development of eye and ear

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    Development of eye and ear

    B.Sc. 2012

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    Embryonic development

    Week 3: Beginning development of the brain, heart, blood cells, circulatorysystem, spinal cord, and digestive system.

    Week 4: Beginning development of bones, facial structures, and limbs(presence of arm and leg buds); continuing development of the heart (whichbegins to beat), brain, and nervous tissue.

    Week 5: Beginning development of eyes, nose, kidneys, lungs; continuingdevelopment of the heart (formation of valves), brain, nervous tissue, and

    digestive tract.

    Week 6: Beginning development of hands, feet, and digits; continuingdevelopment of brain, heart, and circulation system.

    Week 7: Beginning development of hair follicles, nipples, eyelids, and sexorgans (testes or ovaries); first formation of urine in the kidneys and firstevidence of brain waves.

    Week 8: Facial features more distinct, internal organs well developed, thebrain can signal for muscles to move, heart development ends, external sexorgans begin to form.

    By the end of the embryonic stage, all essential external and internalstructures have been formed. The embryo is now referred to as a fetus

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    Development of eye

    The eyes of all vertebrates develop in a patternwhich produces an "inverted" retina, in which theinitial detection of light rays takes place at theoutermost portion.

    The eye is derived from the neural tube(neuroectoderm), from which arise the retinaproper and its associated pigment cell layer;the mesoderm of the head region, whichproduce the corneoscleral and uveal tunics;and the surface ectoderm, from which comesthe lens.

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    Eye development

    Earliest stage of eye development is the formation ofthe paired optic vesicles on either side of theforebrain.

    These growing diverticula expand laterally into themesoderm of the head and develop a stalk-likeconnection to the main portion of the rudimentarycentral nervous system.

    In humans, this process begins at about 22 days ofdevelopment; as the vesicles continue to grow, theirconnection to the brain becomes progressively

    narrower and more stalk-like. The forming stalks will eventually become the

    rudiments of the optic nerves.

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    Eye development

    As this is occurring, the surface ectoderm thickensto form a lens placode, a region visible on thesurface of the embryo. This transformation istriggered by the proximity of the optic vesicle, in atypical example of induction.

    Once the formation of the lens placode has begun,the expanding optic vesicle begins to invaginate toform a cup-shaped structure, and also to fold alongits centerline, enclosing a small amount ofangiogenic mesenchyme as it does so.

    This mesenchyme forms the hyaloid artery and vein,which supply the forming lens; and later, in the fullyformed stage will become the central artery and veinof the retina.

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    Eye development

    As the cup invaginates and folds, it is forming twodistinct layers.

    The inner layer of the optic cup will eventually formthe retinal tunic, including its light-sensitiveelements.

    The outer layer of the optic cup will form the pigmentepithelium layer, which lies outside the sensitiveportion.

    The uveal and corneoscleral tunics eventually willdifferentiate from the surrounding mesoderm.

    Meanwhile, the surface ectoderm of the lens placodehas thickened and is beginning to differentiate twodistinct areas.

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    As the optic cup from the lateral wall of the forebrain comes close to the

    overlying non-neural ectoderm, the terminal induction of the lens placode takes

    place. The lens placode invaginates to form the lens vesicle. Fiber cells form

    and the crystalline gene family is activated.

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    http://eyemakeart.files.wordpress.com/2009/06/eye-stages_in_the_evolution_of_the_eye.png
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    Eye development SUMMARY OF MAIN EVENTS

    The optic vesicles extend from the Diencephalon

    Optic vesicles come into close proximity to epithelialectoderm

    Optic vesicle thickens & folds as optic cup

    Lens placode forms from epithelial ectoderm Lens placode infolds as future lens

    TIME LINE 22 Days--Optic Groove Appears

    24 Days--Optic Vesicle

    26 Days --Optic Cup & Lens Placode

    28 Days--Further folding OC & LP

    33 Days--Sensory & Pigmented Retina

    33 & 36 Days--Lens distinct

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    Eye development

    Formation of lens

    The first is the lens vesicle, and invagination of the surface placode that will separateand form the lens proper.

    As the lens rudiment detaches and drops into position, a space forms external to itthat will become the anterior chamber of the mature eye.

    The surface ectoderm and the mesoderm beneath it differentiate into the cornea andthe eyelids.

    The process of forming the eye by inversion of the optic vesicle and formationof the optic cup is common to all of the vertebrates. It produces an eye that hasan inverted retina. The light-sensitive elements are located at the outer regions,and the neural connection to the brain must consequently come from the innerregion, perforating through the rest of the layers in its course. The enclosure ofthe hyaloid artery and vein in the stalk provides for blood supply to the retina.

    The formation of the sclera, the cornea and the uveal tunic also haveimplications for the nature of the fully-formed eye. Since mesoderm is the onlyembryonic tissue with angiogenic potential, i.e., the capacity to form bloodvessels, its participation in the formation of the uveal tunic and the sclera isnecessary.

    The cornea, although it fuses with the sclera, is derived solely from ectoderm,and it is therefore avascular in its final form. This has clinical significancebecause it isolates the cornea from the immune system, creating a "privilegedsite" suitable for transplantation..

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    Induction in eye development

    Chordamesoderm Induces Neural Tube

    Optic Cup Induces Lens

    Lens with or without the Optic CupInduces Cornea

    Other combinations induce othercomponents

    Eyes form at proper Place & Time Eye components appear in proper position

    & orientation

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    Lens Differentiation

    Lens needs to clarify to allow light to pass

    Lens needs to be proper shape to focus light

    Cells at posterior side elongate

    Cells need to multiply to fill in this space

    Cells fill with clear crystallin proteins so light can passthrough as well as be focused by the lens

    Germinative Region: Cells are actively dividing

    Region of Cell Elongation: Cells are beginning to change

    shape & Differentiate Cells differentiate into Fibre Cells

    Fibre Cells Densely Packed in Centre as Lens Nucleus

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    Pattern Generation

    migration of retinal ganglion axonsStudied in non-mammalian vertebrates

    does apply to mammals

    neurons synapse in optic tectum of midbrain

    position of ganglion cells in inner margin of

    retina determined by cadherins on cell

    membranes

    growth of axon is directed both by presence

    oflaminin and N-CAM on migratory surface

    netrin-1 made by cells at optic disc also aidmigration (cells form optic nerve at disc)

    axons growing on glial cells toward optic

    tectum oriented by N-CAM, cadherins and

    integrins

    when reaching optic chiasma decision to go

    straight or cross brain governed by ephrin

    proteins and L1 and laminin adhesionmolecules

    each axon goes to a specific site in tectum

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    Pattern Generation

    Axons distinguish between different

    regions of optic tectum BY gradient of

    transcription factors which specify cells

    along a dorsal ventral axis

    dorsal retinal cells have high levels of

    TF Tbx5 while ventral cells have high

    levels ofPax2 TF induced byparacrine factors BMP4 and retinoic

    acid from nearby tissues

    in tectum gradient ofephrin proteins

    exists highest in posterior tectum

    ephrin protein repulse axons from

    temporal retina but not nasal retina

    activity dependent synapse formation

    also important here

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    Neural crest formation

    Muscle & dermis

    Tissue-specific transcription factors act in a hierarchical and

    combinatorial network to specify different cell fates.

    Pax3 in Neural crest and

    Dermatomyotome

    expression of MITF

    depends on PAX3

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    Embryonic induction and PAX6

    expressionlink between experimental embryology and genetics

    Signal from optic cup induces placode to develop

    Pax6 initiates and orchestrates program of eye development in

    optic cup and lens placode

    Human heterozygous-aniridia

    homozygotes-complete absence of eye

    Eyeless gene in Drosophila

    DNA binding domainschanges in gene expression

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    Ear development

    Externally the pharyngeal archesare initially the most obviousexternal feature.

    Sensory placodes are present onthe surface (but not obvious).

    Otic placode lies in themesenchyme in the neck region

    The middle ear ossicles (bones)are derived from 1st and 2nd archmesenchyme.

    The space in which these bones

    sit is derived from the 1stpharyngeal pouch

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    Origin of Three Ear Parts

    inner ear otic placode then otocyst

    middle ear 1st pharyngeal pouch

    1st and 2nd arch mesenchyme

    outer ear 1st pharyngeal cleft

    6 surface hillocks

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    Origin of inner Ear

    Otocyst

    pair of surface sensory placodes (otic placodes) in the head region

    placodes fold inwards forming a depression

    pinches off entirely from the surface

    fluid-filled sac or vesicle (otic vesicle, otocyst)

    vesicle sinks into the head mesenchyme

    Inner Ear Development

    vesicle then extends and folds

    membranous labrynth cochlea

    utricle and saccule

    endolymphatic duct

    semicircular canals

    Then innervated by CN VIII

    embedded in developing temporal bone

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    Middle Ear development

    ossicles (bones)

    arch 1 - malleus, incus

    arch 2 - stapes

    muscles (mesoderm)

    arch 1 - tensor tympani

    arch 2 - stapedius

    tympanic cavity

    first pharyngeal pouchextends as tubotympanic

    recess

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    Outer Ear development

    surface hillocks (auricular hillocks)

    three on pharyngeal arch 1

    three on pharyngeal arch 2

    external auditory meatus (canal)

    1st pharyngeal cleft (groove)

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    Development Timing

    Week 3 - otic placode, otic vesicle

    Week 5 - cochlear part of otic vesicle elongates

    (humans 2.5 turns)

    Week 9 - Mesenchyme surroundingmembranous labrynth (otic capsule) chondrifies

    Week 12-16 - Capsule adjacent to membranous

    labrynth undegoes vacuolization to form a cavity(perilymphatic space) around membranous

    labrynth and fills with perilymph

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    http://embryology.med.unsw.edu.au/Notes/images/senses/hearingcartoon.jpg