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Visually dependent dim-light foraging has evolved repeatedly, broadening the ecological niches of some species. Many dim-light foraging lineages evolved from diurnal ancestors, requiring immense visual sensitivity increases to compensate for light levels a billion times dimmer than daylight. Some taxa, such as bees, are anatomically constrained by apposition compound eyes, which function well in daylight but not in starlight. Even with this constraint, the bee genus Megalopta has incredibly sensitive eyes, foraging in light levels up to nine orders of magnitude dimmer than diurnal relatives. Despite many behavioural studies, variation in visual sensitivity and eye morphometry has not been investigated within and across Megalopta species. Here we quantify external eye morphology (corneal area and facet size) for sympatric species of Megalopta, M. genalis and M. amoena, which forage during twilight. We use electroretinograms to show that males, despite being smaller than females, have equivalent visual sensitivity and increased retinal responsivity. Although males have relatively larger eyes compared with females, corneal area and facet size were not correlated with retinal responsivity, suggesting that males have additional non-morphological adaptations to increase retinal responsiveness. These findings provide the foundation for future work into the neural and physiological mechanisms that interface with morphology to influence visual sensitivity, with implications for understanding niche exploitation.

Background Differences in morphology, ecology, and behavior through ontogeny can result in opposing selective pressures at different life stages. Most animals, however, transition through two or more distinct phenotypic phases, which is hypothesized to allow each life stage to adapt more freely to its ecological niche. How this applies to sensory systems, and in particular how sensory systems adapt across life stages at the molecular level, is not well understood. Here, we used whole-eye transcriptomes to investigate differences in gene expression between tadpole and juvenile southern leopard frogs (Lithobates sphenocephalus), which rely on vision in aquatic and terrestrial light environments, respectively. Because visual physiology changes with light levels, we also tested the effect of light and dark exposure. Results We found 42% of genes were differentially expressed in the eyes of tadpoles versus juveniles and 5% for light/dark exposure. Analyses targeting a curated subset of visual genes revealed significant differential expression of genes that control aspects of visual function and development, including spectral sensitivity and lens composition. Finally, microspectrophotometry of photoreceptors confirmed shifts in spectral sensitivity predicted by the expression results, consistent with adaptation to distinct light environments. Conclusions Overall, we identified extensive expression-level differences in the eyes of tadpoles and juveniles related to observed morphological and physiological changes through metamorphosis and corresponding adaptive shifts to improve vision in the distinct aquatic and terrestrial light environments these frogs inhabit during their life cycle. More broadly, these results suggest that decoupling of gene expression can mediate the opposing selection pressures experienced by organisms with complex life cycles that inhabit different environmental conditions throughout ontogeny.

Many animals have complex life cycles where larval and adult forms have distinct ecologies and habitats that impose different demands on their sensory systems. While the adaptive decoupling hypothesis predicts reduced genetic correlations between life stages, how sensory systems adapt across life stages at the molecular level is not well understood. Frogs are a compelling system to study this question in because most species rely on vision as both aquatic tadpoles and terrestrial adults, but these habitats present vastly different light environments. Here we used whole eye transcriptome sequencing to investigate differential expression between aquatic tadpoles and terrestrial juveniles of the southern leopard frog (Lithobates sphenocephalus). Because visual physiology changes with light levels, we also tested how constant light or dark exposure affected gene expression. We found 42% of genes were differentially expressed in the eyes of tadpoles versus juveniles, versus 5% for light/dark exposure. Analyses targeting a curated set of visual genes revealed significant differential expression between life stages in genes that control aspects of visual function and development, including spectral sensitivity and lens composition. Light/dark exposure had a significant effect on a smaller set of visual genes. Finally, microspectrophotometry of photoreceptors confirmed shifts in spectral sensitivity predicted by the expression results, consistent with adaptation to distinct light environments. Overall, we identified extensive expression-level differences in the eyes of tadpole and juvenile frogs related to observed morphological and physiological changes through metamorphosis, and corresponding adaptive shifts to optimize vision in aquatic versus terrestrial environments.

Since all forms of mimicry are based on perceptual deception, the sensory ecology of the intended receiver is of paramount importance to test the necessary precondition for mimicry to occur, that is, model‐mimic misidentification, and to gain insight in the origin and evolutionary trajectory of the signals. Here we test the potential for aggressive mimicry by a group of coral reef fishes, the color polymorphic Hypoplectrus hamlets, from the point of view of their most common prey, small epibenthic gobies and mysid shrimp. We build visual models based on the visual pigments and spatial resolution of the prey, the underwater light spectrum and color reflectances of putative models and their hamlet mimics. Our results are consistent with one mimic‐model relationship between the butter hamlet H. unicolor and its model the butterflyfish Chaetodon capistratus but do not support a second proposed mimic‐model pair between the black hamlet H. nigricans and the dusky damselfish Stegastes adustus. We discuss our results in the context of color morphs divergence in the Hypoplectrus species radiation and suggest that aggressive mimicry in H. unicolor might have originated in the context of protective (Batesian) mimicry by the hamlet from its fish predators rather than aggressive mimicry driven by its prey., Hamlets are colour polymorphic reef fishes believed to be diverging by aggressive mimicking of one of a number of unrelated non‐predatory fishes. We study the visual system of the intended receiver of hamlets' mimicry, their prey, and test whether model‐mimic mis‐identification by the prey is likely to occur. We find that only one hamlet is likely to be an efficient mimic in the eyes of their prey but we observe that the mimic colour morph might in fact have evolved by Batesian mimicry first and only subsequently recruited for aggressive mimicry.

Since all forms of mimicry are based on perceptual deception, the sensory ecology of the intended receiver is of paramount importance to test the necessary precondition for mimicry to occur, i.e. model-mimic misidentification, and to gain insight in the origin and evolutionary trajectory of the signals. Here we test the potential for aggressive mimicry by a group of coral reef fishes, the color polymorphicHypoplectrushamlets, from the point of view of their most common prey, small epibenthic gobies and mysid shrimp. We build visual models based on the visual pigments and spatial resolution of the prey, the underwater light spectrum and color reflectances of putative models and their hamlet mimics. Our results are consistent with one mimic-model relationship between the butter hamletH. unicolorand its model the butterflyfishChaetodon capistratusbut do not support a second proposed mimic-model pair between the black hamletH. nigricansand the dusky damselfishStegastes adustus. We discuss our results in the context of color morphs divergence in theHypoplectrusspecies radiation and suggest that aggressive mimicry inH. unicolormight have originated in the context of protective (Batesian) mimicry by the hamlet from its fish predators.

Although congenital stationary night blindness (CSNB) has been described in a Japanese beagle dog research colony, certain clinical correlates with human CSNB have not yet been described, nor has an estimate of frequency of the condition been made in inbred and outbred beagle populations. A beagle with CSNB obtained from a commercial research dog supplier in the USA and matched control dogs (n = 3) underwent examination, refraction, ocular imaging, assessment of visual navigation ability and detailed electroretinography (ERG). Retrospective review of ERGs in two independent groups of inbred (n = 15 and 537, respectively) and one group of outbred dogs (n = 36) was used to estimate CSNB frequency in these populations. In the affected dog, there were absent dark-adapted b-waves in response to dim-light flashes, severely reduced dark-adapted b-waves in response to bright-light flashes, and normal light-adapted b-waves with a-waves that had broadened troughs. Long-flash ERGs confirmed a markedly reduced b-wave with a preserved d-wave, consistent with cone ON-bipolar cell dysfunction. There was evidence of normal rod photoreceptor a-wave dark adaptation, and rapid light adaptation. In the wider beagle populations, five inbred beagles had a b/a wave ratio of

How animals visually perceive the environment is key to understanding important ecological behaviors, such as predation, foraging, and mating. This study focuses on the visual system properties and visual perception of color in the largemouth bass Micropterus salmoides. This study (1) documents the number and spectral sensitivity of photoreceptors, (2) uses these parameters to model visual perception, and (3) tests the model of color perception using a behavioral assay. Bass possess single cone cells maximally sensitive at 535 nm, twin cone cells maximally sensitive at 614 nm, and rod cells maximally sensitive at 528 nm. A simple model of visual perception predicted that bass should not be able to discern between chartreuse yellow and white nor between green and blue. In contrast, bass should be able to discern red from all achromatic (i.e., gray scale) stimuli. These predictions were partially upheld in behavioral trials. In behavioral trials, bass were first trained to recognize a target color to receive a food reward, and then tested on their ability to differentiate between their target color and a color similar in brightness. Bass trained to red and green could easily discern their training color from all other colors for target colors that were similar in brightness (white and black, respectively). This study shows that bass possess dichromatic vision and do use chromatic (i.e., color) cues in making visual-based decisions.

Vertebrate retinas are generally composed of rod (dim-light) and cone (bright-light) photoreceptors with distinct morphologies that evolved as adaptations to nocturnal/crepuscular and diurnal light environments. Over 70 years ago, the "transmutation" theory was proposed to explain some of the rare exceptions in which a photoreceptor type is missing, suggesting that photoreceptors could evolutionarily transition between cell types. Although studies have shown support for this theory in nocturnal geckos, the origins of all-cone retinas, such as those found in diurnal colubrid snakes, remain a mystery. Here we investigate the evolutionary fate of the rods in a diurnal garter snake and test two competing hypotheses: (i) that the rods, and their corresponding molecular machinery, were lost or (ii) that the rods were evolutionarily modified to resemble, and function, as cones. Using multiple approaches, we find evidence for a functional and unusually blue-shifted rhodopsin that is expressed in small single "cones." Moreover, these cones express rod transducin and have rod ultrastructural features, providing strong support for the hypothesis that they are not true cones, as previously thought, but rather are modified rods. Several intriguing features of garter snake rhodopsin are suggestive of a more cone-like function. We propose that these cone-like rods may have evolved to regain spectral sensitivity and chromatic discrimination as a result of ancestral losses of middle-wavelength cone opsins in early snake evolution. This study illustrates how sensory evolution can be shaped not only by environmental constraints but also by historical contingency in forming new cell types with convergent functionality.

Female mate preferences, male mate preferences and male–male competition can all potentially play roles in promoting speciation by sexual selection. The orangethroat darter, Etheostoma spectabile, is a sexually dichromatic fish in which males compete for access to females. We quantified male and female responses to conspecific versus heterospecific fish under normal light, dim light and blue light, the last of which impaired colour perception. We found that behavioural isolation was likely mediated primarily by male behavioural discrimination against heterospecific fish, with females showing no evident mate preferences. Furthermore, male aggression towards conspecific rivals was reduced in blue light, suggesting that the evolution of male coloration may contribute to speciation through male–male interactions.

The dominant hypothesis for the evolutionary origin of snakes from ‘lizards’ (non-snake squamates) is that stem snakes acquired many snake features while passing through a profound burrowing (fossorial) phase. To investigate this, we examined the visual pigments and their encoding opsin genes in a range of squamate reptiles, focusing on fossorial lizards and snakes. We sequenced opsin transcripts isolated from retinal cDNA and used microspectrophotometry to measure directly the spectral absorbance of the photoreceptor visual pigments in a subset of samples. In snakes, but not lizards, dedicated fossoriality (as in Scolecophidia and the alethinophidian Anilius scytale) corresponds with loss of all visual opsins other than RH1 (kmax 490– 497 nm); all other snakes (including less dedicated burrowers) also have functional sws1 and lws opsin genes. In contrast, the retinas of all lizards sampled, even highly fossorial amphisbaenians with reduced eyes, express functional lws, sws1, sws2 and rh1 genes, and most also express rh2 (i.e. they express all five of the visual opsin genes present in the ancestral vertebrate). Our evidence of visual pigment complements suggests that the visual system of stem snakes was partly reduced, with two (RH2 and SWS2) of the ancestral vertebrate visual pigments being eliminated, but that this did not extend to the extreme additional loss of SWS1 and LWS that subsequently occurred (probably independently) in highly fossorial extant scolecophidians and A. scytale. We therefore consider it unlikely that the ancestral snake was as fossorial as extant scolecophidians, whether or not the latter are para- or monophyletic.

Visual sensory demands vary substantially across vertebrates. Different visual sensory components have evolved to meet these sensory demands and enhance visual behavioral performance. One of these components is the retinal specialization, which is a portion of the retina with generally high ganglion cell densities, which increase spatial resolving power. Retinal specializations are relevant from a functional perspective because animals can align these “acute zones” with objects of interest within a localized region of their visual space, consequently affecting different behavioral dimensions. In this chapter, we reviewed the different types of retinal specializations found in vertebrates (retinal area , fovea, visual streak, radial anisotropy, area gigantocellularis) by discussing the different hypotheses proposed over decades to explain their function. Empirical tests on the functional properties of these different retinal specializations have been limited, which constrains our ability to understand the functional evolution of the vertebrate eye. We derive specific predictions from each of the hypotheses put forward to identify their degree of overlap. Finally, we provide some future directions as to how to test these functional hypotheses by integrating physiological and behavioral approaches. Testing these functional hypotheses will enhance our understanding of the relationship between the eye and the physical environment, and ultimately the visual ecology of vertebrates.

The potential for color vision in elasmobranchs has been studied in detail; however, a high degree of variation exists among the group. Evidence for ultraviolet (UV) vision is lacking, despite the presence of UV vision in every other vertebrate class. An integrative physiological approach was used to investigate color and ultraviolet vision in cownose rays and yellow stingrays, two batoids that inhabit different spectral environments. Both species had peaks in UV, short, medium, and long wavelength spectral regions in dark-, light-, and chromatic-adapted electroretinograms. Although no UV cones were found with microspectrophotometric analysis, both rays had multiple cone visual pigments with λ max at 470 and 551 nm in cownose rays (Rhinoptera bonasus) and 475, 533, and 562 nm in yellow stingrays (Urobatis jamaicensis). The same analysis demonstrated that both species had rod λ max at 500 and 499 nm, respectively. The lens and cornea of cownose rays maximally transmitted wavelengths greater than 350 nm and greater than 376 nm in yellow stingrays. These results support the potential for color vision in these species and future investigations should reveal the extent to which color discrimination is significant in a behavioral context.

Biofluorescence has recently been found to be widespread in marine fishes, including sharks. Catsharks, such as the Swell Shark (Cephaloscyllium ventriosum) from the eastern Pacific and the Chain Catshark (Scyliorhinus retifer) from the western Atlantic, are known to exhibit bright green fluorescence. We examined the spectral sensitivity and visual characteristics of these reclusive sharks, while also considering the fluorescent properties of their skin. Spectral absorbance of the photoreceptor cells in these sharks revealed the presence of a single visual pigment in each species. Cephaloscyllium ventriosum exhibited a maximum absorbance of 484 ± 3 nm and an absorbance range at half maximum (λ1/2max) of 440–540 nm, whereas for S. retifer maximum absorbance was 488 ± 3 nm with the same absorbance range. Using the photoreceptor properties derived here, a “shark eye” camera was designed and developed that yielded contrast information on areas where fluorescence is anatomically distributed on the shark, as seen from other sharks’ eyes of these two species. Phylogenetic investigations indicate that biofluorescence has evolved at least three times in cartilaginous fishes. The repeated evolution of biofluorescence in elasmobranchs, coupled with a visual adaptation to detect it; and evidence that biofluorescence creates greater luminosity contrast with the surrounding background, highlights the potential importance of biofluorescence in elasmobranch behavior and biology.

The Trinidadian pike cichlid (Crenicichla frenata) is a major predator of the guppy (Poecilia reticulata), a model system for visual ecology research, and visual predation by the pike cichlid is known to select for male guppies with reduced short-wavelength reflectance. However, an early study of the pike cichlid's visual system suggested a lack of short-wavelength-sensitive cone photoreceptors, a surprising finding as many African cichlids have highly developed short-wavelength vision. In this study, we found evidence for only four expressed cone opsins (LWS, RH2a, SWS2a, and SWS2b), plus one pseudogene (RH2b). Taken together with our microspectrophotometry data, which revealed the presence of three types of cone photoreceptor, including one sensitive to short-wavelength light, this would indicate a broader spectral capacity than previously believed from earlier visual studies of this fish. Relative to the highly diverse African cichlids, however, this Neotropical cichlid appears to have a greatly reduced opsin complement, reflecting both gene loss along the Neotropical lineage (lacking functional RH2b and, possibly, SWS1 opsins) and gene duplication within the African clade (which possesses paralogous RH2aα and RH2aβ opsins). Molecular evolutionary analyses show that positive selection has shaped the SWS2b and RH1 opsins along the Neotropical lineage, which may be indicative of adaptive evolution to alter nonspectral aspects of opsin biology. These results represent the first molecular evolutionary study of visual pigments in a Neotropical cichlid and thus provide a foundation for further study of a morphologically and ecologically diverse clade that has been understudied with respect to the link between visual ecology and diversification.

The nature and distributions of photoreceptor cell types were investigated in the retinas of 12 species (5 families) of elopomorph anguilliform leptocephalus larvae. Anti-opsin immunofluorescence, light microscopy and transmission electron microscopy (TEM) were used to assess opsin distribution across the retinas and to associate photoreceptor morphology and opsin content. Retinas of all species were immunoreactive with anti-rhodopsin throughout, while anti-cone opsin immunoreactivity was restricted only to the ventral region of the retina in all specimens. Rod and cone photoreceptors were morphologically indistinguishable at low magnifications; TEM revealed that nearly all photoreceptors had rod-like ultrastructure, with only rare examples of cone-like cells identified in the ventral retina. These results indicate a rhodopsin/rod-dominated retina in leptocephalus larvae of anguilliform eels in the teleost subdivision Elopomorpha, contrasting with the cone-dominated retinas of nearly all other species of teleost larvae. This distinctive developmental pattern shared among elopomorph larvae has important evolutionary and ecological implications, indicating a shared ancestor and/or ecological characteristics that are very different from most other teleost larvae.

Progress in developing animal communication theory is frequently constrained by a poor understanding of sensory systems. For example, while lizards have been the focus of numerous studies in visual signalling, we only have data on the spectral sensitivities of a few species clustered in two major clades (Iguania and Gekkota). Using electroretinography and microspectrophotometry, we studied the visual system of the cordylid lizardPlatysaurus broadleyibecause it represents an unstudied clade (Scinciformata) with respect to visual systems and because UV signals feature prominently in its social behaviour. The retina possessed four classes of single and one class of double cones. Sensitivity in the ultraviolet region (UV) was approximately three times higher than previously reported for other lizards. We found more colourless oil droplets (associated with UV-sensitive (UVS) and short wavelength-sensitive (SWS) photoreceptors), suggesting that the increased sensitivity was owing to the presence of more UVS photoreceptors. Using the Vorobyev–Osorio colour discrimination model, we demonstrated that an increase in the number of UVS photoreceptors significantly enhances a lizard's ability to discriminate conspecific male throat colours. Visual systems in diurnal lizards appear to be broadly conserved, but data from additional clades are needed to confirm this.

An understanding of the effect of light on predator–prey interactions in aquatic systems requires the integration of sensory physiology, behavioral ecology, and spatial distributions of predator and prey in the field. Here, we present such an integrative approach to a study on the interactions between the alewife, Alosa pseudoharengus, and the mysid shrimp, Mysis diluviana, (formerlyM. relicta) in Lake Ontario at night, when it is unknown whether visual feeding is possible. Visual pigment analyses of alewife rod photoreceptors were used to derive an alewife-specific unit of brightness—the ‘alelux’ (wavelength of maximum absorbance, lmax 5 505 nm)—which formed the basic unit of light intensity in alewife feeding-rate experiments and field applications. At light levels of 1027 alelux (, 1024.1 lux) and greater in the laboratory, alewives engaged in visual search and strike behaviors and fed at rates that were significantly higher than those under completely dark conditions. Field observations from Lake Ontario showed that light levels at the upper edge of the mysid distribution were within the range of those required for visual feeding in the laboratory on a full moon night, but not on a new moon night. These increased light levels translated into feeding rates that were . 30 times higher on the full moon night, despite a larger degree of spatial separation of the two trophic levels. We hypothesize that observed increased water clarity in Lake Ontario in recent years has led to increased consumption of mysids by alewife at night and associated food-web changes. Of the many factors that affect fish feeding success, light may be the most dynamic because light intensity changes over many orders of magnitude throughout the day–night

Selection should favour coloration in organisms that is more conspicuous to their own visual system than to those of their predators or prey. We tested this prediction in Dickerson’s collared lizard (Crotaphytus dickersonae), a sexually dichromatic desert reptile that preys on insects and smaller lizard species, and which in turn is prey for birds and snakes. We modelled the spectral sensitivities of the lizards and their avian and snake predators, and compared the conspicuousness of the lizards’ entire colour patterns with each class of viewers. Almost all comparisons involving females strongly supported our prediction for greater chromatic and brightness conspicuousness against local terrestrial visual backgrounds to their own modelled visual system than to those of avian and snake predators. Males, in contrast, exhibited far fewer cases of greater conspicuousness to their own visual system than to those of their predators. Our own perception of spectral similarity between blue C. dickersonae males and a local nonterrestrial visual background (i.e. the Sea of Cortez) prompted a further investigation.We compared sea (and sky) radiance with dorsum radiance of C. dickersonae males and with males from two distantly-related Crotaphytus collaris populations in which males possess blue bodies. In all three visual models, C. dickersonae males exhibited significantly lower chromatic contrast with the sea (and sky) than did their noncoastal, blue-bodied congeners. Among potential explanations, the blue body coloration that is unique to male C. dickersonae may offset, if only slightly, the cost of visibility to predators (and to prey) through reduced contrast against the extensive, local, nonterrestrial blue backgrounds of the sea and sky

Since its introduction to Harp Lake, Ontario, Canada, summer abundance of the cladoceran zooplanktivore Bythotrephes longimanus has fluctuated substantially both among and within years. The principal planktivorous fish in Harp Lake is the cold-water Coregonus artedi (cisco). Previous studies hypothesized that Bythotrephes abundance was affected by the thickness of an ephemeral, dark, daytime refuge from cisco that potentially established at the bottom of the metalimnion. During summer of 2003, we estimated peak daytime refuge thickness by simulating light energy visible to cisco and found it was always negative and did not correlate with Bythotrephes death rates. Direct observations using gill-netting and acoustical methods suggested that cisco had frequent metalimnetic forays. Additionally, including years 2000–2004, the previous correlation between mean Bythotrephes abundance and refuge thickness no longer held. The refuge hypothesis appears to fail, as the amount of metalimnetic illumination was always above the level at which cisco reaction distance to prey is maximal. Selection of Bythotrephes by cisco instead appeared to increase once Bythotrephes became abundant, remaining consistent and nontrivial even after Bythotrephes population declined.

Light and temperature strongly influence the vertical distribution of the mysid shrimp, Mysis relicta . We monitored the vertical movements and depth selection behavior of mysids exposed to different light intensities and light–temperature gradients in the laboratory and derived a mysid light preference function in units relevant to mysid vision: “mylux”. Mysids preferred light levels between 10−8 and 10−7 mylux (∼10−6 to 10−5 lux) and rarely moved into waters of 10−3 mylux (∼0.1 lux) and greater. A model that assumed equal weight and independence of mysid light and temperature preference functions successfully predicted the proportion of mysids found in two different temperature–light combinations in the laboratory. This model also predicted the depth of maximum mysid density to within 2 m on two spring nights and within 5 m on two summer nights of varying moon phase and thermal conditions in Lake Ontario. This study provides novel insights into how temperature and light interact to influence the vertical distribution of mysids. Our model may be used to predict mysid vertical distribution in any deepwater system inhabited by mysids in which the primary mysid predators are visual feeders.

Lake sturgeon ( Acipenser fulvescens Rafinesque, 1817) photoreceptors were studied with scanning electron microscopy and microspectrophotometry. The retina contains both rods and cones, with cones estimated composing about 30% of the photoreceptor population. Only large single cones were identified and they are similar to those found in other species of the order Acipenseriformes. The rods are large, with long, broad outer segments, and are similar to the dominant rod found in other sturgeons and the North American paddlefish ( Polyodon spathula (Walbaum, 1792)). Mean (SD) rod packing density at 22 624 ± 3 509 rods/mm2 is low compared with those of other animals that function primarily in dim light. The visual pigment of the rods has a mean (SD) peak absorbance (λmax) at 541 ± 2 nm. Three different cone populations were identified: a long wavelength sensitive cone containing a visual pigment with λmax at 619 ± 3 nm; middle wavelength sensitive cone with λmax at 538 ± 1 nm; and short wavelength sensitive cone with λmax at 448 ± 1 nm. All the visual pigments are based on the vitamin A2 chromophore.

In 1934, Gordon Walls forwarded his radical theory of retinal photoreceptor ‘transmutation’. This proposed that rods and cones used for scotopic and photopic vision, respectively, were not fixed but could evolve into each other via a series of morphologically distinguishable intermediates. Walls' prime evidence came from series of diurnal and nocturnal geckos and snakes that appeared to have pure-cone or pure-rod retinas (in forms that Walls believed evolved from ancestors with the reverse complement) or which possessed intermediate photoreceptor cells. Walls was limited in testing his theory because the precise identity of visual pigments present in photoreceptors was then unknown. Subsequent molecular research has hitherto neglected this topic but presents new opportunities. We identify three visual opsin genes,rh1,sws1andlws, in retinal mRNA of an ecologically and taxonomically diverse sample of snakes central to Walls' theory. We conclude that photoreceptors with superficially rod- or cone-like morphology are not limited to containing scotopic or photopic opsins, respectively. Walls' theory is essentially correct, and more research is needed to identify the patterns, processes and functional implications of transmutation. Future research will help to clarify the fundamental properties and physiology of photoreceptors adapted to function in different light levels.

Unlike the mammalian retina, the teleost fish retina undergoes persistent neurogenesis from intrinsic stem cells. In marine teleosts, most cone photoreceptor genesis occurs early in the embryonic and larval stages, and rods are added primarily during and after metamorphosis. This study demonstrates a developmental paradigm in elopomorph fishes in which retinas are rod-dominated in larvae, but undergo periods of later cone genesis. Retinal characteristics were compared at different developmental stages among three ecologically distinct elopomorph fishes—ladyfish (Elops saurus), bonefish (Albula vulpes), and speckled worm eel (Myrophis punctatus). The objectives were to improve our understanding of (1) the developmental strategy in the elopomorph retina, (2) the functional architecture of the retina as it relates to ecology, and (3) how the light environment influences photoreceptor genesis. Photoreceptor morphologies, distributions, and spectral absorption were studied at larval, juvenile, and adult stages. Premetamorphic retinas in all three species are rod-dominated, but the retinas of these species undergo dramatic change over the course of development, resulting in juvenile and adult retinal characteristics that correlate closely with ecology. AdultE. saurushas high rod densities, grouped photoreceptors, a reflective tapetum, and longer-wavelength photopigments, supporting vision in turbid, low-light conditions. AdultA. vulpeshas high cone densities, low rod densities, and shorter-wavelength photopigments, supporting diurnal vision in shallow, clear water.M. punctatusloses cones during metamorphosis, develops new cones after settlement, and maintains high rod but low cone densities, supporting primarily nocturnal vision.M. punctatussecondary cone genesis occurs rapidly throughout the retina, suggesting a novel mechanism of vertebrate photoreceptor genesis. Finally, in postsettlementM. punctatus, the continuous presence or absence of visible light modulates rod distribution but does not affect secondary cone genesis, suggesting some degree of developmental plasticity influenced by the light environment.

The visual pigments of cones and rods in three species of mollies, Poecilia mexicana, Poecilia latipinna and their asexual hybrid Poecilia formosa, were examined using microspectrophotometry. In P. mexicana, populations from extreme photic habitats were used: one population originated from a clear water habitat, one from a milky water habitat and another from a completely dark cave. Ultraviolet-sensitive cones were found in all species. Differences in the lmax values of the visual pigments were small between species and among the three P. mexicana populations, but dark-reared cave fishes showed appreciably higher variance. The hybrid species P. formosa showed a highly variable long wavelength cone absorbance, ranging from 528! 9 to 598! 5 nm, suggesting multiple opsin expression or chromophore mixing. # 2006 The Authors Journal compilation # 2006 The Fisheries Society of the British Isles

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).

The body colouration extending into ultraviolet wavelengths in three species of Lake Malaŵi rock-dwelling cichlids, Metriaclima zebra, Metriaclima benetos and Metriaclima barlowi was quantified. All three species were ultraviolet reflective with interspecific differences in reflectivity. In addition, individuals were able to behaviourally mediate their reflectivity.

SUMMARYPoison frogs in the anuran family Dendrobatidae use bright colors on their bodies to advertise toxicity. The species Dendrobates pumilio Schmidt 1858, the strawberry poison frog, shows extreme polymorphism in color and pattern in Panama. It is known that females of D. pumiliopreferentially choose mates of their own color morph. Nevertheless, potential predators must clearly see and recognize all color morphs if the aposematic signaling system is to function effectively. We examined the ability of conspecifics and a model predator to discriminate a diverse selection of D. pumilio colors from each other and from background colors. Microspectrophotometry of isolated rod and cone photoreceptors of D. pumilio revealed the presence of a trichromatic photopic visual system. A typical tetrachromatic bird system was used for the model predator. Reflectance spectra of frog and background colors were obtained, and discrimination among spectra in natural illuminants was mathematically modeled. The results revealed that both D. pumilio and the model predator discriminate most colors quite well, both from each other and from typical backgrounds, with the predator generally performing somewhat better than the conspecifics. Each color morph displayed at least one color signal that is highly visible against backgrounds to both visual systems. Our results indicate that the colors displayed by the various color morphs of D. pumilio are effective signals both to conspecifics and to a model predator.

Intraspecific visual communication was studied quantitatively by testing the behavior of Sepia officinalis, the common cuttlefish, as senders and receivers of body pattern signals. These signals can be achromatic and/or linearly polarized and are produced by specialized dermal cells. Experiment 1 addressed whether the presence of conspecifics affected the visible, achromatic body patterns of males. These patterns tended to vary among conditions, suggesting limited sensitivity to audience. One set of body patterns varied with the number of conspecifics viewed while an uncorrelated set of body patterns varied with the sex of conspecifics viewed. Males showed high-contrast zebra banding when viewing another male, supporting previous studies of body patterns in male-male agonistic behavior. Experiment 2 addressed the relationship of polarized light patterns with visible body patterns of males and females, and tested whether senders modified their polarization patterns in response to conspecifics. Polarization patterning was only weakly associated with visible body patterns. Females showed more polarized body patterns than did males, but polarized patterns did not differ among conditions; thus, no sensitivity to audience by senders was found. Experiment 3 addressed whether conspecific receivers used information from polarized body patterns. Limited evidence was found for changes in the behavior of female but not male observers, suggesting that female receivers may use polarized patterns as a source of information about conspecifics. The information contained in polarization patterning may complement that contained in zebra patterning such that both males and females advertise their species, sex, location, and size to conspecific receivers.

Studies of visual ecology have typically focused on differences among species while paying less attention to variation among populations and/or individuals. Here, we show that the relative abundance of UV, violet, yellow, and red cones varies between two populations of bluefin killifish, Lucania goodei. Animals from a spring population (high-transmission UV/blue light) have a higher frequency of UV and violet cones and a lower frequency of yellow and red cones than animals from a swamp population (low-transmission UV/blue light). Visual sensitivity does not vary significantly between the populations, but spring animals tend to be more sensitive in the UV/blue wavelengths (360-440 nm) and less sensitive in longer wavelengths (560-600 nm) than swamp animals. The results have two important implications. First, the tight conservation of functional regions of opsin genes across taxa does not imply that visual systems are constrained in their evolution; differential sensitivity can arise through differential expression of cone classes within the retina. Second, intraspecific visual signals in this species may evolve to maximize contrast between the signaler and the background (as opposed to brightness); males with blue anal fins are most abundant in swamp habitats where animals express fewer UV and violet cones.

SUMMARY We report microspectrophotometric (MSP) data for the visual pigments and oil droplets of 17 species of Caribbean anoline lizard known to live in differing photic habitats and having distinctly different dewlap colors. The outgroup Polychrus marmoratus was also examined to gain insight into the ancestral condition. Except for Anolis carolinensis, which is known to use vitamin A2 as its visual pigment chromophore, all anoline species examined possessed at least four vitamin-A1-based visual pigments with maximum absorbance (λmax) at 564, 495,455 and 365 nm. To the previously reported visual pigments for A. carolinensis we add an ultraviolet-sensitive one withλ max at 365 nm. Five common classes of oil droplet were measured, named according to apparent color and associated with specific cone classes — yellow and green in long-wavelength-sensitive (LWS) cones,green only in medium-wavelength-sensitive (MWS) cones and colorless in short-wavelength-sensitive (SWS) and ultraviolet-sensitive (UVS) cones. MSP data showed that the colorless droplet in the SWS cone had significant absorption between 350 and 400 nm, while the colorless droplet in the UVS cone did not. The pattern for Polychrus marmoratus was identical to that for the anoles except for the presence of a previously undescribed visual cell with a rod-like outer segment, a visual pigment with a λmaxof 497 nm and a colorless oil droplet like that in the UVS cones. These findings suggest that anoline visual pigments, as far as they determine visual system spectral sensitivity, are not necessarily adapted to the photic environment or to the color of significant visual targets (e.g. dewlaps).

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 co...

Males and females from Lacertid lizard species often display conspicuous colourations involved in intraspecific communication. However, visual systems of Lacertidae have rarely been studied and the spectral sensitivity of their retinal photoreceptors remains unknown. Here, we characterised spectral sensitivity of two Lacertid species from contrasted habitats, the wall lizard Podarcis muralis and the common lizard Zootoca vivipara. Both species possess a pure-cone retina with one spectral class of double cones and four spectral classes of single cone photoreceptors. The two species differ in the spectral sensitivity of the LWS cones, the relative abundance of UVS single cones (potentially more abundant in Z. vivipara), and the colouration of oil droplets. Wall lizards have pure vitamin A1-based photopigments while common lizards possess mixed vitamin A1- and A2- photopigments extending spectral sensitivity into near infrared, a rare feature in terrestrial vertebrates. We found that spectral sensitivity in the UV and in the near infrared improves discrimination of small variation in throat colouration among Z. vivipara. Thus, retinal specialisations optimise chromatic resolution in common lizards, which indicates that visual system and visual signals may co-evolve.

SUMMARY Microspectrophotometry was used to measure the visual pigments in the rods and cones of 22 species of marine fish larvae netted from the surface waters off Friday Harbor Laboratories, Washington, USA. 13 species had rods, 12 of which contained visual pigments with a wavelength of maximum absorbance near 500nm, while one, the sand lance (Ammodytes hexapterus), had its absorbance maximum at 482nm. The 22 species of fish larvae possessed varied combinations of single, double and twin cones, ranging in peak absorbance from 353nm to 584nm. Of these, green-sensitive single cones were present in 20 of the 22 species, and were the dominant cone type. Double and twin cones were present in 13 of the species. Most common were identical green-sensitive (twin) cones (in 11 species). Green/yellow-sensitive double cones occurred in four species. In a single instance (Hemilepidotus hemilepidotus) twin blue-sensitive, twin green-sensitive and double blue/yellow-sensitive cones were recorded. Of particular interest was the finding that 18 of the species had ultraviolet- and/or violet-absorbing single cones. It has been suggested that short-wavelength photosensitivity may be beneficial for planktivory by extending the spectral range over which vision can occur. The high percentage (82%) of ultraviolet and violet visual pigments in Pacific northwest fish larvae supports the prediction that short-wavelength sensitivity may be common in marine fish larvae.

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.

Squids have a wide repertoire of body patterns; these patterns contain visual signals assembled from a highly diverse inventory of chromatic, postural, and locomotor components. The chromatic components reflect the activity of dermal chromatophore organs that, like the postural and locomotor muscles, are controlled directly from the central nervous system. Because a thorough knowledge of body patterns is fundamental to an understanding of squid behavior, we have compiled and described an ethogram (a catalog of body patterns and associated behaviors) for Loligo pealei. Observations of this species were made over a period of three years (or = 440 h) and under a variety of behavioral circumstances. The natural behavior of the squid was filmed on spawning grounds off Cape Cod (northwestern Atlantic), and behavioral trials in the laboratory were run in large tanks. The body pattern components--34 chromatic (including 4 polarization components), 5 postural, and 12 locomotor--are each described in detail. Eleven of the most common body patterns are also described. Four of them are chronic, or long-lasting, patterns for crypsis; an example is Banded Bottom Sitting, which produces disruptive coloration against the substrate. The remaining seven patterns are acute; they are mostly used in intraspecific communication among spawning squids. Two of these acute patterns--Lateral Display and Mate Guarding Pattern--are used during agonistic bouts and mate guarding; they are visually bright and conspicuous, which may subject the squids to predation; but we hypothesize that schooling and diurnal activity may offset the disadvantage presented by increased visibility to predators. The rapid changeability and the diversity of body patterns used for crypsis and communication are discussed in the context of the behavioral ecology of this species.

Scanning electron microscopy, microspectrophotometry, and spectrophotometry of digitonin extracts were employed to characterize the photoreceptors and visual pigments of two freshwater Acipenseriformes. The retinas of the shovelnose sturgeon, Scaphirhynchus platorynchus (Acipenseridae), and the paddlefish, Polyodon spathula (Polyodontidae) are dominated by large rods with long, broad outer segments. A second rod, rare and much narrower than the dominant rod, is present in Scaphirhynchus but not seen in Polyodon. The absorbance maximum of the visual pigment in the rods of Polyodon is near 540 nm; that of Scaphirhynchus near 534 nm. The retinas of both species contain substantial numbers of large, single cones, about 33% of the photoreceptors in Scaphirhynchus; 37% in Polyodon. Scaphirhynchus cone pigments have absorbance maxima near 610 nm, 521 nm and 470 nm, respectively. Polyodon cone pigments absorb maximally near 607 nm and 535 nm, respectively. All visual pigments are based on vitamin A2. The data are compared to those from other Acipenseriformes and are discussed in terms of lifestyle and behavior.

Newly hatched white sturgeon (Acipenser transmontanus) yolk-sac larvae are negatively phototactic, photokinetic and geotactic—seeking cover in the substrate during daylight hours. If cover is unavailable and the light intensity is low, they move into the water column and alternate periods of rapid, upward swimming with periods of quiescence during which they sink. In the dark they swim almost continuously. A flash of light inhibits swimming activity totally for a period dependent on the light intensity. Using this observation, we obtained an action spectrum for swimming inhibition for 30 newly hatched larvae. This action spectrum is best fit by a single visual pigment with absorbance maximum at 537 nm. The lack of red sensitivity fits well with previous work showing that red-sensitive photoreceptor cells are absent in young sturgeon. The possible significance of red insensitivity is discussed from both ecological and practical standpoints.

The visual ecology of six closely related species of Puerto Rican anoline lizards was investigated and they were found to occupy four distinct habitat types in terms of light conditions: “full shade”, “partial shade”, “no shade”, and “forest canopy.”The habitats differed substantially in total radiance and irradiance as well as in the shape of the irradiance spectrum. The shape of the radiance spectrum was similar in all of the habitats. We used electroretinogram (ERG) flicker photometry to measure spectral sensitivity and found the curves for all six species to be similar. The spectral sensitivity peaked in the range 550–560 nm, which matched the peak in spectral radiance for all of the habitats. The shape of the spectral-sensitivity curve was similar to those of a number of other terrestrial vertebrates. We suggest that the convergence of the shape of the photopic ERG-determined spectral-sensitivity curve in many terrestrial vertebrates may, in part, be due to the fact that the background radiance of many terrestrial habitats is dominated by the reflectance spectrum of green vegetation which peaks at 550 nm.

The retinal photoreceptors of brown smoothhound sharks (Mustelus henlei) and leopard sharks (Triakis semifasciata) were studied by spectrophotometric analysis of visual pigment extracts, microspectrophotometric examination of single cells, and scanning electron microscopy. Juvenile and adult brown smoothhounds have one, identical visual pigment in their rod photoreceptors. The visual pigment is based on the vitamin A1 chromophore and has a peak absorbance (λmax) (at 496 ± 2 nm. Juvenile and adult leopard sharks also have a single, vitamin A1 based visual pigment, this one with λmax at 502 ± 1 nm. No evidence was found for a shift from a vitamin A2 based visual pigment in the juveniles to a vitamin A1 based pigment in the adults, as is known to occur in the lemon shark, Negaprion brevirostris (Cohen et al. [1990] Vision Res., 30:1949–1953). The retinas of brown smoothhounds and leopard sharks contain both rods and cones, with the rods dominating. The rod outer segments of the brown smoothhound are much longer than are those of the leopard shark. The greater length of brown smoothhound rod outer segments and the shorter wavelength λmax of the brown smoothhound's rod pigment support the idea that brown smoothhounds are either more nocturnal than leopard sharks or are active at greater depths. Cones are very rare in both species with only three identified with the scanning electron microscope and none with the microspectrophotometer. The apparent paucity of cones most likely relates to the bottom dwelling lifestyle of these sharks, although the possibility remains that their retinas contain small, specialized regions of high cone density, which were simply missed. © 1996 Wiley-Liss, Inc.

Retinas of the nocturnal geckos, Hemidactylus turcicus, Hemidactylus garnotii, and Teratoscincus scincus, were studied with microspectrophotometry and immunocytochemistry against various visual pigment epitopes to reveal UV-sensitive photoreceptors. From 6–20% of the thinner members of type C double photoreceptors, earlier believed to be blue-sensitive, were found to contain a UV-absorbing visual pigment with λmax at 363–366 nm. The pigment had bleaching and dichroic properties typical of other photoreceptor cell types of the retina. Presumptive UV-sensitive cells in retinal sections were “negatively” labeled as they did not react with either the cone-specific monoclonal antibody COS-1 or with the anti-rhodopsin polyclonal serum AO, which together labeled all of the remaining photoreceptor types (green-sensitive A singles, B doubles, and thicker members of C doubles, as well as the blue-sensitive majority of thinner members of C doubles). UV cells were moderately stained with the mAb K42–41 produced against the 5–6 loop of bovine rhodopsin, which also moderately labeled blue-sensitive cells. mAb OS-2 strongly stained all outer segments, including the UV-sensitive ones. Similarities between gecko UV visual pigments, and UV visual pigments of other vertebrates, as well as possible functional significance of these cells are discussed.

SUMMARYColor vision is not uniform across the retina because of differences in photoreceptor density and distribution. Retinal areas with a high density of cone photoreceptors may overlap with those with a high density of ganglion cells, increasing hue discrimination. However, there are some exceptions to this cell distribution pattern, particularly in species with horizontal visual streaks (bands of high ganglion cell density across the retina) that live in open habitats. We studied the spectral sensitivity and distribution of cone photoreceptors involved in chromatic and achromatic vision in the Canada goose (Branta canadiensis), which possesses an oblique rather than horizontal visual streak at the ganglion cell layer. Using microspectrophotometry, we found that the Canada goose has a violet-sensitive visual system with four visual pigments with absorbance peaks at 409, 458, 509 and 580 nm. The density of most cones involved in chromatic and achromatic vision peaked along a band across the retina that matched the oblique orientation of the visual streak. With the information on visual sensitivity, we calculated chromatic and achromatic contrasts of different goose plumage regions. The regions with the highest visual saliency (cheek, crown, neck and upper tail coverts) were the ones involved in visual displays to maintain flock cohesion. The Canada goose oblique visual streak is the retinal center for chromatic and achromatic vision, allowing individuals to sample the sky and the ground simultaneously or the horizon depending on head position. Overall, our results show that the Canada goose visual system has features that make it rather different from that of other vertebrates living in open habitats.

Near-UV visual pigments have been reported in single cones of several freshwater and euryhaline fishes. The presence of UV visual pigments in stenohaline marine fishes have, as yet, not been identified. In the pomacentridae near-UV visual pigments are present in single cones from the three species we examined--the tropical coral fishes Dascyllus trimaculatus and Pomacentrus coelestis, and the temperate Chromis punctipinnis. Maximum absorption of the UV pigments is centered around 360 nm. In juvenile Chromis, however, the UV visual pigment is not present. Instead there is a single cone containing a violet-sensitive pigment absorbing maximally around 420 nm. All three species are obligate diurnal planktivores. The UV sensitivity may function to enhance their ability to forage on zooplankton.