Classes Of Photoreceptors Research Articles

Behavioural evidence of spectral opponent processing in the visual system of stomatopod crustaceans.

Stomatopods, commonly known as mantis shrimps, possess an intricate colour vision with up to 12 photoreceptor classes arranged in four specialised ommatidia rows (rows 1-4 in the midband region of the eye) for colour perception. While 2-4 spectral sensitivities suffice for most visual systems, the function and mechanism behind stomatopods' 12-channel colour vision remains unclear. Previous anatomical and behavioural studies have suggested that binning and opponent processing mechanisms may coexist in stomatopods' colour vision. However, direct evidence of colour opponency has been lacking. We hypothesised that if colour opponency exists in stomatopods' vision, they would be able to distinguish colour from grey under coloured illumination. Conversely, if only the binning system is used, they would not. By examining the colour vision of the stomatopod Haptosquilla trispinosa with modified von Frisch grey card experiments, we found that they can differentiate between colour and grey under various coloured illuminations. Our results provide the first direct behavioural evidence of spectral opponency in stomatopods, suggesting that they use a hybrid colour processing system combining opponent and binning mechanisms for colour vision. This study advances our understanding of the complex visual system in stomatopods and highlights the importance of further research into the processing mechanisms, function and evolution of their unique visual system.

What can the eye see with melanopsin?

A subpopulation of human retinal ganglion cells contains the melanopsin photopigment, allowing them to act as a fifth photoreceptor class. These ganglion cells project to the visual cortex, but to reveal its intrinsic contribution to conscious vision is technically challenging as it requires melanopsin to be separated from the responses originating in the rods and three cone classes. Using a display engineered to isolate the melanopic visual response, we show that it detects lowpass spatial (≤0.35 cycles per degree) and temporal image content (≤1 Hz) but cannot reconstruct the stimulus form necessary for object recognition. We demonstrate that a model of the spatially diffuse intrinsically-photosensitive retinal ganglion cells' sampling structure is predictive of the measured image reconstruction limits of melanopic spatial vision. Separately, we find that under five-photoreceptor silent substitution conditions, rod pathways alone can support form vision in bright lighting when typically thought to be in saturation. Form vision that is absent from melanopsin can be only perceived in mixtures of both melanopsin and rod signals because it is the rod pathway that sees the form. Our findings show that melanopsin's unique tuning to the diffuse and slow-changing elements in the world provides a stabilized reference point for vision.

Ips typographus vision system: a comprehensive study.

Aggressive bark beetle species such as the Eurasian spruce bark beetle Ips typographus play a fundamental role in forest ecosystems but can also lead to extensive forest mortality and massive economic damage during outbreaks. Currently I. typographus' eyes, visual perception of the world and recognition of specific targets like host plants are understudied topics. Studying its visual sense can open the way to novel efficient monitoring and management methods, particularly important in avoiding the switch from an endemic to an epidemic condition. In addition, the integration of visual cues in trapping systems may offer new opportunities for surveillance. Vision in I. typographus was investigated by means of morphological analysis, electroretinography (ERG), molecular analysis of opsin genes and behavioural tests. ERG has revealed that the compound eyes contain two classes of photoreceptors, maximally sensitive to UV and green at 370 and 530nm, respectively. The result was further supported by the identification of two relevant opsin genes. Finally, the innate wavelength sensitivity was tested in a Y-maze. Ips typographus consistently preferred UV over non-UV (VIS) light, irrespective of their intensity ratios, but preferred high over low intensity VIS light, consistent with a UV-VIS dichromatic visual system. Overall, the results may open the way to better understand the navigation pattern in tree canopies and the host selection processes of this ecologically and economically important beetle species.

Morphology and spectral sensitivity of long visual fibers and lamina monopolar cells in the butterfly Papilio xuthus.

Extensive analysis of the flower-visiting behavior of a butterfly,Papilio xuthus, has indicated complex interaction between chromatic, achromatic, and motion cues. Their eyes are spectrally rich with six classes of photoreceptors, respectively sensitive in the ultraviolet, violet, blue, green, red, and broad-band wavelength regions. Here, we studied the anatomy and physiology of photoreceptors and second-order neurons ofP. xuthus, focusing on their spectral sensitivities and projection terminals to address where the early visual integration takes place. We thus found the ultraviolet, violet, and blue photoreceptors and all second-order neurons terminate in the distal region of the second optic ganglion, the medulla. We identified five types of second-order neurons based on the arborization in the first optic ganglion, the lamina, and the shape of the medulla terminals. Their spectral sensitivity is independent of the morphological types but reflects the combination of pre-synaptic photoreceptors. The results indicate that the distal medulla is the most plausible region for early visual integration.

One More for Light-triggered Conformational Changes in Cryptochromes: CryP from Phaeodactylum tricornutum

Cryptochromes are a ubiquitously occurring class of photoreceptors. Together with photolyases, they form the Photolyase Cryptochrome Superfamily (PCSf) by sharing a common protein architecture and binding mode of the FAD chromophore. Despite these similarities, PCSf members exert different functions. Photolyases repair UV-induced DNA damage by photocatalytically driven electron transfer between FADH¯ and the DNA lesion, whereas cryptochromes are light-dependent signaling molecules and trigger various biological processes by photoconversion of their FAD redox and charge states. Given that most cryptochromes possess a C-terminal extension (CTE) of varying length, the functions of their CTE have not yet been fully elucidated and are hence highly debated. In this study, the role of the CTE was investigated for a novel subclass of the PCSf, the CryP-like cryptochromes, by hydrogen/deuterium exchange and mass-spectrometric analysis. Striking differences in the relative deuterium uptake were observed in different redox states of CryP from the diatom Phaeodactylum tricornutum. Based on these measurements we propose a model for light-triggered conformational changes in CryP-like cryptochromes that differs from other known cryptochrome families like the insect or plant cryptochromes.

Contributed Session II: Binocular combination of the pupil response depends on photoreceptor pathway.

The pupillary light response is driven by three classes of retinal photoreceptor. Cones and rods are involved in the initial constriction of the pupil, whereas melanopsin-containing intrinsically photosensitive Retinal Ganglion Cells (ipRGCs) maintain constriction over longer timescales. Previous work has characterized the contributions of photoreceptor signals to pupil control, but relatively little is known about binocular combination of these signals when simultaneously stimulating the retina in both eyes. We measured changes in pupil size in 48 participants using a binocular eye-tracker, targeting specific photoreceptor classes with a binocular 10-primary light engine and the silent substitution method. We stimulated the periphery of the retina using light flickering at 0.4 and 0.5 Hz. Participants viewed a disc of either achromatic flickering light, or contrast modulations that targeted the ipRGCs, or the opponent colour pathways L-M or S-(L+M). Using a modified virtual reality headset, we presented the stimuli at a range of modulation amplitudes in three different ocular configurations: monocular, binocular, and dichoptic. We obtained clear pupil responses at both the first and the second harmonic frequencies. Suppression levels differed across conditions with the strongest suppression measured for the L-M condition. We account for the results in a single modelling framework where the weight of interocular suppression determines the binocular combination properties.

Spectral sensitivity of retinal photoreceptors of tortricid moths is not tuned to diel activity period.

Leafrollers (Lepidoptera: Tortricidae) are a large family of small moths containing over 10,000 species, many of which are crop pests. Grapholita molesta, Lobesia botrana and Cydia pomonella adults are sexually active before, during and after sunset, respectively. We wanted to determine whether being active at different times of the day and night is associated with differences in their visual system. Spectral sensitivity (SS) was measured with electroretinograms and selective adaptation with green, blue and ultraviolet light. SS curves could be fitted with a triple nomogram template which indicated the existence of three photoreceptor classes peaking at 355, 440 and 525 nm. The retinae showed clear regionalization, with fewer blue receptors dorsally. No differences among species or between sexes were found. Intracellular recordings in C. pomonella also revealed three photoreceptor classes with sensitivities peaking at 355, 440 and 525 nm. The blue photoreceptors showed inhibitory responses in the green part of the spectrum, indicating the presence of a colour-opponent system. Flicker fusion frequency experiments showed that the response speed was similar between sexes and species and fused at around 100 Hz. Our results indicate that the three species have the ancestral insect retinal substrate for a trichromatic colour vision, based upon the UV, blue and green-sensitive photoreceptors, and lack any prominent adaptations related to being active under different light conditions.

Trichromatic vision in toads: evidence from preference for colour objects during mate choice

Abstract Input of different spectral types of photoreceptors in amphibian colour vision has been assessed by studying the behaviour of male toads Bufo bufo and B. gargarizans in a laboratory. This method is based on the males’ innate reaction of approaching and clasping any visual object during the breeding season. A pair of colour stimuli made of paper or displayed on a LCD monitor has been selected to match the toad’s RGB colour space so that they differ by excitation of only a single spectral class of retinal photoreceptors. With pairwise presentation, the males are found to prefer a more ‘bluish’, or a less ‘reddish’, or a less ‘greenish’ stimulus from each pair. A microspectrophotometry of isolated rod and cone photoreceptors and an observation of behavioural responses in toads have revealed a trichromatic mesopic visual system based on a positive input from blue-sensitive photoreceptors and negative inputs from red-sensitive cones and green-sensitive rods. It is supposed that the observed colour preference and involvement of different chromatic mechanisms determine social interactions of anurans during mate choice.

PySilSub: An open-source Python toolbox for implementing the method of silent substitution in vision and nonvisual photoreception research.

The normal human retina contains several classes of photosensitive cell-rods for low-light vision, three cone classes for daylight vision, and intrinsically photosensitive retinal ganglion cells (ipRGCs) expressing melanopsin for non-image-forming functions, including pupil control, melatonin suppression, and circadian photoentrainment. The spectral sensitivities of the photoreceptors overlap significantly, which means that most lights will stimulate all photoreceptors to varying degrees. The method of silent substitution is a powerful tool for stimulating individual photoreceptor classes selectively and has found much use in research and clinical settings. The main hardware requirement for silent substitution is a spectrally calibrated light stimulation system with at least as many primaries as there are photoreceptors under consideration. Device settings that will produce lights to selectively stimulate the photoreceptor(s) of interest can be found using a variety of analytic and algorithmic approaches. Here we present PySilSub (https://github.com/PySilentSubstitution/pysilsub), a novel Python package for silent substitution featuring flexible support for individual colorimetric observer models (including human and mouse observers), multiprimary stimulation devices, and solving silent substitution problems with linear algebra and constrained numerical optimization. The toolbox is registered with the Python Package Index and includes example data sets from various multiprimary systems. We hope that PySilSub will facilitate the application of silent substitution in research and clinical settings.

The Color Vision Disorder in Diabetic Patients

Introduction: The visible spectrum (about 400 to 700 nm) with a mosaic of three classes of photoreceptors compose the human visual system. They are sensitive to different wavelength ranges with overlapping. The receivers have high sensitivities at short wavelengths (~440 nm), medium (~535 nm), or longer (~565 nm), which are S, M, and L cones, respectively. The study aimed to determine blue color vision defects in diabetes mellitus.
 Methods: A cross-sectional study was done at an ophthalmology clinic on 200 eyes (right and left) of patients with DM for a period of 5 one years (2021). Ishihara plates were used initially for screening. The D-15 test was performed for the evaluation of color vision.
 Results: The mean age of the sample was 45.66±15.65 years. The most frequent disorder visualized was tritanomaly in 63%, followed by trichromate in 27%, and the least disorder was deuteranomaly in 2%. In addition, mixed disorder is seen in 8% of cases. In relation to laterality, right eye tritanomaly was found in 34%, while the left eye was recorded in 29%. The right eye trichromate was observed in 12%, whereas the left eye was reported in 15%. The left eye deuteranomaly was reported more than the right (1.5% vs. 0.5%).
 Conclusion: Color vision evaluation with good screening color vision test can be detected even before clinically visible diabetic retinopathy. Early detection was helpful in the prevention of vascular changes in the retina. All diabetic patients should be given proper awareness and health education regarding color vision deficiency. Timely assessment of color vision may detect tritanomaly earlier in diabetics.

Molecular bases of signaling processes regulated by cryptochrome sensory photoreceptors in plants

The blue-light sensors cryptochromes compose the widespread class of flavoprotein photoreceptors, regulating in plants signaling processes underlying their development, growth, and metabolism. In several algae cryptochromes may act not only as sensory photoreceptors but also as photolyases, catalyzing the repair of UV-induced DNA lesions. Cryptochromes bind FAD as the chromophore in the photolyase homologous region (PHR) domain and contain the cryptochrome C-terminal extension (CCE), which lacks in photolyases. Photosensory process in cryptochrome is initiated by photochemical chromophore conversions, including the formation of FAD redox forms. In a state with the chromophore reduced to neutral radical (FADH•), photoreceptory protein undergoes phosphorylation, conformational changes, and disengagement of the PHR domain and CCE with the subsequent formation of oligomers of cryptochrome molecules. Photooligomerization is a structural basis of functional activity of cryptochromes, which determines the formation of their complexes with variety of signaling proteins, including transcriptional factors and transcription regulators. Interactions in such complexes change the protein signaling activity, leading to gene expression regulation and plant photomorphogenesis. In recent years, multiple papers, reporting novel, more detailed information about molecular mechanisms of above-mentioned processes were published. The present review largely focuses on the analysis of data contained in these publications, particularly regarding structural aspects of cryptochrome transitions into photoactivated states and regulatory signaling processes mediated by cryptochrome photoreceptors in plants.

Molecular Bases of Signaling Processes Regulated by Cryptochrome Sensory Photoreceptors in Plants.

The blue-light sensors, cryptochromes, compose the extensive class of flavoprotein photoreceptors, regulating signaling processes in plants underlying their development, growth, and metabolism. In several algae, cryptochromes may act not only as sensory photoreceptors but also as photolyases, catalyzing repair of the UV-induced DNA lesions. Cryptochromes bind FAD as the chromophore at the photolyase homologous region (PHR) domain and contain the cryptochrome C-terminal extension (CCE), which is absent in photolyases. Photosensory process in cryptochrome is initiated by photochemical chromophore conversions, including formation of the FAD redox forms. In the state with the chromophore reduced to neutral radical (FADH×), the photoreceptor protein undergoes phosphorylation, conformational changes, and disengagement from the PHR domain and CCE with subsequent formation of oligomers of cryptochrome molecules. Photooligomerization is a structural basis of the functional activities of cryptochromes, since it ensures formation of their complexes with a variety of signaling proteins, including transcriptional factors and regulators of transcription. Interactions in such complexes change the protein signaling activities, leading to regulation of gene expression and plant photomorphogenesis. In recent years, multiple papers, reporting novel, more detailed information about the molecular mechanisms of above-mentioned processes were published. The present review mainly focuses on analysis of the data contained in these publications, particularly regarding structural aspects of the cryptochrome transitions into photoactivated states and regulatory signaling processes mediated by the cryptochrome photoreceptors in plants.

Physiological and behavioral evidence for multiple spectral channels in the larval stomatopod visual system.

Larval stomatopods have generally been described as having a typical larval crustacean compound eye, which lacks the visual pigment diversity and morphological specializations of the well-studied stomatopod adult eye. However, recent work has suggested that larval stomatopod eyes are more complex than previously described. In this study we provide physiological and behavioral evidence of at least three distinct photoreceptor classes in three species of larval stomatopods: Gonodactylellus n. sp., Gonodactylaceus falcatus, and Pullosquilla n. sp. First, electroretinogram recordings were used to measure the spectral sensitivity of each species. Evidence for at least three spectral classes were identified in each: a UV peaking at 340-376 nm, a short wavelength blue peaking at 455-464 nm, and a long wavelength orange peaking at 576-602 nm. Next, the behavioral response to light was investigated. We found that each species demonstrated positive phototactic responses to monochromatic stimuli across the ultraviolet-visible spectrum. In wavelength preference trials, distinct preferences among species were identified when different colored light stimuli were presented simultaneously. All species displayed a strong response to the UV stimulus, as well as responses to blue and orange stimuli, although at different response strengths, but no response to green. The results of this study demonstrate that larval stomatopods not only have multiple physiologically active spectral classes, but they also display clear and distinct responses to colors across the spectrum. We propose that the spectral classes demonstrated in each are related to visually guided ecological tasks of the larvae, which may differ between species.

Blue light promotes ascorbate synthesis by deactivating the PAS/LOV photoreceptor that inhibits GDP-L-galactose phosphorylase.

Ascorbate (vitamin C) is an essential antioxidant in fresh fruits and vegetables. To gain insight into the regulation of ascorbate metabolism in plants, we studied mutant tomato plants (Solanum lycopersicum) that produce ascorbate-enriched fruits. The causal mutation, identified by a mapping-by-sequencing strategy, corresponded to a knock-out recessive mutation in a class of photoreceptor named PAS/LOV protein (PLP), which acts as a negative regulator of ascorbate biosynthesis. This trait was confirmed by CRISPR/Cas9 gene editing and further found in all plant organs, including fruit that accumulated 2 to 3 times more ascorbate than in the WT. The functional characterization revealed that PLP interacted with the 2 isoforms of GDP-L-galactose phosphorylase (GGP), known as the controlling step of the L-galactose pathway of ascorbate synthesis. The interaction with GGP occurred in the cytoplasm and the nucleus, but was abolished when PLP was truncated. These results were confirmed by a synthetic approach using an animal cell system, which additionally demonstrated that blue light modulated the PLP-GGP interaction. Assays performed in vitro with heterologously expressed GGP and PLP showed that PLP is a noncompetitive inhibitor of GGP that is inactivated after blue light exposure. This discovery provides a greater understanding of the light-dependent regulation of ascorbate metabolism in plants.

Non-image-forming functional roles of OPN3, OPN4 and OPN5 photopigments

Detecting different wavelengths and intensities of environmental light is crucial for the survival of many animals. In response, a multiplicity of opsins (a special group of photosensitive G protein-coupled receptors), when combined with a retinal chromophore, is able to directly detect light and initiate different downstream phototransduction signaling cascades. Although avian studies from the 1930s suggested the presence of deep brain photoreceptors that could respond to seasonal changes in the light/dark cycle, it was only a few decades ago that photopigments other than those found in the visual system (i.e. rods and cones) were identified as functional photoreceptors. It is now established that several classes of non-visual photoreceptors and the photopigments they express, in lower vertebrates to higher mammals alike, can regulate a plethora of mechanisms that function outside of vision. These include the synchronization of light/dark cycles with biological/cellular rhythms of the body (i.e. photoentrainment); melanogenesis in dermal tissues; thermoregulation in adipose tissue; embryonic eye development; smooth muscle relaxation; and the development of certain cancers. These and other mechanisms have been shown, in part at least, to be controlled by the expression of three important non-visual opsin genes, namely OPN3, OPN4 and OPN5, although other vertebrate opsin classes exist, many with unknown or unclear functional roles assigned to them presently. Specifically, these three opsins have been shown to be expressed during early embryogenesis and throughout adulthood, which will be discussed here. Moreover, this review highlights recent studies that focus on several key non-image-forming functional roles of OPN3, OPN4 and OPN5, and in particular those that impact photoreception in developing structures and pathways, as well as in adulthood.

OsCRY2 and OsFBO10 co-regulate photomorphogenesis and photoperiodic flowering in indica rice

Cryptochromes (CRYs) are a class of photoreceptors that perceive blue/ultraviolet-A light of the visible spectrum to mediate a vast number of physiological responses in bacteria, fungi, animals and plants. In the present study, we have characterized OsCRY2 in a photoperiod sensitive indica variety, Basmati 370, by generating and analyzing overexpression (OE) and knock-down (KD) transgenic lines. The OsCRY2OE lines displayed dwarfism as shown in their reduced plant height and leaf length, attributed largely by an overall reduction in their cell size. The OsCRY2OE lines flowered significantly earlier and showed shorter and broader seeds with an overall reduced seed weight. The OsCRY2KD lines showed contrasting phenotypes, such as increased plant height and delayed flowering, however, decreased seed size and weight were also observed in the KD lines, along with reduced spikelet fertility and high seed shattering rate in mature panicles. Novel interactions were confirmed between OsCRY2 and members of ZEITLUPE family of blue/ultraviolet-A light photoreceptors, encoded by OsFBO8, OsFBO9 and OsFBO10 which are orthologous to ZEITLUPE (ZTL), LOV KELCH PROTEIN2 (LKP2) and FLAVIN BINDING, KELCH REPEAT F-BOX1 (FKF1), respectively, of Arabidopsis thaliana. Since FKF1 is known to play a role in regulating photoperiodic flowering, OsFBO10 was chosen for further studies. OsCRY2 and OsFBO10 interacted in the nucleus and cytoplasm of the cell and cross-regulated the expression of each other. They were also found to regulate the expression of several genes involved in photoperiodic flowering in rice. Both OsCRY2 and OsFBO10 played a positive role in photomorphogenic responses in different light conditions. The physical interaction of OsCRY2 with OsFBO10, their involvement in common physiological and developmental pathways and their cross-regulation of each other suggest that the two photoreceptors may regulate common developmental pathways in plants, either jointly or redundantly.

Circadian rhythm entrainment of the jewel wasp, Nasonia vitripennis, by antagonistic interactions of multiple spectral inputs.

Circadian light entrainment in some insects is regulated by blue-light-sensitive cryptochrome (CRY) protein that is expressed in the clock neurons, but this is not the case in hymenopterans. The hymenopteran clock does contain CRY, but it appears to be light-insensitive. Therefore, we investigated the role of retinal photoreceptors in the photic entrainment of the jewel wasp Nasonia vitripennis. Application of monochromatic light stimuli at different light intensities caused phase shifts in the wasp's circadian activity from which an action spectrum with three distinct peaks was derived. Electrophysiological recordings from the compound eyes and ocelli revealed the presence of three photoreceptor classes, with peak sensitivities at 340 nm (ultraviolet), 450 nm (blue) and 530 nm (green). An additional photoreceptor class in the ocelli with sensitivity maximum at 560-580 nm (red) was found. Whereas a simple sum of photoreceptor spectral sensitivities could not explain the action spectrum of the circadian phase shifts, modelling of the action spectrum indicates antagonistic interactions between pairs of spectral photoreceptors, residing in the compound eyes and the ocelli. Our findings imply that the photic entrainment mechanism in N. vitripennis encompasses the neural pathways for measuring the absolute luminance as well as the circuits mediating colour opponency.

Jewel Beetle Opsin Duplication and Divergence Is the Mechanism for Diverse Spectral Sensitivities.

The evolutionary history of visual genes in Coleoptera differs from other well-studied insect orders, such as Lepidoptera and Diptera, as beetles have lost the widely conserved short-wavelength (SW) insect opsin gene that typically underpins sensitivity to blue light (∼440 nm). Duplications of the ancestral ultraviolet (UV) and long-wavelength (LW) opsins have occurred in many beetle lineages and have been proposed as an evolutionary route for expanded spectral sensitivity. The jewel beetles (Buprestidae) are a highly ecologically diverse and colorful family of beetles that use color cues for mate and host detection. In addition, there is evidence that buprestids have complex spectral sensitivity with up to five photoreceptor classes. Previous work suggested that opsin duplication and subfunctionalization of the two ancestral buprestid opsins, UV and LW, has expanded sensitivity to different regions of the light spectrum, but this has not yet been tested. We show that both duplications are likely unique to Buprestidae or the wider superfamily of Buprestoidea. To directly test photopigment sensitivity, we expressed buprestid opsins from two Chrysochroa species in Drosophila melanogaster and functionally characterized each photopigment type as UV- (356-357 nm), blue- (431-442 nm), green- (507-509 nm), and orange-sensitive (572-584 nm). As these novel opsin duplicates result in significantly shifted spectral sensitivities from the ancestral copies, we explored spectral tuning at four candidate sites using site-directed mutagenesis. This is the first study to directly test opsin spectral tuning mechanisms in the diverse and specious beetles.

Ultrafast protein response in the Pfr state of Cph1 phytochrome

Photoisomerization is a fundamental process in several classes of photoreceptors. Phytochromes sense red and far-red light in their Pr and Pfr states, respectively. Upon light absorption, these states react via individual photoreactions to the other state. Cph1 phytochrome shows a photoisomerization of its phycocyanobilin (PCB) chromophore in the Pfr state with a time constant of 0.7 ps. The dynamics of the PCB chromophore has been described, but whether or not the apoprotein exhibits an ultrafast response too, is not known. Here, we compare the photoreaction of 13C/15N labeled apoprotein with unlabeled apoprotein to unravel ultrafast apoprotein dynamics in Cph1. In the spectral range from 1750 to 1620 cm−1 we assigned several signals due to ultrafast apoprotein dynamics. A bleaching signal at 1724 cm−1 is tentatively assigned to deprotonation of a carboxylic acid, probably Asp207, and signals around 1670 cm−1 are assigned to amide I vibrations of the capping helix close to the chromophore. These signals remain after photoisomerization. The apoprotein dynamics appear upon photoexcitation or concomitant with chromophore isomerization. Thus, apoprotein dynamics occur prior to and after photoisomerization on an ultrafast time-scale. We discuss the origin of the ultrafast apoprotein response with the ‘Coulomb hammer’ mechanism, i.e. an impulsive change of electric field and Coulombic force around the chromophore upon excitation.Graphical abstract

A five-primary Maxwellian-view display for independent control of melanopsin, rhodopsin, and three-cone opsins on a fine spatial scale.

Independent spatiotemporal control of the stimulation of the five photoreceptor classes requires a display with as many primary lights to probe their isolated spatial and temporal responses. No such system exists with suitable performance properties. We present a system to construct a five-primary display from commercially available three-primary digital light processing projectors. It optimizes the set of five primary lights required to maximize the achievable contrast of a single photoreceptor class in a silent substitution protocol, including where the background chromaticity is first specified. From these chosen five primaries, we describe a method to convert complex three-primary (RGB) images to five-primary representations with complete specification of the photoreceptor excitations at each pixel. Key to enabling this multiple display system with a single HDMI connection is a novel control protocol implemented in a deterministic field-programmable gate array controller that splits the data into five video streams to allow nearly synchronous presentation of primary image data through multiple displays. Each pixel is controlled over 9.5 bits for each primary over a single frame for measurement of threshold-level vision. In addition to a large contrast gamut, the Maxwellian view offers high retinal illumination to support the investigation of five opsin-based responses to complex spatiotemporal images with a truly silent substitution protocol, while avoiding the confounding effects of uncontrolled photoreceptor excitations as occurs in four-primary systems. The customizable primaries facilitate this display translation to species with different photoreceptor spectral responses, and the optics are designed for integration into microscopes for use as a stimulus generator in physiological experiments.