Development]


'''Research Labs:''' [http://www.weizmann.ac.il/Biological_Regulation/tzahor/research.html tzahor lab]
'''stapedius muscle''' - (innervated by CN VII tympanic branch) one of the two muscles in the middle ear, contraction of this muscle pulls the stapes and dampens auditory ossicle movement.


==Terms==
'''stapes''' - (stirrup) a middle ear auditory ossicle (bone).stapes footplate - startle response -


===palate===
'''stereocilia''' -finger-like projections from the apical surface of sensory hair cells forming the hair bundle in the cochlea. Formed by tightly cross-linked parallel actin filaments in a paracrystalline array with cell surface specializations (tip links, horizontal top connectors, and tectorial membrane attachment crowns).


:The roof of the mouth (oral cavity) a structure which separates the oral from the nasal cavity. Develops as two lateral palatal shelves which grow and fuse in the midline. Initally a primary palate forms with fusion of the maxillary processes with the nasal processes in early face formation. Later the secondary palate forms the anterior [[H#hard_palate|hard palate]] which will ossify and separate the oral and nasal cavities. The posterior part of the palate is called the soft palate (velum, muscular palate) and contains no bone. Abnormalities of palatal shelf fusion can lead to [[C#cleft_palate|cleft palate]]. (More?  [[Head Development|Head]] | [[Head_Development_-_Abnormalities|Head Abnormalities]] | [http://www.nlm.nih.gov/medlineplus/cleftlipandpalate.html Medline Plus - Cleft Lip and Palate])
'''stratified squamous epithelia''' - classification of epithelium which transiently forms a plug in external ear canal to the outer eardrum.


===palatogenesis===
'''stria vascularis''' - forms the outer wall of the cochlear duct of the mammalian cochlea is composed primarily of three types of cells. Marginal cells line the lumen of the cochlear duct and are of epithelial origin. Basal cells also form a continuous layer and they may be mesodermal or derived from the neural crest. Intermediate cells are melanocyte-like cells, presumably derived from the neural crest, and are scattered between the marginal and basal cell layers. The stria forms endolymph and also contains a rich supply of blood vessels.


:The process of palate formation, divided into primary and secondary palate development. (More? [[Head Development|Head]] | [[Head_Development_-_Abnormalities|Head Abnormalities]] | [http://www.nlm.nih.gov/medlineplus/cleftlipandpalate.html Medline Plus - Cleft Lip and Palate])
sulcus -  


===pharyngeal arch===
'''synostotically''' - anatomically normally separate skeletal bones fused together.


:([[B#branchial arch|branchial arch]], Greek, ''branchial'' = gill) These are a series of externally visible anterior tissue bands lying under the early brain that give rise to the structures of the head and neck. In humans, five arches form (1,2,3,4 and 6) but only four are externally visible on the [[E#embryo|embryo]]. Each arch has initially identical structures: an internal endodermal pouch, a mesenchymal ([[M#mesoderm|mesoderm]] and [[N#neural crest|neural crest]]) core, a membrane ([[E#endoderm|endoderm]] and [[E#ectoderm|ectoderm]]) and external cleft ([[E#ectoderm|ectoderm]]). Each arch mesenchymal core also contains similar components: blood vessel, nerve, muscular, cartilage. Each arch though initially formed from similar components will differentiate to form different head and neck structures. (More? | [[Head Development]] | [[Endocrine System Development|Endocrine]] | [[Neural Crest Development|Neural Crest]])
'''tectorial membrane''' - extracellular matrix that cover the sensory epithelial hair cells of the organ of corti within the cochlea.


===pharyngeal arch artery===
'''alpha-tectorin and beta'''- (TECTA, TECTB) major non-collagenous protein component of the tectorial membrane forming a striated-sheet matrix. Synthesized as glycosylphosphatidylinositol-linked, membrane bound precursors.


:Each early developing pharyngeal arch contains a lateral pair of arteries arising from the aortic sac, above the heart, and running into the dorsal aorta. later in development these arch arteries are extensively remodelled to form specific components of the vascular system. Pharyngeal Arch 1 arteries are mainly lost and forms part of maxillary artery. Pharyngeal Arch 2 arteries remains to form the stapedial arteries. Pharyngeal Arch 3 arteries forms the common carotid arteries, internal carotid arteries in the neck. Pharyngeal Arch 4 arteries will form part of aortic arch (left arch artery) and part right subclavian artery (right arch artery) Pharyngeal Arch 6 arteries form part of left pulmonary artery (left arch artery) and part of right pulmonary artery (right arch artery).  (More? | [[Head Development]] | [[Cardiovascular System Development|Cardiovascular]])
temporal bone -


===pharyngeal arch cartilage===
'''tensor tympani '''- (innervated by CN V mandibular nerve) one of the two muscles in the middle ear, contraction of this muscle pulls the malleus and tenses the tympanic membrane, dampening auditory ossicle movement. The muscle arises from auditory tube (cartilaginous portion) and is inserted into the malleus (manubrium near the root).


:Each early developing pharyngeal arch contains a horseshoe shaped band of cartilage that acts as a template and contributes to the development of head and neck bony and cartilagenous features, including the middle ear bones. Pharyngeal Arch 1 cartilage (Meckel‚Äôs cartilage) dorsal ends form malleus and incus midpart forms ligaments (ant. malleus, sphenomandibular) ventral part forms mandible template. Pharyngeal Arch 2 cartilage (Reichert‚Äôs cartilage) dorsal ends form stapes and Temporal bone styloid process, ventral part ossifies to form hyoid bone components, lesser cornu and superior body. Pharyngeal Arch 3 cartilage forms hyoid components, greater cornu and inferior part of hyoid. Pharyngeal Arch 4 and 6 cartilage forms laryngeal cartilages except epiglottis (from hypobranchial eminence). (More? [[Head Development]] |  [[Hearing_-_Middle_Ear_Development|Middle Ear]])
teratogens - trilaminar embryo -  


===pharyngeal arch nerve===
'''tonotopy''' - term describing the mapping along the tectorial membrane within the cochlea of the different sound frequencies.


:Each early developing pharyngeal arch contains the developing cranial nerves, as a pair, within the arch mesenchyme. Each cranial nerve is numbered (roman numeral) in rostrocaudal sequence and also has a specific name. The cranial nerve within each arch often relates to the other structures formed from taht arch. Pharyngeal Arch 1 contains the trigeminal nerve (CN V, cranial nerve 5). Pharyngeal Arch 2 contains the facial nerve (CN VII, cranial nerve 7). Pharyngeal Arch 3 contains the glossopharyngeal nerve (CN IX, cranial nerve 9) Pharyngeal Arch 4 and 6 contains the Vagus (CN X cranial nerve 10), forming the adult superior laryngeal and recurrent laryngeal branches. (More? | [[Head Development]] | [[Neural System Development|Neural]] | [[Neural Crest Development|Neural Crest]])
tympanic cavity - tympanic membrane -Utricle -Vacuolization - Vesicle - vestibular apparatus - vestibular evoked myogenic potential (VEMP) test


===pharyngeal arch pouch===
'''vestibular ganglion''' - (Scarpa's ganglion) primary afferent vestibular neuron ganglion of the vestibular nerve. Located within the internal auditory meatus.


:An out-pocketing of the [[E#endoderm|endoderm]] lined pharynx occurring between each developing pharyngeal arch. Each of the pharyngeal arch pouches contributes different components of the head and neck, either cavities or endocrine tissues. Pharyngeal Arch 1 pouch elongates to form tubotympanic recess tympanic cavity, mastoid antrum and auditory tube (Eustachian tube). Pharyngeal Arch 2 pouch forms the tonsillar sinus and is later mostly oblierated by palatine tonsil. Pharyngeal Arch 3 pouch forms the inferior parathyroid and thymus. Pharyngeal Arch 4 pouch forms the superior parathyroid, parafollicular cells of Thyroid. (More? [[Hearing_-_Middle_Ear_Development|Middle Ear]] |  [[Endocrine - Thyroid Development‎|Thyroid]] | [[Endocrine - Parathyroid Development|Parathyroid]] | [[Endocrine - Thymus Development|Thymus‎]] |  [[Endocrine System Development|Endocrine]] | [[Head Development]]
'''vestibular membrane''' - (Reissner's) extends from the spiral lamina to the outer wall and divides the cochlea into an upper scala vestibuli, a lower scala tympani.
===pharyngotympanic tube===


:([[A#auditory tube|auditory tube]], [[E#eustachian tube|eustachian tube]], [[O#otopharyngeal tube|otopharyngeal tube]]) A narrow canal connecting the [[M#middle ear|middle ear]] space to the back of the oral cavity. The tube allows ventilation, protection and clearance for the middle ear cavity. Ventilation is the pressure equalization in the middle ear. Clearance is to allow fluid drainage from the middle ear. Embryonic origin is from the first pharyngeal pouch. In development, the canal is initially both horizontal, short and very narrow leading to poor drainage and easy blockage. (More? [[Hearing_-_Middle_Ear_Development|Middle Ear]] | [[Sensory_-_Hearing_and_Balance_Development|Hearing]] | [[Sensory_-_Hearing_Abnormalities|Hearing Abnormalities]])
'''Vestibulocochlear Nerve''' - Cranial Nerve VIII


===pharynx===
'''Whirlin''' - A PDZ scaffold protein expressed in hair cells at the stereocilia tips, essential for the stereocilia elongation process. The DFNB31 gene mutations cause hearing loss in human and mouse. This protein can interact with membrane-associated guanylate kinase (MAGUK) protein, erythrocyte protein p55 (p55).


:(throat) Forms the initial segment of the upper respiratory tract divided anatomically into three regions: nasopharynx, oropharynx, and laryngopharynx (hypopharynx). Anatomically extends from the base of the skull to the level of the sixth cervical vertebra. (More? [[Respiratory System Development]])
'''Wnt7a''' - signaling through the Wnt pathway regulates the development of hair cell unidirectional stereociliary bundle orientation.

Latest revision as of 22:34, 23 January 2019

2019

ILI redevelopment.


2016

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Original movie media player format

<mediaplayer width='410' height='340' image="http://php.med.unsw.edu.au/embryology/images/7/7d/Chicken-neural-crest-migration-01.jpg">File:Chicken-neural crest migration 01.mp4</mediaplayer>

2013

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Stage16-18 face.jpg


Pharyngeal arch structure cartoon.gifStage13 pharyngeal arch excerpts.gif

Textbooks

  • Before We Are Born (5th ed.) Moore and Persaud Chapter 20: p460-479
  • Essentials of Human Embryology, Larson Chapter 12: p252-272

Online Textbooks

Search

Links

Terms

altricial animal - Term used to describe an animal born in a helpless state, with incomplete development of sensory systems at birth. For example rats and mice are born with incomplete development of visual and auditory systems.

ampulla - Term used to describe an anatomical dilation of a tube or canal lumen. Anatomical description of the opening end of the uterine tube lying above the ovary and the enlarged initial segmeny of the semicircular canals of the inner ear vestibular system. (More? [ear6.htm Inner Ear] | [genitalXXuterus.htm Genital System - Female Uterus])

aneurism - (Greek, aneurysma = a widening, aneurysm) A term used to describe an abnormal widening of a vessel or anatomical tubal structure.

aquaeductus vestibuli - see vestibular aqueduct

auditory neuropathy - (AN) abnormality of transmission of sound information to the brain.

auditory tube - (eustachian tube) between the middle ear and oral cavity, has a bony (tympanic 1/3) and cartilaginous (pharyngeal 2/3) portion. The main role is equalization of pressure and fluid drainage in the middle ear.

auricular hillock - see hillock

atresia - narrowing, usually of an anatomical tube or cavity.

autophagocytosis - (Greek, auto = self, phagy = eating, also called autophagy) a cell death mechanism that uses the cell's own lysosomes to self digest.

border cells - columnar cells within the organ of Corti on the medial portion of the basilar membrane.

canalis reuniens - (ductus reuniens, canaliculus reuniens, canalis reuniens, Hensen's canal, Hensen's duct, uniting canal, canalis reuniens of Hensen) short narrow canal connecting the cochlea duct to the saccule. (Victor Hensen, 1835-1924)

cerumen - (ear wax) produced by glands in the skin of the outer portion of the ear canal.

chondrified - the developmental differentiation of cartilage from mesenchye, an embryonic connective tissue.

cristae ampullaris - located in the ampulla of the membranous semicircular canals a region with both supporting and hair cells. The hair cell cilia are embedded in the gelatinous cupula.

claudius cells - (cells of Claudius) columnar cells with microvilli overlying the basilar membrane and extend from Hensen's cells to the spiral prominence. Barrier cells that lie external to the organ of corti in endolymph.

cochlear sac - embryonic structure, which will form the coiled cochlear duct and contribute to the saccule.

cochlear aqueduct - a bony channel containing the fibrous periotic duct. It connects the basal turn of the cochlea perilymphatic space with the subarachnoid space of the posterior cranial cavity.

cochlin - major constituent of the inner ear extracellular matrix.

collagen type II - major constituent of the inner ear extracellular matrix.

conductive loss - term used to describe one of the two major classes of hearing loss involving external and middle ear abnormalities (other form is Sensorineural loss).

connexins - channel proteins of the gap junctions that allow rapid communication between adjacent cells. The two connexins Cx26 and Cx30 are the major proteins of cochlear gap junctions.

connexin 26 - A strikingly high proportion (50%) of congenital bilateral nonsyndromic sensorineural deafness cases have been linked to mutations in the GJB2 coding for the connexin26

cupular deposits - basophilic material on the cupulae of the semicircular ducts, an postnatal ageing phenomenon seen in some vestibular labyrinth.

clinical weeks - taken from last menstrual period (LMP) and therefore approximately two weeks before fertilization occurs.

Deiters' cells

discoidin domain receptor 1 - (DDR1) a tyrosine kinase receptor activated by native collagen, expressed in the basement membrane and with fibrillar collagens. Found in basal cells of the stria vascularis, type III fibrocytes, and cells lining the basilar membrane of the organ of Corti. {Meyer zum Gottesberge, 2008 #1877}

ductus utriculosaccularis -

endochondral ossification - the process of bone formation from a pre-existing cartilage template.

endoderm -

endolymphatic fluid -

endolymphatic sac - inner ear structure that has anatomically both an intraosseous and extraosseous component. Th e sac has functions regulating endolymph that are both secretory and absorptive. Also the site of endolymphatic sac tumors either sporadical occurring or associated with the autosomal-dominant von Hippel-Lindau (VHL) disease, due to a germ line mutation.

embryological weeks - taken from the time of fertilization which typically occurs around the middle (day 14), or just after, of the typical 28 day menstrual cycle.

Emx2 - homeobox gene affecting middle ear and inner ear development.

eustachian tube - (auditory tube) A cavity linking the pharynx to the middle ear, which develops from the first pharyngeal pouch. Named after Bartolomeo Eustachi (1500 - 1574) an Italian anatomist.

external auditory meatus - (ear canal) develops from the first pharyngeal cleft.

ear wax - see cerumen.

epithelia -

espins - calcium-resistant actin-bundling proteins enriched in hair cell stereocilia and sensory cell microvilli and spiral ganglion neurons (SGNs)

eustachian tube - (auditory tube) between the middle ear and oral cavity, equalization of pressure in the middle ear.

external auditory canal -

fenestra ovalis - (oval window) separates the tympanic cavity from the vestibule of the osseous labyrinth.

fenestra rotunda - (round window) separates the tympanic cavity from the scala tympani of the cochlea.

fetus - (foetus) term used to describe human development after the 8th week (10th clinical week, LPM) and covers the developmental periods of second and third trimester.

fibroblast growth factor 1 - (Fgf-1) a growth factor released from cochlea sensory epithelium which stimulates spiral ganglion neurite branching.

fibroblast growth factor 8 - (Fgf-8) a growth factor released by inner hair cells which regulates pillar cell number, position and rate of development.

fibroblast growth factor receptor 3 - (Fgfr-3) a tyrosine kinase receptor with a role in the commitment, differentiation and position of pillar cells in the organ of corti

fundamental frequency - (natural frequency) the lowest frequency in a harmonic series, for the female voice this is about 225 Hz.

helicotrema - term used to describe the cochlear apex.

Hes - (hairy and enhancer of split) family of factors, which has been shown to be a general negative regulator of neurogenesis {Zheng, 2000 #1936}.

hillock - a small hill, used to describe the six surface elevations on pharyngeal arch one and two.

Hindbrain - Invaginate -

Incus - (anvil) auditory ossicle

inner phalangeal cells

inner pillar cells - organ of Corti cells arranged in rows and form a boundary between the single row of inner hair cells and three rows of outer hair cells. These cells have surface-associated microtubule bundles.

inner sulcus - area of the cochlear duct

interdental region -

internal auditory meatus - (internal acoustic meatus, IAM) Anatomical canal in which CN VII and CN VIII ganglia reside and pass through to the brainstem. This bony canal lies between the posterior surface of the petrous pyramid and the bony labyrinth within the dense petrous bone. Also associated clinically with the site where acoustic neuromas may occur.

Kolliker's organ - (Kollicker's organ, greater epithelial ridge) Developing cochlear structure consisting of columnar-shaped supporting cells filling the inner sulcus and lying directly under the tectorial membrane. This transient organ regresses and generates the space of the inner sulcus. Rudolph Albert von Kolliker (1817-1905)??

lateral semicircular duct - Limbus -

LMP - acronym for last menstrual period, used to clinically measure gestation.

malleus - (hammer) auditory ossicle

mastoid process - of temporal bone

Math1 - homolog of the Drosophila proneural gene atonal, necessary and sufficient for the production of hair cells in the mouse inner ear. {Chen, 2002 #1932}Negatively regulated by Hes1 and Hes5

meatal plug - temporary blockage of the external auditory meatus which forms at the end of the embryonic period and remains present until the seventh month.

meatus - anatomical opening, cavity or space (external acoustic meatus,internal auditory meatus)

Meckel's cartilage - first pharyngeal ach cartilage, located within the mandibular prominence. This cartilage first appears at stage 16, stage 20 the beginning of membranous ossification. Named after Johann Friedrich Meckel, (1781 - 1833) a German anatomist. (http://www.whonamedit.com/doctor.cfm/1840.html)

membranous labyrinth - Mesenchyme - Mesoderm - Microtia - Modiolus -

mucopolysaccharidosis - (MPS IIIB, Sanfilippo Syndrome type B) abnormality caused by a deficiency in the lysosomal enzyme N-acetyl-glucosaminidase (Naglu). Children with MPS IIIB develop abnormal hearing, and mental functioning culminating in early death.

netrin-1 - secreted growth factor, expressed in the organ of Corti and spiral ganglion cells, role in process outgrowth.

neural tube -

olivocochlear - brainstem cholinergic and GABAergic efferent system that innervates sensory cells and sensory neurons of the inner ear.

organ of Corti - organ of Corti protein II - (OCP-II) cytosolic protein or transcription factor?

otolithic membrane - extracellular matrix that cover the sensory epithelia of the inner ear.

ossicle - (small bone) the individual bone of the three middle ear bones (auditory ossicles), which reduce vibrational amplitude but increase force to drive fluid-filled inner ear.

ossify -

otic capsule -

otic cup

otic placode -

otic vesicle -

otoconin - inner ear biominerals required for vestibular apparatus function.

otogelin - (Otog) an inner ear specific glycoprotein expressed in cochlea cells at different developmental times.

otolithic membrane - a membrane within the utricle and saccule containing embedded hair cell cilia and small crystalline bodies of calcium carbonate (otoliths). Functions to detect head motion.

otoliths - small crystalline bodies of calcium carbonate found within the otolitic membrane of the utricle and saccule.

ototoxic - compound or drug causing temporary or permanent hearing loss.

outer hair cells - (OHCs) three rows of hair cells that function to increase basilar membrane motion through a local mechanical feedback process within the cochlea, the "cochlear amplifier".

outer pillar cells - arranged in rows and form a boundary between the single row of inner hair cells and three rows of outer hair cells.

paratubal musculature - muscles lying beside the auditory (Eustachian) tube. The tensor veli, palatini (TVP) and tensor tympani muscles.

perilymph - perilymphatic space - Periotic Capsule - petrous portion - of temporal bone

pejvakin gene - in humans, two missense mutations in this gene cause nonsyndromic recessive deafness (DFNB59) by affecting the function of auditory neurons.

pharyngeal archpharyngeal pouchpharyngeal membranePharynx

pillar cells - (PC) form an inner and outer row of support cells that form a boundary between inner and outer hair cells.

Placode

preyer reflex - ear flick in mouse in response to sound.

presbyacusis

prestin - a motor protein structurally similar to the anion transporter family expressed in cochlear outer hair cells.

preauricular tag - skin tags located in front of the external ear opening, are common in neonates and in most cases are normal, though in some cases are indicative of other associated abnormalities.

primordium-

protocadherin 15 - (Pcdh15) required for initial formation of stereocilia bundles and changes in the actin meshwork within hair cells. The Ames waltzer (av) mouse mutant has both auditory and vestibular abnormalities from a mutation in this gene.

Reichert's cartilage - pharyngeal ach 2 cartilage, named after Karl Bogislaus Reichert (1811 - 1883) a German anatomist.

Reissner's membrane - (vestibular membrane, vestibular wall) is a membrane located inside the cochlea separating the scala media from scala vestibuli. Named after Ernst Reissner (1824-1878) a German anatomist. “It primarily functions as a diffusion barrier, allowing nutrients to travel from the perilymph to the endolymph of the membranous labyrinth.

rhombomere -

Saccular macula -

Saccule - (Latin, sacculus = a small pouch)

sacculocollic reflex -

scala tympani - one of the three Cochlea cavities, it is filled with perilymph.

Scarpa's ganglion - (vestibular ganglion) primary afferent vestibular neuron ganglion of the vestibular nerve. Located within the internal auditory meatus.

semicircular canals - series of fluid-filled loops of the inner ear required for balance and sensing acceleration.

sensorineural - term used to describe one of the two major classes of hearing loss involving the central pathway from the cochlear (other form is conductive loss).

space of Nuel - within the cochlea, an organ of Corti space between the outer pillar cells and the phalangeal and hair cells. Named after Jean-Pierre Nuel (1847-1920) a Belgian ophthalmologist.

spiral ganglion neurons - (SGN) innervate the inner (Type I) and outer (Type II) hair cells of the cochlea.

stapedius muscle - (innervated by CN VII tympanic branch) one of the two muscles in the middle ear, contraction of this muscle pulls the stapes and dampens auditory ossicle movement.

stapes - (stirrup) a middle ear auditory ossicle (bone).stapes footplate - startle response -

stereocilia -finger-like projections from the apical surface of sensory hair cells forming the hair bundle in the cochlea. Formed by tightly cross-linked parallel actin filaments in a paracrystalline array with cell surface specializations (tip links, horizontal top connectors, and tectorial membrane attachment crowns).

stratified squamous epithelia - classification of epithelium which transiently forms a plug in external ear canal to the outer eardrum.

stria vascularis - forms the outer wall of the cochlear duct of the mammalian cochlea is composed primarily of three types of cells. Marginal cells line the lumen of the cochlear duct and are of epithelial origin. Basal cells also form a continuous layer and they may be mesodermal or derived from the neural crest. Intermediate cells are melanocyte-like cells, presumably derived from the neural crest, and are scattered between the marginal and basal cell layers. The stria forms endolymph and also contains a rich supply of blood vessels.

sulcus -

synostotically - anatomically normally separate skeletal bones fused together.

tectorial membrane - extracellular matrix that cover the sensory epithelial hair cells of the organ of corti within the cochlea.

alpha-tectorin and beta- (TECTA, TECTB) major non-collagenous protein component of the tectorial membrane forming a striated-sheet matrix. Synthesized as glycosylphosphatidylinositol-linked, membrane bound precursors.

temporal bone -

tensor tympani - (innervated by CN V mandibular nerve) one of the two muscles in the middle ear, contraction of this muscle pulls the malleus and tenses the tympanic membrane, dampening auditory ossicle movement. The muscle arises from auditory tube (cartilaginous portion) and is inserted into the malleus (manubrium near the root).

teratogens - trilaminar embryo -

tonotopy - term describing the mapping along the tectorial membrane within the cochlea of the different sound frequencies.

tympanic cavity - tympanic membrane -Utricle -Vacuolization - Vesicle - vestibular apparatus - vestibular evoked myogenic potential (VEMP) test

vestibular ganglion - (Scarpa's ganglion) primary afferent vestibular neuron ganglion of the vestibular nerve. Located within the internal auditory meatus.

vestibular membrane - (Reissner's) extends from the spiral lamina to the outer wall and divides the cochlea into an upper scala vestibuli, a lower scala tympani.

Vestibulocochlear Nerve - Cranial Nerve VIII

Whirlin - A PDZ scaffold protein expressed in hair cells at the stereocilia tips, essential for the stereocilia elongation process. The DFNB31 gene mutations cause hearing loss in human and mouse. This protein can interact with membrane-associated guanylate kinase (MAGUK) protein, erythrocyte protein p55 (p55).

Wnt7a - signaling through the Wnt pathway regulates the development of hair cell unidirectional stereociliary bundle orientation.