Sensory - Vision Development

Embryology - 5 Dec 2023        Expand to Translate
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Introduction

Adult Human Retina histology

These notes introduce vision development of the eye: induction and regional specification of the eye structures, maturation and formation of retina and optic tectum neuronal connections.

The adult eye has contributions from several different embryonic layers eventually forming neuronal, supportive connective tissue, optical structures, and muscular tissues. Historically determined that at birth, the human eyeball volume is about 3.25 cc and the estimated weight is about 3.40 grams.

There are additional pages shown in the vision links, covering specific topics of vision development.

Some Recent Findings

Vision-dependent specification of cell types and function in the developing cortex "The role of postnatal experience in sculpting cortical circuitry, while long appreciated, is poorly understood at the level of cell types. We explore this in the mouse primary visual cortex (V1) using single-nucleus RNA sequencing, visual deprivation, genetics, and functional imaging. We find that vision selectively drives the specification of glutamatergic cell types in upper layers (L) (L2/3/4), while deeper-layer glutamatergic, GABAergic, and non-neuronal cell types are established prior to eye opening. L2/3 cell types form an experience-dependent spatial continuum defined by the graded expression of ∼200 genes, including regulators of cell adhesion and synapse formation. One of these genes, Igsf9b, a vision-dependent gene encoding an inhibitory synaptic cell adhesion molecule, is required for the normal development of binocular responses in L2/3. In summary, vision preferentially regulates the development of upper-layer glutamatergic cell types through the regulation of cell-type-specific gene expression programs." Endocrine regulation of multichromatic color vision. "Vertebrate color vision requires spectrally selective opsin-based pigments, expressed in distinct cone photoreceptor populations. In primates and in fish, spectrally divergent opsin genes may reside in head-to-tail tandem arrays. Mechanisms underlying differential expression from such arrays have not been fully elucidated. Regulation of human red (LWS) vs. green (MWS) opsins is considered a stochastic event, whereby upstream enhancers associate randomly with promoters of the proximal or distal gene, and one of these associations becomes permanent. We demonstrate that, distinct from this stochastic model, the endocrine signal thyroid hormone (TH) regulates differential expression of the orthologous zebrafish lws1/lws2 array, and of the tandemly quadruplicated rh2-1/rh2-2/rh2-3/rh2-4 array. TH treatment caused dramatic, dose-dependent increases in abundance of lws1, the proximal member of the lws array, and reduced lws2 Fluorescent lws reporters permitted direct visualization of individual cones switching expression from lws2 to lws1 Athyroidism increased lws2 and reduced lws1, except within a small ventral domain of lws1 that was likely sustained by retinoic acid signaling. Changes in lws abundance and distribution in athyroid zebrafish were rescued by TH, demonstrating plasticity of cone phenotype in response to this signal. TH manipulations also regulated the rh2 array, with athyroidism reducing abundance of distal members. Interestingly, the opsins encoded by the proximal lws gene and distal rh2 genes are sensitive to longer wavelengths than other members of their respective arrays; therefore, endogenous TH acts upon each opsin array to shift overall spectral sensitivity toward longer wavelengths, underlying coordinated changes in visual system function during development and growth." Fetal ocular development in the second trimester of pregnancy documented by 7.0 T postmortem Magnetic Resonance Imaging "Few investigators have analyzed fetal ocular growth with Magnetic Resonance Imaging (MRI) of high magnetic strength. Our purpose is to obtain normative biometrics for fetal ocular development in the second trimester of pregnancy. Sixty specimens with a gestational age (GA) of 12-23 weeks were scanned using a 7.0 T MRI scanner. The linear interocular and binocular distances (IOD and BOD, respectively), globe diameter (GD) and lens diameter (LD) were measured on the transverse section of the largest diameter of the eyeballs. The three dimensional (3D) visualization model of the eyeball was reconstructed with Amira software. Then, the globe and lens volumes (GV and LV, respectively) were obtained. All the measurements were plotted as a function of GA. The fetal ocular structures in the second trimester of pregnancy could be clearly delineated on 7.0 T postmortem MRI images. All the linear measurements logarithmically increased with GA, while, the volumetric measurements linearly increased with GA. Postmortem MRI of high magnetic strength can clearly document fetal ocular growth in the second trimester of pregnancy. These quantitative data may be a valuable reference for the assessment of normal fetal eyeball development in clinical settings and may be considered a supplement to anatomical investigations."

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: Vision Development | Vision Embryology

Older papers

These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table. See also the Discussion Page for other references listed by year and References on this current page. The relationship between eye movement and vision develops before birth "While the visuomotor system is known to develop rapidly after birth, studies have observed spontaneous activity in vertebrates in visually excitable cortical areas already before extrinsic stimuli are present. Resting state networks and fetal eye movements were observed independently in utero, but no functional brain activity coupled with visual stimuli could be detected using fetal fMRI. This study closes this gap and links in utero eye movement with corresponding functional networks. BOLD resting-state fMRI data were acquired from seven singleton fetuses between gestational weeks 30-36 with normal brain development. During the scan time, fetal eye movements were detected and tracked in the functional MRI data. We show that already in utero spontaneous fetal eye movements are linked to simultaneous networks in visual- and frontal cerebral areas. In our small but in terms of gestational age homogenous sample, evidence across the population suggests that the preparation of the human visuomotor system links visual and motor areas already prior to birth." Activation of c-Jun N-Terminal Kinase (JNK) during Mitosis in Retinal Progenitor Cells "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." Rearrangement of retinogeniculate projection patterns after eye-specific segregation in mice "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 "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."

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

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 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

See below the drawings of sections of the whole eye from week 8 of development.

Optic Nerve

• Carnegie stage 19 - Optic nerve small, slender. Lumen practically whole length of stalk. Few or no fibers.
• Carnegie stage 20 - Ependymal arrangement partially retained along stalk. Remnant of ependyma along whole length of stalk. Hyaloid groove at bulbar end. A few fibers arriving at brain.
• Carnegie stage 21 - Remnant of ependyma present.
• Carnegie stage 22 - Sheath layer beginning to form. Vascular canal present.
• Carnegie stage 23 - Early nerve sheath. Reticular spongioblastic framework, striate arrangement of nuclei, bundles of fibers. Definite nerve sheath.

• Eye and Optic Nerve
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  Carnegie stage 19

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  Carnegie stage 20

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  Carnegie stage 21

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  Carnegie stage 22

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  Carnegie stage 23

Lens

Human Lens (Carnegie stage Template:CS222)

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

Stage 22 Eye

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  ‎‎Mobile | Desktop | Original Stage 22 | Embryo Slides

Stage 22 - Eye  ‎‎Mobile | Desktop | Original Stage 22 | Embryo Slides

Virtual Slide - Regions of Interest Eye Overview Lens and Cornea Cornea and Anterior Chamber Lens - anterior Lens - posterior Lens -equator Iris and Lens Retina - Neural and Pigmented Retina and Nerve fibre layer Optic Nerve Head and Hyaloid Vessel Hyaloid Vessel Optic Nerve Head Optic Nerve and Sphenoid (alar orbitalis) Extraocular Muscle - Medial Rectus

Links: Embryo Virtual Slides

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):

• 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.

EphA/ephrin-A molecular signaling also thought to have a role in establishing the initial retinotopic map.

Neural Crest

Mouse eye neural crest

Mouse eye TGF-beta model

Links: Image - Mouse eye neural crest | Image - Mouse eye TGF-beta model | Vision Development | Neural Crest Development | Head Development

Schlemm's canal

Schematic showing the stages of Schlemm's canal development in the postnatal mouse by the novel process of canalogenesis. (Cartoons have been drawn for clarity and are not intended to suggest that most early sprouts arise from the LVP.)

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.

About the Muscles	Legend
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.	IR - inferior rectus SR - superior rectus LR - lateral rectus MR - medial rectus SO - superior oblique IO - inferior oblique

Links: Extraocular Muscles

Additional Images

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  Human stage 22 developing iris region

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  Human stage 22 developing iris region

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  Human stage 22 overview of optic nerve

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  Human stage 22 overview of eye

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  Human stage 22 lens and hyaloid vessels

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  Human stage 22 optic nerve (stalk)

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  Human stage 22 retina

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  Mouse adult optic nerve axons

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  Pax6 eye phenotypes

Historic Images

Historic Disclaimer - information about historic embryology pages

Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

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  Frog eye development.

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  Fig. 456. Location of optic areas before the closure of the neural groove.

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  Fig. 457. Location of areas shown in Fig. 456 after the formation of the neural canal.

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  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.

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  Fig. 460. Section through head of chick of two days' incubation.

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  Fig. 461. Section through head of chick of three days' incubation.

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  Fig. 462. Later stage in development of optic cup and lens than is shown in Fig. 461.

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  Fig. 463. Developing lens and optic cup.

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

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

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

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

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  Fig. 468. Development of the retinal cells.

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  Fig. 469. Vertical section through retina of a four months' human embryo.

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  Fig. 470. Vertical section through retina of a five and one-half months' human embryo.

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

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

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

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

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

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

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

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

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

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  Fig. 11. colobomba of the fundus in the adult and means a lack of development.

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/

• Gilbert SF. Developmental Biology. 6th edition. Sunderland (MA): Sinauer Associates; 2000. Development of the Vertebrate Eye. Available from: https://www.ncbi.nlm.nih.gov/books/NBK10024/
  • Evolution of the mammalian middle ear bones from the reptilian jaw | Chick embryo rhombomere neural crest cells | Some derivatives of the pharyngeal arches | Formation of the Neural Tube | Differentiation of the Neural Tube | Tissue Architecture of the Central Nervous System | Neuronal Types | 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 The Auditory System | The Inner Ear | The Middle Ear | The External Ear | Early Brain Development | Construction of Neural Circuits | 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. Neural Development | The three phases of neural development

• Clinical Methods 63. Cranial Nerves IX and X: The Glossopharyngeal and Vagus Nerves | The Tongue | 126. The Ear and Auditory System | An Overview of the Head and Neck - Ears and Hearing | Audiometry

• Health Services/Technology Assessment Text (HSTAT) Bethesda (MD): National Library of Medicine (US), 2003 Oct. Developmental Disorders Associated with Failure to Thrive

• Eurekah Bioscience Collection Cranial Neural Crest and Development of the Head Skeleton
• 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

Beby F & Lamonerie T. (2013). The homeobox gene Otx2 in development and disease. Exp. Eye Res. , 111, 9-16. PMID: 23523800 DOI.

Sung CH & Chuang JZ. (2010). The cell biology of vision. J. Cell Biol. , 190, 953-63. PMID: 20855501 DOI.

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

Articles

Paquette LB, Jackson HA, Tavaré CJ, Miller DA & Panigrahy A. (2009). In utero eye development documented by fetal MR imaging. AJNR Am J Neuroradiol , 30, 1787-91. PMID: 19541779 DOI.

Bookshelf vision development

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Cite this page: Hill, M.A. (2023, December 5) Embryology Sensory - Vision Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Sensory_-_Vision_Development

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