440:—Ronald A. Schachar has proposed in 1992 what has been called a "rather bizarre geometric theory" which claims that focus by the human lens is associated with increased tension on the lens via the equatorial zonules; that when the ciliary muscle contracts, equatorial zonular tension is increased, causing the central surfaces of the crystalline lens to steepen, the central thickness of the lens to increase (anterior-posterior diameter), and the peripheral surfaces of the lens to flatten. While the tension on equatorial zonules is increased during accommodation, the anterior and posterior zonules are simultaneously relaxing. The increased equatorial zonular tension keeps the lens stable and flattens the peripheral lens surface during accommodation. As a consequence, gravity does not affect the amplitude of accommodation and primary
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309:, 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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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 precise fluid movement of the lens front only rather than trying to change the shape of the lens as a whole. While this concept may be involved in the focusing it has been shown by
Scheimpflug photography that the rear of the lens also changes shape in the living eye.
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701:, 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.
655:, 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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contraction. This process can occur in as little as 224 ± 30 milliseconds in bright light. The amplitude of accommodation declines with age. By the fifth decade of life the accommodative amplitude can decline so that the near point of the eye is more remote than the reading distance. When this occurs
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What is less well understood is how the subtle, precise and very quick changes in lens shape are made. 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 vertebrates, aspects of all
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The small difference in refractive index between water and the hydrated cornea means fish and amphibians need to bend the light more using the internal structures of the eye. Therefore, eyes evolved in water have a mechanism involving changing the distance between a rigid rounder more refractive lens
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Aquatic animals include some that also thrive in the air so focusing mechanisms vary more than in those that are only land based. Some whales and seals are able to focus above and below water having two areas of retina with high numbers of rods and cones rather than one as in humans. Having two high
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proposes that the lens, zonule and anterior vitreous comprise a diaphragm between the anterior and vitreous chambers of the eye. Ciliary muscle contraction initiates a pressure gradient between the vitreous and aqueous compartments that support the anterior lens shape. It is in this lens shape that
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is large; i.e. in dim light. The age-related decline in accommodation occurs almost universally to less than 2 dioptres by the time a person reaches 45 to 50 years, by which time most of the population will have noticed a decrease in their ability to focus on close objects and hence require glasses
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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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occurs. The anterior capsule and the zonule form a trampoline shape or hammock shaped surface that is totally reproducible depending on the circular dimensions, i.e. the diameter of the ciliary body (MĂĽeller's muscle). The ciliary body thus directs the shape like the pylons of a suspension bridge,
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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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involving water, air and often both. The eyes are therefor required to bend light different amounts leading to different mechanisms of focus being used in different environments. The air/cornea interface involves a larger difference in refractive index than hydrated structures within the eye. As a
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is the condition where amplitude of accommodation of a person is lesser compared to physiological limits for their age. Premature sclerosis of lens or ciliary muscle weaknesses due to systemic or local cases may cause accommodative insufficiency. Accommodative insufficiency is further categorised
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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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being pulled tight by the pressure of the eyeball. At short focal distance the ciliary muscle contracts, stretching the ciliary body and relieving some of the tension on the suspensory ligaments, allowing the elastic lens to become more spherical, increasing refractive power. Changing focus to an
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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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3D reconstruction based on measurements taken from a 20 year old human male focusing from 26mm to infinity (4.85 dioptre change). Side & back views shown. Most image distortions near the center are due to model being limited to 512 faces to make up the lens. Peripheral distortions are also
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There are many types of accommodation anomalies. It can be broadly classified into two, decreased accommodation and increased accommodation. Decreased accommodation may occur due to physiological (presbyopia), pharmacological (cycloplegia) or pathological. Excessive accommodation and spasm of
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Focusing the light scattered by objects in a three dimensional environment into a two dimensional collection of individual bright points of light requires the light to be bent. To get a good image of these points of light on a defined area requires a precise systematic bending of light called
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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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models may play varying roles in lens focus. The models can be broadly divided into two camps. Those models that stress the importance of external forces acting on a more passively elastic lens and other models that include forces that may be generated by the lens internally.
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The young human eye can change focus from distance (infinity) to as near as 6.5 cm from the eye. This dramatic change in focal power of the eye of approximately 15 dioptres (the reciprocal of focal length in metres) occurs as a consequence of a reduction in
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object at a greater distance requires a thinner less curved lens. This is achieved by relaxing some of the sphincter-like ciliary muscles allowing the ciliarly body to spring back, pulling harder on the lens making it less curved and thinner, so increasing the
536:, physiological insufficiency of accommodation due to age related changes in lens (decreased elasticity and increased hardness) and ciliary muscle power is the commonest form of accommodative dysfunction. It will cause gradual decrease in near vision.
106:
The path of light through the eye calculated using four refractive indexes, cornea and lens curvatures approximating components of real eyes. Note objects in some size ranges and distances do not require the light path to bend noticeably to achieve
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is also known as accommodative inertia. In this condition there will be difficulty in changing accommodation from one point to other. There may be difficulty in adjusting focus from distance from near. It is a comparatively rare condition.
908:(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.
434:. When viewing a near object, the ciliary muscles contract (resisting the outward pressure on the sclera) causing the lens zonules to slacken which allows the lens to spring back into a thicker, more convex, form.
674:, 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
1977:
Chen, Ai Hong; O'Leary, Daniel J.; Howell, Edwin R. (2000). "Near visual function in young children. Part I: near point of convergence. Part II: amplitude of accommodation. Part III: near heterophoria".
150:
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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)".
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Ill-sustained accommodation is a condition similar to accommodative insufficiency. In this, range of accommodation will be normal, but after excessive near work accommodative power will decrease.
1440:
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".
682:, 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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the mechanically reproducible state of a steep radius of curvature in the center of the lens with slight flattening of the peripheral anterior lens, i.e. the shape, in cross section, of a
1773:
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".
219:
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
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occurs when an individual uses more than normal accommodation for performing certain near work. Modern definitions simply regard it as an inability to relax accommodation readily.
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Varying forms of direct experimental proof outlined in this article show that most non-aquatic vertebrates achieve focus, at least in part, by changing the shapes of their lenses.
167:
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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".
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for reading or bifocal lenses. Accommodation decreases to about 1 dioptre at the age of 70 years. The dependency of accommodation amplitude on age is graphically summarized by
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Duane's classical curves showing the amplitude or width of accommodation as changing with age. Mean (B) and approximate lower (A) and upper (C) standard deviations are shown.
1207:
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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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
232:. 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.
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Due to the nature of optics the focused image on the retina is always inverted relative to the object. Different animals live in different environments having different
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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.
390:(nearsighted and require an optical correction for distance or far vision), will find that they see better at near without their distance correction; and those who are
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also known as ciliary spasm is a condition of abnormally excessive accommodation which is out of voluntary control of the person. Vision may be blurred due to induced
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and suspensory ligaments to pull on the lens, flattening it. The source of the tension is the pressure that the vitreous and aqueous humours exert outwards onto the
667:, 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.
568:). It may occur due to ciliary muscle paralysis or occulomotor nerve paralysis. Parasympatholytic drugs like atropine will also cause paralysis of accommodation.
1857:
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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result, animals living in air have most of the bending of light achieved at the air/cornea interface with the lens being involved in finer focus of the image.
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Generally mammals, birds and reptiles living in air vary their eyes' optical power by subtly and precisely changing the shape of the elastic lens using the
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The previous video of the eye lens changing shape with focus is placed into context as the lens in this video is placed into the context of a virtual eye.
1623:
Hermans, EA; Dubbelman, M; Van der Heijde, R; Heethaar, RM (December 2008). "Equivalent refractive index of the human lens upon accommodative response".
1687:
Stahnke, T.; Hadlich, S.; Wree, A.; Guthoff, R.; Stachs, O.; Langner, S. (16 December 2016). "Magnetresonanzmikroskopie des
Akkommodationsapparats".
1057:
Shao, Yilei; Tao, Aizhu; Jiang, Hong; Mao, Xinjie; Zhong, Jianguang; Shen, Meixiao; Lu, Fan; Xu, Zhe; Karp, Carol L.; Wang, Jianhua (1 June 2015).
1288:
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).
394:(farsighted) will find that they may need a correction for both distance and near vision. Note that these effects are most noticeable when the
2079:
Baumeister, M.; Kohnen, T. (June 2008). "Akkommodation und
Presbyopie: Teil 1: Physiologie der Akkommodation und Entwicklung der Presbyopie".
1561:
Gruijters, WT; Kistler, J; Bullivant, S (October 1987). "Formation, distribution and dissociation of intercellular junctions in the lens".
2758:
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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".
1290:"Constant Volume of the Human Lens and Decrease in Surface Area of the Capsular Bag during Accommodation: An MRI and Scheimpflug Study"
971:
Schachar, Ronald A.; Bax, Andrew J. (June 2001). "Mechanism of human accommodation as analyzed by nonlinear finite element analysis".
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If there is amplitude of accommodation between the eyes differ 0.5 dioptre or more, it is considered as unequal. Organic diseases,
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Bony fish eye. Note the harder more spherical lens than in land based animals and a none circular muscle to pull the lens backward
512:) between how much convergence takes place because of accommodation (AC/A ratio, CA/C ratio). Abnormalities with this can lead to
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in a lecture on the 27th Nov 1800. Others such as
Helmholtz and Huxley refined the model in the mid-1800s explaining how the
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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
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Two horse lenses suspended on water by cling wrap with four approximately parallel lasers directed through them. The 1
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690:, there are two muscles, one above and one below the lens, while other amphibians have only the lower muscle.
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842:"Focusing by shape change in the lens of the eye: a commentary on Young (1801) 'On the mechanism of the eye'"
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In paralysis of accommodation, amplitude of accommodation is either markedly reduced or completely absent (
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Gruijters, WT (July 1989). "A non-connexon protein (MIP) is involved in eye lens gap-junction formation".
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of the eye, and thus reduce the amount of accommodation needed to bring the image in focus on the retina.
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resolution area of retina presumably allows two axis of vision one for above and one for below water. In
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shifts in the negative direction during accommodation. The theory has not found much independent support.
2387:
Wang, B.; Ciuffreda, K. J. (2006). "Depth-of-Focus of the Human Eye: Theory and
Clinical Implications".
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497:, the functional role of the pupillary constriction remains less clear. Arguably, it may increase 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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That value follows from the maximum accommodative power and can be calculated as 100 cm/15 dpt.
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While not vertebrate, brief mention is made here of the convergent evolution of vertebrate and
678:, by contrast, a muscle projects from a vascular structure in the floor of the eye, called the
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509:
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Coleman, DJ; Fish, SK (September 2001). "Presbyopia, accommodation, and the mature catenary".
1818:"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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306:
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Schachar, RA (March 1994). "Zonular function: a new hypothesis with clinical implications".
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rather than subtly changing the shape of the lens itself using circularly arranged muscles.
65:—the maximum distance from the eye for which a clear image of an object can be seen, to the
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382:(i.e., do not require optical correction for distance vision) will need an optical aid for
8:
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Khurana, AK (September 2008). "Asthenopia, anomalies of accommodation and convergence".
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Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
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2184:"Wavefront aberration changes caused by a gradient of increasing accommodation stimuli"
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Mulders, SM; Preston, GM; Deen, PM; Guggino, WB; van Os, CH; Agre, P (14 April 1995).
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on an object as its distance varies. In this, distances vary for individuals from the
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1940:"Studies in Monocular and Binocular Accommodation with their Clinical Applications"
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American Journal of Physiology. Regulatory, Integrative and Comparative Physiology
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The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology
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Coleman, D. Jackson (June 1970). "Unified Model for Accommodative Mechanism".
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Coleman, D. Jackson (June 1970). "Unified Model for Accommodative Mechanism".
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493:. While it is well understood that proper convergence is necessary to prevent
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but does not need to support an equatorial traction force to flatten the lens.
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formed from millions of these points of light is what animals see using their
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1999:
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Huggert, Arne (27 May 2009). "The Intracapsular Mechanism of Accommodation".
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Klinische Monatsblätter für Augenheilkunde und für augenärztliche Fortbildung
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2528:(2nd ed.). St. Louis Mo.: Butterworth Heinemann/Elsevier. p. 112.
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1059:"Age-Related Changes in the Anterior Segment Biometry During Accommodation"
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921:"The force of contraction of the human ciliary muscle during accommodation"
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477:. The combination of these three movements (accommodation, convergence and
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69:—the minimum distance for a clear image. Accommodation usually acts like a
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William J., Benjamin (2006). "Accommodation, the Pupil, and Presbyopia".
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893:(5th ed.). London and New York: MACMILLAN AND CO. pp. 256–258.
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Wrinkled lens fibers in picture below compared to straight fibers above
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Virtual eye showing the contribution to focus of different components.
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Suzuki-Kerr, H; Walker, KL; Han, MH; Lim, JC; Donaldson, PJ (2022).
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of the lens. As more is learned about mammalian lens structure from
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2607:. Philadelphia, PA: Holt-Saunders International. pp. 463–464.
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415:—The most widely held theory of accommodation is that proposed by
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Schaeffel, F.; Murphy, C.J.; Howland, H.C. (15 November 1999).
2426:(2nd ed.). St. Louis Mo.: Butterworth Heinemann/Elsevier.
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1143:"The zonular insertion: a scanning electron microscopic study"
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cm spaced grid indicates a near focus focal length of around 6
1477:"Water channel properties of major intrinsic protein of lens"
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The model of a shape changing lens of humans was proposed by
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present in animal lenses though are typically covered by the
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in 1855. When viewing a far object, the circularly arranged
124:. Very even systematic curvature of parts of the cornea and
2114:
Atchison, David A. (1995). "Accommodation and presbyopia".
687:
652:
1401:"Gullstrand Intracapsular Accommodation Mechanism Revised"
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Atchison, DA (July 1995). "Accommodation and presbyopia".
128:
produces this systematic bending of light onto the retina.
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Changing the position of the lens relative to the retina.
50:
2422:
William J., Benjamin (2006). "Fusion and binocularity".
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2673:"Accommodation in the cuttlefish (Sepia officinalis)"
2286:"On the hydraulic suspension theory of accommodation"
2182:
Zhou, X-Y; Wang, L; Zhou, X-T; Yu, Z-Q (2014-10-24).
1976:
2490:
Duke, Elder's (1969). "Anomalies of accommodation".
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accommodation are types of increased accommodation.
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When humans accommodate to a near object, they also
2603:Romer, Alfred Sherwood; Parsons, Thomas S. (1977).
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464:
1525:
2549:Mass, Alla M.; Supin, Alexander YA. (June 2007).
23:Minimum (top) and maximum accommodation (bottom).
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1294:Investigative Ophthalmology & Visual Science
1147:Investigative Ophthalmology & Visual Science
1108:Investigative Ophthalmology & Visual Science
1063:Investigative Ophthalmology & Visual Science
1056:
406:
2455:
2453:
2451:
2449:
2447:
2445:
2443:
2181:
584:may be responsible for unequal accommodation.
551:
539:
520:Anomalies of accommodation described in humans
195:Land based animals and the shape changing lens
2752:
2485:
2483:
2481:
2386:
2159:The Mechanism of Accommodation and Presbyopia
559:
77:, but it can also be consciously controlled.
2766:
2464:(2nd ed.). Elsevier. pp. 100–107.
2462:Theory and practice of optics and refraction
1970:
80:The main ways animals may change focus are:
2627:
2551:"Adaptive features of aquatic mammals' eye"
2440:
2362:"EyeRounds.org: Tutorial: Binocular Vision"
2324:
2029:
1209:Biomechanics and Modeling in Mechanobiology
970:
587:
2759:
2745:
2523:
2478:
2421:
1689:Klinische Monatsblätter für Augenheilkunde
722:Disorders of and relating to accommodation
2729:at the U.S. National Library of Medicine
2688:
2566:
2548:
2519:
2517:
2515:
2513:
2511:
2301:
2217:
2199:
2055:
2032:"Effects of Age on Dynamic Accommodation"
1833:
1591:
1502:
1492:
1424:
1398:
1305:
1082:
944:
903:
865:
599:
90:Changing the axial length of the eyeball.
2156:
2113:
1933:
1931:
1179:
1140:
1105:
1013:
708:. The most complex Molluscan eye is the
637:
629:
571:
378:. Once presbyopia occurs, those who are
355:
271:
263:
234:
164:
146:
130:
101:
26:
18:
2459:
2415:
2283:
2240:
1355:
1249:
614:
3115:
2508:
918:
888:
97:
2740:
2598:
2596:
2594:
2359:
1937:
1928:
1016:Ophthalmic & Physiological Optics
2489:
1775:Progress in Retinal and Eye Research
1399:LĂłpez-Gil, Norberto (3 March 2022).
839:
316:
2494:(8th ed.). London: Churchill.
2128:10.1046/j.1475-1313.1995.9500020e.x
2116:Ophthalmic and Physiological Optics
1980:Ophthalmic and Physiological Optics
1481:The Journal of Biological Chemistry
1328:
1028:10.1046/j.1475-1313.1995.9500020e.x
13:
2723:—"Presbyopia: Cause and Treatment"
2591:
2161:. The Hague: Kugler Publications.
1669:, 2nd ed. (1987), Addison Wesley,
1370:10.1111/j.1755-3768.1964.tb03627.x
625:
259:
251:" model of lens focus proposed by
206:
187:and the retina using less uniform
14:
3149:
2708:
2401:10.1016/j.survophthal.2005.11.003
2243:American Journal of Ophthalmology
2030:Lockhart, T. E.; Shi, W. (2010).
1944:American Journal of Ophthalmology
1252:American Journal of Ophthalmology
693:In the simplest vertebrates, the
93:Changing the shape of the cornea.
3010:
2360:Bhola, Rahul (23 January 2006).
1992:10.1046/j.1475-1313.2000.00498.x
1787:10.1016/j.preteyeres.2022.101152
906:Treatise on physiological optics
891:Lessons in Elementary Physiology
465:Induced effects of accommodation
321:
75:accommodation-convergence reflex
2677:Journal of Experimental Biology
2664:
2621:
2542:
2380:
2353:
2318:
2277:
2234:
2175:
2150:
2107:
2072:
2023:
2014:
1885:
1850:
1809:
1766:
1723:
1680:
1659:
1616:
1585:
1554:
1528:The American Journal of Anatomy
1519:
1468:
1433:
1392:
1349:
1322:
1243:
1200:
1173:
1134:
919:Fisher, R. F. (1 August 1977).
796:Positive relative accommodation
791:Negative relative accommodation
473:their eyes and constrict their
84:Changing the shape of the lens.
1141:Streeten, B. W. (April 1977).
1099:
1050:
1007:
937:10.1113/jphysiol.1977.sp011938
912:
897:
882:
833:
807:
481:) is under the control of the
1:
2642:10.1016/S0042-6989(99)00012-7
2339:10.1016/s0161-6420(01)00691-1
1956:10.1016/s0002-9394(22)90793-7
801:
686:in either type of animal. In
528:
508:There is a measurable ratio (
312:
57:to maintain a clear image or
2526:Borish's clinical refraction
2424:Borish's Clinical Refraction
2255:10.1016/0002-9394(70)91057-3
2048:10.1080/00140139.2010.489968
1906:10.1016/j.visres.2005.03.008
1637:10.1097/OPX.0b013e31818e8d57
1625:Optometry and Vision Science
1454:10.1016/0014-4835(76)90135-4
1264:10.1016/0002-9394(70)91057-3
407:Theories on how humans focus
239:Schachar model of lens focus
46:is the process by which the
7:
716:
552:Ill-sustained accommodation
548:into different categories.
545:Accommodative insufficiency
540:Accommodative insufficiency
330:It has been suggested that
10:
3154:
2492:The practice of refraction
1744:10.1152/ajpregu.00173.2011
1596:. 93 ( Pt 3) (3): 509–13.
1221:10.1007/s10237-021-01417-9
904:Helmholtz, H. von (1962).
766:Amplitude of accommodation
560:Paralysis of accommodation
485:and is referred to as the
333:Amplitude of accommodation
3074:
3019:
3008:
2779:
2093:10.1007/s00347-008-1761-8
1938:Duane, Alexander (1922).
1859:Experimental Eye Research
1565:. 88 ( Pt 3) (3): 351–9.
1442:Experimental Eye Research
985:10.1007/s12019-996-0006-5
925:The Journal of Physiology
889:Huxley, Thomas H (1871).
840:Land, M (19 April 2015).
347:Proposed since July 2024.
3048:Ascending and Descending
2731:Medical Subject Headings
2157:Schachar, R. A. (2012).
1426:10.3390/photonics9030152
781:Edinger-Westphal nucleus
749:
593:Accommodative infacility
588:Accommodative infacility
483:Edinger-Westphal nucleus
73:, including part of the
2690:10.1242/jeb.202.22.3127
2389:Survey of Ophthalmology
2290:Trans Am Ophthalmol Soc
1594:Journal of Cell Science
1563:Journal of Cell Science
1494:10.1074/jbc.270.15.9010
1182:Annals of Ophthalmology
728:Accommodative esotropia
307:Scheimpflug photography
16:Focusing ability of eye
2366:webeye.ophth.uiowa.edu
1871:10.1006/exer.1999.0747
1540:10.1002/aja.1001590406
858:10.1098/rstb.2014.0308
643:
635:
605:Spasm of accommodation
600:Spasm of accommodation
361:
277:
269:
240:
176:
162:
136:
108:
40:
24:
1701:10.1055/s-0042-118599
1120:10.1167/iovs.15-17452
1075:10.1167/iovs.15-16825
973:Comprehensive Therapy
756:Accommodation in fish
641:
633:
572:Unequal accommodation
423:relaxes allowing the
417:Hermann von Helmholtz
403:'s classical curves.
359:
275:
267:
238:
174:
156:
134:
105:
30:
22:
3082:Accidental viewpoint
2727:Ocular+Accommodation
2368:. University of Iowa
2284:Coleman, DJ (1986).
2201:10.1038/eye.2014.244
1602:10.1242/jcs.93.3.509
1571:10.1242/jcs.88.3.351
1358:Acta Ophthalmologica
1307:10.1167/iovs.08-2124
620:Accommodative excess
615:Accommodative excess
491:accommodation reflex
442:spherical aberration
340:into this section. (
221:suspending ligaments
2987:Vertical–horizontal
2605:The Vertebrate Body
1417:2022Photo...9..152L
1329:PAU, H (1952). "".
510:Matthiessen's ratio
369:tension induced by
98:Focusing mechanisms
3087:Auditory illusions
2882:Impossible trident
672:cartilaginous fish
644:
636:
452:D. Jackson Coleman
362:
278:
270:
241:
177:
163:
141:refractive indexes
137:
109:
41:
25:
3110:
3109:
3102:Temporal illusion
3097:Tactile illusions
3067:(2015 photograph)
2768:Optical illusions
2683:(22): 3127–3134.
2636:(17): 2841–2852.
2614:978-0-03-910284-5
2535:978-0-7506-7524-6
2471:978-81-312-1132-8
2433:978-0-7506-7524-6
2168:978-90-6299-233-1
2081:Der Ophthalmologe
1900:(18): 2352–2366.
1695:(12): 1320–1323.
786:Mandelbaum Effect
680:falciform process
354:
353:
349:
172:
154:
3145:
3014:
2967:Schroeder stairs
2942:Peripheral drift
2937:Penrose triangle
2761:
2754:
2747:
2738:
2737:
2703:
2702:
2692:
2668:
2662:
2661:
2625:
2619:
2618:
2600:
2589:
2588:
2570:
2568:10.1002/ar.20529
2546:
2540:
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2506:
2505:
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2378:
2377:
2375:
2373:
2357:
2351:
2350:
2322:
2316:
2315:
2305:
2281:
2275:
2274:
2249:(6): 1063–1079.
2238:
2232:
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2221:
2203:
2179:
2173:
2172:
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2148:
2147:
2111:
2105:
2104:
2076:
2070:
2069:
2059:
2027:
2021:
2018:
2012:
2011:
1974:
1968:
1967:
1935:
1926:
1925:
1889:
1883:
1882:
1854:
1848:
1847:
1837:
1822:Molecular Vision
1813:
1807:
1806:
1770:
1764:
1763:
1727:
1721:
1720:
1684:
1678:
1663:
1657:
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1390:
1389:
1353:
1347:
1346:
1326:
1320:
1319:
1309:
1285:
1276:
1275:
1258:(6): 1063–1079.
1247:
1241:
1240:
1204:
1198:
1197:
1177:
1171:
1170:
1138:
1132:
1131:
1103:
1097:
1096:
1086:
1069:(6): 3522–3530.
1054:
1048:
1047:
1011:
1005:
1004:
968:
959:
958:
948:
916:
910:
909:
901:
895:
894:
886:
880:
879:
869:
837:
831:
830:
828:
826:
811:
761:Adaptation (eye)
514:binocular vision
501:by reducing the
386:; those who are
345:
325:
324:
317:
292:refractive index
173:
155:
38:
34:
3153:
3152:
3148:
3147:
3146:
3144:
3143:
3142:
3113:
3112:
3111:
3106:
3070:
3020:Popular culture
3015:
3006:
2977:Spinning dancer
2797:Ambiguous image
2775:
2765:
2711:
2706:
2669:
2665:
2630:Vision Research
2626:
2622:
2615:
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2592:
2547:
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2180:
2176:
2169:
2155:
2151:
2112:
2108:
2077:
2073:
2028:
2024:
2019:
2015:
1975:
1971:
1950:(11): 865–877.
1936:
1929:
1894:Vision Research
1890:
1886:
1855:
1851:
1814:
1810:
1771:
1767:
1728:
1724:
1685:
1681:
1665:Hecht, Eugene.
1664:
1660:
1631:(12): 1179–84.
1621:
1617:
1590:
1586:
1559:
1555:
1524:
1520:
1487:(15): 9010–16.
1473:
1469:
1438:
1434:
1397:
1393:
1354:
1350:
1327:
1323:
1286:
1279:
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1201:
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969:
962:
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883:
838:
834:
824:
822:
819:www.bio.vobs.at
813:
812:
808:
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752:
724:
719:
628:
626:Aquatic animals
617:
602:
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574:
562:
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531:
522:
467:
409:
374:the patient is
350:
326:
322:
315:
262:
260:Internal forces
209:
207:External forces
197:
165:
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100:
36:
32:
17:
12:
11:
5:
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3099:
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3076:
3072:
3071:
3069:
3068:
3060:
3059:(1961 drawing)
3052:
3051:(1960 drawing)
3044:
3036:
3029:
3023:
3021:
3017:
3016:
3009:
3007:
3005:
3004:
2999:
2994:
2989:
2984:
2979:
2974:
2972:Shepard tables
2969:
2964:
2959:
2954:
2949:
2944:
2939:
2934:
2932:Penrose stairs
2929:
2924:
2919:
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2909:
2904:
2899:
2894:
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2829:
2827:Checker shadow
2824:
2819:
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2809:
2807:Autostereogram
2804:
2799:
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2710:
2709:External links
2707:
2705:
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2613:
2590:
2561:(6): 701–715.
2541:
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2414:
2379:
2352:
2333:(9): 1544–51.
2317:
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2194:(1): 115–121.
2174:
2167:
2149:
2122:(4): 255–212.
2106:
2087:(6): 597–610.
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2042:(7): 892–903.
2022:
2013:
1986:(3): 185–198.
1969:
1927:
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1765:
1738:(2): R335-42.
1722:
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710:Cephalopod eye
706:Molluscan eyes
661:vision in fish
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580:or functional
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499:depth of field
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421:ciliary muscle
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371:ciliary muscle
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296:index gradient
261:
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230:focal distance
217:ciliary muscle
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160:iris (anatomy)
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2501:0-7000-1410-1
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2327:Ophthalmology
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2083:(in German).
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55:optical power
52:
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44:Accommodation
29:
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3033:Trompe-l'Ĺ“il
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2897:Lilac chaser
2867:Gravity hill
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2395:(1): 75–85.
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2372:11 September
2370:. Retrieved
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1865:(6): 663–9.
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825:11 September
823:. Retrieved
818:
809:
744:Pseudomyopia
703:
692:
679:
669:
645:
618:
609:pseudomyopia
603:
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363:
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246:
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213:Thomas Young
210:
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185:
181:ciliary body
178:
138:
110:
79:
43:
42:
3043:(1864 book)
2947:Poggendorff
2922:Oppel-Kundt
2917:Necker cube
2912:MĂĽller-Lyer
2887:Irradiation
1828:: 245–256.
1504:2066/216433
1188:(2): 36–8.
821:(in German)
771:Cycloplegia
578:head trauma
566:cycloplegia
384:near vision
3117:Categories
3040:Spectropia
2957:Rubin vase
2907:McCollough
2902:Mach bands
2852:Ehrenstein
2847:Ebbinghaus
2812:Barberpole
2787:Afterimage
2296:: 846–68.
2036:Ergonomics
1781:: 101152.
1411:(3): 152.
802:References
776:Cyclospasm
665:amphibians
534:Presbyopia
529:Presbyopia
516:problems.
487:near triad
380:emmetropic
376:presbyopic
313:Human eyes
283:aquaporins
118:real image
114:refraction
67:near point
48:vertebrate
3128:Optometry
3092:Illusions
3064:The dress
3056:Waterfall
2857:Flash lag
2837:Cornsweet
2822:Café wall
2802:Ames room
2780:Illusions
2721:eMedicine
2263:0002-9394
2210:0950-222X
2000:0275-5408
1803:254243790
1677:. p. 178.
1653:205907383
1405:Photonics
1237:231704221
1159:0146-0404
734:hyperopia
582:amblyopia
413:Helmholtz
392:hyperopic
287:Connexins
63:far point
2842:Delboeuf
2792:Ambigram
2699:10539961
2658:17808919
2650:10492814
2585:39925190
2577:17516421
2409:16414364
2347:11535447
2228:25341432
2144:24282106
2101:18594896
2066:20582770
2008:10897340
1964:43172462
1914:15979462
1879:10620395
1844:36284672
1795:36470825
1752:21593426
1717:78808282
1709:27984837
1645:19050472
1386:37325357
1378:14213923
1343:14955961
1316:18676625
1229:33491156
1128:26393665
1093:26030106
1044:24282106
1001:71369369
993:11430259
876:25750232
852:(1666).
717:See also
695:lampreys
676:teleosts
649:reptiles
503:aperture
495:diplopia
471:converge
457:catenary
448:Catenary
438:Schachar
249:catenary
53:changes
3075:Related
3002:Zöllner
2992:White's
2927:Orbison
2892:Jastrow
2716:oph/723
2312:3590482
2303:1298753
2271:5423772
2219:4289835
2136:7667018
2057:2908311
1922:8894700
1835:9514545
1760:9525037
1610:2691517
1579:3448099
1548:7223675
1513:7536742
1413:Bibcode
1272:5423772
1194:8010701
1084:4464043
1036:7667018
946:1353417
867:4360117
815:"Augen"
732:Latent
699:hagfish
684:similar
659:. With
657:mammals
428:zonules
367:zonular
342:Discuss
304:in situ
253:Coleman
225:capsule
189:muscles
122:retinas
3138:Vision
3027:Op art
2982:Ternus
2962:Sander
2877:Hering
2817:Bezold
2733:(MeSH)
2697:
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1667:Optics
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999:
991:
955:915798
953:
943:
874:
864:
739:Myopia
479:miosis
475:pupils
432:sclera
388:myopic
338:merged
116:. The
107:focus.
71:reflex
37:
33:
2997:Wundt
2952:Ponzo
2832:Chubb
2654:S2CID
2581:S2CID
2140:S2CID
1960:S2CID
1918:S2CID
1799:S2CID
1756:S2CID
1713:S2CID
1649:S2CID
1382:S2CID
1233:S2CID
1040:S2CID
997:S2CID
750:Other
688:frogs
653:birds
489:, or
401:Duane
396:pupil
247:The "
59:focus
2872:Grid
2772:list
2695:PMID
2646:PMID
2609:ISBN
2573:PMID
2530:ISBN
2496:ISBN
2466:ISBN
2428:ISBN
2405:PMID
2374:2020
2343:PMID
2308:PMID
2267:PMID
2259:ISSN
2224:PMID
2206:ISSN
2163:ISBN
2132:PMID
2097:PMID
2062:PMID
2004:PMID
1996:ISSN
1910:PMID
1875:PMID
1840:PMID
1791:PMID
1748:PMID
1705:PMID
1671:ISBN
1641:PMID
1606:PMID
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1509:PMID
1458:PMID
1374:PMID
1339:PMID
1312:PMID
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1225:PMID
1190:PMID
1163:PMID
1155:ISSN
1124:PMID
1089:PMID
1032:PMID
989:PMID
951:PMID
872:PMID
827:2020
697:and
663:and
651:and
425:lens
290:The
223:and
126:lens
3123:Eye
2719:at
2685:doi
2681:202
2638:doi
2563:doi
2559:290
2397:doi
2335:doi
2331:108
2298:PMC
2251:doi
2214:PMC
2196:doi
2188:Eye
2124:doi
2089:doi
2085:105
2052:PMC
2044:doi
1988:doi
1952:doi
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1830:PMC
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1740:doi
1736:301
1697:doi
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1633:doi
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1567:doi
1536:doi
1532:159
1499:hdl
1489:doi
1485:270
1450:doi
1421:doi
1366:doi
1335:121
1302:doi
1260:doi
1217:doi
1116:doi
1079:PMC
1071:doi
1024:doi
981:doi
941:PMC
933:doi
929:270
862:PMC
854:doi
850:370
670:In
336:be
51:eye
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344:)
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