Duplex Retina in the Mesopelagic Deep-Sea Teleost Lestidiops affinis (Ege, 1930)

The eye of the deep-sea teleost Lestidiops affinis has been examined primarily by light microscopy and found to possess a duplex retina consisting of two main divisions, a pure-cone and a pure-rod region, with a narrow zone of transition, possessing both cones and rods, joining the two. The pure-cone region is located in the temporal (caudal) part of the retina subserving binocular vision in the rostral direction. It has an area temporalis retinae with particularly long and densely packed single cones arranged in a regular hexagonal mosaic. Joined (double or twin) cones have not been recognized with certainty in the pure-cone region. The pure-rod region, comprising the larger part of the retina, contains rods grouped in bundles separated by retinal pigment epithelium (RPE) processes with pigmented cores. The synaptic endings of the rods are arranged in separate clusters in the outer plexiform layer, there being apparently a separate rod pedicle cluster beneath (vitread to) each rod bundle. Structural comparisons with certain other deep-sea teleosts suggest the likely presence of a retinal tapetum in L. affinis, i.e. each single cone or rod bundle is situated in a reflecting pit formed by the RPE, with a discrete reflector apposed to the tip of each cone outer segment and the tips of the outer segments of each square-cut rod bundle.     

Peculiarities of the eye morphology and the spectral sensitivity of the retinal photoreceptors of the Pacific saury <Emphasis Type="Italic">Cololabis saira</Emphasis>

This study examines some peculiarities of the eye organization and spectral properties of retinal photoreceptors of the Pacific saury Cololabis saira. The saury has relatively large eyes with a developed accomodation apparatus and an area of enhanced visual acuity (the fovea) in the retina. A specialized pigmented septum is observed in the vitreal cavity, which is supposed to function as a light-shading screen. The retina contains numerous rods and single and double cones arranged in a square mosaic pattern. Microspectrophotometric measurements indicated that their max occurs at 502 (rods), 380 (single cones), and 478/565 (double cones) nm. Such properties can provide color vision in a broad spectral range, including UV light. The peripheral visual apparatus of the Pacific saury is typical of active diurnal predatory fish that inhabit shallow and upper pelagic water layers.     

Retinal photoreceptors and visual pigments in Boa constrictor imperator

The photoreceptors of Boa constrictor, a boid snake of the subfamily Boinae, were examined with scanning electron microscopy and microspectrophotometry. The retina of B. constrictor is duplex but highly dominated by rods, cones comprising 11% of the photoreceptor population. The rather tightly packed rods have relatively long outer segments with proximal ends that are somewhat tapered. There are two morphologically distinct, single cones. The most common cone by far has a large inner segment and a relatively stout outer segment. The second cone, seen only infrequently, has a substantially smaller inner segment and a finer outer segment. The visual pigments of B. constrictor are virtually identical to those of the pythonine boid, Python regius. Three different visual pigments are present, all based on vitamin A(1.) The visual pigment of the rods has a wavelength of peak absorbance (lambda(max)) at 495 +/- 2 nm. The visual pigment of the more common, large cone has a lambda(max) at 549 +/- 1 nm. The small, rare cone contains a visual pigment with lambda(max) at 357 +/- 2 nm, providing the snake with sensitivity in the ultraviolet. We suggest that B. constrictor might employ UV sensitivity to locate conspecifics and/or to improve hunting efficiency. The data indicate that wavelength discrimination above 430 nm would not be possible without some input from the rods.     

The visual cells of the corsula mullet

The eye of Rhinomugil corsula has a duplex retina differentiated into dorsal and ventral halves, with the ventral retina 116·4 μm thicker than the dorsal retina. The rods of the ventral retina are relatively longer, with longer outer segments. The nuclei of the outer nuclear layer of the dorsal and ventral halves are in four and six to seven rows respectively. The rod outer segment bears a single incision. The mitochondria of cone and rod inner segments has a vitreal-scleral gradient. Single and double cones are present in both halves, with triple cones in the dorsal half only. The outer segments of double cones are equal and united. The single cones have two connecting cilia. The cone cells are arranged in a square mosaic with four double cones and five single cones to each unit in the dorsal half, and in a rhombic pattern in the ventral half.     

Regional variations in the outer retina of atherinomorpha (Beloniformes,Atheriniformes, Cyprinodontiformes: Teleostei): photoreceptors,cone patterns,and cone densities

The outer retinae of adults of 13 atherinomorph species, representing nine different families, were examined by both light and electron microscopy. The retinae were investigated with respect to photoreceptor types, cone densities, and cone patterns. All data were composed to eye maps. This procedure allows an interspecific comparison of the regional differences within the outer retina among these shallow-water fish. Furthermore, for a more detailed pattern analysis nitro-blue tetrazolium chloride- (NBT)-stainings in the retina of Melanotaenia maccullochi are presented. Apart from rods, eight morphologically different cone types could be identified: short, intermediate, and long single cones, double cones (equal and unequal), triple cones (triangular and linear), and in Ameca splendens one quadruple cone. Dimensions and occurrence of photoreceptors vary among the respective species and within the retinal regions. In the light-adapted state, the cones are arranged in highly ordered mosaics. Five different cone tessellation types were found: row patterns, twisted row patterns, square patterns, pentagonal patterns, and, exclusively in Belone belone, a hexagonal pattern. In Melanotaenia maccullochi the different spectral photoreceptor classes correspond well with the distribution of morphological photoreceptor classes within the mosaic. Double cone density maxima together with a highly ordered cone arrangement usually occur in the nasal and/or ventral to ventrotemporal retina. In most of the species that were examined these high-density regions are presumed to process visual stimuli from the assumed main directions of vision, which mainly depend on feeding behavior and predator pressure. Our findings are discussed with respect to the variable behavioral and visual ecology and phylogeny of the respective species.     

Fine structure of the retina of black bass, Micropterus salmoides (Centrarchidae, Teleostei).

The structure of light- and dark-adapted retina of the black bass, Micropterus salmoides has been studied by light and electron microscopy. This retina lacks blood vessels at all levels. The optic fiber layer is divided into fascicles by the processes of Müller cells and the ganglion cell layer is represented by a single row of voluminous cells. The inner nuclear layer consists of two layers of horizontal cells and bipolar, amacrine and interplexiform cells. In the outer plexiform layer we observed the synaptic terminals of photoreceptor cells, rod spherules and cone pedicles and terminal processes of bipolar and horizontal cells. The spherules have a single synaptic ribbon and the pedicles possess multiple synaptic ribbons. Morphologically, we have identified three types of photoreceptors: rods, single cones and equal double cones which undergo retinomotor movements in response to changes in light conditions. The cones are arranged in a square mosaic whereas the rods are dispersed between the cones.     

Comparative morphology and cytology of the eye,with particular reference to the retina,in lizardfishes (Synodontidae,Teleostei)

Fishelson, L., Delarea, Y. and Goren, M. 2012. Comparative morphology and cytology of the eye, with particular reference to the retina, in lizardfishes (Synodontidae, Teleostei). —Acta Zoologica (Stockholm) 93 : 68–79. The retinas of nine species of lizardfishes (Synodontidae) are composed of double cones, single cones, and rods. The cones are 16–28 μm long, and their number in the fundus of adult Synodus variegatus reaches ca. 32,900 mm² (varying from ca. 300,000 to ca. 390,000 in a 10 mm² of the retina), while in Saurida spp., they number ca. 12,000–14,000/mm². The cone ellipsoids are with up to 600 mitochondria, 0.5–1.6 μm in diameter. The rods are 30–50 μm long; their outer segments 0.6–1.2 μm thick and 15–18 μm long; their inner segments elongated. Their number varies from 15 to 128 million/retina. In fish of similar dimensions but of different species, the number of visual cells in the retina differs. In all species, the eyes increase from 2.0 mm in diameter in the smallest fish studied to 12 mm in the largest one. With eye growth, the retina in the various species increases from ca. 3.8 mm² in the smallest fish to ca.160.0 mm² in the large Saurida macrolepis. The possible ecological aspects of the observed phenomena are discussed.     

Morphology of the retina in deep‐water fish Nezumia sclerorhynchus (Valenciennes, 1838) (Gadiformes: Macrouridae)

Using light microscopy, we examined the retina of benthopelagic fish Nezumia sclerorhynchus. Although the retina is typical of other vertebrates, having three nuclear and two synaptic layers, it presents some features associated with the animal's deep‐sea habitat. A stratum argenteum containing iridescent crystals is located in the choroid. The pigment cell layer shows bulky cells filled with melanin granules but without the typical apical processes. The visual cells, consisting of a big population of rods, are arranged in several banks. No cones were observed. The outer segments are very long and cylindrical, and the inner segments are constituted by a small ellipsoid at the proximal end. The outer nuclear layer contains several rows of oval nuclei, and the spherules in the outer plexiform layer have less regular outlines than nuclei. The inner retina is characterized by very large horizontal cells, and presumable bipolar and amacrine cells separated by large spaces that are occupied by neuronal processes. Finally, the low density of ganglion cells produces a thin nerve fibre layer. The results of this study suggest that the retina of Nezumia sclerorhyncus exhibits high visual sensitivity and that vision is a sense that plays an important role in its behaviour.     

Morphology and density distribution of cone photoreceptor in the retina of the atlantic stingray,Dasyatis sabina

Light miscroscopy of the retina in the Atlantic stingray, Dasyatis sabina, reveal two distinct photoreceptor types. Histological criteria establish the two cell types as morphological rods and cones. The retina was studied through a sequential series of vertical sections in a protocol designed to evaluate the entire retina. By performing differential counts in multiple regions and expressing cones as a percent of the total photoreceptors in a domain, it was possible to determine relative cone density and distribution. Cone distribution varies throughout the retina in two patterns. First, relative cone density gradually decreases peripherally. Second, a cone-rich band occurs along the horizontal axis of the eye, extending from ora serrata to ora serrata. This structure appears to be a visual streak. Both distribution patterns are statistically significant and are consistent among animals regardless of age. © 1994 Wiley-Liss, Inc.     

Ontogeny of Double Cones in the Retina of Perch Fry (Perca fluviatilis,Teleostei)

The formation of double cones in the retina of fry of Perca fluviatilis has been investigated by light and electron microscopy. The retina of newly hatched fry is provided with single cones and rods, single cones being the predominant receptor type. Double cones are seen for the first time 22 days after hatching. Mitoses are observed in the periphery of the retina, but are also seen in more central parts of the retina containing differentiated receptors and a cone mosaic. The fate of the cells resulting from the centrally located mitoses is not known. No signs of longitudinal fission of differentiated single cones are seen. It is suggested that double cones in the retina of perch fry arise by fusion of single cones which associate closely and develop subsurface cisterns coextensive with the region of intimate contact in the ellipsoid. During the first few weeks after hatching, there is a gradual shift in arrangement of the cones. In the newly hatched fry, the single cones are arranged in rows. When double cones are first seen, square-pattern units appear, built up from four double cones and a single cone.     

Photoreceptor topography in the duplex retina of the paddlefish (Polyodon spathula)

Retinal whole-mount preparations from the eyes of the North American paddlefish, Polyodon spathula, were examined with a combination of bright field and differential interference contrast microscopy. The entire retina was mapped and population counts of rod and cone photoreceptors were made at regular intervals throughout the retina. The retina is dominated by rods, but a significant percentage (ca. 38%) of the photoreceptors are cones. Mean cone packing density for the entire retina is 6,402+/-1,216 cones/mm2. There is a small (16%) but statistically significant difference between cone packing density in the dorsal retina (6,674+/-1,168 cones/mm2) and the ventral retina (5,745+/-1,076 cones/mm2). There is no region of unusually high cone concentration that might be construed as a fovea or a visual streak. Mean rod packing density for the entire retina is 10,271+/-1,205 rods/mm2. Except in the far periphery, where rods are less numerous, the density of rods is fairly uniform throughout the retina. The data are discussed with regard to paddlefish habitat and behavior.     

The photoreceptors and visual pigments of the garter snake (Thamnophis sirtalis): a microspectrophotometric, scanning electron microscopic and immunocytochemical study

Scanning electron microscopy, immunocytochemistry, and single cell microspectrophotometry were employed to characterize the photoreceptors and visual pigments in the retina of the garter snake, Thamnophis sirtalis. The photoreceptor population was found to be comprised entirely of cones, of which four distinct types were identified. About 45.5% of the photoreceptors are double cones consisting of a large principal member joined near the outer segment with a much smaller accessory member. About 40% of the photoreceptors are large single cones, and about 14.5% are small single cones forming two subtypes. The outer segments of the large single cones and both the principal and accessory members of the doubles contain the same visual pigment, one with peak absorbance near 554 nm. The small single cones contain either a visual pigment with peak absorbance near 482 nm or one with peak absorbance near 360 nm. Two classes of small single cones could be distinguished also by immunocytochemistry and scanning electron microscopy. The small single cones with the 360-nm pigment provide the garter snake with selective sensitivity to light in the near ultraviolet region of the spectrum. This ultraviolet sensitivity might be important in localization of pheromone trails. Accepted: 10 March 1997     

Ontogeny of the photoreceptors in the embryonic retina of the viviparous guppy,Poecilia reticulata P. (Teleostei)

Summary Ultrastructural analyses of retinal development in the guppy embryo show that at midgestation all types of photoreceptors are differntiated in the fundus, and at birth differentiation extends over the whole retina. Formation of discs of outer segments is more rapid in rods than in cones. Double cones differentiate simultaneously with long single cones and are formed by the adhesion of two primordial inner segments; short single cones develop last. Wherever cones are differentiated, they are arranged in an adult-type square mosaic. The rods in the embryo, as opposed to the adult, are likewise regularly arranged within the mosaic unit.These results are at variance with the generally held opinion that adult teleosts which possess duplex retinae have larvae with pure cone retinae, and that rods, double cones and mosaics appear in late larval life or only at metamorphosis.In the double cones of the guppy embryo subsurface cisternae develop along the adjoining primordial inner segments. Additionally, regularly distributed subsurface cisternae are formed in the regions of intimate contact of long single cones with double cones and rods.We suggest that the early development of rods and double cones, and a square-mosaic with regular distribution of rods and subsurface cisternae, provide the newly born with a fully functional optical apparatus, especially suited to perception of movements. This is necessary for its survival against predatory, especially maternal, attacks.This work was in part subsidized by a grant from the Medical Research Council of Ireland to Y.W.K.     

Developmental Changes in the Visual Pigments of the Yellowfin Tuna,Thunnus Albacares

Previous studies have suggested that adult tunas have only two visual pigments in their retinas - a rod pigment with a wavelength at maximum absorbance (u max) around 485 nm and one with similar u max in both twin and single cones inferred from extraction data. Using microspectrophotometry we confirm the presence of a u max 483 nm visual pigment in the rods of adult yellowfin tuna and a u max 485 nm pigment in both members of the twin cones. However, all single cones contain a previously undetected violet visual pigment with u max 426 nm making the adult yellowfin tuna a photopic dichromat. The situation for larvae and early juveniles is different from that of the adults. The all single-cone retina of preflexion larvae shows a wide distribution in individual cone absorbances suggesting not only mixtures of the two adult cone pigments, but the presence of at least a third visual pigment with u max greater than 560 nm. With growth, the variation in cone absorbances decreases with convergence to the adult condition coincident with cone twinning. The significance of u max variability, multiple visual pigment expression and age-related differences are discussed in terms of the visual ecology of larval, juvenile and adult tunas.     

Photoreceptors and visual pigments in the retina of the fully anadromous green sturgeon (<Emphasis Type="Italic">Acipenser medirostrus</Emphasis>) and the potamodromous pallid sturgeon (<Emphasis Type="Italic">Scaphirhynchus albus</Emphasis>)

Green sturgeon and pallid sturgeon photoreceptors were studied with scanning electron microscopy (SEM), microspectrophotometry and, in the case of the green sturgeon, retinal whole-mounts. The retinas of both species contain both rods and cones: cones comprise between 23% (whole-mount) and 36% (SEM) of the photoreceptors. The cone population of both species is dominated by large single cones, but a rare small single cone is also present. In both species, most rods have long outer segments of large diameter. A rod with a relatively thin outer segment is present in the pallid sturgeon retina. Mean cone packing density for the entire green sturgeon retina is 4,690±891 cones/mm², with the dorsal retina 14% more dense than the ventral. There is evidence for a horizontal visual streak just above and including the optic disc. Mean rod packing density is 16,006±1,668 rods/mm² for the entire retina, and fairly uniform throughout. Both species have rods with peak absorbance near 540 nm, as well as short-wavelength-sensitive cones (green: 464.5±0.7 nm; pallid: 439.7±3.5 nm); middle-wavelength-sensitive cones (green: 538.0±1.4 nm; pallid: 537.0±1.7 nm); and long-wavelength-sensitive cones (green: 613.9±3.0 nm; pallid: 617.8±7.6 nm).     

Developmental Changes in the Visual Pigments of the Yellowfin Tuna, Thunnus Albacares

Previous studies have suggested that adult tunas have only two visual pigments in their retinas - a rod pigment with a wavelength at maximum absorbance ( λmax ) around 485 nm and one with similar λmax in both twin and single cones inferred from extraction data. Using microspectrophotometry we confirm the presence of a λmax 483 nm visual pigment in the rods of adult yellowfin tuna and a λmax 485 nm pigment in both members of the twin cones. However, all single cones contain a previously undetected violet visual pigment with λmax 426 nm making the adult yellowfin tuna a photopic dichromat. The situation for larvae and early juveniles is different from that of the adults. The all single-cone retina of preflexion larvae shows a wide distribution in individual cone absorbances suggesting not only mixtures of the two adult cone pigments, but the presence of at least a third visual pigment with λmax greater than 560 nm. With growth, the variation in cone absorbances decreases with convergence to the adult condition coincident with cone twinning. The significance of λmax variability, multiple visual pigment expression and age-related differences are discussed in terms of the visual ecology of larval, juvenile and adult tunas.     

Light-induced rod and cone cell death and regeneration in the adult albino zebrafish (Danio rerio) retina

Light-induced photoreceptor cell degeneration has been studied in several species, but not extensively in the teleost fish. Furthermore, the continual production of rods and cones throughout the teleost's life may result in regeneration of lost rods and cones. We exposed adult albino zebrafish to 7 days of constant darkness, followed by 7 days of constant 8000 lux light, followed by 28 days of recovery in a 14-h light:10-h dark cycle. We characterized the resulting photoreceptor layer cell death and subsequent regeneration using immunohistochemistry and light microscopy. Within the first 24 h of constant light, the zebrafish retina exhibited widespread rod and cone cell apoptosis. High levels of cell proliferation within the inner nuclear layer (INL) were observed within the first 3 days of constant light, as assessed by immunodetection of proliferating cell nuclear antigen and BrdU labeling. The proliferating cells within the INL were closely associated with the radial processes of Müller glia, similar to the pluripotent retinal stem cells observed during embryonic development. Using antibodies generated against the individual zebrafish opsins, we determined that rods and the green, blue, and ultraviolet cone cells were replaced within the 28 day recovery period. While both rods and cones were replaced, the well-ordered cone cell mosaic was not reestablished.     

Tetrachromatic colour vision in the duck (Anas platyrhynchos L.):microspectrophotometry of visual pigments and oil droplets

Summary Microspectrophotometric examination of the visual receptors of the duck,Anas platyrhynchos, revealed four types of single cone containing visual pigments absorbing maximally at about 420 nm, 452 nm, 502 nm and 570 nm. A single population of double cones contained the P570 in both members. Rods absorbed maximally at 505 nm.Within the single cones, three types of oil droplet, acting as cut-off filters, were identified by the wavelength at which 50% transmission occurred, approximately 580, 515 and 450 nm. A further droplet, transparent throughout the visible spectrum, was also found in a small population of single cones. A fifth droplet type with a variable cutoff between 475–500 nm was located in the principal member of the double cones.The optical density of the anterior half of the eye, established by spectrophotometry, was used, in conjunction with the visual pigment and oil droplet combinations found within intact cones, to estimate the relative spectral sensitivities of the major cone types within the retina.     

Fine structure and development of the retina of the grenadier anchovy Coilia nasus (Engraulididae, Clupeiformes)

A study of the morphogenesis of the grenadier anchovy retina was undertaken using light and electron microscopy. Five developmental stages from prelarvae 3 days after fertilization to adult fish were studied. In addition to the general morphology of the eye and retina, special emphasis was given to the development of the photoreceptors and pigment epithelium (PE). The earliest retinae showing structural features indicative of a functioning eye are pure cone retinae composed of rows of alternating long and short cones forming a transient, tesselated pattern. At this stage there is a conventional PE containing melanin. In older stages cone rows are separated by the newly formed rods and by PE wedges filled with diffusely reflecting guanine crystallites. The findings are compared with the retinae of other engraulidids and with the development of teleost retinae in general. Moreover, the observed structural changes are discussed with respect to the photic habitat conditions of these anadromous fish that move between coastal waters, estuary, and river.     

Ultrastructure of the distal retina of the adult zebrafish, Danio rerio

The organization, morphological characteristics, and synaptic structure of photoreceptors in the adult zebrafish retina were studied using light and electron microscopy. Adult photoreceptors show a typical ordered tier arrangement with rods easily distinguished from cones based on outer segment (OS) morphology. Both rods and cones contain mitochondria within the inner segments (IS), including the large, electron-dense megamitochondria previously described (Kim et al.) Four major ultrastructural differences were observed between zebrafish rods and cones: (1) the membranes of cone lamellar disks showed a wider variety of relationships to the plasma membrane than those of rods, (2) cone pedicles typically had multiple synaptic ribbons, while rod spherules had 1-2 ribbons, (3) synaptic ribbons in rod spherules were ∼2 times longer than ribbons in cone pedicles, and (4) rod spherules had a more electron-dense cytoplasm than cone pedicles. Examination of photoreceptor terminals identified four synaptic relationships at cone pedicles: (1) invaginating contacts postsynaptic to cone ribbons forming dyad, triad, and quadrad synapses, (2) presumed gap junctions connecting adjacent postsynaptic processes invaginating into cone terminals, (3) basal junctions away from synaptic ribbons, and (4) gap junctions between adjacent photoreceptor terminals. More vitread and slightly farther removed from photoreceptor terminals, extracellular microtubule-like structures were identified in association with presumed horizontal cell processes in the OPL. These findings, the first to document the ultrastructure of the distal retina in adult zebrafish, indicate that zebrafish photoreceptors have many characteristics similar to other species, further supporting the use of zebrafish as a model for the vertebrate visual system.