When looking into someone's eyes, we can easily see several structures:
• A black-looking aperture, the pupil, that allows light to enter the eye (it appears dark because of the absorbing pigments in the retina).
• A colored circular muscle, the iris, which is beautifully pigmented giving us our eye's color (the central aperture of the iris is the pupil). This circular muscle controls the size of the pupil so that more or less light, depending on conditions, is allowed to enter the eye. Eye color, or more correctly, iris color is due to variable amounts of eumelanin (brown/black melanins) and pheomelanin (red/yellow melanins) produced by melanocytes. More of the former is in brown eyed people and of the latter in blue and green-eyed people. The Melanocortin-1 Receptor Gene is a regulator of eumelanin production and is located on chromosome(MCIR) 16q24.3. Point mutations in the MCIR gene will affect melanogenesis. The presence of point mutations in the MCIR gene alleles is a common feature in light skinned and blue/green eyed people (J.A.W. Metzelaar-Blok et al., Invest. Ophthal. Vis. Sci. 42,1951-4, 2001; P. Valverde et al. Nat. Genet. 11, 328-330, 1995)).
• A transparent external surface, the cornea, that covers both the pupil and the iris. This is the first and most powerful lens of the optical system of the eye and allows, together with the crystalline lens the production of a sharp image at the retinal photoreceptor level.
• The "white of the eye", the sclera,which forms part of the supporting wall of the eyeball. The sclera is continuous with the cornea. Furthermore this external covering of the eye is in continuity with the dura of the central nervous system.
When we remove the eye from the orbit, we can see that the eye is a slightly asymmetrical sphere with an approximate sagittal diameter or length of 24 to 25 mm. and a transverse diameter of 24 mm. It has a volume of about 6.5 cc.
A cross-sectional view of the eye shows:
• Three different layers
1. The external layer, formed by the sclera and cornea
2. The intermediate layer, divided into two parts: anterior (iris and ciliary body) and posterior (choroid)
3. The internal layer, or the sensory part of the eye, the retina
• Three chambers of fluid: Anterior chamber (between cornea and iris), Posterior chamber (between iris, zonule fibers and lens) and the Vitreous chamber (between the lens and the retina). The first two chambers are filled with aqueous humor whereas the vitreous chamber is filled with a more viscous fluid, the vitreous humor.
• The sagittal section of the eye also reveals the lens which is a transparent body located behind the iris. The lens is suspended by ligaments (called zonule fibers) attached to the anterior portion of the ciliary body. The contraction or relaxation of these ligaments as a consequence of ciliary muscle actions, changes the shape of the lens, a process called accommodation that allows us to form a sharp image on the retina.
Light rays are focussed through the transparent cornea and lens upon the retina. The central point for image focus (the visual axis) in the human retina is the fovea. Here a maximally focussed image initiates resolution of the finest detail and direct transmission of that detail to the brain for the higher operations needed for perception. Slightly more nasally than the visual axis is the optic axis projecting closer to the optic nerve head. The optic axis is the longest sagittal distance between the front or vertex of the corna and the furthest posterior part of the eyeball. It is about the optic axis that the eye is rotated by the eye muscles. Some vertebrate retinas have instead of a fovea, another specialization of the central retina, known as an area centralis or a visual streak.
Extraocular muscles.
Each eyeball is held in position in the orbital cavity by various ligaments, muscles and fascial expansions that surround it (see Fig. 3).
Inserted into the sclera are three pairs of muscles (6 muscles altogether). Two pairs are rectus muscles running straight to the bony orbit of the skull orthogonal to each other (the superior rectus, the inferior rectus, the lateral rectus and the medial rectus muscles). A further pair of muscles, the oblique muscles (superior oblique and inferior oblique) are angled as the name implies obliquely. These muscles, named extraocular muscles rotate the eyeball in the orbits and allow the image to be focussed at all times on the fovea of central retina.
Development of the eye.
The retina is a part of the central nervous system and an ideal region of the vertebrate brain to study, because like other regions of the central nervous system, it derives from the neural tube. The retina is formed during development of the embryo from optic vesicles outpouching from two sides of the developing neural tube. The primordial optic vesicles fold back in upon themselves to form the optic cup with the inside of the cup becoming the retina and the outside remaining a single monolayer of epithelium known as the retinal pigment epithelium. Initially both walls of the optic cup are one cell thick, but the cells of the inner wall divide to form a neuroepithelial layer many cells thick: the retina
CLICK HERE to see a Morph of development (683 K quicktime movie)
Sensory retinal development begins as early as the optic vesicle stage, with the migration of cell nuclei to the inner surface of the sensory retina. Additional retinal development is characterized by the formation of further layers arising from cell division and subsequent cell migration. The retina develops in an inside to outside manner: ganglion cells are formed first and photoreceptors cells become fully mature last.
Further changes in retinal morphology are accomplished by simultaneous formation of multiple complex intercellular connections. Thus by 5 months of gestation most of the basic neural connections of the retina have been established (Mann, 1964).
The functional synapses are made almost exclusively in the two plexiform layers and the perikarya of the nerve cells are distributed in the three nuclear layers.
Photoreceptor cell maturation begins with the formation of outer segments (OS) containing visual pigment from multiple infoldings of the plasma membrane of each cell. Outer segment formation proceeds, and the eye becomes sensitive to light at about 7 months' gestation.
The final portion of the sensory retina to mature is the fovea, where the ganglion cell layer thickening begins during midgestation. The outer nuclear layer is also wider here than elsewhere in the retina and consists almost entirely of developing cone cells. The ganglion cell nuclei migrate radially outwards in a circle, leaving the fovea free of ganglion cell nuclei. Cell-cell attachments persist, however and foveal cone cells alter their shape to accomodate the movement of ganglion cells. Foveal development continues with cell rearrangements and alteration in cone shape until about 4 years after birth (Hendrickson and Yondelis, 1984; Curcio and Hendrickson, 1991).
Surface membranes cover the eye cup and develop into lens, iris and cornea with the three chambers of fluid filled with aqueous and vitreous humors.
Surface membranes cover the eye cup and develop into lens, iris and cornea with the three chambers of fluid filled with aqueous and vitreous humors.
Summary.
In following chapters, we will describe in greater detail the individual nerve cells that make up the retina and the functional pathways into which these neurons are organized. Eventually, we will progress to a stage where we can appreciate the summary diagrams below that show the functional wiring of two well-understood mammalian retinas, namely cat and primate retinas.
References.
Hendrickson, A.E. and Youdelis, C. (1984). The morphological development of the human fovea. Ophthalmology 91, 603-612.
Mann, I. (1964). The development of the human eye. Grune and Stralton, New York.
Ogden, T.E. (1989) Retina: Basic Science and inherited retinal disease, vol 1. The CV Mosby Co. St. Louis.
Curcio, C.A. and Hendrickson, A.E. (1991). Organization and development of the primate photoreceptor mosaic. Prog. Ret. Ret. 10, 89-120.
Types of eyes
There are ten different eye layouts — indeed every way of capturing an image known to man, with the exceptions of zoom and Fresnel lenses. Eye types can be categorized into "simple eyes", with one concave chamber, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface. Note that "simple" does not imply a reduced level of complexity or acuity. Indeed, any eye type can be adapted for almost any behavior or environment. The only limitations specific to eye types are that of resolution — the physics of compound eyes prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eyes, so are better suited to dark-dwelling creatures. Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being cilliated (as in the vertebrates) or rhabdomeric. These two groups are not monophyletic; the cnidaria also possess cilliated cells, and some annelids possess both.
Normal eyes
Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times in vertebrates, cephalopods, annelids, crustacea and cubozoa.
Pit eyes
Pit eyes, also known as stemma, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eyespot, to allow the organism to deduce the angle of incoming light. Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eye". They are small, comprising up to about 100 cells covering about 100 µm. The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material.
Spherical lensed eye
The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index to form a lens, which may greatly reduce the blur radius encountered — hence increasing the resolution obtainable. The most basic form, still seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index, decreasing to the edges; this decreases the focal length and thus allows a sharp image to form on the retina. This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing spherical aberration. Such an inhomogeneous lens is necessary in order for the focal length to drop from about 4 times the lens radius, to 2.5 radii.
Heterogeneous eyes have evolved at least eight times: four or more times in gastropods, once in the copepods, once in the annelids and once in the cephalopods. No aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".
This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimize the effect of eye motion while the animal moves, most such eyes have stabilizing eye muscles.
The ocelli of insects bear a simple lens, but their focal point always lies behind the retina; consequently they can never form a sharp image. This capitulates the function of the eye. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the whole visual field; this fast response is further accelerated by the large nerve bundles which rush the information to the brain. Focusing the image would also cause the sun's image to be focused on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity. This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above).
Multiple lenses
Some marine organisms bear more than one lens; for instance the copepod Pontella has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod, Copilia's eyes have two lenses, arranged like those in a telescope. Such arrangements are rare and poorly understood, but represent an interesting alternative construction. An interesting use of multiple lenses is seen in some hunters such as eagles and jumping spiders, which have a refractive cornea (discussed next): these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution.
Refractive cornea
In the eyes of most mammals, birds, reptiles, and most other terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air, relieving the lens of the function of reducing the focal length. This has freed it up for fine adjustments of focus, allowing a very high resolution to be obtained. As with spherical lenses, the problem of spherical aberration caused by the lens can be countered either by using an inhomogeneous lens material, or by flattening the lens. Flattening the lens has a disadvantage: the quality of vision is diminished away from the main line of focus, meaning that animals requiring all-round vision are detrimented. Such animals often display an inhomogeneous lens instead.
As mentioned above, a refractive cornea is only useful out of water; in water, there is no difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures which have returned to the water — penguins and seals, for example — lose their refractive cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strong cornea.
Reflector eyes
An alternative to a lens is to line the inside of the eye with " mirrors", and reflect the image to focus at a central point. The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out.
Many small organisms such as rotifers, copeopods and platyhelminths use such organs, but these are too small to produce usable images. Some larger organisms, such as scallops, also use reflector eyes. The scallop Pecten has up to 100 millimeter-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses.
There is at least one vertebrate, the spookfish, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from the above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of guanine crystals.
Compound eyes
An image of a house fly compound eye surface by using Scanning Electron Microscope at X457 magnification
Arthropods such as this carpenter bee have compound eyes
A compound eye may consist of thousands of individual photoreceptor units. The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the polarization of light. Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained. This can only be countered by increasing lens size and number. To see with a resolution comparable to our simple eyes, humans would require compound eyes which would each reach the size of their head.
Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image. Compound eyes are common in arthropods, and are also present in annelids and some bivalved molluscs.
Compound eyes, in arthropods at least, grow at their margins by the addition of new ommatidia.
Apposition eyes
Apposition eyes are the most common form of eye, and are presumably the ancestral form of compound eye. They are found in all arthropod groups, although they may have evolved more than once within this phylum. Some annelids and bivalves also have apposition eyes. They are also possessed by Limulus, the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point. (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.)
Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information.
The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is absorbed by the dark wall of the ommatidium. In the other kind of apposition eye, found in the Strepsiptera, lenses are not fused to one another, and each forms an entire image; these images are combined in the brain. This is called the schizochroal compound eye or the neural superposition eye. Because images are combined additively, this arrangement allows vision under lower light levels.
Superposition eyes
The second type is named the superposition eye. The superposition eye is divided into three types; the refracting, the reflecting and the parabolic superposition eye. The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This kind is used mostly by nocturnal insects. In the parabolic superposition compound eye type, seen in arthropods such as mayflies, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied decapod crustaceans such as shrimp, prawns, crayfish and lobsters are alone in having reflecting superposition eyes, which also has a transparent gap but uses corner mirrors instead of lenses.
Parabolic superposition
This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.
Other
The compound eye of a dragonfly
Good fliers like flies or honey bees, or prey-catching insects like praying mantis or dragonflies, have specialized zones of ommatidia organized into a fovea area which gives acute vision. In the acute zone the eyes are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution.
There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the mysid shrimp Dioptromysis paucispinosa. The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialized retina. The resulting eye is a mixture of a simple eye within a compound eye.
Another version is the pseudofaceted eye, as seen in Scutigera. This type of eye consists of a cluster of numerous ocelli on each side of the head, organized in a way that resembles a true compound eye.
The body of Ophiocoma wendtii, a type of brittle star, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chitons.
References.
1-^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab Land, M F; Fernald, R D (1992). "The Evolution of Eyes". Annual Review of Neuroscience 15: 1–29. doi:10.1146/annurev.ne.15.030192.000245. PMID 1575438.
2-^ a b c d Cronin, T. W.; Porter, M. L. (2008). "Exceptional Variation on a Common Theme: the Evolution of Crustacean Compound Eyes". Evolution Education and Outreach 1: 463–475. doi:10.1007/s12052-008-0085-0. edit
3-Kozmik, Zbynek; Ruzickova, Jana; Jonasova, Kristyna; Matsumoto, Yoshifumi; Vopalensky, Pavel; Kozmikova, Iryna; Strnad, Hynek; Kawamura, Shoji et al. (2008). "Assembly of the cnidarian camera-type eye from vertebrate-like components" (PDF). Proceedings of the National Academy of Sciences 105 (26): 8989–8993. doi:10.1073/pnas.0800388105. PMID 18577593. PMC 2449352. http://www.pnas.org/cgi/reprint/0800388105v1.pdf.
4-Fernald, Russell D. (September 2006). "Casting a Genetic Light on the Evolution of Eyes". Science 313 (5795): 1914–1918. doi:10.1126/science.1127889. PMID 17008522.
5-"Vision optics and evolution" BioScience 39 (5): 298–307. 1 May 1989. doi:10.2307/1311112. ISSN 00063568. http://jstor.org/stable/1311112. edit
6-^ a b c Wilson, M. (1978). "The functional organisation of locust ocelli". Journal of Comparative Physiology 124 (4): 297–316. doi:10.1007/BF00661380.
7-^ Wagner, H.J., Douglas, R.H., Frank, T.M., Roberts, N.W., and Partridge, J.C. (Jan. 27, 2009). "A Novel Vertebrate Eye Using Both Refractive and Reflective Optics". Current Biology 19 (2): 108–114. doi:10.1016/j.cub.2008.11.061. PMID 19110427.
8-^ Völkel, R; Eisner, M; Weible, K. J (June 2003). "Miniaturized imaging systems" (PDF). Microelectronic Engineering 67-68 (1): 461–472. doi:10.1016/S0167-9317(03)00102-3. http://www.suss-microoptics.com/downloads/Publications/Miniaturized_Imaging_Systems.pdf.
9-^ Gaten, Edward (1998). "Optics and phylogeny: is there an insight? The evolution of superposition eyes in the Decapoda (Crustacea)". Contributions to Zoology 67 (4): 223–236. http://dpc.uba.uva.nl/ctz/vol67/nr04/art01#FIGURE1.
10-^ Ritchie, Alexander (1985). "Ainiktozoon loganense Scourfield, a protochordate? from the Silurian of Scotland". Alcheringa 9: 137. doi:10.1080/03115518508618961.
11-^ Mayer, G. (2006). "Structure and development of onychophoran eyes: What is the ancestral visual organ in arthropods?". Arthropod Structure and Development 35 (4): 231–245. doi:10.1016/j.asd.2006.06.003. PMID 18089073.
• A black-looking aperture, the pupil, that allows light to enter the eye (it appears dark because of the absorbing pigments in the retina).
• A colored circular muscle, the iris, which is beautifully pigmented giving us our eye's color (the central aperture of the iris is the pupil). This circular muscle controls the size of the pupil so that more or less light, depending on conditions, is allowed to enter the eye. Eye color, or more correctly, iris color is due to variable amounts of eumelanin (brown/black melanins) and pheomelanin (red/yellow melanins) produced by melanocytes. More of the former is in brown eyed people and of the latter in blue and green-eyed people. The Melanocortin-1 Receptor Gene is a regulator of eumelanin production and is located on chromosome(MCIR) 16q24.3. Point mutations in the MCIR gene will affect melanogenesis. The presence of point mutations in the MCIR gene alleles is a common feature in light skinned and blue/green eyed people (J.A.W. Metzelaar-Blok et al., Invest. Ophthal. Vis. Sci. 42,1951-4, 2001; P. Valverde et al. Nat. Genet. 11, 328-330, 1995)).
• A transparent external surface, the cornea, that covers both the pupil and the iris. This is the first and most powerful lens of the optical system of the eye and allows, together with the crystalline lens the production of a sharp image at the retinal photoreceptor level.
• The "white of the eye", the sclera,which forms part of the supporting wall of the eyeball. The sclera is continuous with the cornea. Furthermore this external covering of the eye is in continuity with the dura of the central nervous system.
When we remove the eye from the orbit, we can see that the eye is a slightly asymmetrical sphere with an approximate sagittal diameter or length of 24 to 25 mm. and a transverse diameter of 24 mm. It has a volume of about 6.5 cc.
• Three different layers
1. The external layer, formed by the sclera and cornea
2. The intermediate layer, divided into two parts: anterior (iris and ciliary body) and posterior (choroid)
3. The internal layer, or the sensory part of the eye, the retina
• Three chambers of fluid: Anterior chamber (between cornea and iris), Posterior chamber (between iris, zonule fibers and lens) and the Vitreous chamber (between the lens and the retina). The first two chambers are filled with aqueous humor whereas the vitreous chamber is filled with a more viscous fluid, the vitreous humor.
• The sagittal section of the eye also reveals the lens which is a transparent body located behind the iris. The lens is suspended by ligaments (called zonule fibers) attached to the anterior portion of the ciliary body. The contraction or relaxation of these ligaments as a consequence of ciliary muscle actions, changes the shape of the lens, a process called accommodation that allows us to form a sharp image on the retina.
Light rays are focussed through the transparent cornea and lens upon the retina. The central point for image focus (the visual axis) in the human retina is the fovea. Here a maximally focussed image initiates resolution of the finest detail and direct transmission of that detail to the brain for the higher operations needed for perception. Slightly more nasally than the visual axis is the optic axis projecting closer to the optic nerve head. The optic axis is the longest sagittal distance between the front or vertex of the corna and the furthest posterior part of the eyeball. It is about the optic axis that the eye is rotated by the eye muscles. Some vertebrate retinas have instead of a fovea, another specialization of the central retina, known as an area centralis or a visual streak.
Extraocular muscles.
Inserted into the sclera are three pairs of muscles (6 muscles altogether). Two pairs are rectus muscles running straight to the bony orbit of the skull orthogonal to each other (the superior rectus, the inferior rectus, the lateral rectus and the medial rectus muscles). A further pair of muscles, the oblique muscles (superior oblique and inferior oblique) are angled as the name implies obliquely. These muscles, named extraocular muscles rotate the eyeball in the orbits and allow the image to be focussed at all times on the fovea of central retina.
Development of the eye.
The retina is a part of the central nervous system and an ideal region of the vertebrate brain to study, because like other regions of the central nervous system, it derives from the neural tube. The retina is formed during development of the embryo from optic vesicles outpouching from two sides of the developing neural tube. The primordial optic vesicles fold back in upon themselves to form the optic cup with the inside of the cup becoming the retina and the outside remaining a single monolayer of epithelium known as the retinal pigment epithelium. Initially both walls of the optic cup are one cell thick, but the cells of the inner wall divide to form a neuroepithelial layer many cells thick: the retina
CLICK HERE to see a Morph of development (683 K quicktime movie)
Sensory retinal development begins as early as the optic vesicle stage, with the migration of cell nuclei to the inner surface of the sensory retina. Additional retinal development is characterized by the formation of further layers arising from cell division and subsequent cell migration. The retina develops in an inside to outside manner: ganglion cells are formed first and photoreceptors cells become fully mature last.
Further changes in retinal morphology are accomplished by simultaneous formation of multiple complex intercellular connections. Thus by 5 months of gestation most of the basic neural connections of the retina have been established (Mann, 1964).
The functional synapses are made almost exclusively in the two plexiform layers and the perikarya of the nerve cells are distributed in the three nuclear layers.
Photoreceptor cell maturation begins with the formation of outer segments (OS) containing visual pigment from multiple infoldings of the plasma membrane of each cell. Outer segment formation proceeds, and the eye becomes sensitive to light at about 7 months' gestation.
The final portion of the sensory retina to mature is the fovea, where the ganglion cell layer thickening begins during midgestation. The outer nuclear layer is also wider here than elsewhere in the retina and consists almost entirely of developing cone cells. The ganglion cell nuclei migrate radially outwards in a circle, leaving the fovea free of ganglion cell nuclei. Cell-cell attachments persist, however and foveal cone cells alter their shape to accomodate the movement of ganglion cells. Foveal development continues with cell rearrangements and alteration in cone shape until about 4 years after birth (Hendrickson and Yondelis, 1984; Curcio and Hendrickson, 1991).
Surface membranes cover the eye cup and develop into lens, iris and cornea with the three chambers of fluid filled with aqueous and vitreous humors.
Surface membranes cover the eye cup and develop into lens, iris and cornea with the three chambers of fluid filled with aqueous and vitreous humors.
Summary.
In following chapters, we will describe in greater detail the individual nerve cells that make up the retina and the functional pathways into which these neurons are organized. Eventually, we will progress to a stage where we can appreciate the summary diagrams below that show the functional wiring of two well-understood mammalian retinas, namely cat and primate retinas.
References.
Hendrickson, A.E. and Youdelis, C. (1984). The morphological development of the human fovea. Ophthalmology 91, 603-612.
Mann, I. (1964). The development of the human eye. Grune and Stralton, New York.
Ogden, T.E. (1989) Retina: Basic Science and inherited retinal disease, vol 1. The CV Mosby Co. St. Louis.
Curcio, C.A. and Hendrickson, A.E. (1991). Organization and development of the primate photoreceptor mosaic. Prog. Ret. Ret. 10, 89-120.
Types of eyes
There are ten different eye layouts — indeed every way of capturing an image known to man, with the exceptions of zoom and Fresnel lenses. Eye types can be categorized into "simple eyes", with one concave chamber, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface. Note that "simple" does not imply a reduced level of complexity or acuity. Indeed, any eye type can be adapted for almost any behavior or environment. The only limitations specific to eye types are that of resolution — the physics of compound eyes prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eyes, so are better suited to dark-dwelling creatures. Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being cilliated (as in the vertebrates) or rhabdomeric. These two groups are not monophyletic; the cnidaria also possess cilliated cells, and some annelids possess both.
Normal eyes
Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times in vertebrates, cephalopods, annelids, crustacea and cubozoa.
Pit eyes
Pit eyes, also known as stemma, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eyespot, to allow the organism to deduce the angle of incoming light. Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eye". They are small, comprising up to about 100 cells covering about 100 µm. The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material.
Spherical lensed eye
The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index to form a lens, which may greatly reduce the blur radius encountered — hence increasing the resolution obtainable. The most basic form, still seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index, decreasing to the edges; this decreases the focal length and thus allows a sharp image to form on the retina. This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing spherical aberration. Such an inhomogeneous lens is necessary in order for the focal length to drop from about 4 times the lens radius, to 2.5 radii.
Heterogeneous eyes have evolved at least eight times: four or more times in gastropods, once in the copepods, once in the annelids and once in the cephalopods. No aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".
This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimize the effect of eye motion while the animal moves, most such eyes have stabilizing eye muscles.
The ocelli of insects bear a simple lens, but their focal point always lies behind the retina; consequently they can never form a sharp image. This capitulates the function of the eye. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the whole visual field; this fast response is further accelerated by the large nerve bundles which rush the information to the brain. Focusing the image would also cause the sun's image to be focused on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity. This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above).
Multiple lenses
Some marine organisms bear more than one lens; for instance the copepod Pontella has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod, Copilia's eyes have two lenses, arranged like those in a telescope. Such arrangements are rare and poorly understood, but represent an interesting alternative construction. An interesting use of multiple lenses is seen in some hunters such as eagles and jumping spiders, which have a refractive cornea (discussed next): these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution.
Refractive cornea
In the eyes of most mammals, birds, reptiles, and most other terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air, relieving the lens of the function of reducing the focal length. This has freed it up for fine adjustments of focus, allowing a very high resolution to be obtained. As with spherical lenses, the problem of spherical aberration caused by the lens can be countered either by using an inhomogeneous lens material, or by flattening the lens. Flattening the lens has a disadvantage: the quality of vision is diminished away from the main line of focus, meaning that animals requiring all-round vision are detrimented. Such animals often display an inhomogeneous lens instead.
As mentioned above, a refractive cornea is only useful out of water; in water, there is no difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures which have returned to the water — penguins and seals, for example — lose their refractive cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strong cornea.
Reflector eyes
An alternative to a lens is to line the inside of the eye with " mirrors", and reflect the image to focus at a central point. The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out.
Many small organisms such as rotifers, copeopods and platyhelminths use such organs, but these are too small to produce usable images. Some larger organisms, such as scallops, also use reflector eyes. The scallop Pecten has up to 100 millimeter-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses.
There is at least one vertebrate, the spookfish, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from the above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of guanine crystals.
Compound eyes
An image of a house fly compound eye surface by using Scanning Electron Microscope at X457 magnification
Arthropods such as this carpenter bee have compound eyes
A compound eye may consist of thousands of individual photoreceptor units. The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the polarization of light. Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained. This can only be countered by increasing lens size and number. To see with a resolution comparable to our simple eyes, humans would require compound eyes which would each reach the size of their head.
Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image. Compound eyes are common in arthropods, and are also present in annelids and some bivalved molluscs.
Compound eyes, in arthropods at least, grow at their margins by the addition of new ommatidia.
Apposition eyes
Apposition eyes are the most common form of eye, and are presumably the ancestral form of compound eye. They are found in all arthropod groups, although they may have evolved more than once within this phylum. Some annelids and bivalves also have apposition eyes. They are also possessed by Limulus, the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point. (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.)
Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information.
The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is absorbed by the dark wall of the ommatidium. In the other kind of apposition eye, found in the Strepsiptera, lenses are not fused to one another, and each forms an entire image; these images are combined in the brain. This is called the schizochroal compound eye or the neural superposition eye. Because images are combined additively, this arrangement allows vision under lower light levels.
Superposition eyes
The second type is named the superposition eye. The superposition eye is divided into three types; the refracting, the reflecting and the parabolic superposition eye. The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This kind is used mostly by nocturnal insects. In the parabolic superposition compound eye type, seen in arthropods such as mayflies, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied decapod crustaceans such as shrimp, prawns, crayfish and lobsters are alone in having reflecting superposition eyes, which also has a transparent gap but uses corner mirrors instead of lenses.
Parabolic superposition
This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.
Other
The compound eye of a dragonfly
Good fliers like flies or honey bees, or prey-catching insects like praying mantis or dragonflies, have specialized zones of ommatidia organized into a fovea area which gives acute vision. In the acute zone the eyes are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution.
There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the mysid shrimp Dioptromysis paucispinosa. The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialized retina. The resulting eye is a mixture of a simple eye within a compound eye.
Another version is the pseudofaceted eye, as seen in Scutigera. This type of eye consists of a cluster of numerous ocelli on each side of the head, organized in a way that resembles a true compound eye.
The body of Ophiocoma wendtii, a type of brittle star, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chitons.
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