Retinal Tapetum Research Articles

Micro-morphology of the retina of the light-adapted African catfish (Clarias gariepinus).

The current study aimed to investigate the ultrastructure of the retinal photoreceptors of the African catfish and to demonstrate their adaptation to nocturnal or diurnal visions or by the two ways. The eyes of eight adult catfish were collected during the daytime, and the retinae were separated and examined by light and transmission electron microscopy. The photoreceptors' layer appeared in contact with the retina's pigmented epithelium. Two photoreceptors were detected in cones and hidden rods. Cones predominate in light-adapted retinae. The outer segments of cones appeared between the retinal pigmented epithelium protrusions, which indicates the movement of melanosomes away from the photoreceptors as a retinomotor response of the catfish. The two types of retinal tapetum were in between cones. The first type, the cored granules, were large, spherical, and had black peripheral parts and central lucent parts, and contained some granules. The second type was Guanine crystallites of tapetum lucidum, which were small electron-lucent, and their shape varied from spherical to rectangular. Melanosomes vary in shape from spherical to elliptical. The Müller cells were darkly stained elongated cells that measured about 5.5-8.5μm in length and 2.2-2.5μm in width, and their microvilli appeared between the inner segments of the rods and cones. Müller cell processes were extended from the photoreceptor layer to the inner limiting membrane. Zonula occludentes appeared between the Müller cell processes and the internal segment of the rods and cones. African catfish have eyes which are adapted not only for nocturnal but also for daytime light.

The eye of the tongue sole Cynoglossus bilineatus (Lacepède, 1802) (Teleostei: Pleuronectiformes)

We report the ocular features of the tongue sole, Cynoglossus bilineatus (Lacepède, 1802), a marine, bottom-dwelling flatfish. In this species, both eyes are located juxtaposed on the same side of the flat head. Histology revealed the sclera to be fibrous (collagenous) in nature. The choroid possesses the choriocapillaris, and adjacent to it, 3–4 rows of iridophores with stacks of cytoplasmic platelets. No choroidal gland is present. The retinal pigment epithelium (RPE) contains scanty melanin granules. Its vitread half is modified into a dense tapetum with lipid spheres (about 0.34 μm in diameter). In juveniles, the tapetal spheres arise by budding from the smooth endoplasmic reticulum of the RPE. There are blood vessels within the retina; the vitreal vessels penetrate the retina and ramify close to the level of the outer limiting membrane. The vessels are capillaries in nature. The photoreceptor layer contains abundant rods, and twin cones and single cones, being arranged into square mosaics. The optic disc is non-pleated and shows pan- cytokeratin immunopositivity, which is related to the bundled cytokeratin filaments detected in astrocytes by electron microscopy. The retinal tapetum and choroidal iridophores help the species to live in a muddy bottom having dim-light environment. The lack of a choroidal gland, hypoxic aquatic condition and presence of a dense retinal tapetum (that limits O2 transport to the photoreceptors) appear to have favored the proliferation of vitreal vessels within the retina in this species. The fibrous sclera has probably arisen to provide structural support to the eye in migration from the lateral to the dorsal aspect of the head during larval metamorphosis.

Colour vision in the tropical giant mottled eel Anguilla marmorata as determined by classical conditioning test

The giant mottled eel Anguilla marmorata Quoy & Gaimard is an important aquaculture candidate in eel farming industry. The high economic value of the species leads to intensive cultivation affecting its welfare under farm condition. Since colour environment affects the welfare of confined fishes, basic information on colour vision is necessary. By means of a behavioural experiment of A. marmorata elvers in the laboratory, we determined colour vision under natural daylight. The elvers were subjected to classical conditioning to associate a reward feed with a blue or a green stimulus placed amid seven shades of grey. The elvers learned this visual task after 30 or 36 trials and thus the ability of colour discrimination was confirmed. Since the elvers have the retinal tapetum, it may be suggested that colour vision in dim light enables crepuscular feeding by the A. marmorata elvers, clearly an advantage in aquaculture.

Colour discrimination in dim light by the larvae of the African catfish Clarias gariepinus

Many demersal fish species undergo vertical shifts in habitats during ontogeny especially after larval metamorphosis. The visual spectral sensitivity shifts with the habitat, indicating a change in colour vision. Colour vision depends on sufficient ambient light and becomes ineffective at a particular low light intensity. It is not known how fishes see colour in dim light. By means of a behavioural experiment on larval African catfish Clarias gariepinus in the laboratory, we determined colour vision and colour discrimination in dim light. Light-adapted larvae were subjected to classical conditioning to associate a reward feed with a green or a red stimulus placed among 7 shades of grey. The larvae learned this visual task after 70 and 90 trials. A different batch of larvae were trained to discriminate between green and red and then tested for the ability to discriminate between these colours, as the light intensity was reduced. The larvae learned this visual task after 110 trials in bright light and were able to discriminate colours, as light was dimmed until 0.01 lx, the minimal illuminance measurable in this study, and similar to starlight. The retinae of the larvae were found to be light adapted at 0.01 lx; thus indicating cone-based colour vision at this illuminance. For comparison, three human subjects were tested under similar conditions and showed a colour vision threshold at between 1.5 and 0.1 lx. For the larvae of C. gariepinus, the ability of colour discrimination in dim light is probably due to its retinal tapetum, which could increase the sensitivity of cones.

Early appearance of the retinal tapetum, cones, and rods in the larvae of the African catfish Clarias gariepinus

In the retina of the African catfish Clarias gariepinus, the pigment epithelium and the tapetum were formed in newly hatched larvae, the cones developed within 2 days, and the rods within 3 days after hatching. The retinal tapetum shone under surface light under a light microscope; the shine was located in the apical projections of the pigment epithelial cells. Early appearance of the retinal elements enables African catfish larvae to see and feed well even in dim light.

The relative eye size, visual cells, cone mosaic and retinal tapetum in the spotted barb Puntius binotatus (Valenciennes, 1842)

The relative eye size, types of visual cell and mosaic, and the existence of retinal tapetum in the eyes of the spotted barb Puntius binotatus were determined to gain baseline information on its visual capability. The P. binotatus acquired relatively larger eye size than the other similar sized freshwater fish species with its retina being contained both cone and rod visual cells (cone ellipsoid to outer nuclei ratio = 1: 5.7). Three types of cone were identified (double-, central single- and corner single- cones), arranging in the square mosaic, and the retinal tapetum was determined to be existed. These results evidenced that the P. binotatus has good visual capability as it possessed both the photopic and scotopic visions. Vision can be the primary sense for the P. binotatus . Further study is needed to gain more information on the vision of this species.

Grouped retinae and tapetal cups in some Teleostian fish: Occurrence, structure, and function

This article presents a summary and critical review of what is known about the 'grouped retina', a peculiar type of retinal organization in fish in which groups of photoreceptor cell inner and outer segments are arranged in spatially separated bundles. In most but not all cases, these bundles are embedded in light-reflective cups that are formed by the retinal pigment epithelial cells. These cups constitute a specialized type of retinal tapetum (i.e., they are biological 'mirrors' that cause eye shine) and appear to be optimized for different purposes in different fishes. Generally, the large retinal pigment epithelial cells are filled with light-reflecting photonic crystals that consist of guanine, uric acid, or pteridine depending on species, and which ensure that the incoming light becomes directed onto the photoreceptor outer segments. This structural specialization has so far been found in representatives of 17 fish families; of note, not all members of a given family must possess a grouped retina, and the 17 families are not all closely related to each other. In many cases (e.g., in Osteoglossomorpha and Aulopiformes) the inner surface of the cup is formed by three to four layers of strikingly regularly shaped and spaced guanine platelets acting as an optical multilayer. It has been estimated that this provides an up to 10fold increase of the incident light intensity. In certain deep-sea fish (many Aulopiformes and the Polymixidae), small groups of rods are embedded in such 'parabolic mirrors'; most likely, this is an adaptation to the extremely low light intensities available in their habitat. Some of these fishes additionally possess similar tapetal cups that surround individual cones and, very likely, also serve as amplifiers of the weak incident light. In the Osteoglossomorpha, however, that inhabit the turbid water of rivers or streams, the structure of the cups is more complex and undergoes adaptation-dependent changes. At dim daylight, probably representing the usual environmental conditions of the fish, the outer segments of up to 30 cone cells are placed at the bottom of the cup where light intensity is maximized. Strikingly, however, a large number of rod receptor cells are positioned behind each mirroring cup. This peculiar arrangement (i) allows vision at deep red wavelenghts, (ii) matches the sensitivity of rod and cone photoreceptors, and (iii) facilitates the detection of low-contrast and color-mixed stimuli, within the dim, turbid habitat. Thus, for these fish the grouped retina appears to aid in reliable and quick detection of large, fast moving, biologically relevant stimuli such as predators. Overall, the grouped retina appears as a peculiar type of general retinal specialization in a variety of fish species that is adaptive in particular habitats such as turbid freshwater but also the deep-sea. The authors were prompted to write this review by working on the retina of Gnathonemus petersii; the data resulting from this work (Landsberger et al., 2008; Kreying et al., 2012) are included in the present review.

Ontogenetic eye development and related behavioural changes in larvae and juveniles of barramundi Lates calcarifer (Bloch)

Larvae and juveniles of barramundi Lates calcarifer (Bloch) were examined for the development of the retina, occurrence of the retinomotor response and retinal tapetum and change in eye size with age in days. The barramundi hatched with unpigmented non-functional eyes in which retinal cells had not yet differentiated into the various elements. Soon it was followed by rapid changes in the histology of the retina. Two-day-old larvae had a well-pigmented retina with area temporalis which would allow acute vision and prey attack in the nasal direction. At 10 days, rod cells and the retinal tapetum first appeared in the retina and the retinomotor response first occurred; these would allow feeding in dim light. The retinal tapetum moved in unison with the cones and the pigment epithelium during the retinomotor response. At 26 days, the horizontal cells were divided into two layers and the twin cones appeared. These changes in the eyes occurred concurrently or in anticipation of behavioural changes, such as the onset of the first feeding at 2 days, the shift of habitat from coastal waters to swamps at the notochord flexion stage at 7–15 days, the abrupt change in feeding behaviour from roving zooplanktivore to lurking predator at 25–30 days and a later shift of habitat from turbid swamps to open coastal or lake areas at the early juvenile stage.

A Novel Vertebrate Eye Using Both Refractive and Reflective Optics

Sunlight is attenuated rapidly in the ocean, resulting in little visually useful light reaching deeper than approximately 1000 m in even the clearest water. To maximize sensitivity to the relatively brighter downwelling sunlight, to view the silhouette of animals above them, and to increase the binocular overlap of their eyes, many mesopelagic animals have developed upward-pointing tubular eyes. However, these sacrifice the ability to detect bioluminescent and reflective objects in other directions. Thus, some mesopelagic fish with tubular eyes extend their visual fields laterally and/or ventrally by lensless ocular diverticula, which are thought to provide unfocused images, allowing only simple detection of objects, with little spatial resolution. Here, we show that a medial mirror within the ventrally facing ocular diverticulum of the spookfish, Dolichopteryx longipes, consisting of a multilayer stack derived from a retinal tapetum, is used to reflect light onto a lateral retina. The reflective plates are not orientated parallel to the surface of the mirror. Instead, plate angles change progressively around the mirror, and computer modeling indicates that this provides a well-focused image. This is the first report of an ocular image being formed in a vertebrate eye by a mirror.

Vision in the southern hemisphere lampreyMordacia mordax: Spatial distribution, spectral absorption characteristics, and optical sensitivity of a single class of retinal photoreceptor

The dorso-laterally located eyes of the southern hemisphere lamprey Mordacia mordax (Agnatha) contain a single morphological type of retinal photoreceptor, which possesses ultrastructural characteristics of both rods and cones. This photoreceptor has a large refractile ellipsosome in the inner segment and a long cylindrical outer segment surrounded by a retinal pigment epithelium that contains two types of tapetal reflectors. The photoreceptors form a hexagonal array and attain their peak density (33,200 receptors/mm2) in the ventro-temporal retina. Using the size and spacing of the photoreceptors and direct measures of aperture size and eye dimensions, the peak spatial resolving power and optical sensitivity are estimated to be 1.7 cycles deg-1 (minimum separable angle of 34'7'') and 0.64 microm2 steradian (white light) and 1.38 microm2 steradian (preferred wavelength or lambdamax), respectively. Microspectrophotometry reveals that the visual pigment located within the outer segment is a rhodopsin with a wavelength of maximum absorbance (lambdamax) at 514 nm. The ellipsosome has very low absorptance (<0.05) across the measured spectrum (350-750 nm) and probably does not act as a spectral filter. In contrast to all other lampreys studied, the optimized receptor packing, the large width of the ellipsosome-bearing inner segment, together with the presence of a retinal tapetum in the photophobic Mordacia, all represent adaptations for low light vision and optimizing photon capture.

Comparative morphology of the tapetum lucidum (among selected species).

The phenomenon of 'eye-shine' is seen in a variety of animal species, and is generally thought to be related to the presence of an intraocular reflecting structure, the tapetum lucidum. The tapetum lucidum is a biologic reflector system that is a common feature in the eyes of vertebrates. It normally functions to provide the light-sensitive retinal cells with a second opportunity for photon-photoreceptor stimulation, thereby enhancing visual sensitivity at low light levels. The tapetum lucidum is presented here according to a classification based on the location, as well as the composition, of this reflective layer. Finally, the physical and chemical properties, as well as the origins of the different tapeta lucida, are discussed and compared. The anatomic and biochemical aspects of the tapetum lucidum in various vertebrates are examined. Morphologic observations were made from paraffin and plastic embedded specimens. Specimens were treated with traditional stains and observed by light and transmission electron microscopy. Some species (primates, squirrels, birds, red kangaroo and pig) do not have this structure and they usually are diurnal animals. In vertebrates, the tapetum lucidum exhibits diverse structure, organization and composition. Therefore, the retinal tapetum (teleosts, crocodilians, marsupials, fruit bat), the choroidal guanine tapetum (elasmobranchs), the choroidal tapetum cellulosum (carnivores, rodents, cetacea), and the choroidal tapetum fibrosum (cow, sheep, goat, horse) are described. The tapetum lucidum represents a remarkable example of neural cell and tissue specialization as an adaptation to a dim light environment and, despite these differences, all tapetal variants act to increase retinal sensitivity by reflecting light back through the photoreceptor layer. These variations regarding both its location and structure, as well as the choice of reflective material, may represent selective visual adaptations associated with their feeding behavior, in response to the use of specific wavelengths and amount of reflectance required.

Evolution of the tapetum. Schwab IR, Yuen CK, Buyukmihci NC, Blankenship TN, Fitzgerald PG. Trans Am Ophthalmol Soc 2002;100:187–200

Ocular specimens of representative fish in piscine families, including Acipenseridae, Cyprinidae, and Chacidae; the reptilian family Corcodylidae; the mammalian family Felidae; and the Lepidopteran family Sphingidae were reviewed and histologically compared. All known varieties of tapeta were examined, classified and compared with the known cladogram representing the evolution of each specific family. Types of tapeta include tapetum cellosum, tapetum fibrosum, retinal tapetum, invertebrate pigmented tapetum, and invertebrate thin-film tapetum. All but the invertebrate pigmented tapetum were examined histologically. Review of the evolutionary cladogram and comparison with known tapeta suggest that the tapetum evolved in the Devonian period 345 to 395 million years ago. Tapeta developed independently in at least three separate orders in invertebrates and vertebrates, and all have surprisingly similar mechanisms of light reflection, including thin-film interference, diffusely reflecting tapeta, Mie scattering, Rayleigh scattering, and possibly orthogonal retroflection. Tapeta are found in invertebrates and vertebrates and display different physical mechanisms of reflection. Each tapetum reflects the wavelengths most relevant to the species’ ecological niche. The authors hypothesize that the tapetum evolved independently in both invertebrates and vertebrates as early as the Devonian period and coincided with an explosion of life forms.—Hans E. Grossniklaus

Guanine-type retinal tapetum in the eye of the Pacific pomfret Brama japonica (Perciformes: Bramidae)

The eye of Brama japonica, which exhibits eyeshine, contains a retinal tapetum composed of guanine. The total amounts of guanine in the eyes of two specimens measuring 19.5 and 51.4 cm in standard length were 7.4 and 70.5 mg, respectively, the respective retinal surface areas being 4.9 and 39.9 cm2. The mean guanine content was almost identical (1.5 and 1.8 mg/cm2, respectively). A locus tapetalis, with guanine values exceeding 2.5 mg/cm2, was developed in the ventrotemporal region of the retina, where it cooccurred with the area centralis. Evidence is presented that sound functional reasons exist for both the development of the locus tapetalis and its position in the retina. A plea is made for future examination of the retinae of additional pelagic and nonpelagic species for the presence and location of a locus tapetalis.

Ultrastructural analysis of hydraulic and abrasive retinal pigment epithelial cell debridements

Differential changes in Bruch's membrane, choriocapillaris, retinal pigment epithelium, retina, and tapetum after hydraulic or abrasive debridement of the retinal pigment epithelium in the cat area centralis were documented by fluorescein angiography, histology, and transmission electron microscopy at 1-hour, 1-day, 3-day, 1-week, or 4-week time points. Abrasive debridement is associated with abnormal fluorescein angiography and incomplete ingrowth of retinal pigment epithelial cells. Transmission electron microscopy shows that abrasive debridement inflicts more long-lasting ultrastructural damage to Bruch's membrane, the choriocapillaris, tapetum, and retina than does hydraulic debridement. Because the retinal pigment epithelium can resurface abrasively debrided Bruch's membrane that is disorganized, split, reduplicated, or missing, we cannot correlate the ultrastructural appearance of Bruch's membrane with the likelihood of complete resurfacing of the debrided area. Primary choriocapillary or retinal damage in abrasive debridements may contribute to the poor outcome. Regions of retinal degeneration with no underlying retinal pigment epithelial cell monolayer were significantly larger in abrasive debridements at the 4-week than at the 1-week time point. Reduced resurfacing at the later time point suggests that not all cells resurfacing abrasively debrided areas survived over the longer term. This finding may mean that retinal pigment epithelial cells are not able to resurface completely and permanently areas showing geographic atrophy of the choriocapillaris.

Guanine-type retinal tapetum and ganglion cell topography in the retina of a carangid fish, Kaiwarinus equula.

A guanine-type retinal tapetum was recorded in the eyes of a carangid fish Kaiwarinus equula (= Carangoides equula), spectrophotometric evidence of such being presented. The total amount of guanine in one eye was about 6.5 mg, the guanine density being ca. 1.3 mg cm(-2) over the retinal surface area. To examine the guanine distribution within the retina, the latter was divided into 21 regions. An area of high guanine density (more than 2.0 mg cm(-2)) was observed in the dorsal fundus of the retina, suggesting that the most sensitive vision was checked downward. Using whole-mount retinal preparations, the distribution of Nissl-stained cells within the retinal ganglion cell layer was examined. The greatest cell density area (area centralis) was observed only in the temporal retina. The visual acuity of the area centralis was 4.3 cycles deg(-1), suggesting that high resolution and binocular vision were directed frontally in this species. The eyes of a related carangid (Pseudocaranx dentex), lacking a tapetum, were also examined for comparison. The possible ecological advantage resulting from the tapetum is discussed in terms of visual threshold.

Does the taurine transporter gene play a role in 3p-syndrome?

Taurine has been shown to be essential for the development and survival of mammalian cells, especially cells of the cerebellum and retina18. Notably, taurine concentration reaches the mM range in the CNS20. The highest intracellular concentration of taurine is found in neonatal and early postnatal brain, suggesting a developmental role. Sturman's group found that the surviving F1 offspring of taurine-deficient female cats have a large number of neurologic defects, including degeneration of retinal pigmented epithelium and ocular tapetum, delayed cerebellar granule cell division and migration, and abnormal cerebral cortical development. Taurine also appears to optimize the proliferation and differentiation of human fetal cerebral brain cells in culture. Addition of taurine to the medium improves neuron growth, neurite expansion, and neuronal survival2. Taurine is considered a conditionally essential amino acid in man, as the enzymatic activity of cysteine sulfonic acid decarboxylase (CSAD), which catalyzes the formation of taurine from cysteine, is low in humans, and even lower or absent in term and preterm infant liver28. This ontogeny of catalytic enzyme activity is also true in other primates21. In children with short gut syndrome, taurine is an essential component of total parenteral nutrition (TPN) solutions to prevent a defined retinopathy, which is reversed only by taurine administration. Human milk also contains high concentrations of taurine11. We have shown previously that taurine is a semi-conditionally essential amino acid in very low birth weight premature infants, especially those who cannot ingest food and are obliged to be fed by TPN28. Plasma taurine values in preterm infants fall to almost non-detectable values,

Grouped, stacked rods and tapeta lucida in the retina of Japanese anchovy Engraulis japonicus

: Morphology of the photoreceptor cells and tapetum of the Japanese anchovy Engraulis japonicus was studied by histologically and by chemical analysis. The Japanese anchovy has a duplex retina. The cones form parallel rows consisting of alternately placed long cones and bifid cones. Both types of cones are intimately associated and form triple units that are regularly spaced along the row of cones. The rods are grouped and stacked. This fish has a retinal tapetum lucidum composed of guanine and hypoxanthine. Three structures of the tapetum lucidum were recognized: platelet, diamond and rod types. Photomechanical changes include movements of the photoreceptor cells and the retinal tapetum. The retina of the Japanese anchovy is thought to be highly sensitive and well adapted to a dim light environment.

Guanine and its retinal distribution in the tapetum of the bigeye tuna, Thunnus obesus

The eye of the bigeye tuna (Thunnus obesus) contains a retinal tapetum composed of guanine. The total amount of the guanine in one eye of the fish (SL=120 cm) was about 88.6 mg. The mean guanine content of the tapetum was approximately 1.25 mg/cm2 of the retinal surface. The highest content of guanine (2.15 mg/cm2) was observed only in the ventro-temporal part of the retina. To distinguish this area from the rest of the eye, we suggested the term ‘locus tapetalis’ for it. The visual accommodation system clearly indicated that the visual axis of the fish is upper-forward and the resulting retinal area for acute vision was suggested to be in the ventro-temporal retina. We discussed that the area centralis of the bigeye tuna may have two functions: to guarantee high visual acuity and to allow for high photo-sensitivity in dim light vision.

The Ocular Morphology of the Southern Hemisphere Lamprey Mordacia mordax Richardson with Special Reference to a Single Class of Photoreceptor and a Retinal Tapetum

A description is given of the morphological and ultrastructural characteristics of the eye of Mordacia mordax, one of the two parasitic species of the southern hemisphere lamprey family Mordaciidae. The avascular retina is well-differentiated, containing two ganglion cell sublaminae, two layers of horizontal cells and a large population of displaced bipolar cells. Intermediate filaments, which stain positively for glial fibrillary acidic protein (GFAP), lie in perpendicularly-arranged bundles between the two layers of horizontal cells. M. mordax contains only a single type of rod-like photoreceptor, in contrast to representatives in the other two lamprey families Geotriidae and Petromyzontidae, which possess three (two cone-like and one rod-like) and two (one cone and one rod) types of photoreceptor, respectively. The large photoreceptors of M. mordax possess a large mitochondrial ellipsosome, a cylindrical outer segment with incisures, and a nucleus immediately abutting the outer limiting membrane. The photoreceptors are arranged in an hexagonal array with their outer segments surrounded by a retinal tapetum which elicits a yellow fundal eyeshine. The tapetum contains both diffuse (vitread spheroidal granules) and specular (sclerad needles) reflectors. The characteristics of the eye of M. mordax are related to the diurnal burrowing of this species during its upstream migration, a feature not found in the other two extant families of lampreys. The absence of cone-like photoreceptors and the presence of a tapetum suggest that although photopic vision is probably greatly diminished, characteristics have been evolved by the eye to maximize light capture and increase sensitivity in the low light intensities encountered during this species’ nocturnal migration.

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

AbstractThe 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.