Vision - Cornea Development: Difference between revisions

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Introduction

These notes introduce the development of the cornea of the eye.

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

Integumentary Development

Some Recent Findings

^[1]

More recent papers
This table allows an automated computer search of the external PubMed database using the listed "Search term" text link. This search now requires a manual link as the original PubMed extension has been disabled. The displayed list of references do not reflect any editorial selection of material based on content or relevance. References also appear on this list based upon the date of the actual page viewing. References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability. More? References \| Discussion Page \| Journal Searches \| 2019 References \| 2020 References Search term: Cornea Development <pubmed limit=5>Cornea Development</pubmed> Search term: Cornea Embryology <pubmed limit=5>Cornea Embryology</pubmed>

Timeline

Embryonic Development

Weeks 3 - 4 Eye Fields-Optic Vesicle
Weeks 5 - 6 Optic Cup, Lens Vesicle, Choroid Fissure, Hyaloid Artery
Weeks 7 - 8 Cornea, Anterior Chamber, Pupillary Membrane, Lens, Retina
Weeks 9 - 15 Iris, Ciliary Body
Weeks 8 - 10 Eyelids

Carnegie Stages - Eye

The following data is from a study of human embryonic carnegie stages^[2] and other sources.

Stage 10 - optic primordia appear.
Stage 11 - right and left optic primordia meet at the optic chiasma forming a U-shaped rim.
Stage 12 - optic neural crest reaches its maximum extent and the optic vesicle becomes covered by a complete sheath,
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.
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.
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.
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.
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.
Stages 18 - Mesenchyme invades the region between the lens epithelium and the surface ectoderm.
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.
Stage 20 - The lens cavity is lost and a lens suture begins to form. The inner canthus is established.
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).

Frog Cornea

This developmental timeline is from a recent Xenopus laevis cornea study^[3]

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)

Links: Frog Development

Additional Images

Historic Images

Fig. 463. Developing lens and 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. 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. 1. Section through head of pig, 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. 4. 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. 7. Section through head of Foetal Pig, 7 mm long.
Fig. 8. Section through head of pig, 8 mm long.
Fig. 9. Section through head of pig, 9 mm long.

References

↑ <pubmed>22496813</pubmed>
↑ <pubmed>7364662</pubmed>
↑ <pubmed>23896054</pubmed>| Exp Eye Res.

Reviews

<pubmed>20855501</pubmed>| JCB

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

Articles

Bookshelf cornea development

Search Pubmed

Search Pubmed: cornea development

Search Entrez: cornea development

Terms

External Links

External Links Notice - The dynamic nature of the internet may mean that some of these listed links may no longer function. If the link no longer works search the web with the link text or name. Links to any external commercial sites are provided for information purposes only and should never be considered an endorsement. UNSW Embryology is provided as an educational resource with no clinical information or commercial affiliation.

UNSW Virtual Slides Eye Development Histology (requires login)

Glossary Links

Glossary: A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y | Z | Numbers | Symbols | Term Link

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

What Links Here?

[PMID-1] <pubmed>22496813</pubmed>

[2] <pubmed>7364662</pubmed>

[PMID23896054-3] <pubmed>23896054</pubmed>| Exp Eye Res.

[1]

[2]

[3]

@@ Line 1: / Line 1: @@
 {{Header}}
 ==Introduction==
-[[File:Historic_retina_drawing.jpg|right|300px]]
+These notes introduce the development of the cornea of the eye.
-These notes introduce the development of the eye cornea.
 {{Vision Links}}
+[[Integumentary Development]]
+== Some Recent Findings ==
-{{Senses Links}}
-== 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>]]
 {|
 |-bgcolor="F5FAFF"
 |
-* '''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."
+* <ref name = PMID><pubmed>22496813</pubmed></ref>
-* '''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."
 |}
 {| class="wikitable mw-collapsible mw-collapsed"
@@ Line 25: / Line 19: @@
 | [[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>Vision Embryology</pubmed>
+<pubmed limit=5>Cornea Development</pubmed>
+Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Cornea+Embryology ''Cornea Embryology'']
+<pubmed limit=5>Cornea Embryology</pubmed>
 |}
 ==Timeline==
@@ Line 56: / Line 54: @@
 * [[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==
+==Frog Cornea==
-[[File:Stage_22_image_155.jpg|thumb|Human Lens (stage 22)]]
+This developmental timeline is from a recent Xenopus laevis cornea study<ref name=PMID23896054><pubmed>23896054</pubmed>| [http://www.sciencedirect.com/science/article/pii/S0014483513002194 Exp Eye Res.]</ref>
-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.
-:'''Links:''' [[Vision - Lens Development]]
-==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.
-The retinotectal map (eye to brain) of birds (lower vertebrates):
+* '''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)
-* temporal (posterior) retina is connected to the rostral (anterior) part of the contralateral optic tectum
-* 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.
+:'''Links:''' [[Frog Development]]
-EphA/ephrin-A molecular signaling also thought to have a role in establishing the initial retinotopic map.
-==Neural Crest==
-{|
-| [[File:Mouse eye neural crest.jpg|400px]]
-Mouse eye neural crest<ref name="PMID16403239"><pubmed>16403239</pubmed>| [http://jbiol.com/content/4/3/11 J Biol.]</ref>
-| [[File:Mouse_eye_TGF-beta_model.jpg|400px]]
-Mouse eye TGF-beta model<ref name="PMID16403239"><pubmed>16403239</pubmed>| [http://jbiol.com/content/4/3/11 J Biol.]</ref>
-|}
-:'''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]]
-==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.
-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>
-{|
-! About the Muscles
-! Legend
-|-
-| valign=top width=400px valign=top|
-* 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).
-** The Annulus of Zinn surrounds the optic nerve, ophthalmic artery, and ophthalmic vein at their entrance through the apex of the orbit.
-* 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==
 <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>
 ===Historic Images===
 <gallery>
-File:Bailey456.jpg|Fig. 456. Location of optic areas before the closure of the neural groove.
-File:Bailey457.jpg|Fig. 457. Location of areas shown in Fig. 456 after the formation of the neural canal.
-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:Bailey460.jpg|Fig. 460. Section through head of chick of two days' incubation.
-File:Bailey461.jpg|Fig. 461. Section through head of chick of three days' incubation.
-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:Bailey464.jpg|Fig. 464. Model showing lens and formation of optic cup.
@@ Line 141: / Line 80: @@
 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: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:Brown002.jpg|Fig. 2. Section through head of chick, 2 mm long.
@@ Line 154: / Line 90: @@
 File:Brown009.jpg|Fig. 9. Section through head of pig, 9 mm long.
 File:Brown010.jpg
-File:Brown011.jpg|Fig. 11. colobomba of the fundus in the adult and means a lack of development.
 </gallery>
@@ Line 160: / Line 95: @@
 <references/>
-===Online Textbooks===
-* 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]
-* '''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]
-* '''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]
-* '''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]
-* '''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===
@@ Line 185: / Line 106: @@
-'''Bookshelf'''  [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=vision%20development vision development]
+'''Bookshelf'''  [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=cornea%20development cornea development]
 ===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==