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Otic vesicle, or auditory vesicle, consists of either of the two sac-like invaginations formed and subsequently closed off during embryonic development. It is part of the neural
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 ...
, which will develop into the membranous labyrinth of the inner ear. This labyrinth is a continuous
epithelium Epithelium or epithelial tissue is a thin, continuous, protective layer of cells with little extracellular matrix. An example is the epidermis, the outermost layer of the skin. Epithelial ( mesothelial) tissues line the outer surfaces of man ...
, giving rise to the
vestibular system The vestibular system, in vertebrates, is a sensory system that creates the sense of balance and spatial orientation for the purpose of coordinating motor coordination, movement with balance. Together with the cochlea, a part of the auditory sys ...
and auditory components of the inner ear. During the earlier stages of
embryogenesis An embryo ( ) is the initial stage of development for 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 ...
, the
otic placode In embryology, the otic placode is a thickening of the ectoderm on the outer surface of a developing embryo from which the ear develops. The ear, including both the vestibular system and the auditory system, develops from the otic placode beginni ...
invaginates to produce the otic cup. Thereafter, the otic cup closes off, creating the otic vesicle. Once formed, the otic vesicle will reside next to the neural tube medially, and on the lateral side will be
paraxial mesoderm Paraxial mesoderm, also known as presomitic or somitic mesoderm, is the area of mesoderm in the neurulating embryo that flanks and forms simultaneously with the neural tube. The cells of this region give rise to somites, blocks of tissue running ...
. Neural crest cells will migrate
rostral Rostral may refer to: Anatomy * Rostral (anatomical term), situated toward the oral or nasal region * Rostral bone, in ceratopsian dinosaurs * Rostral organ, of certain fish * Rostral scale The rostral scale, or rostral, in snakes and other sca ...
and
caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into th ...
to the placode. The general sequence in formation of the otic vesicle is relatively conserved across
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s, although there is much variation in timing and stages. Patterning during morphogenesis into the distinctive inner ear structures is determined by
homeobox A homeobox is a Nucleic acid sequence, DNA sequence, around 180 base pairs long, that regulates large-scale anatomical features in the early stages of embryonic development. Mutations in a homeobox may change large-scale anatomical features of ...
transcription factors In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The fun ...
including
PAX2 Paired box gene 2, also known as Pax-2, is a protein which in humans is encoded by the ''PAX2'' gene. Function The Pax Genes, or Paired-Box Containing Genes, play important roles in the development and proliferation of multiple cell lines, de ...
,
DLX5 Homeobox protein DLX-5 is a protein that in humans is encoded by the distal-less homeobox 5 gene, or ''DLX5'' gene. DLX5 is a member of the DLX gene family. Function This gene encodes a member of a homeobox transcription factor gene family si ...
and
DLX6 Homeobox protein DLX-6 is a protein that in humans is encoded by the ''DLX6'' gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is tr ...
, with the former specifying for ventral otic vesicle derived auditory structures and the latter two specifying for dorsal vestibular structures.


Development

The otic vesicle is derived from the cranial placode. The early otic vesicle is characterized as having broad competence and can be subdivided into sensory, non-sensory, and neurogenic components. Sensory epithelial cells and neurons are derived from the proneurosensory domain. This domain can be further sub-categorized into the neurogenic sub-domain and prosensory sub-domain. Prosensory sub-domain eventually gives rise to the support cells and hair cells while the neurogenic sub-domain gives rise to the auditory neuron and vestibular neuron. The middle part of the otic vesicle develops into the ductus and
saccus endolymphaticus From the posterior wall of the saccule a canal, the endolymphatic duct, is given off; this duct is joined by the utriculosaccular duct, and then passes along the vestibular aqueduct and ends in a blind pouch, the endolymphatic sac, on the posterio ...
. The anterior end of the otic vesicle gradually elongates as a tube and coils upon itself forming the beginnings of the
cochlear duct Cochlear, the adjective form of cochlea, may refer to: * Cochlear implant, a sensory aid for the deaf * Cochlear nuclei, the ventral cochlear nucleus and the dorsal (or lateral) cochlear nucleus * Vestibular-cochlear or Vestibulocochlear nerve, ...
. The vestibular extremity subsequently constricts to form the canalis reuniens. The central part of the otic vesicle represents the membranous vestibule, and is subdivided by a constriction into a smaller ventral part, the
saccule The saccule (Latin: sacculus) is a bed of sensory cells in the inner ear that detects linear acceleration and head tilting in the vertical plane, and converts these vibrations into electrical impulses to be interpreted by the brain. When the he ...
, and a larger dorsal and posterior part, the utricle. The dorsal component of the inner ear also consists of what will become the
semicircular canals The semicircular canals are three semicircular interconnected tubes located in the innermost part of each ear, the inner ear. The three canals are the lateral, anterior and posterior semicircular canals. They are the part of the bony labyrinth, ...
. The utricle and saccule communicate with each other by means of a Y-shaped canal.


Gene signaling

The FGF, Bmp, Wnt and
Pax genes In evolutionary developmental biology, Paired box (Pax) genes are a family of genes coding for tissue specific transcription factors containing an N-terminal paired domain and usually a partial, or in the case of four family members (PAX3, PAX4, P ...
are likely to be involved in otic induction. FGF and BMP signals help control patterning in the early otic vesicle. Fgf3 and Fgf10 are suggested to play a role in otic induction in mice, as were Msx genes suggested to play a role in otic vesicle formation in chicks. Pax8 is expressed during the entirety of otic vesicle formation. Other genes found in the otic vesicle across species that may play a role in patterning include Hmx, Fox, Dlx, and Gbx genes.


Other animals

Formation of the otic vesicle has been studied extensively in developmental
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
s including
chicken The chicken (''Gallus gallus domesticus'') is a domesticated subspecies of the red junglefowl (''Gallus gallus''), originally native to Southeast Asia. It was first domesticated around 8,000 years ago and is now one of the most common and w ...
,
Xenopus ''Xenopus'' () (Gk., ξενος, ''xenos'' = strange, πους, ''pous'' = foot, commonly known as the clawed frog) is a genus of highly aquatic frogs native to sub-Saharan Africa. Twenty species are currently described with ...
,
zebrafish The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...
,
axolotl The axolotl (; from ) (''Ambystoma mexicanum'') is a neoteny, paedomorphic salamander, one that Sexual maturity, matures without undergoing metamorphosis into the terrestrial adult form; adults remain Aquatic animal, fully aquatic with obvio ...
, and
mouse A mouse (: mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus'' ...
. The transition from the otic placode to the otic vesicle occurs during the 19th
somite The somites (outdated term: primitive segments) are a set of bilaterally paired blocks of paraxial mesoderm that form in the embryogenesis, embryonic stage of somitogenesis, along the head-to-tail axis in segmentation (biology), segmented animals. ...
stage in zebrafish, Xenopus, and the chick. In the chick, invagination of the otic placode occurs passively due to the movements of the surrounding placode. The otic placode in zebrafish, on the other hand, occurs by cavitation; the ectodermal placode condenses and forms an ovoid ball directly below the embryo surface. Otic vesicle formation occurs later, during the 25-30 somite stage in mice.


Additional images

File:Gray900.png, Lateral views of membranous labyrinth and acoustic complex. X 25 dia. File:Gray901.png, Median views of membranous labyrinth and acoustic complex in human embryos. X 25 dia.


References

{{Authority control Embryology of nervous system