Vision - Cornea Development: Difference between revisions

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==Introduction==
==Introduction==
[[File:Historic_retina_drawing.jpg|right|300px]]
[[File:Mouse eye neural crest cornea 01.jpg||thumb|300px|alt=Cornea structure]]
These notes introduce the development of the eye cornea.
[[File:Stage_22_image_208.jpg|thumb|300px|alt=Human embryonic cornea|Human embryonic cornea ([[Week 8]], [[Carnegie stage 22]])]]
[[File:Stage_22_image_212.jpg|thumb|300px|alt=Human embryonic cornea|Human embryonic cornea detail ([[Week 8]], [[Carnegie stage 22]])]]
[[File:Gray0883.jpg|thumb|300px|alt=Section through front of Eyeball|Section through human cornea]]
These notes introduce the development of the cornea of the eye. The adult cornea has three layers: an outer epithelium layer (ectoderm), a middle stromal layer of collagen-rich extracellular matrix between stromal keratocytes (neural crest) and an inner layer of endothelial cells (neural crest).  


The cornea is a vision-specific specialised sensory epithelia that in humans differentiates mainly in the postnatal period. It arises initially from cranial ectoderm adjacent to the lens placode and forms a presumptive corneal epithelium. Later neural crest cells migrate between the lens and presumptive structure to form both the corneal endothelium and the stromal fibroblasts (keratocytes). Neural crest development in humans, reptiles and birds differs from that seen in rodents, cats, rabbits, and cattle.


{{Vision Links}}




{{Senses Links}}
{{Vision Links}}


:'''Links:''' [[:Category:Cornea|Category:Cornea]] | [[Neural Crest Development]] | [[Integumentary Development]]
== Some Recent Findings ==
== Some Recent Findings ==
[[File:Human-retina-01.jpg|thumb|300px|Adult Human Retina histology<ref><pubmed>12186651</pubmed>| [http://genomebiology.com/content/3/8/REVIEWS1022 Genome Biol.]
 
</ref>]]
{|
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* '''Activation of c-Jun N-Terminal Kinase (JNK) during Mitosis in Retinal Progenitor Cells'''<ref><pubmed>22496813</pubmed></ref> "Most studies of c-Jun N-terminal Kinase (JNK) activation in retinal tissue were done in the context of neurodegeneration. In this study, we investigated the behavior of JNK during mitosis of progenitor cells in the retina of newborn rats. ... The data show, for the first time, that JNK is activated in mitotic progenitor cells of developing retinal tissue, suggesting a new role of JNK in the control of progenitor cell proliferation in the retina."
* '''Review - Corneal Development Different Cells from a Common Progenitor'''{{#pmid:26310148|PMID26310148}} "Development of the vertebrate cornea is a multistep process that involves cellular interactions between various ectodermal-derived tissues. Bilateral interactions between the neural ectoderm-derived optic vesicles and the cranial ectoderm give rise to the presumptive corneal epithelium and other epithelia of the ocular surface. Interactions between the neural tube and the adjacent ectoderm give rise to the neural crest cells, a highly migratory and multipotent cell population. Neural crest cells migrate between the lens and presumptive corneal epithelium to form the corneal endothelium and the stromal keratocytes. The sensory nerves that abundantly innervate the corneal stroma and epithelium originate from the neural crest- and ectodermal placode-derived trigeminal ganglion."
* '''Rearrangement of retinogeniculate projection patterns after eye-specific segregation in mice'''<ref><pubmed>20544023</pubmed>| [http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0011001 PLoS ONE]</ref> "When monocular enucleation was performed after eye-specific segregation, rearrangement of retinogeniculate axons in the dorsal lateral geniculate nucleus (dLGN) was observed within 5 days. ...We also examined the critical period for this rearrangement and found that the rearrangement became almost absent by the beginning of the critical period for ocular dominance plasticity in the primary visual cortex."
 
* '''The long noncoding RNA RNCR2 directs mouse retinal cell specification'''<ref><pubmed>20459797</pubmed></ref>"We find that the RNCR2 is selectively expressed in a subset of both mitotic progenitors and postmitotic retinal precursor cells. ShRNA-mediated knockdown of RNCR2 results in an increase of both amacrine cells and Müller glia, indicating a role for this lncRNA in regulating retinal cell fate specification."
* '''Bovine cornea extracellular matrix structure'''{{#pmid:25019467|PMID25019467}} "Electron microscopy and X-ray fibre diffraction were used to ascertain collagen fibril architecture. The bovine cornea was 1021±5.42μm thick at its outer periphery, defined as 9-12mm from the corneal centre, compared to 844±8.10μm at the centre. The outer periphery of the cornea was marginally, but not significantly, more hydrated than the centre (H=4.3 vs. H=3.7), and was more abundant in hydroxyproline (0.12 vs. 0.06mg/mg dry weight of cornea). DMMB assays indicated no change in the total amount of sulphated GAG across the cornea. Immunohistochemistry revealed the presence of both high- and low-sulphated epitopes of KS, as well as DS, throughout the cornea, and CS only in the peripheral cornea before the limbus. Quantification by ELISA, disclosed that although both high- and low-sulphated KS remained constant throughout stromal depth at different radial positions, high-sulphated epitopes remained constant from the corneal centre to outer-periphery, whereas low-sulphated epitopes increased significantly.
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{| class="wikitable mw-collapsible mw-collapsed"
{| class="wikitable mw-collapsible mw-collapsed"
! More recent papers
! More recent papers &nbsp;
|-
|-
| [[File:Mark_Hill.jpg|90px|left]] {{Most_Recent_Refs}}
| [[File:Mark_Hill.jpg|90px|left]] {{Most_Recent_Refs}}


Search term: ''Vision Embryology''
Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Cornea+Development ''Cornea Development'']
 
<pubmed limit=5>Cornea Development</pubmed>
 
Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Cornea+Embryology ''Cornea Embryology'']


<pubmed limit=5>Vision Embryology</pubmed>
<pubmed limit=5>Cornea Embryology</pubmed>
|}
|}
==Timeline==
==Carnegie Stages - Eye==
{{Eye Timeline table}}
 
==Human Cornea==
 
===Week 8 Stage 22===
The images below link to virtual slides of the human developing eye at [[Carnegie stage 22]]. Click on the image to open or select specific regions from the regions of interest links.
 
{|
{|
|  
| {{SlideStage22-08}}
Embryonic Development
| {{SlideStage22-08-eye}}
* Weeks 3 - 4 Eye Fields-Optic Vesicle
|
* Weeks 5 - 6 Optic Cup, Lens Vesicle, Choroid Fissure, Hyaloid Artery
====Virtual Slide - Regions of Interest====
* Weeks 7 - 8 Cornea, Anterior Chamber, Pupillary Membrane, Lens, Retina
* [http://embryology.med.unsw.edu.au/embryology/Slides/Embryo_Stages/Stage22/08-eye/Stage22-08-eye.html?zoom=3&lat=-3544&lon=4688&layers=B Eye Overview]
* Weeks 9 - 15 Iris, Ciliary Body
* [http://embryology.med.unsw.edu.au/embryology/Slides/Embryo_Stages/Stage22/08-eye/Stage22-08-eye.html?zoom=4&lat=-2030&lon=2572&layers=B Lens and Cornea]
* Weeks 8 - 10 Eyelids
* [http://embryology.med.unsw.edu.au/embryology/Slides/Embryo_Stages/Stage22/08-eye/Stage22-08-eye.html?zoom=5&lat=-1208&lon=2473&layers=B Cornea and Anterior Chamber]
| [[File:Eye_and_retina_cartoon.jpg|600px]]
* [http://embryology.med.unsw.edu.au/embryology/Slides/Embryo_Stages/Stage22/08-eye/Stage22-08-eye.html?zoom=6&lat=-2410&lon=1843&layers=B Iris and Lens]
|-
|}
 
'''Links:''' [[Embryo Virtual Slides]]
 
==Cornea Epithelia==
{|
| The cornea ocular surface is composed of three epithelia, conjunctival, limbal and corneal.
 
* Limbal stem cells are located in the palisades of Vogt, the transitional zone between the cornea and the conjunctiva.
* Limbal stem cells are close to blood vessels.
* They generate transient amplifying cells that terminally differentiate after a discrete number of cell divisions to corneal epithelial cells and undergo both centripetal migration and vertical migration.
 
| [[File:Corneal_epithelial_cells_01.jpg|alt=Corneal Epithelial Cells cartoon|300px]]
 
Corneal epithelial cells cartoon{{#pmid:19668514|PMID19668514}}
|-
| The Adult Human Limbal Palisades of Vogt
 
* '''A''' - Palisades of Vogt (arrow) are readily recognized in the human limbus.
* '''B''' - Such a unique pigmented structure can be identified on the flat mount preparation of Dispase-isolated human limbal epithelial sheets.
* '''C''' - In donors with a darker skin, these palisades of Vogt are pigmented (arrow).
* '''D''' - Under higher magnification, these limbal areas show undulated epithelial papillae (stars).
* '''E''' - Hematoxyline staining highlights higher stratification and more undulation of the limbal epithelium, and the underlying limbal stroma has high cellularity and vascularity (arrow shows blood vessel).
 
Bar represents 500 μm in A and B, 200 μm in C and E, and 50 μm in D
 
 
| [[File:Limbal palisades of Vogt PMID17211449.jpg|alt=Adult human limbal palisades of Vogt|300px]]
 
Adult human limbal palisades of Vogt{{#pmid:17211449|PMID17211449}}
|}
|}


===Carnegie Stages - Eye===
===Limbal Stem Cells===
The following data is from a study of human embryonic carnegie stages<ref><pubmed>7364662</pubmed></ref> and other sources.
[[File:Limbal_stem_cell_niche_cartoon_PMID17211449.jpg|500px]]
*  [[Carnegie_stage_10|Stage 10]] - optic primordia appear.
 
*  [[Carnegie_stage_11|Stage 11]] - right and left optic primordia meet at the optic chiasma forming a U-shaped rim.
Cartoon showing the location of limbal stem cells at the limbal basal layer.{{#pmid:17211449|PMID17211449}}
*  [[Carnegie_stage_12|Stage 12]] - optic neural crest reaches its maximum extent and the optic vesicle becomes covered by a complete sheath,
 
* [[Carnegie_stage_13|Stage 13]] - By the end of the fourth week the optic vesicle lies close to the surface ectoderm. Optic evagination differentiation allows identification of optic part of retina, future pigmented layer of retina, and optic stalk. The surface ectoderm overlying the optic vesicle, in response to this contact, has thickened to form the lense placode.
* [[Carnegie_stage_14|Stage 14]] - (about 32 days) the lens placode is indented by the lens pit, cup-shaped and still communicates with the surface by a narrowing pore.
* [[Carnegie_stage_15|Stage 15]] - (about 33 days) the lens pit is closed. The lens vesicle and optic cup lie close to the surface ectoderm and appear to press against the surface.
* [[Carnegie_stage_16|Stage 16]] - (37 days) Growth of the lens body results in a D-shaped lens cavity. Perilental blood vessels (tunica vasculosa lentis) are visible. Prior to the development of the eyelids, one small sulcus or groove forms above the eye (eyelid groove) and another below it.
* [[Carnegie_stage_17|Stages 17 - 19]] - Retinal pigment is visible  and the retinal fissure is largely closed. Eyelids grooves deepen, eyelid folds develop, first below, and then above, the eye.
* [[Carnegie_stage_18|Stages 18]] - Mesenchyme invades the region between the lens epithelium and the surface ectoderm.
* [[Carnegie_stage_19|Stages 19 - 22]] - the eyelid folds develop into the eyelids and cover more of the eye as the palpebral fissure takes shape. The upper and the lower eyelids meet at the outer canthus in Stage 19.
* [[Carnegie_stage_20|Stage 20]] - The lens cavity is lost and a lens suture begins to form. The inner canthus is established.
* [[Carnegie_stage_23|Stage 23]] - The retina comprises the pigmented layer, external limiting membrane, proliferative zone, external neuroblastic layer, transient fiber layer, internal neuroblastic layer, nerve fiber layer, and internal limiting membrane. Eyelids closure is complete (Note - shown as still open in the Kyoto embryo).


==Lens==
:'''Links:''' [[Stem Cells]]
[[File:Stage_22_image_155.jpg|thumb|Human Lens (stage 22)]]
The lens or crystalline lens or aquula (Latin, aquula = a little stream) has a key role in focussing light (with the cornea) upon the neural retina. The lens embryonic origin is from surface ectoderm of the sensory placodes that form in the head region (More? Week 4 - Placodes). The lens focusses by refracting light as it passes through the biconvex lens, which can be altered in shape (accommodation) by surrounding ciliary muscles. These ciliary muscles are activated (contracted) by parasympathetic innervation from the ciliary ganglion itself innervated by the oculomotor nerve (Cranial Nerve III) (More? Cranial Nerves).


surface ectoderm -> lens placode -> lens pit -> lens vesicle -> lens fibres -> lens capsule and embryonic/fetal nucleus.
==Descemet Membrane==


Corneal endothelium basement membrane beginning in children at 3 μm thick and increases in adults to 10 μm. Consists of collagen type IV and VIII fibrils.


:'''Links:''' [[Vision - Lens Development]]
Composed of two layers:
# anterior banded layer - commencing in week 10 ({{GA}} week 12) as collagen lamellae and proteoglycans.
# posterior non-banded layer - deposited by endothelial cells over time and thickens postnatally over decades.


==Retinotopic Map==


This neuroscience term describes how the developing retina is precisely "mapped" onto the visual cortex through a series of signaling and activity dependent mechanisms. This follows from Hubel and Wiesel (1981 Nobel Prize in Physiology or Medicine) key discoveries (1959-70) of how in development system matching occurs in the visual system. The topographic map establishes an ordered neuronal connection between sensory structures and the central nervous system.
Descemet membrane was historically named after Jean Descemet (1732–1810) a French physician.


The retinotectal map (eye to brain) of birds (lower vertebrates):
==Palisades of Vogt==


* temporal (posterior) retina is connected to the rostral (anterior) part of the contralateral optic tectum
The palisades of Vogt are a series of radially oriented fibrovascular ridges concentrated along the upper and lower corneoscleral limbus, the vasculature component consists of radially oriented hairpin loops of narrow arterial and venous vessels. Named by Vogt in 1921. (for review see{{#pmid:7182957|PMID7182957}})
* nasal (anterior) retina to the caudal (posterior) tectum
* ventral retina to the dorsal (medial) tectum
* dorsal ventral (lateral) tectum


Retinal waves a form of coordinated spontaneous activity that occurs in the developing retina. These waves of electrical activity (action potentials) are thought to have a role in establishing the initial retinotopic map by correlating/coordinating the activity of neighbouring retinal ganglion cells.


EphA/ephrin-A molecular signaling also thought to have a role in establishing the initial retinotopic map.
Aggregate into distinct crescentic zones and lie peripheral to the terminal capillary loops of the limbus and central to Schlemm’s canal. Lying between the connective tissue palisades are intervening radial zones of thickened conjunctival epithelium, the so-called inter-palisades or epithelial rete ridges.


==Neural Crest==


==Mouse Cornea==
{|
{|
| [[File:Mouse eye neural crest.jpg|400px]]
| [[File:Mouse eye neural crest.jpg|alt=histology Mouse eye neural crest|400px]]
 
Neural crest-derived cells contribute to mouse cornea development.{{#pmid:16403239|PMID16403239}}
|
* '''a''' Toluidine blue staining of an adult eye. The boxed areas correspond to b and c
* '''b''' A detailed view of the corneal assembly, including outer epithelium, stroma, and inner endothelium
* '''c''' The chamber angle at the irido-corneal transition which includes the trabecular meshwork (tm).
* '''d-j''' In vivo fate mapping of NC-derived, β-galactosidase (βGal)-expressing cells (blue)
* '''d''' The NC origin of corneal keratocytes (arrows) and of corneal endothelium (arrowhead).
* '''e''' Structures of the chamber angle, including the trabecular meshwork are seen to be NC-derived.
* '''f''' At E10, the optic cup is surrounded by NC-derived cells expressing βGal.
* '''g-i''' The majority of the cells in the periocular mesenchyme (arrows), which forms the anterior eye segment, are of NC origin, as assessed from E11.5 to E13.5.
* '''j''' The primary vitreous at E13.5 (arrowheads) shows a strong NC contribution.
|}
<gallery>
File:Mouse cornea E12.5.jpg|Mouse E12.5
File:Mouse cornea E13.5.jpg|Mouse E13.5
File:Mouse cornea E16.5.jpg|Mouse E16.5
File:Mouse cornea P0.jpg|Mouse P0
</gallery>
 
==Frog Cornea==
 
This developmental timeline is from a recent frog (Xenopus laevis) cornea study{{#pmid:23896054|PMID23896054}}


Mouse eye neural crest<ref name="PMID16403239"><pubmed>16403239</pubmed>| [http://jbiol.com/content/4/3/11 J Biol.]</ref>
* '''stage 25''' - cornea starts from a simple embryonic epidermis overlying the developing optic vesicle.
| [[File:Mouse_eye_TGF-beta_model.jpg|400px]]
* '''stage 30''' - detachment of the lens placode, cranial neural crest cells start to invade the space between the lens and the embryonic epidermis to construct the corneal endothelium.
* '''stage 41''' - a second wave of migratory cells containing presumptive keratocytes invades the matrix leading to the formation of inner cornea and outer cornea. A unique cell mass (stroma attracting center) connects the two layers like the center pole of a tent.
* '''stage 48''' - many secondary stromal keratocytes individually migrate to the center and form the stroma layer.
* '''stage 60''' - the stroma space is filled by collagen lamellae and keratocytes, and the stroma attracting center disappears. At early metamorphosis, the embryonic epithelium gradually changes to the adult corneal epithelium, which is covered by microvilli.  
* '''stage 62''' - the embryonic epithelium thickens and cell death is observed in the epithelium, coinciding with eyelid opening.  
* '''After metamorphosis''' - cornea has attained the adult structure of three cellular layers, epithelium, stroma, and endothelium, and between the cellular layers lie two acellular layers (Bowman's layer and Descemet's membrane)


Mouse eye TGF-beta model<ref name="PMID16403239"><pubmed>16403239</pubmed>| [http://jbiol.com/content/4/3/11 J Biol.]</ref>
[[File:Xenopus_cornea_development_timeline.jpg|800px||alt=Xenopus cornea development timeline]]


|}


:'''Links:''' [[:File:Mouse eye neural crest.jpg|Image - Mouse eye neural crest]] | [[:File:Mouse_eye_TGF-beta_model.jpg|Image - Mouse eye TGF-beta model]] | [[Sensory - Vision Development|Vision Development]] | [[Neural Crest Development]] | [[Head Development]]
:'''Links:''' [[Frog Development]]  
==Extraocular Muscles==
 
Extraocular muscles are required to move the eye within the orbit. Their embryonic origin requires an interaction between the cranial mesoderm and the migrating neural crest cells.
==Molecular==
 
[[File:Mouse eye TGF-beta model.jpg|600px]]


The following is from a recent paper comparing human to zebrafish muscle development.<ref><pubmed>22132088</pubmed>| [http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0027095 PLoS One.]</ref>
Mouse Eye TGF-beta Model - Summary of the TGFβ-dependent development of anterior and posterior ocular structures.{{#pmid:16403239|PMID16403239}}
{|
{|
! About the Muscles
| valign=top|'''a''' Neural crest-derived cells (NC, blue) contribute to structures of the anterior eye segment and the primary vitreous (PV).  
! Legend
* TGFβ signaling is involved in the formation of the ciliary body (CB) and the trabecular meshwork (TM), and in control of PV growth.  
|-
* Moreover, normal PV development and/or TGFβ signaling are important for correct retinal patterning.  
| valign=top width=400px valign=top|
| valign=top|'''b''' In the cornea, prospective stromal keratocytes and endothelial cells are of neural crest origin.
* Five of the six muscles (inferior rectus, superior rectus, lateral rectus, medial rectus, and superior oblique) originate at a common tendinous ring of fibrous tissue (the Annulus of Zinn).
* Here, TGFβ signaling is needed for the expression of the transcription factors Foxc1 and Pitx2 and for normal differentiation of NC-derived cells into collagen-synthesizing stromal keratocytes.
** The Annulus of Zinn surrounds the optic nerve, ophthalmic artery, and ophthalmic vein at their entrance through the apex of the orbit.  
* Moreover, in forming corneal endothelial cells (and in the TM), expression of Foxc1 and cell survival requires TGFβ signalling.
* The sixth muscle (inferior oblique) has a separate origin point on the orbital side of the bony maxilla at the anterior inferomedial strut.
| valign=top width=200px|
* '''IR''' - inferior rectus
* '''SR''' - superior rectus
* '''LR''' - lateral rectus
* '''MR''' - medial rectus
* '''SO''' - superior oblique
* '''IO''' - inferior oblique
| valign=top| [[File:Human_extraocular_muscles_01.jpg|200px]]
|}
|}


==Additional Images==
==Additional Images==
<gallery>
<gallery>
File:Stage 22 image 212.jpg|Human stage 22 developing iris region
File:Stage 22 image 211.jpg|Human stage 22 developing iris region
File:Stage 22 image 209.jpg|Human stage 22 overview of optic nerve
File:Stage 22 image 208.jpg|Human stage 22 overview of eye
File:Stage 22 image 207.jpg|Human stage 22 lens and hyaloid vessels
File:Stage 22 image 206.jpg|Human stage 22 optic nerve (stalk)
File:Stage 22 image 154.jpg|Human stage 22 retina
File:Mouse-optic nerve axons.jpg|Mouse adult optic nerve axons
File:Pax6 eye phenotypes.jpg|Pax6 eye phenotypes
</gallery>
</gallery>


===Historic Images===
===Historic Images===
<gallery>
<gallery>
File:Bailey456.jpg|Fig. 456. Location of optic areas before the closure of the neural groove.
File:Foster052.jpg|Fig. 52. Eye of a Fowl on the day 8
File:Bailey457.jpg|Fig. 457. Location of areas shown in Fig. 456 after the formation of the neural canal.
File:Foster128.jpg|Fig. 128. Eye of a Rabbit Embryo 12 Days
File:Bailey458-459.jpg|Fig. 458. Location of the optic area after the beginning of the formation of the optic cup and optic stalk. Fig. 459. Dorsal view of head of chick of 58 hours' incubation.
File:Kollmann713.jpg|
File:Bailey460.jpg|Fig. 460. Section through head of chick of two days' incubation.
File:Kollmann715.jpg|
File:Bailey461.jpg|Fig. 461. Section through head of chick of three days' incubation.
File:Kollmann717.jpg|
File:Bailey462.jpg|Fig. 462. Later stage in development of optic cup and lens than is shown in Fig. 461.
File:Bailey463.jpg|Fig. 463. Developing lens and optic cup.
File:Bailey463.jpg|Fig. 463. Developing lens and optic cup.
File:Bailey464.jpg|Fig. 464. Model showing lens and formation of optic cup.
File:Bailey464.jpg|Fig. 464. Model showing lens and formation of optic cup.
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File:Bailey466.jpg|Fig. 466. Section through optic cup and lens invagination of chick of fifty-four hours' incubation.
File:Bailey466.jpg|Fig. 466. Section through optic cup and lens invagination of chick of fifty-four hours' incubation.
File:Bailey467.jpg|Fig. 467. Section through eye of human embryo of 13-14 weeks.
File:Bailey467.jpg|Fig. 467. Section through eye of human embryo of 13-14 weeks.
File:Bailey468.jpg|Fig. 468. Development of the retinal cells.
File:Bailey469.jpg|Fig. 469. Vertical section through retina of a four months' human embryo.
File:Bailey470.jpg|Fig. 470. Vertical section through retina of a five and one-half months' human embryo.
File:Brown001.jpg|Fig. 1. Section through head of pig, 2 mm long.
File:Brown001.jpg|Fig. 1. Section through head of pig, 2 mm long.
File:Brown002.jpg|Fig. 2. Section through head of chick, 2 mm long.
File:Brown002.jpg|Fig. 2. Section through head of chick, 2 mm long.
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File:Brown009.jpg|Fig. 9. Section through head of pig, 9 mm long.
File:Brown009.jpg|Fig. 9. Section through head of pig, 9 mm long.
File:Brown010.jpg
File:Brown010.jpg
File:Brown011.jpg|Fig. 11. colobomba of the fundus in the adult and means a lack of development.
</gallery>
</gallery>


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<references/>
<references/>


===Online Textbooks===
===Journals===
* Kolb H, Fernandez E, Nelson R, editors. '''Webvision: The Organization of the Retina and Visual System''' [Internet]. Salt Lake City (UT): University of Utah Health Sciences Center; 1995-. Available from: http://www.ncbi.nlm.nih.gov/books/NBK11530/
* '''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]


* '''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]
* [http://journals.lww.com/corneajrnl/pages/default.aspx Cornea] "For corneal specialists and for all general ophthalmologists with an interest in this exciting subspecialty, Cornea brings together the latest clinical and basic research on the cornea and the anterior segment of the eye." [http://www.ncbi.nlm.nih.gov/pubmed?term=%22Cornea%22[jour] PuMed Listing]
===Reviews===
{{#pmid:26310148}}


* '''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]
{{#pmid:23819758}}


* '''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]
{{#pmid:20599432}}


* '''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]
{{#pmid:19343693}}


* '''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]
* [http://www.ncbi.nlm.nih.gov/books/NBK11530 Webvision: The Organization of the Retina and Visual System]. Kolb H, Fernandez E, Nelson R, editors. Salt Lake City (UT): University of Utah Health Sciences Center; 1995-.
===Reviews===
<pubmed>20855501</pubmed>|  [http://jcb.rupress.org/content/190/6/953.full JCB]


The International Journal of Developmental Biology [http://www.ijdb.ehu.es/web/contents.php?vol=48&issue=8-9 Vol. 48 Nos. 8/9 (2004) Eye Development]
The International Journal of Developmental Biology [http://www.ijdb.ehu.es/web/contents.php?vol=48&issue=8-9 Vol. 48 Nos. 8/9 (2004) Eye Development]


===Articles===
===Articles===
<pubmed>19541779</pubmed>
{{#pmid:13429485}}




 
'''Bookshelf'''  [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=cornea%20development cornea development]
'''Bookshelf'''  [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=vision%20development vision development]


===Search Pubmed===
===Search Pubmed===




'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/pubmed?term=vision%20development vision development] | [http://www.ncbi.nlm.nih.gov/pubmed?term=eye%20development eye development] | [http://www.ncbi.nlm.nih.gov/pubmed?term=eye%20embryology eye embryology] | [http://www.ncbi.nlm.nih.gov/pubmed?term=retina%20embryology retina embryology] | [http://www.ncbi.nlm.nih.gov/pubmed?term=lens%20embryology lens embryology]
'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/pubmed?term=cornea%20development cornea development]




'''Search Entrez:''' [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=vision%20development vision development] | [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=eye%20development eye development] | [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=eye%20embryology eye embryology] | [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=retina%20embryology retina embryology] | [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=lens%20embryology lens embryology]
'''Search Entrez:''' [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=cornea%20development cornea development]  


==Terms==
==Terms==
* '''Limbal epithelial stem cells''' - cells located at the limbal basal layer.
* '''palisades of Vogt''' - series of radially oriented fibrovascular ridges concentrated along the upper and lower corneoscleral limbus, the vasculature component consists of radially oriented hairpin loops of narrow arterial and venous vessels. Named by Vogt in 1921. [https://www.ncbi.nlm.nih.gov/pubmed/7182957 PMID 7182957]


.
==External Links==
==External Links==


{{External Links}}
{{External Links}}


* UNSW SoMS research - [https://medicalsciences.med.unsw.edu.au/research/groups/mechanisms-disease-and-translational-research MDTR]
* '''UNSW Virtual Slides''' [http://vslides.unsw.edu.au/VirtualSlideV2.nsf/id/D0C49C Eye Development Histology] (requires login)
* '''UNSW Virtual Slides''' [http://vslides.unsw.edu.au/VirtualSlideV2.nsf/id/D0C49C Eye Development Histology] (requires login)




{{Glossary}}
{{Glossary}}


{{Footer}}
{{Footer}}


[[Category:Vision]]
[[Category:Vision]][[Category:Cornea]][[Category:Neural Crest]]

Latest revision as of 10:34, 9 August 2018

Embryology - 25 May 2024    Facebook link Pinterest link Twitter link  Expand to Translate  
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Introduction

Cornea structure
Human embryonic cornea
Human embryonic cornea (Week 8, Carnegie stage 22)
Human embryonic cornea
Human embryonic cornea detail (Week 8, Carnegie stage 22)
Section through front of Eyeball
Section through human cornea

These notes introduce the development of the cornea of the eye. The adult cornea has three layers: an outer epithelium layer (ectoderm), a middle stromal layer of collagen-rich extracellular matrix between stromal keratocytes (neural crest) and an inner layer of endothelial cells (neural crest).

The cornea is a vision-specific specialised sensory epithelia that in humans differentiates mainly in the postnatal period. It arises initially from cranial ectoderm adjacent to the lens placode and forms a presumptive corneal epithelium. Later neural crest cells migrate between the lens and presumptive structure to form both the corneal endothelium and the stromal fibroblasts (keratocytes). Neural crest development in humans, reptiles and birds differs from that seen in rodents, cats, rabbits, and cattle.


Vision Links: vision | lens | retina | placode | extraocular muscle | cornea | eyelid | lacrima gland | vision abnormalities | Student project 1 | Student project 2 | Category:Vision | sensory
Historic Embryology - Vision 
Historic Embryology: 1906 Eye Embryology | 1907 Development Atlas | 1912 Eye Development | 1912 Nasolacrimal Duct | 1917 Extraocular Muscle | 1918 Grays Anatomy | 1921 Eye Development | 1922 Optic Primordia | 1925 Eyeball and optic nerve | 1925 Iris | 1927 Oculomotor | 1928 Human Retina | 1928 Retina | 1928 Hyaloid Canal | Historic Disclaimer
Links: Category:Cornea | Neural Crest Development | Integumentary Development

Some Recent Findings

  • Review - Corneal Development Different Cells from a Common Progenitor[1] "Development of the vertebrate cornea is a multistep process that involves cellular interactions between various ectodermal-derived tissues. Bilateral interactions between the neural ectoderm-derived optic vesicles and the cranial ectoderm give rise to the presumptive corneal epithelium and other epithelia of the ocular surface. Interactions between the neural tube and the adjacent ectoderm give rise to the neural crest cells, a highly migratory and multipotent cell population. Neural crest cells migrate between the lens and presumptive corneal epithelium to form the corneal endothelium and the stromal keratocytes. The sensory nerves that abundantly innervate the corneal stroma and epithelium originate from the neural crest- and ectodermal placode-derived trigeminal ganglion."
  • Bovine cornea extracellular matrix structure[2] "Electron microscopy and X-ray fibre diffraction were used to ascertain collagen fibril architecture. The bovine cornea was 1021±5.42μm thick at its outer periphery, defined as 9-12mm from the corneal centre, compared to 844±8.10μm at the centre. The outer periphery of the cornea was marginally, but not significantly, more hydrated than the centre (H=4.3 vs. H=3.7), and was more abundant in hydroxyproline (0.12 vs. 0.06mg/mg dry weight of cornea). DMMB assays indicated no change in the total amount of sulphated GAG across the cornea. Immunohistochemistry revealed the presence of both high- and low-sulphated epitopes of KS, as well as DS, throughout the cornea, and CS only in the peripheral cornea before the limbus. Quantification by ELISA, disclosed that although both high- and low-sulphated KS remained constant throughout stromal depth at different radial positions, high-sulphated epitopes remained constant from the corneal centre to outer-periphery, whereas low-sulphated epitopes increased significantly.
More recent papers  
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Carnegie Stages - Eye

Human Eye Development
Carnegie Stage Event
10 optic primordia appear.
11 Right and left optic primordia meet at the optic chiasma forming a U-shaped rim.
12 optic neural crest reaches its maximum extent and the optic vesicle becomes covered by a complete sheath,
13 By the end of the fourth week the optic vesicle lies close to the surface ectoderm. Optic evagination differentiation allows identification of optic part of retina, future pigmented layer of retina, and optic stalk. The surface ectoderm overlying the optic vesicle, in response to this contact, has thickened to form the lense placode.
14 (about 32 days) Lens placode is indented by the lens pit, cup-shaped and still communicates with the surface by a narrowing pore.
15 (about 33 days) Lens pit is closed. The lens vesicle and optic cup lie close to the surface ectoderm and appear to press against the surface.
16 (37 days) Growth of the lens body results in a D-shaped lens cavity. Perilental blood vessels (tunica vasculosa lentis) are visible. Prior to the development of the eyelids, one small sulcus or groove forms above the eye (eyelid groove) and another below it.
17 - 19 Retinal pigment is visible and the retinal fissure is largely closed. Eyelids grooves deepen, eyelid folds develop, first below, and then above, the eye.
18 Mesenchyme invades the region between the lens epithelium and the surface ectoderm.
19 - 22 Eyelid folds develop into the eyelids and cover more of the eye as the palpebral fissure takes shape. The upper and the lower eyelids meet at the outer canthus in Stage 19.
20 Lens cavity is lost and a lens suture begins to form. The inner canthus is established.
23 retina comprises the pigmented layer, external limiting membrane, proliferative zone, external neuroblastic layer, transient fiber layer, internal neuroblastic layer, nerve fiber layer, and internal limiting membrane. Eyelids closure is complete (Note - shown as still open in the Kyoto embryo).
Data from a study of human embryonic carnegie stages[3] and other sources.
Week: 1 2 3 4 5 6 7 8
Carnegie stage: 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

Human Cornea

Week 8 Stage 22

The images below link to virtual slides of the human developing eye at Carnegie stage 22. Click on the image to open or select specific regions from the regions of interest links.

Stage 22 - Eye and Nose

Stage 22 image 008.jpg

 ‎‎Mobile | Desktop | Original

Stage 22 | Embryo Slides
Stage 22 - Eye

Stage 22 image 008-eye.jpg

 ‎‎Mobile | Desktop | Original

Stage 22 | Embryo Slides

Virtual Slide - Regions of Interest

Links: Embryo Virtual Slides

Cornea Epithelia

The cornea ocular surface is composed of three epithelia, conjunctival, limbal and corneal.
  • Limbal stem cells are located in the palisades of Vogt, the transitional zone between the cornea and the conjunctiva.
  • Limbal stem cells are close to blood vessels.
  • They generate transient amplifying cells that terminally differentiate after a discrete number of cell divisions to corneal epithelial cells and undergo both centripetal migration and vertical migration.
 Corneal Epithelial Cells cartoon

Corneal epithelial cells cartoon[4]

The Adult Human Limbal Palisades of Vogt
  • A - Palisades of Vogt (arrow) are readily recognized in the human limbus.
  • B - Such a unique pigmented structure can be identified on the flat mount preparation of Dispase-isolated human limbal epithelial sheets.
  • C - In donors with a darker skin, these palisades of Vogt are pigmented (arrow).
  • D - Under higher magnification, these limbal areas show undulated epithelial papillae (stars).
  • E - Hematoxyline staining highlights higher stratification and more undulation of the limbal epithelium, and the underlying limbal stroma has high cellularity and vascularity (arrow shows blood vessel).

Bar represents 500 μm in A and B, 200 μm in C and E, and 50 μm in D


Adult human limbal palisades of Vogt

Adult human limbal palisades of Vogt[5]

Limbal Stem Cells

Limbal stem cell niche cartoon PMID17211449.jpg

Cartoon showing the location of limbal stem cells at the limbal basal layer.[5]


Links: Stem Cells

Descemet Membrane

Corneal endothelium basement membrane beginning in children at 3 μm thick and increases in adults to 10 μm. Consists of collagen type IV and VIII fibrils.

Composed of two layers:

  1. anterior banded layer - commencing in week 10 (GA week 12) as collagen lamellae and proteoglycans.
  2. posterior non-banded layer - deposited by endothelial cells over time and thickens postnatally over decades.


Descemet membrane was historically named after Jean Descemet (1732–1810) a French physician.

Palisades of Vogt

The palisades of Vogt are a series of radially oriented fibrovascular ridges concentrated along the upper and lower corneoscleral limbus, the vasculature component consists of radially oriented hairpin loops of narrow arterial and venous vessels. Named by Vogt in 1921. (for review see[6])


Aggregate into distinct crescentic zones and lie peripheral to the terminal capillary loops of the limbus and central to Schlemm’s canal. Lying between the connective tissue palisades are intervening radial zones of thickened conjunctival epithelium, the so-called inter-palisades or epithelial rete ridges.


Mouse Cornea

histology Mouse eye neural crest

Neural crest-derived cells contribute to mouse cornea development.[7]

  • a Toluidine blue staining of an adult eye. The boxed areas correspond to b and c
  • b A detailed view of the corneal assembly, including outer epithelium, stroma, and inner endothelium
  • c The chamber angle at the irido-corneal transition which includes the trabecular meshwork (tm).
  • d-j In vivo fate mapping of NC-derived, β-galactosidase (βGal)-expressing cells (blue)
  • d The NC origin of corneal keratocytes (arrows) and of corneal endothelium (arrowhead).
  • e Structures of the chamber angle, including the trabecular meshwork are seen to be NC-derived.
  • f At E10, the optic cup is surrounded by NC-derived cells expressing βGal.
  • g-i The majority of the cells in the periocular mesenchyme (arrows), which forms the anterior eye segment, are of NC origin, as assessed from E11.5 to E13.5.
  • j The primary vitreous at E13.5 (arrowheads) shows a strong NC contribution.
  • Mouse E12.5

    Mouse E12.5

  • Mouse E13.5

    Mouse E13.5

  • Mouse E16.5

    Mouse E16.5

  • Mouse P0

    Mouse P0

Frog Cornea

This developmental timeline is from a recent frog (Xenopus laevis) cornea study[8]

  • stage 25 - cornea starts from a simple embryonic epidermis overlying the developing optic vesicle.
  • stage 30 - detachment of the lens placode, cranial neural crest cells start to invade the space between the lens and the embryonic epidermis to construct the corneal endothelium.
  • stage 41 - a second wave of migratory cells containing presumptive keratocytes invades the matrix leading to the formation of inner cornea and outer cornea. A unique cell mass (stroma attracting center) connects the two layers like the center pole of a tent.
  • stage 48 - many secondary stromal keratocytes individually migrate to the center and form the stroma layer.
  • stage 60 - the stroma space is filled by collagen lamellae and keratocytes, and the stroma attracting center disappears. At early metamorphosis, the embryonic epithelium gradually changes to the adult corneal epithelium, which is covered by microvilli.
  • stage 62 - the embryonic epithelium thickens and cell death is observed in the epithelium, coinciding with eyelid opening.
  • After metamorphosis - cornea has attained the adult structure of three cellular layers, epithelium, stroma, and endothelium, and between the cellular layers lie two acellular layers (Bowman's layer and Descemet's membrane)

Xenopus cornea development timeline


Links: Frog Development

Molecular

Mouse eye TGF-beta model.jpg

Mouse Eye TGF-beta Model - Summary of the TGFβ-dependent development of anterior and posterior ocular structures.[7]

a Neural crest-derived cells (NC, blue) contribute to structures of the anterior eye segment and the primary vitreous (PV).
  • TGFβ signaling is involved in the formation of the ciliary body (CB) and the trabecular meshwork (TM), and in control of PV growth.
  • Moreover, normal PV development and/or TGFβ signaling are important for correct retinal patterning.
 b In the cornea, prospective stromal keratocytes and endothelial cells are of neural crest origin.
  • Here, TGFβ signaling is needed for the expression of the transcription factors Foxc1 and Pitx2 and for normal differentiation of NC-derived cells into collagen-synthesizing stromal keratocytes.
  • Moreover, in forming corneal endothelial cells (and in the TM), expression of Foxc1 and cell survival requires TGFβ signalling.

Additional Images

Historic Images

  • Fig. 52. Eye of a Fowl on the day 8

    Fig. 52. Eye of a Fowl on the day 8

  • Fig. 128. Eye of a Rabbit Embryo 12 Days

    Fig. 128. Eye of a Rabbit Embryo 12 Days

  • Kollmann713.jpg
  • Kollmann715.jpg
  • Kollmann717.jpg
  • Fig. 463. Developing lens and optic cup.

    Fig. 463. Developing lens and optic cup.

  • Fig. 464. Model showing lens and formation of optic cup.

    Fig. 464. Model showing lens and formation of optic cup.

  • Fig. 465. Stages in the development of the lens in the rabbit embryo.

    Fig. 465. Stages in the development of the lens in the rabbit embryo.

  • Fig. 466. Section through optic cup and lens invagination of chick of fifty-four hours' incubation.

    Fig. 466. Section through optic cup and lens invagination of chick of fifty-four hours' incubation.

  • Fig. 467. Section through eye of human embryo of 13-14 weeks.

    Fig. 467. Section through eye of human embryo of 13-14 weeks.

  • Fig. 1. Section through head of pig, 2 mm long.

    Fig. 1. Section through head of pig, 2 mm long.

  • Fig. 2. Section through head of chick, 2 mm long.

    Fig. 2. Section through head of chick, 2 mm long.

  • Fig. 3. Section through head of Foetal Pig, 2 mm long.

    Fig. 3. Section through head of Foetal Pig, 2 mm long.

  • Fig. 4. Section through head of Foetal Pig, 3 mm long.

    Fig. 4. Section through head of Foetal Pig, 3 mm long.

  • Fig. 5. Section through head of Foetal Pig, 3 mm long.

    Fig. 5. Section through head of Foetal Pig, 3 mm long.

  • Fig. 6. Section through head of Foetal Pig, 4 mm long.

    Fig. 6. Section through head of Foetal Pig, 4 mm long.

  • Fig. 7. Section through head of Foetal Pig, 7 mm long.

    Fig. 7. Section through head of Foetal Pig, 7 mm long.

  • Fig. 8. Section through head of pig, 8 mm long.

    Fig. 8. Section through head of pig, 8 mm long.

  • Fig. 9. Section through head of pig, 9 mm long.

    Fig. 9. Section through head of pig, 9 mm long.

  • Brown010.jpg

References

  1. ↑ Lwigale PY. (2015). Corneal Development: Different Cells from a Common Progenitor. Prog Mol Biol Transl Sci , 134, 43-59. PMID: 26310148 DOI.
  2. ↑ Ho LT, Harris AM, Tanioka H, Yagi N, Kinoshita S, Caterson B, Quantock AJ, Young RD & Meek KM. (2014). A comparison of glycosaminoglycan distributions, keratan sulphate sulphation patterns and collagen fibril architecture from central to peripheral regions of the bovine cornea. Matrix Biol. , 38, 59-68. PMID: 25019467 DOI.
  3. ↑ Pearson AA. (1980). The development of the eyelids. Part I. External features. J. Anat. , 130, 33-42. PMID: 7364662
  4. ↑ Kayama M, Kurokawa MS, Ueno H & Suzuki N. (2007). Recent advances in corneal regeneration and possible application of embryonic stem cell-derived corneal epithelial cells. Clin Ophthalmol , 1, 373-82. PMID: 19668514
  5. ↑ 5.0 5.1 Li W, Hayashida Y, Chen YT & Tseng SC. (2007). Niche regulation of corneal epithelial stem cells at the limbus. Cell Res. , 17, 26-36. PMID: 17211449 DOI.
  6. ↑ Goldberg MF & Bron AJ. (1982). Limbal palisades of Vogt. Trans Am Ophthalmol Soc , 80, 155-71. PMID: 7182957
  7. ↑ 7.0 7.1 Ittner LM, Wurdak H, Schwerdtfeger K, Kunz T, Ille F, Leveen P, Hjalt TA, Suter U, Karlsson S, Hafezi F, Born W & Sommer L. (2005). Compound developmental eye disorders following inactivation of TGFbeta signaling in neural-crest stem cells. J. Biol. , 4, 11. PMID: 16403239 DOI.
  8. ↑ Hu W, Haamedi N, Lee J, Kinoshita T & Ohnuma S. (2013). The structure and development of Xenopus laevis cornea. Exp. Eye Res. , 116, 109-28. PMID: 23896054 DOI.

Journals

  • Cornea "For corneal specialists and for all general ophthalmologists with an interest in this exciting subspecialty, Cornea brings together the latest clinical and basic research on the cornea and the anterior segment of the eye." [jour PuMed Listing]

Reviews

Lwigale PY. (2015). Corneal Development: Different Cells from a Common Progenitor. Prog Mol Biol Transl Sci , 134, 43-59. PMID: 26310148 DOI.

Maycock NJ & Marshall J. (2014). Genomics of corneal wound healing: a review of the literature. Acta Ophthalmol , 92, e170-84. PMID: 23819758 DOI.

Hassell JR & Birk DE. (2010). The molecular basis of corneal transparency. Exp. Eye Res. , 91, 326-35. PMID: 20599432 DOI.

Masters BR. (2009). Correlation of histology and linear and nonlinear microscopy of the living human cornea. J Biophotonics , 2, 127-39. PMID: 19343693 DOI.


The International Journal of Developmental Biology Vol. 48 Nos. 8/9 (2004) Eye Development

Articles

MAURICE DM. (1957). The structure and transparency of the cornea. J. Physiol. (Lond.) , 136, 263-86. PMID: 13429485


Bookshelf cornea development

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Terms

  • Limbal epithelial stem cells - cells located at the limbal basal layer.
  • palisades of Vogt - series of radially oriented fibrovascular ridges concentrated along the upper and lower corneoscleral limbus, the vasculature component consists of radially oriented hairpin loops of narrow arterial and venous vessels. Named by Vogt in 1921. PMID 7182957


.

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Cite this page: Hill, M.A. (2024, May 25) Embryology Vision - Cornea Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Vision_-_Cornea_Development

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