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and air. The fiber cells of fish are generally considerably thinner than those of land vertebrates and it appears crystallin proteins are transported to the organelle free cells at the lens exterior to the inner cells through many layers of cells. Some vertebrates need to see well both above and below water at times. One example is diving birds which have the ability to change focus by 50 to 80 dioptres. Compared with animals adapted for only one environment diving birds have a somewhat altered lens and cornea structure with focus mechanisms to allow for both environments. Even among terrestrial animals the lens of
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803:, MRI and physiological investigations it is becoming apparent the lens itself is not responding entirely passively to the surrounding ciliary muscle but may be able to change its overall refractive index through mechanisms involving water dynamics in the lens still to be clarified. The accompanying micrograph shows wrinkled fibers from a relaxed sheep lens after it is removed from the animal indicating shortening of the lens fibers during near focus accommodation. The age related changes in the human lens may also be related to changes in the water dynamics in the lens.
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muscles in the ciliary body. In this model the ligaments may pull to varying degrees on the lens at the equator using the radial muscles while the ligaments offset from the equator to the front and back are relaxed to varying degrees by contracting the circular muscles. These multiple actions operating on the elastic lens allows it to change lens shape at the front more subtly. Not only changing focus, but also correcting for lens aberrations that might otherwise result from the changing shape while better fitting mathematical modeling.
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Allvar
Gullstrand spoke on "How I found the intracapsular mechanism of accommodation" and this aspect of lens focusing continues to be investigated. Young spent time searching for the nerves that could stimulate the lens to contract without success. Since that time it has become clear the lens is not a simple muscle stimulated by a nerve so the 1909 Helmholtz model took precedence. Pre-twentieth century investigators did not have the benefit of many later discoveries and techniques. Membrane proteins such as
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623:. The whole process of differentiation of the epithelial cells into crystallin filled fiber cells without organelles occurs within the confines of the lens capsule. Older cells cannot be shed and are instead internalized towards the center of the lens. This process results in a complete temporally layered record of the differentiation process from the start at the lens surface to the end at the lens center. The lens is therefore valuable to scientists studying the process of cell differentiation.
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like a globe of the world. The front and back of the lens are referred to as the anterior and posterior "poles", like the North and South poles. The "equator" is the outer edge of the lens often hidden by the iris and is the area of most cell differentiation. As the equator is not generally in the light path of the eye, the structures involved with metabolic activity avoid scattering light that would otherwise affect vision.
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580:. As the name suggests the crystallins can form a clear highly refractive jelly. These elongating cells eventually fill in the center of the vesicle with cells, that are long and thin like a strand of hair, called fibers. These primary fibers become the nucleus in the mature lens. The epithelial cells that do not form into fibers nearest the lens front give rise to the lens epithelium.
205:. Relatively long, thin fiber cells make up the majority of the lens. These cells vary in architecture and are arranged in concentric layers. New layers of cells are recruited from a thin epithelium at the front of the lens, just below the basement membrane surrounding the lens. As a result the vertebrate lens grows throughout life. The surrounding lens membrane referred to as the
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reach as far towards the front and back of the lens. The lens fibers that do not reach the poles form tight, interdigitating seams with neighboring fibers. These seams being less crystalline than the bulk of the lens are more visible and are termed "sutures". The suture patterns become more complex as more layers of lens fibers are added to the outer portion of the lens.
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Na/K-ATPase pumps located in the equatorially positioned cells of the lens epithelium. The interaction of these pumps with water channels into cells called aquaporins, molecules less than 100 daltons in size among cells via gap junctions, and calcium using transporters/regulators (TRPV channels) results in a flow of nutrients throughout the lens.
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concentric layers rather like the layers of an onion. If cut along the equator, cells have a hexagonal cross section, appearing as a honeycomb. The approximate middle of each fiber lies around the equator. These tightly packed layers of lens fibers are referred to as laminae. The lens fiber cytoplasms are linked together via
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front can then reform its shape between the suspensory ligaments in a similar way to a slack chain hanging between two poles might change its curve when the poles are moved closer together. This model requires fluid movement of the lens front only rather than trying to change the shape of the lens as a whole.
422:. The capsule is very elastic and so allows the lens to assume a more spherical shape when the tension of the suspensory ligaments is reduced. The human capsule varies from 2 to 28 micrometres in thickness, being thickest near the equator (peri-equatorial region) and generally thinner near the posterior pole.
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881:, the lens is not attached to the outer surface of the eyeball at all. There is no aqueous humor in these fish, and the vitreous body simply presses the lens against the surface of the cornea. To focus its eyes, a lamprey flattens the cornea using muscles outside of the eye and pushes the lens backwards.
835:, the ciliary body which supports the lens via suspensory ligaments also touches the lens with a number of pads on its inner surface. These pads compress and release the lens to modify its shape while focusing on objects at different distances; the suspensory ligaments usually perform this function in
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With the advent of other ways of looking at cellular structures of lenses while still in the living animal it became apparent that regions of fiber cells, at least at the lens anterior, contain large voids and vacuoles. These are speculated to be involved in lens transport systems linking the surface
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The lens continues to grow after birth, with the new secondary fibers being added as outer layers. New lens fibers are generated from the equatorial cells of the lens epithelium, in a region referred to as the "germinative zone" and "bow region". The lens epithelial cells elongate, lose contact with
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The lens fibers form the bulk of the lens. They are long, thin, transparent cells, firmly packed, with diameters typically 4β7 micrometres and lengths of up to 12mm long in humans. The lens fibers stretch lengthwise from the posterior to the anterior poles and, when cut horizontally, are arranged in
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Schachar has proposed a model for land based vertebrates that was not well received. The theory allows mathematical modeling to more accurately reflect the way the lens focuses while also taking into account the complexities in the suspensory ligaments and the presence of radial as well as circular
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In many aquatic vertebrates, the lens is considerably thicker, almost spherical resulting in increased light refraction. This difference helps compensate for the smaller angle of refraction between the eye's cornea and the watery environment, as they have more similar refractive indices than cornea
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Additional fibers are derived from lens epithelial cells located at the lens equator. These cells lengthen towards the front and back wrapping around fibers already laid down. The new fibers need to be longer to cover earlier fibers but as the lens gets larger the ends of the newer fibers no longer
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also grows in a systematic way ensuring the lens maintains an optically suitable shape in concert with the underlying fiber cells. Thousands of suspensory ligaments are embedded into the capsule at its largest diameter which suspend the lens within the eye. Most of these lens structures are derived
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that compose over 90% of the protein within the lens. The three main crystallin types found in the human eye are Ξ±-, Ξ²-, and Ξ³-crystallins. Crystallins tend to form soluble, high-molecular weight aggregates that pack tightly in lens fibers, thus increasing the index of refraction of the lens while
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which allow electrical coupling of cells are also prevalent. Electron microscopy and immunofluorescent microscopy show fiber cells to be highly variable in structure and composition. Magnetic resonance imaging confirms a layering in the lens that may allow for different refractive plans within it.
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being pulled tight by the pressure of the eyeball. At short focal distance the ciliary muscle contracts relieving some of the tension on the ligaments, allowing the lens to elastically round up a bit, increasing refractive power. Changing focus to an object at a greater distance requires a thinner
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of the lens epithelium form a thin layer between the lens capsule and the outermost layer of lens fibers at the front of the lens but not the back. The lens itself lacks nerves, blood vessels, or connective tissue. Anatomists will often refer to positions of structures in the lens by describing it
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1:posterior segment 2:ora serrata 3:ciliary muscle 4:ciliary zonules 5:Schlemm's canal 6:pupil 7:anterior chamber 8:cornea 9:iris 10:lens cortex 11:lens nucleus 12:ciliary process 13:conjunctiva 14:inferior oblique muscule 15:inferior rectus muscule 16:medial rectus muscle 17:retinal arteries and
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When Thomas Young proposed the changing of the human lens's shape as the mechanism for focal accommodation in 1801 he thought the lens may be a muscle capable of contraction. This type of model is termed intracapsular accommodation as it relies on activity within the lens. In a 1911 Nobel lecture
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demands less tension on the ligaments suspending the lens. Rather than the lens as a whole being stretched thinner for distance vision and allowed to relax for near focus, contraction of the circular ciliary muscles results in the lens having less hydrostatic pressure against its front. The lens
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After regression of the hyaloid artery, the lens receives all its nourishment from the aqueous humor. Nutrients diffuse in and waste diffuses out through a constant flow of fluid from the anterior/posterior poles of the lens and out of the equatorial regions, a dynamic that is maintained by the
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The photos from electron and light microscopes show an area of the capsule lens equator where the capsule grows and adjacent to where thousands of suspensory ligaments attach. Attachment must be strong enough to stop the ligaments being detached from the lens capsule. Forces are generated from
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The lens is split into regions depending on the age of the lens fibers of a particular layer. Moving outwards from the central, oldest layer, the lens is split into an embryonic nucleus, the fetal nucleus, the adult nucleus, the inner and outer cortex. New lens fibers, generated from the lens
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which is superficially similar structure and function to a vertebrate eye, including accommodation, while differing in basic ways such as having a two part lens and no cornea. The fundamental requirements of optics must be filled by all eyes with lenses using the tissues at their disposal so
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are opacities of the lens. While some are small and do not require any treatment, others may be large enough to block light and obstruct vision. Cataracts usually develop as the aging lens becomes more and more opaque, but cataracts can also form congenitally or after injury to the lens.
677:, is often referred to as a "model". Direct experimental proof of any lens model is necessarily difficult as the vertebrate lens is transparent and only functions well in the living animals. When considering all vertebrates aspects of all models may play varying roles in lens focus.
251:). In many fully aquatic vertebrates, such as fish, other methods of accommodation are used, such as changing the lens's position relative to the retina rather than changing the shape of the lens. Accommodation is analogous to the focusing of a photographic camera via changing its
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less curved lens. This is achieved by relaxing some of the sphincter like ciliary muscles. While not referenced this presumably allows the pressure in the eyeball to again expand it outwards, pulling harder on the lens making it less curved and thinner, so increasing the
600:, and then finally lose their nuclei (enucleate) as they become mature lens fibers. In humans, as the lens grows by laying down more fibers through to early adulthood, the lens becomes more ellipsoid in shape. After about age 20 the lens grows rounder again and the
925:, and so it is believed that the crystallin proteins were evolutionarily recruited from chaperone proteins for optical purposes. The chaperone functions of Ξ±-crystallin may also help maintain the lens proteins, which must last a human for their entire lifetime.
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is the age-related loss of accommodation, which is marked by the inability of the eye to focus on nearby objects. The exact mechanism is still unknown, but age-related changes in the hardness, shape, and size of the lens have all been linked to the
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maintaining its transparency. Ξ² and Ξ³ crystallins are found primarily in the lens, while subunits of Ξ± -crystallin have been isolated from other parts of the eye and the body. Ξ±-crystallin proteins belong to a larger superfamily of molecular
455:
and volume, with equatorially positioned lens epithelium cells contributing most to this current. The activity of the Na/K-ATPases keeps water and current flowing through the lens from the poles and exiting through the equatorial regions.
315:, biconvex shape. The front surface is less curved than the back. In a human adult, the lens is typically about 10mm in diameter and 4mm thick, though its shape changes with accommodation and its size grows throughout a person's lifetime.
2361:(English translation edited by JPC Southall. The Optical Society of America. From the third German Edition of Handbuch der Physiologischen Optik (1909), Leopold Voss, Leipzig. Dover reprint ed.). New York, NY: Dover Publications Inc.
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holding the lens in place and the forces added to during focusing. While the capsule is thinnest at the equator where its area is increasing, the anterior and posterior capsule is thinner than the area of ligament attachment.
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constricts and the lens cells bud off from the embryo's skin to form a sphere of cells known as the "lens vesicle". When the embryo is about 10mm long the lens vesicle has completely separated from the skin of the embryo.
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Paidi, Santosh Kumar; Zhang, Qinrong; Yang, Yuhan; Xia, Chun-Hong; Ji, Na; Gong, Xiaohua (19 January 2023). "Adaptive optical two-photon fluorescence microscopy probes cellular organization of ocular lenses in vivo".
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veins 18:optic disc 19:dura mater 20:central retinal artery 21:central retinal vein 22:optic nerve 23:vorticose vein 24:bulbar sheath 25:macula 26:fovea 27:sclera 28:choroid 29:superior rectus muscle 30:retina
270:, roughly one-third of the eye's total power of about 60 dioptres. By 25 years of age the ability of the lens to alter the light path has reduced to 10 dioptres and accommodation continues to decline with age.
854:, the suspensory ligaments are replaced by a membrane, including a small muscle at the underside of the lens. This muscle pulls the lens forward from its relaxed position when focusing on nearby objects. In
708:
Two horse lenses suspended on water by cling wrap with 4 approximately parallel lasers directed through them. The 1 cm spaced grid indicates an accommodated, i.e. relaxed, near focus, focal length of around
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from the developing retina, inducing the cells closest to the posterior end of the lens vesicle to elongate toward the anterior end of the vesicle. These signals also induce the synthesis of proteins called
438:
is a single layer of cells at the front of the lens between the lens capsule and the lens fibers. By providing the lens fibers with nutrients and removing waste, the cells of the epithelium maintain lens
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The lens is metabolically active and requires nourishment in order to maintain its growth and transparency. Compared to other tissues in the eye, however, the lens has considerably lower energy demands.
513:
of the lens to deeper regions. Very similar looking structures also indicate cell fusion in the lens. The cell fusion is shown by micro-injection to form a stratified syncytium in whole lens cultures.
688:
3322:
Jones, C.E.; Atchison, D.A.; Meder, R.; Pope, J.M. (August 2005). "Refractive index distribution and optical properties of the isolated human lens measured using magnetic resonance imaging (MRI)".
1397:"Macromolecular organization of basement membranes. Characterization and comparison of glomerular basement membrane and lens capsule components by immunochemical and lectin affinity procedures"
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The model of a shape changing lens of humans was proposed by Young in a lecture on the 27th Nov 1800. Others such as
Helmholtz and Huxley refined the model in the mid-1800s explaining how the
2829:
Broekhuyse, R. M.; Kuhlmann, E. D.; Stols, A. L. (September 1976). "Lens membranes II. Isolation and characterization of the main intrinsic polypeptide (MIP) of bovine lens fiber membranes".
862:, and serves to pull the lens backwards from the relaxed position to focus on distant objects. While amphibians move the lens forward, as do cartilaginous fish, the muscles involved are not
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3203:
Donaldson, Paul J.; Chen, Yadi; Petrova, Rosica S.; Grey, Angus C.; Lim, Julie C. (December 2022). "Regulation of lens water content: Effects on the physiological optics of the lens".
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contracts rounding the lens to focus near and this model was popularized by
Helmholtz in 1909. The model may be summarized like this. Normally the lens is held under tension by its
500:
Cellular and supercellular structure in the mouse lens. Photos at increasing depth: A-Epithelium B-Broadening fiber ends C-Fiber ends lock together D-F- Voids G-Vacuoles I-Sutures
3160:
Vaghefi, E; Pontre, BP; Jacobs, MD; Donaldson, PJ (August 2011). "Visualizing ocular lens fluid dynamics using MRI: manipulation of steady state water content and water fluxes".
1321:"equator of the crystalline lens - definition of equator of the crystalline lens in the Medical dictionary - by the Free Online Medical Dictionary, Thesaurus and Encyclopedia"
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Knaus, Katherine R.; Hipsley, AnnMarie; Blemker, Silvia S. (June 2021). "The action of ciliary muscle contraction on accommodation of the lens explored with a 3D model".
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987:. Beginning in the fourth month of development, the hyaloid artery and its related vasculature begin to atrophy and completely disappear by birth. In the postnatal eye,
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Tracing of
Scheimpflug photographs of 20 year old human lens being thicker focusing near and thinner when focusing far. Internal layering of the lens is also significant
726:. There is a problem with the Helmholtz model in that despite mathematical models being tried none has come close enough to working using only the Helmholtz mechanisms.
2017:
Limi, Saima; Senecal, Adrien; Coleman, Robert; Lopez-Jones, Melissa; Guo, Peng; Polumbo, Christina; Singer, Robert H.; Skoultchi, Arthur I.; Cvekl, Ales (August 2018).
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superficially eyes all tend to look similar. It is the way optical requirements are met using different cell types and structural mechanisms that varies among animals.
847:, the lens is fixed in shape, and focusing is instead achieved by moving the lens forwards or backwards within the eye using a muscle called the retractor lentus.
536:
is about 4mm long. The accompanying picture shows the process in a more easily studied chicken embryo. Unlike the rest of the eye which is derived mostly from the
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The cells of the lens epithelium also divide into new lens fibers at the lens equator. The lens lays down fibers from when it first forms in embryo until death.
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Moffat, BA; Landman, KA; Truscott, RJ; Sweeney, MH; Pope, JM (December 1999). "Age-related changes in the kinetics of water transport in normal human lenses".
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Schachar, Ronald A. (22 September 2015). "Human
Accommodative Ciliary Muscle Configuration Changes Are Consistent With Schachar's Mechanism of Accommodation".
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Left to right we have a smooth capsule, a small patch of epithelium next to fused lens fibers or perhaps a void, straighter fibers, and finally wrinkled fibers
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that maintains the precise shape and packing of the lens fibers; disruptions/mutations in certain cytoskeletal elements can lead to the loss of transparency.
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formed by budding of the inner embryo layers comes close to the embyro's outer skin. The sphere of cells induces nearby outer skin to start changing into the
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Hermans, EA; Dubbelman, M; Van der Heijde, R; Heethaar, RM (December 2008). "Equivalent refractive index of the human lens upon accommodative response".
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Stahnke, T.; Hadlich, S.; Wree, A.; Guthoff, R.; Stachs, O.; Langner, S. (16 December 2016). "Magnetresonanzmikroskopie des
Akkommodationsapparats".
552:. The lens placode is the first stage of transformation of a patch of skin into the lens. At this early stage, the lens placode is a single layer of
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Shao, Yilei; Tao, Aizhu; Jiang, Hong; Mao, Xinjie; Zhong, Jianguang; Shen, Meixiao; Lu, Fan; Xu, Zhe; Karp, Carol L.; Wang, Jianhua (1 June 2015).
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Katzir, Gadi; Howland, Howard C. (1 March 2003). "Corneal power and underwater accommodation in great cormorants( Phalacrocorax carbo sinensis )".
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Maddala, Rupalatha; Chauhan, Bharesh K.; Walker, Christopher; Zheng, Yi; Robinson, Michael L.; Lang, Richard A.; Rao, Ponugoti V. (December 2011).
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light in the wavelength range of 300β400 nm; shorter wavelengths are blocked by the cornea. The pigment responsible for blocking the light is
406:. It is presumed to be synthesized by the lens epithelium and its main components in order of abundance are heparan sulfate proteoglycan (sulfated
335:
Sheep eye lens para-formaldehyde fixed front view. Small lenses are about 1cm in diameter. Small bumps at edge are remnants of suspensory ligaments
496:
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Hermans, Erik A.; Pouwels, Petra J. W.; Dubbelman, Michiel; Kuijer, Joost P. A.; van der Heijde, Rob G. L.; Heethaar, Rob M. (1 January 2009).
255:. In land vertebrates the lens is flatter on its anterior side than on its posterior side, while in fish the lens is often close to spherical.
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695:
3D reconstruction of lens in a living 20 year old human male focusing from 0 dioptres (infinity) to 4.85 dioptres (26mm) side & back views
2019:"Transcriptional burst fraction and size dynamics during lens fiber cell differentiation and detailed insights into the denucleation process"
138:
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Gruijters, WT; Kistler, J; Bullivant, S (October 1987). "Formation, distribution and dissociation of intercellular junctions in the lens".
4093:
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Jagger, W. S; Sands, P. J (1 August 1999). "A wide-angle gradient index optical model of the crystalline lens and eye of the octopus".
3078:"Constant Volume of the Human Lens and Decrease in Surface Area of the Capsular Bag during Accommodation: An MRI and Scheimpflug Study"
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Schachar, Ronald A.; Bax, Andrew J. (June 2001). "Mechanism of human accommodation as analyzed by nonlinear finite element analysis".
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such as humans is unusually flat going some way to explain why our vision, unlike diving birds, is particularly blurry under water.
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Another important factor in maintaining the transparency of the lens is the absence of light-scattering organelles such as the
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Section through the margin of the lens, showing the transition of the epithelium into the lens fibers known as the bow region.
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2228:
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Sivak, J.G.; Hildebrand, T.; Lebert, C. (January 1985). "Magnitude and rate of accommodation in diving and nondiving birds".
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Bloemendal, Hans; De Jong, Wilfried; Jaenicke, Rainer; Lubsen, Nicolette H.; Slingsby, Christine; Tardieu, Annette (2004).
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Kuszak, J; Alcala, J; Maisel, H (December 1980). "The surface morphology of embryonic and adult chick lens-fiber cells".
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of human lens varies from approximately 1.406 in the central layers down to 1.386 in less dense layers of the lens. This
1968:
CantΓΉ, Claudio; Zimmerli, Dario; Hausmann, George; Valenta, Tomas; Moor, Andreas; Aguet, Michel; Basler, Konrad (2014).
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Kardong, K. (2008). Vertebrates: Comparative anatomy, function, evolution (5th ed.). (pp. 676β677). Boston: McGraw-Hill
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forming the outermost layer of the lens. Inside the capsule, much thinner lens fibers form the bulk of the lens. The
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Download and open with
Inkscape 9.1. The separate components reside on different "layers" to facilitated editing.
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Whikehart, David R. (2003). Biochemistry of the Eye, 2nd ed. 2003. Philadelphia: Butterworth
Heinemann, p. 107β8
1868:"Rac1 GTPase-deficient mouse lens exhibits defects in shape, suture formation, fiber cell migration and survival"
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is the absence of the lens from the eye. Aphakia can be the result of surgery or injury, or it can be congenital.
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Shi, Yanrong; Barton, Kelly; De Maria, Alicia; Petrash, J. Mark; Shiels, Alan; Bassnett, Steven (15 May 2009).
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870:, there are two muscles, one above and one below the lens, while other amphibians have only the lower muscle.
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is considered the master regulator gene of this organ. Other effectors of proper lens development include the
2295:"Focusing by shape change in the lens of the eye: a commentary on Young (1801) 'On the mechanism of the eye'"
2246:"Focusing by shape change in the lens of the eye: a commentary on Young (1801) 'On the mechanism of the eye'"
195:
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Candia, Oscar A. (2004). "Electrolyte and fluid transport across corneal, conjunctival and lens epithelia".
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of the eye, enabling them to focus on objects at various distances. This adjustment of the lens is known as
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4522:
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3748:
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Gruijters, WT (July 1989). "A non-connexon protein (MIP) is involved in eye lens gap-junction formation".
909:(OD) of the human crystalline lens for newborn, 30-year-old, and 65-year-old from wavelengths 300-1400 nm.
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Andrew M.Wood and Roger J.W.Truscott (March 1993). "UV Filters in Human Lenses: Tryptophan
Catabolism".
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are therefore manufactured to also block ultraviolet light. People lacking a lens (a condition known as
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catabolism in the lens epithelium. High intensity ultraviolet light can harm the retina, and artificial
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Accommodation in humans is well studied and allows artificial means of supplementing our focus such as
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3540:"Apoptosis in mammalian eye development: lens morphogenesis, vascular regression and immune privilege"
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1018:
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Streeten, B W (1977). "B W Streeten; The zonular insertion: a scanning electron microscopic study".
1797:"Lens placode planar cell polarity is dependent on Cdc42-mediated junctional contraction inhibition"
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By nine weeks into human development, the lens is surrounded and nourished by a net of vessels, the
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which allow water to flow into and out of cells are the most abundant membrane protein in the lens.
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This svg file was configured so that the rays, diaphragm and crystalline lens are easily modified
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1970:"Pax6-dependent, but Ξ²-catenin-independent, function of Bcl9 proteins in mouse lens development"
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Shi, Yanrong; Tu, Yidong; De Maria, Alicia; Mecham, Robert P.; Bassnett, Steven (1 April 2013).
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that completely surrounds the lens. The capsule is elastic and its main structural component is
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4331:
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2079:"Constant lens fiber cell thickness in fish suggests crystallin transport to denucleated cells"
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While not vertebrate, brief mention is made here of the convergent evolution of vertebrate and
858:, by contrast, a muscle projects from a vascular structure in the floor of the eye, called the
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3D lens model from sheep with parts labeled and images of cells from different parts overlayed
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which bathes the lens with nutrients and other things. Land vertebrate lenses usually have an
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3248:"Hyposmotic stress causes ATP release in a discrete zone within the outer cortex of rat lens"
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Diving bird (Cormorant) lens focusing can be up to 80 dioptres for clearer underwater vision.
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Giannone, Adrienne A.; Li, Leping; Sellitto, Caterina; White, Thomas W. (23 December 2021).
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Schachar, RA (March 1994). "Zonular function: a new hypothesis with clinical implications".
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3657:"Violet and blue light blocking intraocular lenses: Photoprotection versus photoreception"
1486:"Development, Composition, and Structural Arrangements of the Ciliary Zonule of the Mouse"
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pumps in the lens epithelial cells pump ions out of the lens to maintain appropriate lens
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Anderson, Robert M. (1983). "Visual
Perceptions and Observations of an Aphakic Surgeon".
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begins to deepen and bow inwards. As the placode continues to deepen, the opening to the
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Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences
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1821:
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1744:
1720:
1703:
1600:
1575:
1510:
1485:
1272:
1247:
988:
922:
863:
851:
746:
704:
407:
343:
Sheep lens fixed side view. Note the largest lens has damaged capsule and iris attached
3378:
2864:
Mulders, SM; Preston, GM; Deen, PM; Guggino, WB; van Os, CH; Agre, P (14 April 1995).
1413:
1396:
237:. In many land animals the shape of the lens can be altered, effectively changing the
4639:
4355:
4259:
4176:
4047:
3973:
3908:
3882:
3831:
3782:
3766:
3721:
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3512:
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3382:
3339:
3304:
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3041:
3029:
2994:
2963:
2932:
2897:
2846:
2842:
2762:
2727:
2700:
2696:
2669:
2657:
2622:
2576:
2541:
2484:
2441:
2403:
2324:
2275:
2224:
2186:
2182:
2143:
2112:
2100:
2050:
1999:
1950:
1916:
1897:
1850:
1826:
1774:
1725:
1662:
1605:
1553:
1515:
1466:
1452:
1418:
1375:
1354:
1277:
1263:
1222:
1039:
399:
360:
331:
252:
3733:
3524:
3478:
3394:
3146:
2774:
2496:
2453:
2198:
4624:
4562:
4399:
4052:
4004:
3872:
3862:
3821:
3811:
3713:
3676:
3668:
3629:
3592:
3551:
3504:
3458:
3421:
3374:
3351:
3331:
3296:
3259:
3212:
3189:
3169:
3126:
3089:
3021:
2986:
2955:
2924:
2887:
2877:
2838:
2809:
2754:
2692:
2649:
2568:
2531:
2523:
2476:
2433:
2393:
2389:
2385:
2314:
2306:
2265:
2257:
2178:
2155:
2135:
2090:
2040:
2030:
1989:
1981:
1940:
1932:
1887:
1879:
1816:
1808:
1764:
1756:
1715:
1652:
1595:
1587:
1545:
1505:
1497:
1456:
1448:
1408:
1267:
1259:
1202:
1047:
960:
785:
564:
541:
415:
359:, the lens epithelium, and the lens fibers. The lens capsule is a relatively thick
263:
3162:
American Journal of Physiology. Regulatory, Integrative and Comparative Physiology
4665:
4537:
4496:
4492:
4414:
3335:
3025:
2095:
2078:
1227:
1050:
involves the removal of the lens and insertion of an artificial intraocular lens.
811:
448:
377:
339:
222:
126:
3717:
1883:
1812:
1760:
758:
661:
An image that is partially in focus, but mostly out of focus in varying degrees.
347:
4542:
4532:
4517:
4512:
4209:
4201:
4181:
4057:
4032:
3173:
2653:
1795:
Muccioli, Maria; Qaisi, Dalya; Herman, Ken; Plageman, Timothy F. (April 2016).
1549:
1207:
1178:
1060:
984:
889:
840:
723:
714:
601:
572:
553:
364:
296:
292:
230:
218:
3867:
3816:
3462:
2683:
Coleman, D. Jackson (June 1970). "Unified Model for Accommodative Mechanism".
2437:
1687:
1657:
1640:
4654:
4567:
4502:
4339:
2814:
2789:
2745:
Huggert, Arne (27 May 2009). "The Intracapsular Mechanism of Accommodation".
2720:
Klinische MonatsblΓ€tter fΓΌr Augenheilkunde und fΓΌr augenΓ€rztliche Fortbildung
2035:
1009:
is the primary energy source for the lens. As mature lens fibers do not have
793:
545:
537:
469:
444:
308:
304:
283:
The lens is located towards the front part of the vertebrate eye, called the
198:
150:
86:
3851:"Ion Transport Regulation by TRPV4 and TRPV1 in Lens and Ciliary Epithelium"
3672:
3449:
Hoehenwarter, W.; Klose, J.; Jungblut, P. R. (2006). "Eye lens proteomics".
3426:
3409:
2882:
2865:
901:
4527:
4351:
4186:
4168:
3886:
3835:
3690:
3633:
3606:
3565:
3556:
3539:
3516:
3470:
3435:
3386:
3343:
3308:
3300:
3273:
3224:
3181:
3138:
3103:
3033:
2928:
2766:
2731:
2661:
2580:
2545:
2512:"Age-Related Changes in the Anterior Segment Biometry During Accommodation"
2445:
2374:"The force of contraction of the human ciliary muscle during accommodation"
2328:
2310:
2279:
2261:
2147:
2104:
2054:
2003:
1985:
1954:
1901:
1830:
1778:
1729:
1666:
1609:
1557:
1519:
1470:
1441:"A new eye lens structure associated with capsule/basement membrane growth"
1281:
1212:
1010:
941:
937:
929:
823:
Bony fish eye. Note the spherical lens and muscle to pull the lens backward
635:
Bony fish eye. Note the spherical lens and muscle to pull the lens backward
560:
549:
485:
356:
307:
which helps hold the lens in place. At the front of the lens is the liquid
300:
238:
206:
81:
3725:
3641:
3581:"Ageing and vision: Structure, stability and function of lens crystallins"
3495:
Andley, Usha P. (2007). "Crystallins in the eye: Function and pathology".
3130:
2998:
2967:
2936:
2901:
2704:
2626:
2572:
2527:
2488:
2190:
1501:
1422:
819:
729:
657:
631:
390:
386:
Sheep lens capsule removed. Decapsulation leads to a nearly formless blob.
382:
4619:
4488:
4445:
4383:
4347:
3094:
3077:
2990:
2959:
2850:
2407:
948:
885:
806:
680:
673:
In humans the widely quoted Helmholtz mechanism of focusing, also called
596:
the capsule and epithelium at the back and front of the lens, synthesize
583:
577:
476:
and interdigitations of the cells that resemble "ball and socket" forms.
440:
2346:(5th ed.). London and New York: MACMILLAN AND CO. pp. 256β258.
2218:
1639:
He, Mingyan; Zhou, Guangkai; Lin, Qinghong; Zhou, Nan (September 2023).
4433:
4191:
2892:
1936:
1591:
1191:
1053:
956:
913:
906:
770:
Wrinkled lens fibers in picture below compared to straight fibers above
766:
597:
481:
299:, attaching the lens at its equator to the rest of the eye through the
266:
of a younger human lens in its natural environment is approximately 18
226:
96:
3011:
2419:
2417:
2250:
Philosophical Transactions of the Royal Society B: Biological Sciences
2139:
1921:"The cellular and molecular mechanisms of vertebrate lens development"
1569:
1567:
940:
within the mature lens fibers. Lens fibers also have a very extensive
607:
Several proteins control the embryonic development of the lens though
480:
epithelium, are added to the outer cortex. Mature lens fibers have no
175:
4409:
4404:
4323:
3977:
1034:
844:
776:
521:
411:
312:
57:
3619:
3246:
Suzuki-Kerr, H; Walker, KL; Han, MH; Lim, JC; Donaldson, PJ (2022).
796:
of the lens. As more is learned about mammalian lens structure from
504:
39:
4327:
2414:
2223:. Philadelphia, PA: Holt-Saunders International. pp. 463β464.
1564:
1043:
994:
780:
742:
665:
403:
1017:. The remaining fraction of glucose is shunted primarily down the
295:
positioned in front of the lens. The lens is held in place by the
4579:
4480:
4424:
4140:
1349:
John Forrester, Andrew Dick, Paul McMenamin, William Lee (1996).
1066:
1006:
964:
917:
878:
874:
855:
828:
718:
641:
419:
394:
Eye lens micrographs and diagram of growth region of the capsule.
323:
267:
259:
74:
3849:
Delamere, Nicholas A.; Shahidullah, Mohammad (31 January 2022).
967:) perceive ultraviolet light as whitish blue or whitish-violet.
4574:
4484:
4238:
4024:
3996:
3578:
3408:
Schaeffel, F.; Murphy, C.J.; Howland, H.C. (15 November 1999).
836:
288:
234:
214:
669:
Eye and detailed ray path including one intraocular lens layer
4214:
3075:
2866:"Water channel properties of major intrinsic protein of lens"
2016:
1147:
Another view of the eye and the structures of the eye labeled
620:
109:
3286:
3159:
2077:
KozΕowski, Tomasz M.; KrΓΆger, Ronald H.H. (September 2019).
1967:
4128:
1794:
1745:"Epithelial morphogenesis: the mouse eye as a model system"
1370:
Yanoff, Myron; Duker, Jay S.; Augsburger, James J. (2008).
867:
832:
616:
608:
525:
Similar to a human, this is a lens forming in a chicken eye
2790:"Gullstrand Intracapsular Accommodation Mechanism Revised"
2467:
Atchison, DA (July 1995). "Accommodation and presbyopia".
3971:
3448:
3116:
2949:
2863:
1865:
1641:"The role of mip in the development of lens in zebrafish"
1627:
Encyclopædia Britannica 2006 Ultimate Reference Suite DVD
1063:
is the displacement of the lens from its normal position.
443:. As ions, nutrients, and liquid enter the lens from the
202:
3202:
1573:
3797:
3245:
2828:
3938: β Histology Learning System at Boston University
3407:
3321:
1742:
1013:, approximately 80% of the glucose is metabolized via
807:
Lenses of birds, reptiles, amphibians, fish and others
681:
The shape changing lens of many land based vertebrates
3410:"Accommodation in the cuttlefish (Sepia officinalis)"
2168:
1790:
1788:
1369:
544:. The first stage of lens formation takes place when
3800:"Physiological Mechanisms Regulating Lens Transport"
1168:
587:
Pattern of lens fibers (anterior and lateral aspect)
3848:
2639:
2219:Romer, Alfred Sherwood; Parsons, Thomas S. (1977).
1483:
2914:
1785:
491:
1914:
1743:Chauhan, B; Plageman, T; Lou, M; Lang, R (2015).
1679:
1576:"The stratified syncytium of the vertebrate lens"
1025:means that the lens consumes very little oxygen.
991:marks the former location of the hyaloid artery.
44:Lens of eye changing shape to focus near and far.
4652:
3082:Investigative Ophthalmology & Visual Science
2561:Investigative Ophthalmology & Visual Science
2516:Investigative Ophthalmology & Visual Science
1490:Investigative Ophthalmology & Visual Science
210:from the epithelium of the embryo before birth.
4086:
2509:
2076:
1374:(3rd ed.). Edinburgh: Mosby. p. 382.
896:
626:
351:Microscope image of lens cell types and capsule
3957:
3781:The Eye: Basic Sciences in Practice, p. 104,
2125:
1638:
532:of the vertebrate lens begins when the human
3585:Progress in Biophysics and Molecular Biology
1353:. London: W. B. Saunders Company Ltd. p. 28
3364:
2642:Biomechanics and Modeling in Mechanobiology
2423:
1842:
1840:
1625:"eye, human". Encyclopædia Britannica from
1434:
1432:
1111:The crystalline lens, hardened and divided.
3964:
3950:
3900:
3898:
3896:
3119:Klinische MonatsblΓ€tter fΓΌr Augenheilkunde
1621:
1619:
1394:
1323:. Medical-dictionary.thefreedictionary.com
398:The lens capsule is a smooth, transparent
50:
38:
3876:
3866:
3825:
3815:
3680:
3596:
3555:
3490:
3488:
3425:
3263:
3093:
2980:
2891:
2881:
2813:
2787:
2535:
2397:
2356:
2318:
2269:
2094:
2044:
2034:
1993:
1944:
1891:
1820:
1768:
1719:
1686:
1656:
1599:
1509:
1460:
1438:
1412:
1271:
3746:
3703:
3654:
2612:
2593:
2558:
2466:
1837:
1701:
1531:
1529:
1429:
1245:
1028:
993:
900:
888:. The most complex Molluscan eye is the
818:
810:
765:
757:
728:
703:
684:
664:
656:
630:
604:is very important for this development.
582:
520:
503:
495:
389:
381:
346:
338:
330:
322:
3893:
2744:
2682:
1749:Current Topics in Developmental Biology
1616:
262:for correction of sight as we age. The
14:
4653:
3494:
3485:
2371:
2341:
1535:
1439:Gruijters, Wouterus TM (1 July 2024).
1395:Mohan, PS; Spiro, RG (25 March 1986).
1345:
1343:
1341:
1339:
1337:
278:
3945:
2469:Ophthalmic & Physiological Optics
2214:
2212:
2210:
2208:
1526:
1363:
1046:is another risk factor for cataract.
297:suspensory ligaments (Zonule of Zinn)
3537:
3497:Progress in Retinal and Eye Research
3205:Progress in Retinal and Eye Research
2788:LΓ³pez-Gil, Norberto (3 March 2022).
2292:
2243:
1298:. Biology-Online.org. Archived from
1252:Progress in Retinal and Eye Research
1099:Interior of anterior chamber of eye.
1073:
303:. Behind the lens is the jelly-like
2870:The Journal of Biological Chemistry
2717:
2481:10.1046/j.1475-1313.1995.9500020e.x
1847:The Eye: Basic Sciences in Practice
1401:The Journal of Biological Chemistry
1351:The Eye: Basic Sciences in Practice
1334:
1296:"Equator of lens - definition from"
1087:MRI scan of human eye showing lens.
1042:is a type of age-related cataract.
355:The lens has three main parts: the
24:
4082:
3058:, 2nd ed. (1987), Addison Wesley,
2759:10.1111/j.1755-3768.1964.tb03627.x
2205:
753:
745:" model of lens focus proposed by
699:
429:
201:structure in most land vertebrate
25:
4677:
3929:
3538:Lang, Richard A. (January 1997).
2685:American Journal of Ophthalmology
1708:Journal of Anatomy and Physiology
1135:The structures of the eye labeled
998:Channels regulate lens transport.
873:In the simplest vertebrates, the
3661:British Journal of Ophthalmology
3598:10.1016/j.pbiomolbio.2003.11.012
3544:Cell Death & Differentiation
3509:10.1016/j.preteyeres.2006.10.003
3217:10.1016/j.preteyeres.2022.101152
2359:Treatise on physiological optics
2344:Lessons in Elementary Physiology
1453:10.17912/micropub.biology.000828
1264:10.1016/j.preteyeres.2020.100902
1171:
1152:
1140:
1128:
1116:
1104:
1092:
1080:
3917:
3842:
3791:
3775:
3759:
3747:Hambling, David (29 May 2002).
3740:
3697:
3648:
3613:
3572:
3531:
3442:
3414:Journal of Experimental Biology
3401:
3358:
3315:
3280:
3239:
3196:
3153:
3110:
3069:
3048:
3005:
2974:
2943:
2917:The American Journal of Anatomy
2908:
2857:
2822:
2781:
2738:
2711:
2676:
2633:
2606:
2587:
2552:
2503:
2460:
2372:Fisher, R. F. (1 August 1977).
2365:
2350:
2335:
2286:
2244:Land, Michael (19 April 2015).
2237:
2162:
2128:Journal of Experimental Biology
2119:
2070:
2061:
2023:Journal of Biological Chemistry
2010:
1961:
1908:
1859:
1736:
1695:
1673:
1632:
559:As development progresses, the
540:, the lens is derived from the
492:Cell fusion, voids and vacuoles
371:
2390:10.1113/jphysiol.1977.sp011938
1477:
1388:
1313:
1288:
1239:
970:
516:
462:
13:
1:
3379:10.1016/S0042-6989(99)00012-7
1414:10.1016/S0021-9258(17)35665-X
1246:Bassnett, Steven (May 2021).
1233:
866:in either type of animal. In
248:
4610:Optical coherence tomography
4364:Photosensitive ganglion cell
3336:10.1016/j.visres.2005.03.008
3026:10.1097/OPX.0b013e31818e8d57
3014:Optometry and Vision Science
2843:10.1016/0014-4835(76)90135-4
2697:10.1016/0002-9394(70)91057-3
2596:Invest. Ophthalmol. Vis. Sci
2183:10.1016/0042-6989(85)90203-2
2096:10.1016/j.visres.2019.06.008
983:, which is derived from the
897:Crystallins and transparency
733:Schachar model of lens focus
627:Variations in lens structure
273:
7:
4360:Giant retina ganglion cells
4149:Capillary lamina of choroid
3718:10.2466/pms.1983.57.3f.1211
3706:Perceptual and Motor Skills
1884:10.1016/j.ydbio.2011.09.004
1813:10.1016/j.ydbio.2016.02.016
1761:10.1016/bs.ctdb.2014.11.011
1702:Mitchell, PC (April 1891).
1164:
652:
647:
10:
4682:
4304:Retinal pigment epithelium
4294:External limiting membrane
3174:10.1152/ajpregu.00173.2011
2985:. 93 ( Pt 3) (3): 509β13.
2654:10.1007/s10237-021-01417-9
2357:Helmholtz, H. von (1962).
1550:10.1016/j.exer.2003.08.015
1199:, for how the lens evolved
375:
318:
4592:
4553:
4467:
4458:
4392:
4312:
4247:
4237:
4200:
4167:
4139:
4127:
4080:
4023:
3995:
3984:
3936:Histology image: 08001loa
3868:10.3389/fphys.2021.834916
3817:10.3389/fphys.2021.818649
3622:Experimental Eye Research
3463:10.1007/s00726-005-0283-9
3289:Experimental Eye Research
2954:. 88 ( Pt 3) (3): 351β9.
2831:Experimental Eye Research
2438:10.1007/s12019-996-0006-5
2378:The Journal of Physiology
2342:Huxley, Thomas H (1871).
2293:Land, M (19 April 2015).
1688:10.1101/2023.01.17.524320
1658:10.1016/j.gep.2023.119330
1538:Experimental Eye Research
1019:pentose phosphate pathway
168:
156:
144:
132:
120:
108:
103:
92:
80:
70:
65:
56:Schematic diagram of the
49:
37:
32:
3749:"Let the light shine in"
3655:Mainster, M. A. (2006).
2815:10.3390/photonics9030152
2036:10.1074/jbc.RA118.001927
1645:Gene Expression Patterns
1445:MicroPublication Biology
955:glucoside, a product of
4299:Layer of rods and cones
4255:Inner limiting membrane
3905:Biochemistry of the Eye
3855:Frontiers in Physiology
3804:Frontiers in Physiology
3673:10.1136/bjo.2005.086553
3427:10.1242/jeb.202.22.3127
2983:Journal of Cell Science
2952:Journal of Cell Science
2883:10.1074/jbc.270.15.9010
2615:Annals of Ophthalmology
1974:Genes & Development
1580:Journal of Cell Science
981:tunica vasculosa lentis
801:Scheimpflug photography
4121:
3712:(3_suppl): 1211β1218.
3634:10.1006/exer.1993.1041
3557:10.1038/sj.cdd.4400211
3301:10.1006/exer.1999.0747
2929:10.1002/aja.1001590406
2311:10.1098/rstb.2014.0308
2262:10.1098/rstb.2014.0308
1986:10.1101/gad.246140.114
999:
910:
824:
816:
771:
763:
734:
710:
696:
670:
662:
636:
588:
571:The embryo then sends
542:skin around the embryo
526:
509:
501:
395:
387:
352:
344:
336:
328:
170:Anatomical terminology
4615:Eye care professional
4420:Foveal avascular zone
4282:Outer plexiform layer
4270:Inner plexiform layer
4225:Iris sphincter muscle
4118:
3131:10.1055/s-0042-118599
2573:10.1167/iovs.15-17452
2528:10.1167/iovs.15-16825
2426:Comprehensive Therapy
1872:Developmental Biology
1801:Developmental Biology
1704:"Double Chick Embryo"
1502:10.1167/iovs.13-11619
1029:Clinical significance
997:
947:The lens blocks most
934:endoplasmic reticulum
904:
822:
814:
769:
761:
732:
707:
694:
668:
660:
634:
586:
524:
507:
499:
474:intercellular bridges
453:osmotic concentration
393:
385:
350:
342:
334:
326:
287:, which includes the
4635:Physiological Optics
4605:Ocular immune system
4344:Retina ganglion cell
3907:, 2nd ed, p. 107β8,
3095:10.1167/iovs.08-2124
2991:10.1242/jcs.93.3.509
2960:10.1242/jcs.88.3.351
2747:Acta Ophthalmologica
1197:Evolution of the eye
1187:Accommodation reflex
1015:anaerobic metabolism
719:suspending ligaments
4459:Anatomical regions
4320:Photoreceptor cells
4287:Outer nuclear layer
4275:Inner nuclear layer
4265:Ganglion cell layer
4220:Iris dilator muscle
4015:Trabecular meshwork
2806:2022Photo...9..152L
2718:PAU, H (1952). "".
2221:The Vertebrate Body
2029:(34): 13176β13190.
1714:(Pt 3): 316β324.1.
1218:Phacoemulsification
1023:aerobic respiration
953:3-hydroxykynurenine
538:inner embryo layers
279:Position in the eye
4122:
2256:(1666): 20140308.
1937:10.1242/dev.107953
1592:10.1242/jcs.045203
1203:Intraocular lenses
1000:
961:intraocular lenses
923:chaperone proteins
916:are water-soluble
911:
852:cartilaginous fish
825:
817:
772:
764:
735:
711:
697:
671:
663:
637:
589:
527:
510:
502:
408:glycosaminoglycans
396:
388:
353:
345:
337:
329:
4661:Human eye anatomy
4648:
4647:
4640:Visual perception
4588:
4587:
4555:Posterior segment
4523:Posterior chamber
4454:
4453:
4356:Bistratified cell
4260:Nerve fiber layer
4233:
4232:
4177:Ciliary processes
4078:
4077:
3420:(22): 3127β3134.
3373:(17): 2841β2852.
3330:(18): 2352β2366.
3125:(12): 1320β1323.
2230:978-0-03-910284-5
2140:10.1242/jeb.00142
1980:(17): 1879β1884.
1931:(23): 4432β4447.
1586:(10): 1607β1615.
1223:Visual perception
1074:Additional images
1040:Nuclear sclerosis
860:falciform process
692:
546:a sphere of cells
400:basement membrane
361:basement membrane
184:
183:
179:
16:(Redirected from
4673:
4625:Refractive error
4563:Vitreous chamber
4508:Anterior chamber
4469:Anterior segment
4465:
4464:
4245:
4244:
4154:Bruch's membrane
4137:
4136:
4129:Uvea / vascular
4085:
4005:Episcleral layer
3993:
3992:
3966:
3959:
3952:
3943:
3942:
3924:
3921:
3915:
3902:
3891:
3890:
3880:
3870:
3846:
3840:
3839:
3829:
3819:
3795:
3789:
3779:
3773:
3763:
3757:
3756:
3744:
3738:
3737:
3701:
3695:
3694:
3684:
3652:
3646:
3645:
3617:
3611:
3610:
3600:
3576:
3570:
3569:
3559:
3535:
3529:
3528:
3492:
3483:
3482:
3446:
3440:
3439:
3429:
3405:
3399:
3398:
3362:
3356:
3355:
3319:
3313:
3312:
3284:
3278:
3277:
3267:
3252:Molecular Vision
3243:
3237:
3236:
3200:
3194:
3193:
3157:
3151:
3150:
3114:
3108:
3107:
3097:
3073:
3067:
3052:
3046:
3045:
3009:
3003:
3002:
2978:
2972:
2971:
2947:
2941:
2940:
2912:
2906:
2905:
2895:
2885:
2861:
2855:
2854:
2826:
2820:
2819:
2817:
2785:
2779:
2778:
2742:
2736:
2735:
2715:
2709:
2708:
2691:(6): 1063β1079.
2680:
2674:
2673:
2637:
2631:
2630:
2610:
2604:
2603:
2591:
2585:
2584:
2556:
2550:
2549:
2539:
2522:(6): 3522β3530.
2507:
2501:
2500:
2464:
2458:
2457:
2421:
2412:
2411:
2401:
2369:
2363:
2362:
2354:
2348:
2347:
2339:
2333:
2332:
2322:
2290:
2284:
2283:
2273:
2241:
2235:
2234:
2216:
2203:
2202:
2166:
2160:
2159:
2123:
2117:
2116:
2098:
2074:
2068:
2065:
2059:
2058:
2048:
2038:
2014:
2008:
2007:
1997:
1965:
1959:
1958:
1948:
1917:Ashery-Padan, R.
1912:
1906:
1905:
1895:
1863:
1857:
1844:
1835:
1834:
1824:
1792:
1783:
1782:
1772:
1740:
1734:
1733:
1723:
1699:
1693:
1692:
1690:
1677:
1671:
1670:
1660:
1636:
1630:
1623:
1614:
1613:
1603:
1571:
1562:
1561:
1533:
1524:
1523:
1513:
1496:(4): 2504β2515.
1481:
1475:
1474:
1464:
1436:
1427:
1426:
1416:
1392:
1386:
1385:
1367:
1361:
1347:
1332:
1331:
1329:
1328:
1317:
1311:
1310:
1308:
1307:
1292:
1286:
1285:
1275:
1243:
1181:
1176:
1175:
1174:
1156:
1144:
1132:
1120:
1108:
1096:
1084:
1048:Cataract surgery
786:refractive index
693:
565:surface ectoderm
416:type IV collagen
285:anterior segment
264:refractive power
223:vitreous humours
192:crystalline lens
176:edit on Wikidata
173:
54:
42:
30:
29:
21:
4681:
4680:
4676:
4675:
4674:
4672:
4671:
4670:
4651:
4650:
4649:
4644:
4584:
4549:
4538:Capsule of lens
4493:Lacrimal system
4460:
4450:
4410:Parafoveal area
4405:Perifoveal area
4388:
4332:Horizontal cell
4308:
4229:
4196:
4163:
4159:Sattler's layer
4130:
4123:
4117:
4074:
4019:
4010:Schlemm's canal
3988:
3980:
3972:Anatomy of the
3970:
3932:
3927:
3922:
3918:
3903:
3894:
3847:
3843:
3796:
3792:
3780:
3776:
3764:
3760:
3745:
3741:
3702:
3698:
3653:
3649:
3618:
3614:
3577:
3573:
3536:
3532:
3493:
3486:
3447:
3443:
3406:
3402:
3367:Vision Research
3363:
3359:
3324:Vision Research
3320:
3316:
3285:
3281:
3244:
3240:
3201:
3197:
3158:
3154:
3115:
3111:
3074:
3070:
3054:Hecht, Eugene.
3053:
3049:
3020:(12): 1179β84.
3010:
3006:
2979:
2975:
2948:
2944:
2913:
2909:
2876:(15): 9010β16.
2862:
2858:
2827:
2823:
2786:
2782:
2743:
2739:
2716:
2712:
2681:
2677:
2638:
2634:
2611:
2607:
2592:
2588:
2557:
2553:
2508:
2504:
2465:
2461:
2422:
2415:
2370:
2366:
2355:
2351:
2340:
2336:
2291:
2287:
2242:
2238:
2231:
2217:
2206:
2171:Vision Research
2167:
2163:
2124:
2120:
2083:Vision Research
2075:
2071:
2066:
2062:
2015:
2011:
1966:
1962:
1913:
1909:
1864:
1860:
1845:
1838:
1793:
1786:
1741:
1737:
1700:
1696:
1678:
1674:
1637:
1633:
1624:
1617:
1572:
1565:
1534:
1527:
1482:
1478:
1437:
1430:
1393:
1389:
1382:
1368:
1364:
1348:
1335:
1326:
1324:
1319:
1318:
1314:
1305:
1303:
1294:
1293:
1289:
1248:"Zinn's zonule"
1244:
1240:
1236:
1228:Zonules of Zinn
1177:
1172:
1170:
1167:
1160:
1157:
1148:
1145:
1136:
1133:
1124:
1121:
1112:
1109:
1100:
1097:
1088:
1085:
1076:
1031:
989:Cloquet's canal
973:
907:optical density
899:
809:
756:
754:Internal forces
702:
700:External forces
685:
683:
655:
650:
629:
519:
494:
465:
432:
430:Lens epithelium
380:
378:Capsule of lens
374:
321:
281:
276:
213:Along with the
180:
115:lens crystallin
61:
45:
28:
23:
22:
15:
12:
11:
5:
4679:
4669:
4668:
4663:
4646:
4645:
4643:
4642:
4637:
4632:
4627:
4622:
4617:
4612:
4607:
4602:
4596:
4594:
4590:
4589:
4586:
4585:
4583:
4582:
4577:
4572:
4571:
4570:
4559:
4557:
4551:
4550:
4548:
4547:
4546:
4545:
4543:Zonule of Zinn
4540:
4530:
4525:
4520:
4515:
4513:Aqueous humour
4510:
4505:
4500:
4473:
4471:
4462:
4456:
4455:
4452:
4451:
4449:
4448:
4443:
4442:
4441:
4431:
4430:
4429:
4428:
4427:
4422:
4412:
4407:
4396:
4394:
4390:
4389:
4387:
4386:
4316:
4314:
4310:
4309:
4307:
4306:
4301:
4296:
4290:
4289:
4284:
4278:
4277:
4272:
4267:
4262:
4257:
4251:
4249:
4242:
4235:
4234:
4231:
4230:
4228:
4227:
4222:
4217:
4212:
4206:
4204:
4198:
4197:
4195:
4194:
4189:
4184:
4182:Ciliary muscle
4179:
4173:
4171:
4165:
4164:
4162:
4161:
4156:
4151:
4145:
4143:
4134:
4125:
4124:
4081:
4079:
4076:
4075:
4073:
4072:
4071:
4070:
4065:
4060:
4055:
4050:
4045:
4035:
4029:
4027:
4021:
4020:
4018:
4017:
4012:
4007:
4001:
3999:
3990:
3982:
3981:
3969:
3968:
3961:
3954:
3946:
3940:
3939:
3931:
3930:External links
3928:
3926:
3925:
3916:
3892:
3841:
3790:
3774:
3758:
3739:
3696:
3667:(6): 784β792.
3647:
3628:(3): 317β325.
3612:
3591:(3): 407β485.
3571:
3530:
3484:
3457:(4): 369β389.
3441:
3400:
3357:
3314:
3279:
3238:
3195:
3168:(2): R335-42.
3152:
3109:
3088:(1): 281β289.
3068:
3047:
3004:
2973:
2942:
2923:(4): 395β410.
2907:
2856:
2837:(3): 365β371.
2821:
2780:
2753:(2): 389β397.
2737:
2710:
2675:
2648:(3): 879β894.
2632:
2605:
2586:
2551:
2502:
2459:
2432:(2): 122β132.
2413:
2364:
2349:
2334:
2285:
2236:
2229:
2204:
2177:(7): 925β933.
2161:
2134:(5): 833β841.
2118:
2069:
2060:
2009:
1960:
1907:
1858:
1836:
1784:
1735:
1694:
1672:
1631:
1615:
1563:
1544:(3): 527β535.
1525:
1476:
1428:
1407:(9): 4328β36.
1387:
1381:978-0323057516
1380:
1362:
1333:
1312:
1287:
1237:
1235:
1232:
1231:
1230:
1225:
1220:
1215:
1210:
1205:
1200:
1194:
1189:
1183:
1182:
1179:Medical portal
1166:
1163:
1162:
1161:
1158:
1151:
1149:
1146:
1139:
1137:
1134:
1127:
1125:
1122:
1115:
1113:
1110:
1103:
1101:
1098:
1091:
1089:
1086:
1079:
1075:
1072:
1071:
1070:
1064:
1061:Ectopia lentis
1058:
1051:
1030:
1027:
1021:. The lack of
985:hyaloid artery
972:
969:
905:Graph showing
898:
895:
890:Cephalopod eye
886:Molluscan eyes
841:vision in fish
808:
805:
790:index gradient
755:
752:
724:focal distance
715:ciliary muscle
701:
698:
682:
679:
654:
651:
649:
646:
628:
625:
518:
515:
493:
490:
464:
461:
431:
428:
376:Main article:
373:
370:
320:
317:
280:
277:
275:
272:
182:
181:
172:
166:
165:
160:
154:
153:
148:
142:
141:
136:
130:
129:
124:
118:
117:
112:
106:
105:
101:
100:
94:
90:
89:
84:
78:
77:
72:
68:
67:
63:
62:
55:
47:
46:
43:
35:
34:
26:
9:
6:
4:
3:
2:
4678:
4667:
4664:
4662:
4659:
4658:
4656:
4641:
4638:
4636:
4633:
4631:
4630:Accommodation
4628:
4626:
4623:
4621:
4618:
4616:
4613:
4611:
4608:
4606:
4603:
4601:
4598:
4597:
4595:
4591:
4581:
4578:
4576:
4573:
4569:
4568:Vitreous body
4566:
4565:
4564:
4561:
4560:
4558:
4556:
4552:
4544:
4541:
4539:
4536:
4535:
4534:
4531:
4529:
4526:
4524:
4521:
4519:
4516:
4514:
4511:
4509:
4506:
4504:
4503:Fibrous tunic
4501:
4498:
4494:
4490:
4486:
4482:
4478:
4475:
4474:
4472:
4470:
4466:
4463:
4457:
4447:
4444:
4440:
4437:
4436:
4435:
4432:
4426:
4423:
4421:
4418:
4417:
4416:
4413:
4411:
4408:
4406:
4403:
4402:
4401:
4398:
4397:
4395:
4391:
4385:
4381:
4377:
4373:
4369:
4365:
4361:
4357:
4353:
4349:
4345:
4341:
4340:Amacrine cell
4337:
4333:
4329:
4325:
4321:
4318:
4317:
4315:
4311:
4305:
4302:
4300:
4297:
4295:
4292:
4291:
4288:
4285:
4283:
4280:
4279:
4276:
4273:
4271:
4268:
4266:
4263:
4261:
4258:
4256:
4253:
4252:
4250:
4246:
4243:
4240:
4236:
4226:
4223:
4221:
4218:
4216:
4213:
4211:
4208:
4207:
4205:
4203:
4199:
4193:
4190:
4188:
4185:
4183:
4180:
4178:
4175:
4174:
4172:
4170:
4166:
4160:
4157:
4155:
4152:
4150:
4147:
4146:
4144:
4142:
4138:
4135:
4132:
4126:
4069:
4066:
4064:
4061:
4059:
4056:
4054:
4051:
4049:
4046:
4044:
4041:
4040:
4039:
4036:
4034:
4031:
4030:
4028:
4026:
4022:
4016:
4013:
4011:
4008:
4006:
4003:
4002:
4000:
3998:
3994:
3991:
3987:
3986:Fibrous tunic
3983:
3979:
3975:
3967:
3962:
3960:
3955:
3953:
3948:
3947:
3944:
3937:
3934:
3933:
3920:
3914:
3913:0-7506-7152-1
3910:
3906:
3901:
3899:
3897:
3888:
3884:
3879:
3874:
3869:
3864:
3860:
3856:
3852:
3845:
3837:
3833:
3828:
3823:
3818:
3813:
3809:
3805:
3801:
3794:
3788:
3787:0-7020-1790-6
3784:
3778:
3772:
3771:0-7506-7152-1
3768:
3762:
3754:
3750:
3743:
3735:
3731:
3727:
3723:
3719:
3715:
3711:
3707:
3700:
3692:
3688:
3683:
3678:
3674:
3670:
3666:
3662:
3658:
3651:
3643:
3639:
3635:
3631:
3627:
3623:
3616:
3608:
3604:
3599:
3594:
3590:
3586:
3582:
3575:
3567:
3563:
3558:
3553:
3549:
3545:
3541:
3534:
3526:
3522:
3518:
3514:
3510:
3506:
3502:
3498:
3491:
3489:
3480:
3476:
3472:
3468:
3464:
3460:
3456:
3452:
3445:
3437:
3433:
3428:
3423:
3419:
3415:
3411:
3404:
3396:
3392:
3388:
3384:
3380:
3376:
3372:
3368:
3361:
3353:
3349:
3345:
3341:
3337:
3333:
3329:
3325:
3318:
3310:
3306:
3302:
3298:
3294:
3290:
3283:
3275:
3271:
3266:
3261:
3257:
3253:
3249:
3242:
3234:
3230:
3226:
3222:
3218:
3214:
3210:
3206:
3199:
3191:
3187:
3183:
3179:
3175:
3171:
3167:
3163:
3156:
3148:
3144:
3140:
3136:
3132:
3128:
3124:
3120:
3113:
3105:
3101:
3096:
3091:
3087:
3083:
3079:
3072:
3065:
3064:0-201-11609-X
3061:
3057:
3051:
3043:
3039:
3035:
3031:
3027:
3023:
3019:
3015:
3008:
3000:
2996:
2992:
2988:
2984:
2977:
2969:
2965:
2961:
2957:
2953:
2946:
2938:
2934:
2930:
2926:
2922:
2918:
2911:
2903:
2899:
2894:
2889:
2884:
2879:
2875:
2871:
2867:
2860:
2852:
2848:
2844:
2840:
2836:
2832:
2825:
2816:
2811:
2807:
2803:
2799:
2795:
2791:
2784:
2776:
2772:
2768:
2764:
2760:
2756:
2752:
2748:
2741:
2733:
2729:
2725:
2721:
2714:
2706:
2702:
2698:
2694:
2690:
2686:
2679:
2671:
2667:
2663:
2659:
2655:
2651:
2647:
2643:
2636:
2628:
2624:
2620:
2616:
2609:
2602:(4): 364β375.
2601:
2597:
2590:
2582:
2578:
2574:
2570:
2566:
2562:
2555:
2547:
2543:
2538:
2533:
2529:
2525:
2521:
2517:
2513:
2506:
2498:
2494:
2490:
2486:
2482:
2478:
2475:(4): 255β72.
2474:
2470:
2463:
2455:
2451:
2447:
2443:
2439:
2435:
2431:
2427:
2420:
2418:
2409:
2405:
2400:
2395:
2391:
2387:
2383:
2379:
2375:
2368:
2360:
2353:
2345:
2338:
2330:
2326:
2321:
2316:
2312:
2308:
2304:
2300:
2296:
2289:
2281:
2277:
2272:
2267:
2263:
2259:
2255:
2251:
2247:
2240:
2232:
2226:
2222:
2215:
2213:
2211:
2209:
2200:
2196:
2192:
2188:
2184:
2180:
2176:
2172:
2165:
2157:
2153:
2149:
2145:
2141:
2137:
2133:
2129:
2122:
2114:
2110:
2106:
2102:
2097:
2092:
2088:
2084:
2080:
2073:
2064:
2056:
2052:
2047:
2042:
2037:
2032:
2028:
2024:
2020:
2013:
2005:
2001:
1996:
1991:
1987:
1983:
1979:
1975:
1971:
1964:
1956:
1952:
1947:
1942:
1938:
1934:
1930:
1926:
1922:
1918:
1911:
1903:
1899:
1894:
1889:
1885:
1881:
1877:
1873:
1869:
1862:
1856:
1855:0-7020-1790-6
1852:
1848:
1843:
1841:
1832:
1828:
1823:
1818:
1814:
1810:
1806:
1802:
1798:
1791:
1789:
1780:
1776:
1771:
1766:
1762:
1758:
1754:
1750:
1746:
1739:
1731:
1727:
1722:
1717:
1713:
1709:
1705:
1698:
1689:
1684:
1676:
1668:
1664:
1659:
1654:
1650:
1646:
1642:
1635:
1628:
1622:
1620:
1611:
1607:
1602:
1597:
1593:
1589:
1585:
1581:
1577:
1570:
1568:
1559:
1555:
1551:
1547:
1543:
1539:
1532:
1530:
1521:
1517:
1512:
1507:
1503:
1499:
1495:
1491:
1487:
1480:
1472:
1468:
1463:
1458:
1454:
1450:
1446:
1442:
1435:
1433:
1424:
1420:
1415:
1410:
1406:
1402:
1398:
1391:
1383:
1377:
1373:
1372:Ophthalmology
1366:
1360:
1359:0-7020-1790-6
1356:
1352:
1346:
1344:
1342:
1340:
1338:
1322:
1316:
1302:on 2012-03-22
1301:
1297:
1291:
1283:
1279:
1274:
1269:
1265:
1261:
1257:
1253:
1249:
1242:
1238:
1229:
1226:
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1216:
1214:
1211:
1209:
1206:
1204:
1201:
1198:
1195:
1193:
1190:
1188:
1185:
1184:
1180:
1169:
1155:
1150:
1143:
1138:
1131:
1126:
1119:
1114:
1107:
1102:
1095:
1090:
1083:
1078:
1077:
1068:
1065:
1062:
1059:
1055:
1052:
1049:
1045:
1041:
1036:
1033:
1032:
1026:
1024:
1020:
1016:
1012:
1008:
1004:
996:
992:
990:
986:
982:
977:
968:
966:
962:
958:
954:
950:
945:
943:
939:
935:
931:
926:
924:
919:
915:
908:
903:
894:
891:
887:
882:
880:
876:
871:
869:
865:
861:
857:
853:
848:
846:
842:
838:
834:
830:
821:
813:
804:
802:
799:
795:
794:optical power
792:enhances the
791:
787:
782:
778:
768:
760:
751:
748:
744:
739:
731:
727:
725:
720:
716:
706:
678:
676:
675:accommodation
667:
659:
645:
643:
633:
624:
622:
618:
614:
613:Wnt signaling
610:
605:
603:
599:
593:
585:
581:
579:
574:
569:
566:
562:
557:
555:
551:
547:
543:
539:
535:
531:
523:
514:
506:
498:
489:
487:
483:
477:
475:
471:
470:gap junctions
460:
457:
454:
450:
446:
445:aqueous humor
442:
437:
427:
423:
421:
417:
413:
409:
405:
401:
392:
384:
379:
369:
366:
362:
358:
349:
341:
333:
325:
316:
314:
310:
309:aqueous humor
306:
305:vitreous body
302:
298:
294:
290:
286:
271:
269:
265:
261:
256:
254:
250:
246:
245:
244:accommodation
240:
236:
232:
228:
224:
220:
216:
211:
208:
204:
200:
197:
193:
189:
177:
171:
167:
164:
161:
159:
155:
152:
149:
147:
143:
140:
137:
135:
131:
128:
125:
123:
119:
116:
113:
111:
107:
102:
98:
95:
91:
88:
87:Visual system
85:
83:
79:
76:
73:
69:
64:
59:
53:
48:
41:
36:
31:
27:Eye structure
19:
4634:
4528:Ciliary body
4368:Diencephalon
4367:
4352:Parasol cell
4336:Bipolar cell
4187:Pars plicata
4169:Ciliary body
4037:
3919:
3904:
3858:
3854:
3844:
3807:
3803:
3793:
3777:
3761:
3753:The Guardian
3752:
3742:
3709:
3705:
3699:
3664:
3660:
3650:
3625:
3621:
3615:
3588:
3584:
3574:
3550:(1): 12β20.
3547:
3543:
3533:
3503:(1): 78β98.
3500:
3496:
3454:
3450:
3444:
3417:
3413:
3403:
3370:
3366:
3360:
3327:
3323:
3317:
3295:(6): 663β9.
3292:
3288:
3282:
3255:
3251:
3241:
3208:
3204:
3198:
3165:
3161:
3155:
3122:
3118:
3112:
3085:
3081:
3071:
3055:
3050:
3017:
3013:
3007:
2982:
2976:
2951:
2945:
2920:
2916:
2910:
2873:
2869:
2859:
2834:
2830:
2824:
2797:
2793:
2783:
2750:
2746:
2740:
2726:(2): 224β6.
2723:
2719:
2713:
2688:
2684:
2678:
2645:
2641:
2635:
2618:
2614:
2608:
2599:
2595:
2589:
2567:(10): 6075.
2564:
2560:
2554:
2519:
2515:
2505:
2472:
2468:
2462:
2429:
2425:
2384:(1): 51β74.
2381:
2377:
2367:
2358:
2352:
2343:
2337:
2302:
2298:
2288:
2253:
2249:
2239:
2220:
2174:
2170:
2164:
2131:
2127:
2121:
2086:
2082:
2072:
2063:
2026:
2022:
2012:
1977:
1973:
1963:
1928:
1924:
1910:
1878:(1): 30β43.
1875:
1871:
1861:
1846:
1807:(1): 32β43.
1804:
1800:
1752:
1748:
1738:
1711:
1707:
1697:
1675:
1648:
1644:
1634:
1583:
1579:
1541:
1537:
1493:
1489:
1479:
1444:
1404:
1400:
1390:
1371:
1365:
1350:
1325:. Retrieved
1315:
1304:. Retrieved
1300:the original
1290:
1255:
1251:
1241:
1213:Lens capsule
1011:mitochondria
1005:
1001:
978:
974:
946:
942:cytoskeleton
938:mitochondria
927:
912:
883:
872:
859:
849:
826:
797:
773:
740:
736:
712:
672:
638:
606:
594:
590:
570:
561:lens placode
558:
550:lens placode
528:
511:
478:
466:
458:
433:
424:
397:
372:Lens capsule
357:lens capsule
354:
301:ciliary body
282:
257:
242:
239:focal length
233:it onto the
212:
207:lens capsule
191:
187:
185:
139:A15.2.05.001
114:
4620:Eye disease
4600:Keratocytes
4489:Conjunctiva
4446:Ora serrata
4384:Muller glia
4348:Midget cell
4068:Endothelium
4058:Dua's layer
3451:Amino Acids
3258:: 245β256.
2893:2066/216433
2621:(2): 36β8.
1925:Development
1915:Cvekl, A.;
971:Nourishment
949:ultraviolet
914:Crystallins
615:components
578:crystallins
530:Development
517:Development
463:Lens fibers
449:Na/K-ATPase
441:homeostasis
196:transparent
104:Identifiers
4655:Categories
4461:of the eye
4434:Optic disc
4192:Pars plana
4063:Descemet's
4043:Epithelium
3861:: 834916.
3810:: 818649.
3211:: 101152.
2800:(3): 152.
1849:, p. 102,
1755:: 375β99.
1651:: 119330.
1327:2012-11-25
1306:2012-11-25
1258:: 100902.
1234:References
1192:Crystallin
1057:condition.
1054:Presbyopia
957:tryptophan
845:amphibians
777:aquaporins
598:crystallin
482:organelles
436:epithelium
247:(see also
4439:Optic cup
4324:Cone cell
3978:human eye
3233:254243790
3066:. p. 178.
3042:205907383
2794:Photonics
2670:231704221
2113:195820065
2089:: 29β34.
1035:Cataracts
781:Connexins
434:The lens
410:(GAGs)),
313:ellipsoid
274:Structure
58:human eye
4328:Rod cell
4133:(middle)
4048:Bowman's
3887:35173627
3836:35002784
3734:20005737
3691:16714268
3607:15302206
3566:16465205
3525:29317220
3517:17166758
3479:19978371
3471:16583312
3436:10539961
3395:17808919
3387:10492814
3344:15979462
3309:10620395
3274:36284672
3225:36470825
3182:21593426
3147:78808282
3139:27984837
3104:18676625
3034:19050472
2775:37325357
2767:14213923
2732:14955961
2662:33491156
2581:26393665
2546:26030106
2497:24282106
2454:71369369
2446:11430259
2329:25750232
2305:(1666).
2280:25750232
2199:42368520
2148:12547938
2105:31278970
2055:29959226
2004:25184676
1955:25406393
1919:(2014).
1902:21945075
1831:26902112
1779:25662266
1730:17231922
1667:37369320
1610:19401333
1558:15106931
1520:23493297
1471:39139582
1462:11320119
1282:32980533
1165:See also
1044:Diabetes
918:proteins
875:lampreys
856:teleosts
829:reptiles
743:catenary
653:Focusing
648:Function
642:primates
412:entactin
404:collagen
268:dioptres
231:focusing
227:refracts
199:biconvex
93:Function
18:Eye lens
4580:Choroid
4481:Eyebrow
4425:Foveola
4241:(inner)
4141:Choroid
3989:(outer)
3976:of the
3878:8841554
3827:8735835
3726:6664798
3682:1860240
3642:8472787
3352:8894700
3265:9514545
3190:9525037
2999:2691517
2968:3448099
2937:7223675
2902:7536742
2802:Bibcode
2705:5423772
2627:8010701
2537:4464043
2489:7667018
2399:1353417
2320:4360117
2271:4360117
2191:4049742
2156:3096767
2046:6109918
1995:4197948
1946:4302924
1893:3215831
1822:7370377
1770:6014593
1721:1328169
1683:bioRxiv
1601:2680101
1511:3621578
1423:3512568
1273:8139560
1067:Aphakia
1007:Glucose
965:aphakia
930:nucleus
879:hagfish
864:similar
839:. With
837:mammals
798:in situ
747:Coleman
573:signals
420:laminin
319:Anatomy
260:glasses
229:light,
219:aqueous
194:, is a
127:D007908
97:Refract
75:Eyeball
71:Part of
66:Details
4666:Lenses
4575:Retina
4485:Eyelid
4477:Adnexa
4400:Macula
4380:K cell
4376:M cell
4372:P cell
4248:Layers
4239:Retina
4210:Stroma
4053:Stroma
4038:layers
4033:Limbus
4025:Cornea
3997:Sclera
3911:
3885:
3875:
3834:
3824:
3785:
3769:
3732:
3724:
3689:
3679:
3640:
3605:
3564:
3523:
3515:
3477:
3469:
3434:
3393:
3385:
3350:
3342:
3307:
3272:
3262:
3231:
3223:
3188:
3180:
3145:
3137:
3102:
3062:
3056:Optics
3040:
3032:
2997:
2966:
2935:
2900:
2851:976377
2849:
2773:
2765:
2730:
2703:
2668:
2660:
2625:
2579:
2544:
2534:
2495:
2487:
2452:
2444:
2408:915798
2406:
2396:
2327:
2317:
2278:
2268:
2227:
2197:
2189:
2154:
2146:
2111:
2103:
2053:
2043:
2002:
1992:
1953:
1943:
1900:
1890:
1853:
1829:
1819:
1777:
1767:
1728:
1718:
1685:
1665:
1608:
1598:
1556:
1518:
1508:
1469:
1459:
1421:
1378:
1357:
1280:
1270:
936:, and
534:embryo
486:nuclei
289:cornea
253:lenses
235:retina
215:cornea
82:System
4593:Other
4497:Orbit
4415:Fovea
4393:Other
4330:) β (
4313:Cells
4215:Pupil
4131:tunic
3974:globe
3730:S2CID
3521:S2CID
3475:S2CID
3391:S2CID
3348:S2CID
3229:S2CID
3186:S2CID
3143:S2CID
3038:S2CID
2771:S2CID
2666:S2CID
2493:S2CID
2450:S2CID
2195:S2CID
2152:S2CID
2109:S2CID
868:frogs
833:birds
741:The "
621:Pygo2
554:cells
365:cells
249:below
190:, or
174:[
163:58241
110:Latin
99:light
4533:Lens
4518:Iris
4366:) β
4342:) β
4334:) β
4202:Iris
3909:ISBN
3883:PMID
3832:PMID
3783:ISBN
3767:ISBN
3722:PMID
3687:PMID
3638:PMID
3603:PMID
3562:PMID
3513:PMID
3467:PMID
3432:PMID
3383:PMID
3340:PMID
3305:PMID
3270:PMID
3221:PMID
3178:PMID
3135:PMID
3100:PMID
3060:ISBN
3030:PMID
2995:PMID
2964:PMID
2933:PMID
2898:PMID
2847:PMID
2763:PMID
2728:PMID
2701:PMID
2658:PMID
2623:PMID
2577:PMID
2542:PMID
2485:PMID
2442:PMID
2404:PMID
2325:PMID
2276:PMID
2225:ISBN
2187:PMID
2144:PMID
2101:PMID
2051:PMID
2000:PMID
1951:PMID
1898:PMID
1851:ISBN
1827:PMID
1775:PMID
1726:PMID
1663:PMID
1629:2009
1606:PMID
1554:PMID
1516:PMID
1467:PMID
1419:PMID
1376:ISBN
1355:ISBN
1278:PMID
1208:Iris
877:and
843:and
831:and
784:The
619:and
617:BCL9
609:PAX6
602:iris
418:and
293:iris
291:and
221:and
203:eyes
188:lens
186:The
151:6798
134:TA98
122:MeSH
33:Lens
4338:β (
3873:PMC
3863:doi
3822:PMC
3812:doi
3714:doi
3677:PMC
3669:doi
3630:doi
3593:doi
3552:doi
3505:doi
3459:doi
3422:doi
3418:202
3375:doi
3332:doi
3297:doi
3260:PMC
3213:doi
3170:doi
3166:301
3127:doi
3123:233
3090:doi
3022:doi
2987:doi
2956:doi
2925:doi
2921:159
2888:hdl
2878:doi
2874:270
2839:doi
2810:doi
2755:doi
2724:121
2693:doi
2650:doi
2569:doi
2532:PMC
2524:doi
2477:doi
2434:doi
2394:PMC
2386:doi
2382:270
2315:PMC
2307:doi
2303:370
2266:PMC
2258:doi
2254:370
2179:doi
2136:doi
2132:206
2091:doi
2087:162
2041:PMC
2031:doi
2027:293
1990:PMC
1982:doi
1941:PMC
1933:doi
1929:141
1888:PMC
1880:doi
1876:360
1817:PMC
1809:doi
1805:412
1765:PMC
1757:doi
1753:111
1716:PMC
1653:doi
1596:PMC
1588:doi
1584:122
1546:doi
1506:PMC
1498:doi
1457:PMC
1449:doi
1409:doi
1405:261
1268:PMC
1260:doi
850:In
827:In
709:6cm
484:or
225:it
158:FMA
146:TA2
4657::
4495:,
4491:,
4487:,
4483:,
4382:,
4378:,
4374:,
4370::
4362:,
4358:,
4354:,
4350:,
4326:,
3895:^
3881:.
3871:.
3859:12
3857:.
3853:.
3830:.
3820:.
3808:12
3806:.
3802:.
3751:.
3728:.
3720:.
3710:57
3708:.
3685:.
3675:.
3665:90
3663:.
3659:.
3636:.
3626:56
3624:.
3601:.
3589:86
3587:.
3583:.
3560:.
3546:.
3542:.
3519:.
3511:.
3501:26
3499:.
3487:^
3473:.
3465:.
3455:30
3453:.
3430:.
3416:.
3412:.
3389:.
3381:.
3371:39
3369:.
3346:.
3338:.
3328:45
3326:.
3303:.
3293:69
3291:.
3268:.
3256:28
3254:.
3250:.
3227:.
3219:.
3209:95
3207:.
3184:.
3176:.
3164:.
3141:.
3133:.
3121:.
3098:.
3086:50
3084:.
3080:.
3036:.
3028:.
3018:85
3016:.
2993:.
2962:.
2931:.
2919:.
2896:.
2886:.
2872:.
2868:.
2845:.
2835:23
2833:.
2808:.
2796:.
2792:.
2769:.
2761:.
2751:42
2749:.
2722:.
2699:.
2689:69
2687:.
2664:.
2656:.
2646:20
2644:.
2619:26
2617:.
2600:16
2598:.
2575:.
2565:56
2563:.
2540:.
2530:.
2520:56
2518:.
2514:.
2491:.
2483:.
2473:15
2471:.
2448:.
2440:.
2430:27
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