Vision - Retina Development

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Retina development (Week 8 stage 22)
Retina development (Week 8 stage 22)


The retina and its development has been the subject of research from the early histology studies of Santiago Ramón y Cajal (1852 - 1934) and Camillo Golgi (1843 - 1926). It has a complex differentiation involving both a neural retina and a pigmented retina, in the embryo separated by a space and in the adult closely apposed to each other.

The retina is also a key focus of clinical research, as genetic or environmental damage and vision loss is generally a result of damage to the retinal cells or their processes. Retinitis pigments is a degenerative retinal disease. Retinoblastoma is retinal cancer that allowed researchers to identify retinoblastoma protein (Rb), the first identified tumour suppressor protein acting as a G1 cell cycle checkpoint.

Vision Links: Introduction | Lens | Retina | Placodes | Extraocular Muscle | Cornea | Eyelid | Abnormalities | Student project 1 | Student project 2 | Category:Vision
Historic Vision Embryology  
1906 Eye Embryology | 1907 Development Atlas | 1912 Eye Development | 1912 Nasolacrimal Duct | 1918 Grays Anatomy | 1921 Eye Development | 1922 Optic Primordia | Historic Disclaimer

Some Recent Findings

  • Analysis of expression of transcription factors in early human retina[1]"Here, we analyze the expression of transcription factors that are involved in the early development of the retina in human embryos at 6-12 weeks post-conception. Our work demonstrates that early developing neural retinas can be divided into two layers, the outer and inner neuroblast layers. Eye-field transcription factors and those related to the early development of the retina have distinct expression patterns in the two layers. Cell-type-specific transcription factors emerge at 8 weeks. These data provide clear and systemic structures for early retinal development in human."
  • Development of Retinal Layers in Prenatal Human Retina[2] "A histology collection of human eyes from fetal week (Fwk) 8 to postnatal (P) 10 weeks was analyzed. The length of the nasal and temporal retina was measured along the horizontal meridian in 20 eyes. The location of the inner plexiform layer (IPL) and outer plexiform layer (OPL) was identified at each age, and its length measured. The human eye retinal length increased from 5.19 mm at Fwk 8 to 20.92 mm at midgestation to 32.88 mm just after birth. The IPL appeared in the presumptive fovea at Fwk 8, reached the eccentricity of the optic nerve by Fwk 12 and was present to both nasal and temporal peripheral edges by Fwk 18-21. By contrast, the OPL developed slowly. A short OPL was first present in the Fwk 11 fovea and did not reach the eccentricity of the optic nerve until midgestation. The OPL reached the retinal edges by Fwk 30. Laminar development of both IPL and OPL occurred before vascular formation In human fetal retina, the IPL reached the far peripheral edge of the retina by midgestation and the OPL by late gestation. Only very early in utero events could affect IPL lamination in central retina, but events occurring after Fwk 20 in peripheral retina would overlap OPL laminar development in outer retina."
  • A complex choreography of cell movements shapes the vertebrate eye[3] "Optic cup morphogenesis (OCM) generates the basic structure of the vertebrate eye. Although it is commonly depicted as a series of epithelial sheet folding events, this does not represent an empirically supported model. Here, we combine four-dimensional imaging with custom cell tracking software and photoactivatable fluorophore labeling to determine the cellular dynamics underlying OCM in zebrafish."
  • Roles of homeobox genes in retinal ganglion cell differentiation and axonal guidance[4] "In this review, we have described four major families of homeobox genes which play important roles in RGC differentiation as well as axonal pathfinding. The mechanism underlying how these HD TFs affect axonal pathfinding is not entirely known. One possibility is that the downstream targets directly regulated by these HD TF are responsible for axonal guidance. Examples of this are the repression of EphB1 by Isl2, and Vax1/Vax2 regulation of EphB2/EphB3 expression. The roles of Dlx homeobox genes in RGC axonal guidance have not yet been reported. However, in the mouse telencephalon, Dlx1 and Dlx2 promote the tangential migration of GABAergic interneurons by repressing axonal growth (Cobos et al. 2007) and inhibiting Neuropilin-2 expression (Le et al. 2007). It is possible that the genetic program defining RGC identity also encodes a unique "sensory" network for their axons, determining how and where RGC axons respond to guidance cues en route to CNS targets."
More recent papers  
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This table shows an automated computer PubMed search using the listed sub-heading term.

  • Therefore the list of references do not reflect any editorial selection of material based on content or relevance.
  • References appear in 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.

Links: References | Discussion Page | Pubmed Most Recent | Journal Searches

Search term: Retina Embryology

Dorota Rogińska, Miłosz P Kawa, Ewa Pius-Sadowska, Renata Lejkowska, Karolina Łuczkowska, Barbara Wiszniewska, Kai Kaarniranta, Jussi J Paterno, Christian A Schmidt, Bogusław Machaliński, Anna Machalińska Depletion of the Third Complement Component Ameliorates Age-Dependent Oxidative Stress and Positively Modulates Autophagic Activity in Aged Retinas in a Mouse Model. Oxid Med Cell Longev: 2017, 2017;5306790 PubMed 28928904

Lihua Liu, Zhongfu Zuo, Sijing Lu, Aihua Liu, Xuezheng Liu Naringin attenuates diabetic retinopathy by inhibiting inflammation, oxidative stress and NF-κB activation in vivo and in vitro. Iran J Basic Med Sci: 2017, 20(7);813-821 PubMed 28852447

S Lule, A I Colpak, B Balci-Peynircioglu, Y Gursoy-Ozdemir, S Peker, U Kalyoncu, A Can, N Tekin, D Demiralp, T Dalkara Behçet Disease serum is immunoreactive to neurofilament medium which share common epitopes to bacterial HSP-65, a putative trigger. J. Autoimmun.: 2017; PubMed 28844827

Neveen E R El-Bakary, Mohamed M A Abumandour Visual adaptations of the eye of the gilthead sea bream (Sparus aurata). Vet. Res. Commun.: 2017; PubMed 28842847

Klaudia Szabó, Anna Énzsöly, Bulcsú Dékány, Arnold Szabó, Rozina I Hajdú, Tamás Radovits, Csaba Mátyás, Attila Oláh, Lenke K Laurik, Gábor M Somfai, Béla Merkely, Ágoston Szél, Ákos Lukáts Histological Evaluation of Diabetic Neurodegeneration in the Retina of Zucker Diabetic Fatty (ZDF) Rats. Sci Rep: 2017, 7(1);8891 PubMed 28827737

Adult Retina

Historic retina drawing.jpg

Eye and retina cartoon.jpg

Vertebrates have ten identifiable layers formed from neurons, their processes (nerve fibers), membranes, photoreceptors and pigmented cells. Light must pass through nearly all these layers to the photoreceptors.

  1. Inner limiting membrane - Müller cell footplates.
  2. Nerve fiber layer - retinal ganglion axons eventually the optic nerve.
  3. Ganglion cell layer - neuronal cell bodies of retinal ganglion cells, their axons form the nerve fiber layer and eventually the optic nerve.
  4. Inner plexiform layer - another layer of neuronal processes.
  5. Inner nuclear layer - neuronal cell bodies
  6. Outer plexiform layer - another layer of neuronal processes.
  7. Outer nuclear layer - neuronal cell bodies
  8. External limiting membrane - layer separating inner segment portions of photoreceptors from their cell nuclei.
  9. Photoreceptor layer - rods and cones that convert light into signals.
  10. Retinal pigment epithelium.

Week 5

Stage 13

Stage 13 image 060.jpgStage 13 image 061.jpg

Week 8

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: Stage 22 | Embryo Virtual Slides

Stage 22 image 208.jpg Stage 22 image 206.jpg

Retinal Pigment Epithelium

Retinal pigment epithelium (RPE) cells are generated directly from the optic neuroepithelium. The choroidal melanocytes, the other pigmented cells, are derived from neural crest cells that have migrated towards the eye.

Retinal pigment epithelium cells:

  • cuboidal cells
  • apical side form multiple villi
    • these villi are in direct contact with the outer segments of the photoreceptor cells
  • lateral sides are joined together by tight, adherens and gap junctions
  • basal side contacts the underlying basal membrane (Bruch’s membrane)

Ciliary Body

A proposed model of ciliary body development begins with specification at the optic vesicle stage, when the neural retina and pigmented epithelium are also specified. The molecular signals could involve overlapping BMP and FGF signals. The lens has recently been shown in the chicken model to not be required for specification of the iris and ciliary body.[5]


  1. Jing Chen, Lixiang Ma, Songtao Wang, Xiaobing Wang, Yan Sun, Lu Gao, Jin Li, Guomin Zhou Analysis of expression of transcription factors in early human retina. Int. J. Dev. Neurosci.: 2017; PubMed 28377129
  2. Anita Hendrickson Development of Retinal Layers in Prenatal Human Retina. Am. J. Ophthalmol.: 2015; PubMed 26410132
  3. Kristen M Kwan, Hideo Otsuna, Hinako Kidokoro, Keith R Carney, Yukio Saijoh, Chi-Bin Chien A complex choreography of cell movements shapes the vertebrate eye. Development: 2012, 139(2);359-72 PubMed 22186726 | PMC3243097 | Development
  4. Qi Zhang, David D Eisenstat Roles of homeobox genes in retinal ganglion cell differentiation and axonal guidance. Adv. Exp. Med. Biol.: 2012, 723;685-91 PubMed 22183394
  5. Magnus R Dias da Silva, Nicola Tiffin, Tatsuo Mima, Takashi Mikawa, Jeanette Hyer FGF-mediated induction of ciliary body tissue in the chick eye. Dev. Biol.: 2007, 304(1);272-85 PubMed 17275804

Online Textbooks


Massimiliano Andreazzoli Molecular regulation of vertebrate retina cell fate. Birth Defects Res. C Embryo Today: 2009, 87(3);284-95 PubMed 19750521

Ales Cvekl, Wei-Lin Wang Retinoic acid signaling in mammalian eye development. Exp. Eye Res.: 2009, 89(3);280-91 PubMed 19427305

Lucia Galli-Resta, Paola Leone, David Bottari, Monica Ensini, Elisa Rigosi, Elena Novelli The genesis of retinal architecture: an emerging role for mechanical interactions? Prog Retin Eye Res: 2008, 27(3);260-83 PubMed 18374618

D Taylor Developmental abnormalities of the optic nerve and chiasm. Eye (Lond): 2007, 21(10);1271-84 PubMed 17914430

Barbara L Finlay The developing and evolving retina: using time to organize form. Brain Res.: 2008, 1192;5-16 PubMed 17692298

Michel Cayouette, Lucia Poggi, William A Harris Lineage in the vertebrate retina. Trends Neurosci.: 2006, 29(10);563-70 PubMed 16920202

Bookshelf retina development


Jihua Yao, Jingyao Zhou, Qiaoling Liu, Daru Lu, Lu Wang, Xiaojing Qiao, William Jia Atoh8, a bHLH transcription factor, is required for the development of retina and skeletal muscle in zebrafish. PLoS ONE: 2010, 5(6);e10945 PubMed 20532172

Henrik Boije, Henrik Ring, Meritxell López-Gallardo, Carmen Prada, Finn Hallböök Pax2 is expressed in a subpopulation of Müller cells in the central chick retina. Dev. Dyn.: 2010, 239(6);1858-66 PubMed 20503381

Sean A Georgi, Thomas A Reh Dicer is required for the transition from early to late progenitor state in the developing mouse retina. J. Neurosci.: 2010, 30(11);4048-61 PubMed 20237275

Fengming Yue, Kohei Johkura, Sakiko Shirasawa, Tadayuki Yokoyama, Yuji Inoue, Daihachiro Tomotsune, Katsunori Sasaki Differentiation of primate ES cells into retinal cells induced by ES cell-derived pigmented cells. Biochem. Biophys. Res. Commun.: 2010, 394(4);877-83 PubMed 20206598

Netta Levin, Serge O Dumoulin, Jonathan Winawer, Robert F Dougherty, Brian A Wandell Cortical maps and white matter tracts following long period of visual deprivation and retinal image restoration. Neuron: 2010, 65(1);21-31 PubMed 20152110

Fredrik Ghosh, Karin Arnér Cell type differentiation dynamics in the developing porcine retina. Dev. Neurosci.: 2010, 32(1);47-58 PubMed 20150723

Patrick W Keeley, Benjamin E Reese Role of afferents in the differentiation of bipolar cells in the mouse retina. J. Neurosci.: 2010, 30(5);1677-85 PubMed 20130177

Ana V Sánchez-Sánchez, Esther Camp, Aránzazu Leal-Tassias, José L Mullor Wnt signaling has different temporal roles during retinal development. Dev. Dyn.: 2010, 239(1);297-310 PubMed 20014102

Laszlo Hackler, Jun Wan, Anand Swaroop, Jiang Qian, Donald J Zack MicroRNA profile of the developing mouse retina. Invest. Ophthalmol. Vis. Sci.: 2010, 51(4);1823-31 PubMed 19933188

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