Retinal ultrastructure may mediate polarization sensitivity in larvae of the Sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae)

A number of invertebrates are known to be sensitive to the polarization of light and use this trait in orientation, communication, or prey detection. In these animals polarization sensitivity tends to originate in rhabdomeric photoreceptors that are more or less uniformly straight and parallel. Typically, polarization sensitivity is based on paired sets of photoreceptors with orthogonal orientation of their rhabdomeres. Sunburst diving beetle larvae are active swimmers and highly visual hunters which could potentially profit from polarization sensitivity. These larvae, like those of most Dytiscids, have a cluster of six lens eyes or stemmata (designated E1 through E6) on each side of the head capsule. We examined the ultrastructure of the photoreceptor cells of the principal eyes (E1 and E2) of first instar larvae to determine whether their rhabdomeric organization could support polarization sensitivity. A detailed electron microscopical study shows that the proximal retinas of E1 and E2 are in fact composed of photoreceptors with predominantly parallel microvilli and that neighboring rhabdomeres are oriented approximately perpendicularly to one another. A similar organization is observed in the medial retina of E1, but not in the distal retinas of E1&2. Our findings suggest that T. marmoratus larvae might be able to analyze polarized light. If so, this could be used by freshly hatched larvae to find water or within the water to break the camouflage of common prey items such as mosquito larvae. Physiological and behavioral tests are planned to determine whether larvae of T. marmoratus can actually detect and exploit polarization signals.     

How aquatic water-beetle larvae with small chambered eyes overcome challenges of hunting under water

A particularly unusual visual system exists in the visually guided aquatic predator, the Sunburst Diving Beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae). The question arises: how does this peculiar visual system function? A series of experiments suggests that their principal eyes (E1 and E2) are highly specialized for hunting. These eyes are tubular and have relatively long focal lengths leading to high image magnification. Their retinae are linear, and are divided into distinct green-sensitive distal and UV and polarization-sensitive proximal portions. Each distal retina, moreover, has many tiers of photoreceptors with rhabdomeres the long axis of which are peculiarly oriented perpendicular to the light path. Based on detailed optical investigations, the lenses of these eyes are bifocal and project focused images onto specific retinal tiers. Behavioral experiments suggest that these larvae approach prey within their eyes’ near-fields, and that they can correctly gauge prey distances even when conventional distance-vision mechanisms are unavailable. In the near-field of these eyes object distance determines which of the many retinal layers receive the best-focused images. This retinal organization could facilitate an unusual distance-vision mechanism. We here summarize past findings and discuss how these eyes allow Thermonectus larvae to be such successful predators.     

Eye and optic lobe metamorphosis in the sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae)

Nearly nothing is known about the transition that visual brain regions undergo during metamorphosis, except for Drosophila in which larval eyes and the underlying neural structure are strongly reduced. We have studied the larvae of the sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae), which are sophisticated visually oriented predators characterized by six elaborate stemmata on each side of the head and an associated large optic lobe. We used general neurohistological staining and 3D reconstruction to determine how the eyes and optic lobe of T. marmoratus change morphologically during metamorphosis. We find that in third (last) instar larvae, the adult neuropils are already forming de novo dorsally and slightly anteriorly to the larval neuropils, while the latter rapidly degenerate. Larval eyes are eventually reduced to distinct areas with dark pigmentation. This complete reorganization, which may be an evolutionarily conserved trait in holometabolous insects, occurs despite the considerable costs that must apply to such a visually complex animal. Our findings are consistent with the concept that stemmata are homologous to the most posterior ommatidia of hemimetabolous insects, an idea also recently supported by molecular data.     

Lunar influences in the diet habits and selectivity of larval clingfish Gobiesox marmoratus

The influence of the lunar cycle on the feeding habits, feeding success and selectivity was tested in situ on larval stages of the clingfish Gobiesox marmoratus (Gobiesocidae). Gobiesox marmoratus larvae and their prey were collected in the water column during the lunar cycle in austral spring 2015 and 2016, in shallow waters (<30 m depth) of El Quisco, central Chile. Feeding incidence was high (80–100%) throughout the moon cycle. The diet was composed of 32 prey items, being gastropod larvae and invertebrate eggs the most important. Prey composition varied among lunar phases with the lowest prey richness during new moon. During 2016, G. marmoratus larvae ingested the lowest number, but the largest prey during new moon. In spring 2015, there were no significant changes in the abundance of each prey taxon in the water column among moon phases, but in spring 2016 there were significant differences in the abundance of cypris and copepod nauplii, particularly between full and new moon. Feeding selectivity index showed that gastropod larvae were positively selected throughout the lunar cycle during spring 2015. In spring 2016, invertebrate eggs were selectively ingested in full moon and third quarter, but at a new moon G. marmoratus larvae selected gastropod larvae. Mean temperature of the water column and its vertical gradient and nocturnal cloud cover influences the feeding success of larval G. marmoratus. Lunar illumination favoured only an increase of richness of prey items. Therefore, nocturnal cloud cover precludes a greater influence of lunar illumination in the larval trophic ecology of this crypto-benthic fish.     

Behavioural and anatomical measures of visual acuity in first-feeding Yellowtail Kingfish (<Emphasis Type="Italic">Seriola lalandi</Emphasis>) larvae

Ontogenetic change in the visual acuity of Seriola lalandi larvae was measured using both behavioural and anatomical techniques. Visual acuity improved over early development (day 4 to day 7 post-hatch), although for all three larval ages examined estimates of anatomical acuity were consistently lower (higher acuity) than estimates of behavioural acuity. At hatching the eyes of larval kingfish were characterized by an undifferentiated retina surrounding a spherical lens, by day 4 post-hatch the eyes appeared to be functional, the retina was fully pigmented and the optic nerve had made contact with the optic tectum. Analysis of prey search behaviour indicated that larvae employ a saltatory type search behaviour in which brief stationary periods are interspersed with repositioning movements. The mean reactive angle increased between day 4 and day 7 post-hatch indicating that the horizontal visual field was expanding with development, thereby increasing the search area of larvae. Pre-strike distances of early larvae were substantially less than one body length, being constantly around a 1/3 of a body length for all larval ages examined.     

Feeding Habits of Two Large Insects from a Desert Stream: Abedus herberti (Hemiptera: Belostomatidae) and Thermonectus marmoratus (Coleoptera: Dytiscidae)

Feeding preference experiments were conducted to determine the feeding habits of Abedus herberti (Heteroptera, Belostomatidae) and Thermonectus marmoratus (Coleoptera, Dytiscidae), two large insects in Sycamore Creek, an intermittent Sonoran desert stream, Arizona, U.S.A. Numbers of live versus dead prey consumed were tested between and across three prey sizes. Five prey species were offered simultaneously (5 live and 5 dead specimens) in each size class. We found that A. herberti preferred live prey of small and medium size, but it chose mainly dead prey in the large size class. These results fitted the model of size-selective predation (Zaret, 1980). Size dependent predators selected prey of increased size, according to their visibility, but only up to where difficulty in handling and probability of escape affect successful consumption. Snails were the most preferred prey of A. herberti. By contrast, T. marmoratus consumed only dead prey of all sizes, but it preferred soft organisms with thin cuticle, such as immature larvae of some mayflies, beetles, dragonflies or fishes.     

Sharing morphospaces: early ontogenetic shape changes in two clingfish larvae (Pisces: Gobiesocidae) from the south‐east Pacific Ocean

Larval body shape changes and developmental timing were examined in two clingfish species from the south‐east Pacific Ocean, Gobiesox marmoratus and Sicyases sanguineus. Ontogenetic allometry showed no interspecific variation and <7 mm standard length (L_S) larvae of both species occupied similar morphospace, but larger G. marmoratus showed increased body depth while larvae of S. sanguineus developed a flattened head and maintained a hydrodynamic body. Estimated developmental timing suggests that larval body shape changes were faster in G. marmoratus than in S. sanguineus prior to settlement.     

Ingested prey increase risks of visual predation in transparent Chaoborus larvae

Summary Transparency reduces the chances of detection of large planktonic animals by visual predators. An important constraint on the transparency of planktonic animals may be ingested food which could be seen through the body, thereby increasing the vulnerability of transparent zooplankton to visual predators. To test this hypothesis, we presented fed and un-fed Chaoborus larvae to juvenile coho salmon (Oncorhynchus kisutch). Overall, the presence of prey in the gut of Chaoborus increased their probability of capture by 68%. Predation risks due to the visibility of ingested food increased in proportion to meal size: larvae with nearly full gut were captured about three times faster on the average than larvae which had little food in their gut. Although Chaoborus larvae may be able to reduce this increased predation risk by migrating downward to low light levels, this behavior would reduce feeding opportunities by removing the larvae from surface waters where prey density is generally high. In this way, visual predators may limit the growth and the maximum size that can be achieved by transparent animals.     

Parasitic copepods affect morphospace and diet of larvae of a temperate reef fish

The effects of ectoparasites on larvae of the clingfish Gobiesox marmoratus were evaluated at the dietary and morphometric levels. The larvae and ectoparasites were collected by nearshore plankton samplings during October, November and December 2013 off El Quisco Bay, central Chile. The standardized abundance of total larvae and those ectoparasitized larvae (PL) was positively related and high parasite prevalence was found throughout the sampling period (up to 38%). Geometric morphometrics analyses indicate main changes in the shape through early ontogeny and subtle but significant variations between PL and non‐parasitized larvae (NPL). Prey composition varied between PL and NPL; small size (<6 mm standard length, L_S) parasitized larval G. marmoratus ate mostly gastropod larvae, whereas small non‐parasitized specimens ate mainly cirripede nauplii. All larger (>8 mm L_S), pre‐settlement stages parasitized by Trifur and, or Caligus copepods had content in their gut, suggesting that ectoparasites did not diminish prey capture in host with larger size. Morphometric and dietary changes occurring during larval development were decoupled, both for PL and NPL. The maintenance of a slender, more hydrodynamic body through pelagic development and the ingestion of less‐mobile prey in PL suggests non‐lethal effects of ectoparasitism on rocky‐reef fish larvae.     

Pupil dilation and visual field in the piked dogfish, <Emphasis Type="Italic">Squalus acanthias</Emphasis>

The relatively large eye and pupil of the Piked Dogfish, Squalus acanthias, is visually arresting. However, knowledge of its basic visual characteristics lags far behind other areas in this generally well studied species. This study quantifies pupil dilation in a species that is naturally exposed to a broad range of light intensities and finds that the pupil area in the dark adapted state is 35.3% of the total eye area, an increase of 12.4% from the light adapted state. The anterior and posterior extents of the horizontal visual field are assessed and compared with both morphological and electrophysiological techniques and the results are integrated with the measured head yaw to derive the anterior convergence distance and blind area. The position of the eyes and the triangular, pointed snout of S. acanthias provides excellent anterior vision, which likely facilitates foraging upon its mobile prey.     

Visual fields, eye movements, and scanning behavior of a sit-and-wait predator, the black phoebe (Sayornis nigricans)

Foraging mode influences the dominant sensory modality used by a forager and likely the strategies of information gathering used in foraging and anti-predator contexts. We assessed three components of visual information gathering in a sit-and-wait avian predator, the black phoebe (Sayornis nigricans): configuration of the visual field, degree of eye movement, and scanning behavior through head-movement rates. We found that black phoebes have larger lateral visual fields than similarly sized ground-foraging passerines, as well as relatively narrower binocular and blind areas. Black phoebes moved their eyes, but eye movement amplitude was relatively smaller than in other passerines. Black phoebes may compensate for eye movement constraints with head movements. The rate of head movements increased before attacking prey in comparison to non-foraging contexts and before movements between perches. These findings suggest that black phoebes use their lateral visual fields, likely subtended by areas of high acuity in the retina, to track prey items in a three-dimensional space through active head movements. These head movements may increase depth perception, motion detection and tracking. Studying information gathering through head movement changes, rather than body posture changes (head-up, head-down) as generally presented in the literature, may allow us to better understand the mechanisms of information gathering from a comparative perspective.     

Diet changes in prey size selectivity in larvae of perch <Emphasis Type="Italic">Perca fluviatilis</Emphasis>

A hypothesis on change in prey size selectivity in relation to illumination level was tested on the basis of data on weight and size composition of the content of the digestive tract of larvae of perch Perca fluviatilis and on zooplankton in the layer 0–6 m (Wallersee Lake, Austria). Larvae foraging in the twilight-night period had almost two times more food in the intestine than those foraging in the daytime. The size composition of perch larvae and concentration and size composition of zooplankton hardly differed in the daytime and twilight-night samples. For the first time, it is shown on field material that more intensive feeding of larvae at twilight is related to selection of significantly larger prey than in daytime feeding. In the day-time, the larvae consumed more prey but their maximum size did not exceed 0.6 of the diameter of mouth opening of the fish; at twilight it was over 0.8. In case of feeding on so large prey, not only the weight of the consumed feed increases but the time used for capture and swallowing also considerably increases. The larvae which in the period of investigation did not yet form schools, which perform the principal defensive function, were especially vulnerable for predators feeding on relatively large prey. The decrease of the part of small-sized prey at twilight is not related to their lesser availability due to a low illumination. It is assumed that feeding on energetically more valuable but less available prey is shifted to the period of low illumination when the larvae are less exposed to predation risk. The obtained results are discussed from positions of the triotroph concept (Manteifel’, 1961).     

Development,consumption rates and reproductive biology of <Emphasis Type="Italic">Orius albidipennis</Emphasis> reared on various prey

The predatory bug Orius albidipennis (Reuter) (Hemiptera: Anthocoridae) has tremendous potential as a biological control agent, especially in its native range around the Mediterranean Basin and East Africa. The need to exploit native biological control agents is growing in importance as concerns over the introduction of non-native species continue to increase. However, little is known of the effects of different prey on development and reproduction of O. albidipennis compared with other species of Orius. Therefore, we compared the development, survival, reproductive biology, and prey consumption of O. albidipennis when fed eggs of Ephestia kuehniella Zeller, Tetranychus urticae Koch, and Trialeurodes vaporariorum (Westwood), and larvae of Gynaikothrips ficorum (Marchal), under laboratory of 26 ± 1°C, 60 ± 10% RH and 16L:8D photoperiod. Individuals were reared from the neonate stage until death on one of the four prey types. The type of prey had profound effects on all measured performance traits. The highest survival rate was recorded for nymphs that were fed on E. kuehniella eggs, while the lowest survival rate was observed for those fed on T. vaporariorum eggs. The shortest nymphal period was recorded for nymphs fed on E. kuehniella eggs, while the longest was measured for those fed on T. urticae eggs. During the nymphal period, O. albidipennis consumed significantly more eggs of T. urticae than other prey types, whereas the lowest number of consumed prey were eggs of E. kuehniella. Adult females and males consumed significantly more T. urticae eggs than other types of prey. However, Orius albidipennis females showed the highest fecundity when fed on E. kuehniella eggs, and the lowest when fed on T. vaporariorum eggs. Adult females and males that fed on G. ficorum larvae had significantly longer life spans compared with those fed other prey. Because of their relatively rapid development and high fecundity, O. albidipennis fed E. kuehniella eggs had a significantly higher net reproductive rate (R_o) and intrinsic rate of increase (r _m ) than O. albidipennis fed other prey types. Overall, eggs of E. kuehniella were the most suitable diet for nymphs and adults of O. albidipennis. Although less suitable, O. albidipennis could survive and reproduce on the other prey types, which is a favourable attribute in biological control agents. These results on the effect of different prey types on development and reproduction of O. albidipennis will also contribute to the development of mass rearing programs for biological control agents in developing countries, such as Egypt.     

Developmental morphology of the cottid fish Pseudoblennius marmoratus

Embryonic, larval, and juvenile development of a Japanese cottid fish Pseudoblennius marmoratus is described from laboratory-reared specimens. Eggs of P. marmoratus resembled those of two congeners, Pseudoblennius cottoides and Pseudoblennius percoides, but differed in their smaller diameter and yolk color. Although the larvae and juveniles of P. marmoratus were also similar to them in general morphology, the former differed in the lengths of newly hatched larvae, pigmentation patterns, number of pectoral fin rays, and dorsal fin spine elongation.     

Chromosome 1 in crested and marbled newts (Triturus)

The heteromorphic chromosomes 1 of Triturus cristatus carnifex and T. marmoratus were studied in mitotic metaphase after staining with the Giemsa C-banding technique and with the fluorochromes, DAPI (AT-specific) and mithramycin (GC-specific). They were also examined in the lampbrush form under phase-contrast before fixation and after fixation and staining with Giemsa. Chromosomes 1 of T.c. carnifex are asynaptic and achiasmatic throughout most of their long arms. They are also heteromorphic in most of their long arms for the patterns of Giemsa and fluorochrome staining and the distribution of distinctive lampbrush loops. The heteromorphic regions correspond to the regions that are asynaptic and achiasmatic. They stain more strongly with mithramycin and more weakly with DAPI than the remainder of the chromosomes, signifying that their DNA is relatively rich in GC. The patterns of staining with Giemsa and fluorochromes and the distributions of distinctive lateral loops vary from one animal to another in the same species and even in the same population. The asynaptic and achiasmatic regions of chromosomes 1 in T. marmoratus extend throughout the whole of the long arms and well beyond the heterochromatic region. Chiasmata form only in the short arm and occasionally in the short euchromatic segment at the tip of the long arms. The staining patterns of chromosomes 1 in T. marmoratus differ from those in T.c. carnifex although, like carnifex, their DNA is relatively GC-rich. The chromosomes 1 of T. marmoratus are more submetacentric than those of T.c. carnifex. In T. marmoratus chromosome 1B is about 12% shorter than 1A. There is a short paracentric inversion heterozygosity in the long arm of chromosome 1B in T. marmoratus which probably accounts for the lack of chiasmata in the euchromatin that separates the centromere from the start of the heterochromatin. In both carnifex and marmoratus, embryos that are homomorphic for chromosome 1 arrest and die at the late tailbud stage of development. The same applies to F₁ hybrid embryos T.c. carnifex x T. marmoratus, and this has permitted identification of chromosomes 1A and 1B in both species. There is no correspondence between patterns of Giemsa or fluorochrome staining of the heteromorphic regions of chromosome 1 and any feature of the lampbrush chromosomes. However, the short euchromatic ends of the long arms of chromosomes 1 in both species are distinguished in the lampbrush form by a series of uniformly small loops of fine texture associated with very small chromomeres. The Giemsa C-staining patterns of both chromosomes 1A and 1B are different in each of the four subspecies of T. cristatus. T.c. karelinii stands out by having unusually large masses of Giemsa C-staining centromeric heterochromatin on all but 1 of its 12 chromosomes. A scheme is proposed for the evolution of chromosome 1 in T. cristatus and T. marmoratus, based on all available cytological and molecular data.     

Prey location,biomechanical constraints,and motor program choice during prey capture in the tomato frog, <Emphasis Type="Italic">Dyscophus guineti</Emphasis>

This study investigated how visual information about prey location and biomechanical constraints of the feeding apparatus influence the feeding behavior of the tomato frog, Dyscophus guineti. When feeding on prey at small azimuths (less than ± 40°), frogs aimed their heads toward the prey but did not aim their tongues relative to their heads. Frogs projected their tongues rapidly by transferring momentum from the lower jaw to the tongue. Storage and recovery of elastic energy by the mouth opening muscles amplified the velocities of mouth opening and tongue projection. This behavior can only occur when the lower jaw and tongue are aligned (i.e., within the range of motion of the neck). When feeding on prey at large azimuths (greater than ± 40°), frogs aimed both the head and tongue toward the prey and used a muscular hydrostatic mechanism to project the tongue. Hydrostatic elongation allows for frogs to capture prey at greater azimuthal locations. Because the tongue moves independently of the lower jaw, frogs can no longer take advantage of momentum transfer to amplify the speed of tongue projection. To feed on prey at different azimuthal locations, tomato frogs switch between alternative strategies to circumvent these biomechanical constraints.     

Consumption rate and functional response of the predaceous mite<Emphasis Type="Italic"> Kampimodromus aberrans</Emphasis> to two-spotted spider mite <Emphasis Type="Italic">Tetranychus urticae</Emphasis> in the laboratory

Prey stage preference of female Kampimodromus aberrans (Oudemans) (Phytoseiidae) at constant densities of different stages of Tetranychus urticae Koch (Tetranychidae), functional response types and parameters of the predator females to the varying densities of eggs, larvae, protonymphs and deutonymps of T. urticae were determined in order to establish its potential for the mite biological control. Experiments were conducted at 25 ± 1°C, 65 ± 10% RH and 16:8 (L:D) photoperiod. Our results indicated that the predator consumed significantly more prey larvae than other prey stages. Functional response type of predator was determined by a logistic regression model. The predator exhibited a Type II response on all prey stages. The attack rate (α) and handling time (T _h ) coefficients of a Type II response were estimated by fitting a “random-predator” equation to the data. The lowest estimated value α and the highest value of T _h (including digestion) were obtanined for the predator feeding on deutonmph. The lowest value of T _h were obtained for the predator feeding on prey larvae, but the attack rate value obtained on larva wasn’t different than that obtained on egg and protonymph. According to our results, K. aberrans could be an efficient biological control agent of T. urticae at least at low prey densities. However, further field based studies are needed to draw firm conclusions.     

Eye movements and target fixation during dragonfly prey-interception flights

The capture of flying insects by foraging dragonflies is a highly accurate, visually guided behavior. Rather than simply aiming at the prey’s position, the dragonfly aims at a point in front of the prey, so that the prey is intercepted with a relatively straight flight trajectory. To better understand the neural mechanisms underlying this behavior, we used high-speed video to quantify the head and body orientation of dragonflies (female Erythemis simplicicollis flying in an outdoor flight cage) relative to an artificial prey object before and during pursuit. The results of our frame-by-frame analysis showed that during prey pursuit, the dragonfly adjusts its head orientation to maintain the image of the prey centered on the “crosshairs” formed by the visual midline and the dorsal fovea, a high acuity streak that crosses midline at right angles about 60° above the horizon. The visual response latencies to drifting of the prey image are remarkably short, ca. 25 ms for the head and 30 ms for the wing responses. Our results imply that the control of the prey-interception flight must include a neural pathway that takes head position into account.     

Use of the lateral line and tactile senses in feeding in four antarctic nototheniid fishes

Synopsis Fishes of the family Nototheniidae (Pisces: Perciformes) dominate antarctic fish communities and have radiated to fill diverse niches. The most southern species must operate under an extended austral night and under thick sea ice, yet have eyes more typical of shallow coastal fishes. In winter, the eyes are probably useless, except for detecting bioluminescence. I compared the responses of four species to hydromechanical and tactile signals: two benthivores,Trematomus bernacchii andT. pennellii, a benthic planktivore,T. nicolai, andPagothenia borchgrevinki, which feeds near the ice undersurface and within ice cracks. The planktivores have dorsal mouths, with eyes oriented dorsally or laterally (Pagothenia); their lateral line canals and receptor organs are larger dorsally. The benthivores have more ventrally oriented mouths and eyes. All species responded to hydromechanical cues to the head, but only the two benthivores responded to trunk hydromechanical stimuli or tactile stimuli to the ventral trunk or pelvic fins. Possibly responses to plankton along the trunk are of little use if a reorientation washes pelagic prey away. In responding to trunk stimuli,T. bernacchii reorients its head to the target in two stages by slowly pivoting on its pelvic fins. In contrast,T. pennellii reorients in a single quick flip. It is argued that, becauseT. bernacchii has wider canals thanT. pennellii, it must move more slowly to reduce self-generated noise. It is likely that further studies of winter diet and prey behavior may reveal the relative advantages of the two repositioning styles.     

Effects of continuous light and darkness on the eyes of the troglobitic salamander Typhlotriton spelaeus

Larval Typhlotriton spelaeus collected from five caves in Pulaski Co., Missouri, were kept as larvae or induced to transform in darkness or continuous fluorescent illumination. Larvae maintained in darkness for 215 and 279 days had smaller eyes, smaller rod inner and outer segments, and fewer metaphase figures in the genninative zone of the neural retina than comparable larvae maintained in light (258 lux). Except for visual cell size, differences were small and for each characteristic exceptions were observed. One larva kept in light showed early retinal degeneration comparable to that in transformed adults of T. spelaeus. All larvae exhibited optomotor behavior both before and after the experiment. Among animals induced to transform by L-thyroxin and maintained in darkness 111 to 366 days, visual cell and pigment epithelium degeneration was more extensive and more frequent than in animals kept for the same length of time in light (237-298 lux). In darkness the frequency of animals with retinal degeneration increased between 111 and 366 days. In light some animals exhibited pigment epithelium reduction with normal visual cells, and others had free, pigmented cells in the subretinal space. These effects were not comparable to degeneration in darkness. Eyelids covered the eyes of only a few animals in both light and dark treatments. The extent of eyelid encroachment over the eye was greater in darkness than in light. Most animals exhibited optomotor responses after experiments, but responses of animals kept in darkness were impaired in comparison to those of animals kept in light.