Colour vision in stomatopod crustaceans: more questions than answers.

Stomatopod crustaceans, or mantis shrimps, are known for their extensive range of spectral sensitivity but relatively poor spectral discrimination. Instead of the colour-opponent mechanism of other colour vision systems, the 12 narrow-band colour channels they possess may underlie a different method of colour processing. We investigated one hypothesis in which the photoreceptors are proposed to act as individual wave-band detectors, interpreting colour as a parallel pattern of photoreceptor activation, rather than a ratiometric comparison of individual signals. This different form of colour detection has been used to explain previous behavioural tests in which low-saturation blue was not discriminated from grey, potentially because of similar activation patterns. Results here, however, indicate that the stomatopod Haptosquilla trispinosa was able to easily distinguish several colours, including blue of both high and low saturation, from greys. The animals did show a decrease in performance over time in an artificially lit environment, indicating plasticity in colour discrimination ability. This rapid plasticity, most likely the result of a change in opsin (visual pigment) expression, has now been noted in several animal lineages (both invertebrate and vertebrate) and is a factor we suggest needs attention and potential re-examination in any colour-based behavioural tests. As for stomatopods, it remains unclear why they achieve poor colour discrimination using the most comprehensive set of spectral sensitivities in the animal kingdom and also what form of colour processing they may utilise.

IMPAIRMENTS IN CONE PIGMENT REGENERATION AND ABSOLUTE THRESHOLD IN MACULAR TELANGIECTASIA TYPE 2.

PURPOSE: To test the hypothesis that Müller cell dysfunction in macular telangiectasia type 2 (MacTel) results in delayed cone adaptation kinetics and to assess absolute cone and rod thresholds in this condition. METHODS: Adaptation after an approximate 63.5% full-field cone photopigment bleach was assessed for Goldmann size V (1.7° diameter) 640 nm (red) and 480 nm (blue) targets presented at a retinal locus corresponding to 2° temporal to fixation. The cone time constant of adaptation and absolute cone and rod thresholds were calculated from exponential functions fitted to the resultant dark adaptation curves. RESULTS: Eighteen eyes with MacTel (from 11 patients) were compared with 19 control eyes (from 16 normal subjects). Cone adaptation kinetics were significantly impaired in MacTel, as was the absolute cone threshold. Final thresholds for blue targets were also significantly elevated in MacTel, consistent with impaired rod absolute threshold. Losses in sensitivity observed in MacTel were consistent with a so-called d1/2 mechanism (i.e., receptoral) site of sensitivity loss. CONCLUSION: In addition to previously documented impairments in rod dark adaptation, MacTel results in a significant elevation in cone thresholds because of pathology at the level of the photoreceptors. The delays in cone adaptation that we found in eyes with MacTel may reflect impairment of the Müller cell-mediated cone-specific visual cycle.

A phase I clinical trial of human embryonic stem cell-derived retinal pigment epithelial cells for early-stage Stargardt macular degeneration: 5-years' follow-up.

OBJECTIVES: To evaluate the long-term biosafety and efficacy of transplantation of human embryonic stem cells-derived retinal pigment epithelial (hESC-RPE) cells in early-stage of Stargardt macular degeneration (STGD1). MATERIALS AND METHODS: Seven patients participated in this prospective clinical study, where they underwent a single subretinal transplantation of 1 × 105 hESC-RPE cells in one eye, whereas the fellow eye served as control. These patients were reassessed for a 60-month follow-up through systemic and ophthalmic examinations. RESULTS: None of the patients experienced adverse reactions systemically or locally, except for two who had transiently high intraocular pressure post-operation. Functional assessments demonstrated that all of the seven operated eyes had transiently increased or stable visual function 1-4 months after transplantation. At the last follow-up visit, two of the seven eyes showed visual function loss than the baseline; however, one of them showed a stable visual acuity when compared with the change of fellow eye. Obvious small high reflective foci in the RPE layer were displayed after the transplantation, and maintained until the last visit. Interestingly, three categories of patients who were classified based on autofluorescence, exhibited distinctive patterns of morphological and functional change. CONCLUSIONS: Subretinal transplantation of hESC-RPE in early-stage STGD1 is safe and tolerated in the long term. Further investigation is needed for choosing proper subjects according to the multi-model image and function assessments.

The spectral sensitivity of Drosophila photoreceptors.

Drosophila melanogaster has long been a popular model insect species, due in large part to the availability of genetic tools and is fast becoming the model for insect colour vision. Key to understanding colour reception in Drosophila is in-depth knowledge of spectral inputs and downstream neural processing. While recent studies have sparked renewed interest in colour processing in Drosophila, photoreceptor spectral sensitivity measurements have yet to be carried out in vivo. We have fully characterised the spectral input to the motion and colour vision pathways, and directly measured the effects of spectral modulating factors, screening pigment density and carotenoid-based ocular pigments. All receptor sensitivities had significant shifts in spectral sensitivity compared to previous measurements. Notably, the spectral range of the Rh6 visual pigment is substantially broadened and its peak sensitivity is shifted by 92 nm from 508 to 600 nm. We show that this deviation can be explained by transmission of long wavelengths through the red screening pigment and by the presence of the blue-absorbing filter in the R7y receptors. Further, we tested direct interactions between inner and outer photoreceptors using selective recovery of activity in photoreceptor pairs.

Visual cells and visual pigments of the river lamprey revisited.

Retinas of the river lamprey Lampetra fluviatilis were studied by microspectrophotometry, electroretinography and single-photoreceptor electrophysiology to reconcile the apparently contradictory conclusions on the nature of lamprey photoreceptor cells drawn in the early work by Govardovskii and Lychakov (J Comp Physiology A 154:279-286, 1984) and in recent studies. In agreement with recent works, we confirmed former identification of short photoreceptors as rods and of long photoreceptors as cones. In line with the results of 1984, we show that within a certain range of light intensities the lamprey retina exhibits "color discrimination". We found that the overlap of working intensity ranges of rods and cones is not a unique feature of lamprey short receptors, and suggest that rod-cone (possibly color) vision may be common among vertebrates. We show that the decay of meta-intermediates in lamprey cones occurs almost 100 times faster than in typical rod metarhodopsins. Rate of decay of metarhodopsins of lamprey rods take an intermediate position between typical rods and cones. This makes lamprey rhodopsin similar to transmuted cone visual pigment in "rods" of nocturnal geckos. We argue that defining various types of photoreceptors as simply "rods" and "cones" may be functionally correct, but neglects their genetic, biochemical and morphological features and evolutionary history.

Polarization perception in humans: on the origin of and relationship between Maxwell's spot and Haidinger's brushes.

Under specific conditions of illumination and polarization, differential absorption of light by macular pigments is perceived as the entoptic phenomena of Maxwell's spot (MS) or Haidinger's brushes (HB). To simulate MS and HB, an existing computational model of polarization-dependent properties of the human macula was extended by incorporating neuronal adaptation to stabilized retinal images. The model predicted that polarized light modifies the appearance of MS leading to the perception of a novel phenomenon. The model also predicted a correlation between the observed diameters of MS and HB. Predictions were tested psychophysically in human observers, whose measured differences in the diameters of each entoptic phenomenon generated with depolarized and linearly polarized light were consistent with the model simulations. These findings support a common origin of each phenomenon, and are relevant to the clinical use of polarization stimuli in detecting and monitoring human eye disorders, including macular degeneration. We conclude: (i) MS and HB both result from differential light absorption through a radial diattenuator, compatible with the arrangement of macular pigments in Henle fibres; (ii) the morphology of MS is dependent on the degree of linear polarization; (iii) perceptual differences between MS and HB result from different states of neural adaptation.

A detailed investigation of the visual system and visual ecology of the Barrier Reef anemonefish, Amphiprion akindynos.

Vision plays a major role in the life of most teleosts, and is assumingly well adapted to each species ecology and behaviour. Using a multidisciplinary approach, we scrutinised several aspects of the visual system and ecology of the Great Barrier Reef anemonefish, Amphiprion akindynos, including its orange with white patterning, retinal anatomy and molecular biology, its symbiosis with anemones and sequential hermaphroditism. Amphiprion akindynos possesses spectrally distinct visual pigments and opsins: one rod opsin, RH1 (498 nm), and five cone opsins, SWS1 (370 nm), SWS2B (408 nm), RH2B (498 nm), RH2A (520 nm), and LWS (554 nm). Cones were arranged in a regular mosaic with each single cone surrounded by four double cones. Double cones mainly expressed RH2B (53%) in one member and RH2A (46%) in the other, matching the prevailing light. Single cones expressed SWS1 (89%), which may serve to detect zooplankton, conspecifics and the host anemone. Moreover, a segregated small fraction of single cones coexpressed SWS1 with SWS2B (11%). This novel visual specialisation falls within the region of highest acuity and is suggested to increase the chromatic contrast of Amphiprion akindynos colour patterns, which might improve detection of conspecifics.

Multichannel spectrometers in animals.

Multispectral, hyperspectral, polarimetric, and other types of multichannel imaging spectrometers are coming into common use for a variety of applications, including remote sensing, material identification, forensics, and medical diagnosis. These instruments are often bulky and intolerant of field abuse, so designing compact, reliable, portable, and robust devices is a priority. In contrast to most engineering designs, animals have been building compact and robust multichannel imaging systems for millennia-their eyes. Biological sensors arise by evolution, of course, and are not designed 'for' a particular use; they exist because the creatures that were blessed with useful mutations were better able to survive and reproduce than their competitors. While this is an inefficient process for perfecting a sensor, it brings unexpected innovations and novel concepts into visual system design-concepts that may be useful in the inspiration of new engineered solutions to problematic challenges, like the ones mentioned above. Here, we review a diversity of multichannel visual systems from both vertebrate and invertebrate animals, considering the receptor molecules and cells, spectral sensitivity and its tuning, and some aspects of the higher-level processing systems used to shape spectral (and polarizational) channels in vision. The eyes of mantis shrimps are presented as potential models for biomimetic multichannel imaging systems. We end with a description of a bioinspired, newly developed multichannel spectral/polarimetric imaging system based on mantis shrimp vision that is highly adaptable to field application.

Photopigments and the dimensionality of animal color vision.

Early color-matching studies established that normal human color vision is trichromatic. Subsequent research revealed a causal link between trichromacy and the presence in the retina of three classes of cone photopigments. Over the years, measurements of the photopigment complements of other species have expanded greatly and these are frequently used to predict the dimensionality of an animal's color vision. This review provides an account of how the linkage between the number of active photopigments and the dimensions of human color vision developed, summarizes the various mechanisms that can impact photopigment spectra and number, and provides an across-species survey to examine cases where the photopigment link to the dimensionality of color vision has been claimed. The literature reveals numerous instances where the human model fails to account for the ways in which the visual systems of other animals exploit information obtained from the presence of multiple photopigments in support of their behavior.

Volumetric integration model of the Stiles-Crawford effect of the first kind and its experimental verification.

The integrated Stiles-Crawford function is commonly used as apodization model for vision through the natural eye pupil. However, this method does not account for possible effects related to the retinal thickness, the large length-to-diameter aspect ratio of the photoreceptors, or the use of nonMaxwellian illumination. Here, we introduce a geometrical optics model to calculate the fraction of overlap between light at the retina and the photoreceptor outer segments where absorption triggers vision. The model, which does not account for photoreceptor waveguiding, is discussed for both Maxwellian and nonMaxwellian illumination. The integrated Stiles-Crawford effect is analyzed experimentally with a uniaxial pupil-size flicker methodology and we find that the psychophysical measurements match better to the geometrical optics predictions than direct integration of a Stiles-Crawford function.

Functional interplay of visual, sensitizing and screening pigments in the eyes of Drosophila and other red-eyed dipteran flies.

Several fly species have distinctly red-coloured eyes, meaning that the screening pigments that provide a restricted angular sensitivity of the photoreceptors may perform poorly in the longer wavelength range. The functional reasons for the red transparency and possible negative visual effects of the spectral properties of the eye-colouring screening pigments are discussed within the context of the photochemistry, arrestin binding and turnover of the visual pigments located in the various photoreceptor types. A phylogenetic survey of the spectral properties of the main photoreceptors of the Diptera indicates that the transition of the brown eye colour of the Nematocera and lower Brachycera to a much redder eye colour of the higher Brachycera occurred around the emergence of the Tabanidae family.

[Physiology of the visual retinal signal: From phototransduction to the visual cycle]. / Physiologie du signal visuel rétinien : de la phototransduction jusqu'au cycle visuel.

The retinal photoreceptors (rods and cones) are responsible for light absorption and transduction of the signal, which is transmitted to the other retinal nerve cells and then to the brain. The chromophore of visual pigments of rods and cones is a particular isomer of a vitamin A derivative. Light absorption by this chromophore leads to its isomerization and to a phototransduction cascade, which results in photoreceptor hyperpolarization and cessation of glutamate secretion at their synaptic terminals. Phototransduction of cones and rods differs in their signal amplification and inactivation, which is consistent with their respective functions. The rods serve for dim light vision, whereas color and detailed vision is provided by cones. The rods are thus much more sensitive than cones, but the time course of cones' photoresponse is ∼10 times faster than that of rods. The orientation of cone visual pigments in the retina is optimized to achieve their function. The isomerized chromophore of visual pigments is regenerated by a mechanism known as the visual cycle. This process takes place mainly in the retinal pigment epithelium for the rods and the glial Müller cells for the cones. Mutations of a large number of proteins involved in visual phototransduction and in the retinoid visual cycle are responsible for hereditary diseases leading to photoreceptor degeneration. However, gene therapy offers quite a bit of hope for treatment.

Relationship between saccadic eye movements and formation of the Krukenberg's spindle-a CFD study.

In this research, a series of numerical simulations for evaluating the effects of saccadic eye movement on the aqueous humour (AH) flow field and movement of pigment particles in the anterior chamber (AC) was performed. To predict the flow field of AH in the AC, the unsteady forms of continuity, momentum balance and conservation of energy equations were solved using the dynamic mesh technique for simulating the saccadic motions. Different orientations of the human eye including horizontal, vertical and angles of 10° and 20° were considered. The Lagrangian particle trajectory analysis approach was used to find the trajectories of pigment particles in the eye. Particular attention was given to the relation between the saccadic eye movement and potential formation of Krukenberg's spindle in the eye. The simulation results revealed that the natural convection flow was an effective mechanism for transferring pigment particles from the iris to near the cornea. In addition, the saccadic eye movement was the dominant mechanism for deposition of pigment particles on the cornea, which could lead to the formation of Krukenberg's spindle. The effect of amplitude of saccade motion angle in addition to the orientation of the eye on the formation of Krukenberg's spindle was investigated.

The evidence informing the surgeon's selection of intraocular lens on the basis of light transmittance properties.

In recent years, manufacturers and distributors have promoted commercially available intraocular lenses (IOLs) with transmittance properties that filter visible short-wavelength (blue) light on the basis of a putative photoprotective effect. Systematic literature review. Out of 21 studies reporting on outcomes following implantation of blue-light-filtering IOLs (involving 8914 patients and 12 919 study eyes undergoing cataract surgery), the primary outcome was vision, sleep pattern, and photoprotection in 9 (42.9%), 9 (42.9%), and 3 (14.2%) respectively, and, of these, only 7 (33.3%) can be classed as high as level 2b (individual cohort study/low-quality randomized controlled trials), all other studies being classed as level 3b or lower. Of the level 2b studies, only one (14.3%) found in favor of blue-light-filtering IOLs vs ultraviolet (UV)-only filtering IOLs on the basis of an association between better post-operative contrast sensitivity (CS) at select frequencies with the former; however, that study did not measure or report CS preoperatively in either group, and the finding may simply reflect better preoperative CS in the eyes scheduled to be implanted with the blue-light-filtering IOL; moreover, that study failed to measure macular pigment, a natural preceptoral filter of blue-light, augmentation of which is now known to improve CS. In terms of photoprotection, there is no level 2b (or higher) evidence in support of blue filtering IOLs vs UV-only filtering IOLs. On the basis of currently available evidence, one cannot advocate for the use of blue-light-filtering IOLs over UV-only filtering IOLs.

Vitamin A and Vision.

Visual systems detect light by monitoring the effect of photoisomerization of a chromophore on the release of a neurotransmitter from sensory neurons, known as rod and cone photoreceptor cells in vertebrate retina. In all known visual systems, the chromophore is 11-cis-retinal complexed with a protein, called opsin, and photoisomerization produces all-trans-retinal. In mammals, regeneration of 11-cis-retinal following photoisomerization occurs by a thermally driven isomerization reaction. Additional reactions are required during regeneration to protect cells from the toxicity of aldehyde forms of vitamin A that are essential to the visual process. Photochemical and phototransduction reactions in rods and cones are identical; however, reactions of the rod and cone visual pigment regeneration cycles differ, and perplexingly, rod and cone regeneration cycles appear to use different mechanisms to overcome the energy barrier involved in converting all-trans- to 11-cis-retinoid. Abnormal processing of all-trans-retinal in the rod regeneration cycle leads to retinal degeneration, suggesting that excessive amounts of the retinoid itself or its derivatives are toxic. This line of reasoning led to the development of various approaches to modifying the activity of the rod visual cycle as a possible therapeutic approach to delay or prevent retinal degeneration in inherited retinal diseases and perhaps in the dry form of macular degeneration (geographic atrophy). In spite of great progress in understanding the functioning of rod and cone regeneration cycles at a molecular level, resolution of a number of remaining puzzling issues will offer insight into the amelioration of several blinding retinal diseases.

Comparative visual ecology of cephalopods from different habitats.

Previous investigations of vision and visual pigment evolution in aquatic predators have focused on fish and crustaceans, generally ignoring the cephalopods. Since the first cephalopod opsin was sequenced in late 1980s, we now have data on over 50 cephalopod opsins, prompting this functional and phylogenetic examination. Much of this data does not specifically examine the visual pigment spectral absorbance position (λmax) relative to environment or lifestyle, and cephalopod opsin functional adaptation and visual ecology remain largely unknown. Here we introduce a new protocol for photoreceptor microspectrophotometry (MSP) that overcomes the difficulty of bleaching the bistable visual pigment and that reveals eight coastal coleoid cephalopods to be monochromatic with λmax varying from 484 to 505 nm. A combination of current MSP results, the λmax values previously characterized using cephalopod retinal extracts (467-500 nm) and the corresponding opsin phylogenetic tree were used for systematic comparisons with an end goal of examining the adaptations of coleoid visual pigments to different light environments. Spectral tuning shifts are described in response to different modes of life and light conditions. A new spectral tuning model suggests that nine amino acid substitution sites may determine the direction and the magnitude of spectral shifts.

Spectral sensitivity of guppy visual pigments reconstituted in vitro to resolve association of opsins with cone cell types.

The guppy (Poecilia reticulata) shows remarkable variation of photoreceptor cells in the retina, especially those sensitive to middle-to-long wavelengths of light. Microspectrophotometry (MSP) has revealed varying "green", "green-yellow" and "yellow" cone cells among guppies in Trinidad and Venezuela (Cumana). In the guppy genome, there are four "long-wave" opsin loci (LWS-1, -2, -3 and -4). Two LWS-1 alleles have potentially differing spectral sensitivity (LWS-1/180Ser and LWS-1/180Ala). In addition, two "middle-wave" loci (RH2-1 and -2), two "short-wave" loci (SWS2-A and -B), and a single "ultraviolet" locus (SWS1) as well as a single "rhodopsin" locus (RH1) are present. However, the absorption spectra of these photopigments have not been measured directly and the association of cell types with these opsins remains speculative. In the present study, we reconstituted these opsin photopigments in vitro. The wavelengths of maximal absorbance (λmax) were 571nm (LWS-1/180Ser), 562nm (LWS-1/180Ala), 519nm (LWS-3), 516nm (LWS-2), 516nm (RH2-1), 476nm (RH2-2), 438nm (SWS2-A), 408nm (SWS2-B), 353nm (SWS1) and 503nm (RH1). The λmax of LWS-3 is much shorter than the value expected (560nm) from the "five-sites" rule. The two LWS-1 alleles could explain difference of the reported MSP λmax values for the yellow cone class between Trinidad and Cumana guppies. Absence of the short-wave-shifted LWS-3 and the green-yellow cone in the green swordtail supports the hypothesis that this cell class of the guppy co-expresses the LWS-1 and LWS-3. These results reveal the basis of variability in the guppy visual system and provide insight into the behavior and ecology of these tropical fishes.

Dimerization of visual pigments in vivo.

It is a deeply engrained notion that the visual pigment rhodopsin signals light as a monomer, even though many G protein-coupled receptors are now known to exist and function as dimers. Nonetheless, recent studies (albeit all in vitro) have suggested that rhodopsin and its chromophore-free apoprotein, R-opsin, may indeed exist as a homodimer in rod disk membranes. Given the overwhelmingly strong historical context, the crucial remaining question, therefore, is whether pigment dimerization truly exists naturally and what function this dimerization may serve. We addressed this question in vivo with a unique mouse line (S-opsin(+)Lrat(-/-)) expressing, transgenically, short-wavelength-sensitive cone opsin (S-opsin) in rods and also lacking chromophore to exploit the fact that cone opsins, but not R-opsin, require chromophore for proper folding and trafficking to the photoreceptor's outer segment. In R-opsin's absence, S-opsin in these transgenic rods without chromophore was mislocalized; in R-opsin's presence, however, S-opsin trafficked normally to the rod outer segment and produced functional S-pigment upon subsequent chromophore restoration. Introducing a competing R-opsin transmembrane helix H1 or helix H8 peptide, but not helix H4 or helix H5 peptide, into these transgenic rods caused mislocalization of R-opsin and S-opsin to the perinuclear endoplasmic reticulum. Importantly, a similar peptide-competition effect was observed even in WT rods. Our work provides convincing evidence for visual pigment dimerization in vivo under physiological conditions and for its role in pigment maturation and targeting. Our work raises new questions regarding a potential mechanistic role of dimerization in rhodopsin signaling.

Visual pigments in a palaeognath bird, the emu Dromaius novaehollandiae: implications for spectral sensitivity and the origin of ultraviolet vision.

A comprehensive description of the spectral characteristics of retinal photoreceptors in palaeognaths is lacking. Moreover, controversy exists with respect to the spectral sensitivity of the short-wavelength-sensitive-1 (SWS1) opsin-based visual pigment expressed in one type of single cone: previous microspectrophotometric (MSP) measurements in the ostrich (Struthio camelus) suggested a violet-sensitive (VS) SWS1 pigment, but all palaeognath SWS1 opsin sequences obtained to date (including the ostrich) imply that the visual pigment is ultraviolet-sensitive (UVS). In this study, MSP was used to measure the spectral properties of visual pigments and oil droplets in the retinal photoreceptors of the emu (Dromaius novaehollandiae). Results show that the emu resembles most other bird species in possessing four spectrally distinct single cones, as well as double cones and rods. Four cone and a single rod opsin are expressed, each an orthologue of a previously identified pigment. The SWS1 pigment is clearly UVS (wavelength of maximum absorbance [λmax] = 376 nm), with key tuning sites (Phe86 and Cys90) consistent with other vertebrate UVS SWS1 pigments. Palaeognaths would appear, therefore, to have UVS SWS1 pigments. As they are considered to be basal in avian evolution, this suggests that UVS is the most likely ancestral state for birds. The functional significance of a dedicated UVS cone type in the emu is discussed.