Shift In Spectral Sensitivity Research Articles

Functional characterization of four opsins and two G alpha subtypes co-expressed in the molluscan rhabdomeric photoreceptor

BackgroundRhabdomeric photoreceptors of eyes in the terrestrial slug Limax are the typical invertebrate-type but unique in that three visual opsins (Gq-coupled rhodopsin, xenopsin, Opn5A) and one retinochrome, all belonging to different groups, are co-expressed. However, molecular properties including spectral sensitivity and G protein selectivity of any of them are not determined, which prevents us from understanding an advantage of multiplicity of opsin properties in a single rhabdomeric photoreceptor. To gain insight into the functional role of the co-expression of multiple opsin species in a photoreceptor, we investigated the molecular properties of the visual opsins in the present study.ResultsFirst, we found that the fourth member of visual opsins, Opn5B, is also co-expressed in the rhabdomere of the photoreceptor together with previously identified three opsins. The photoreceptors were also demonstrated to express Gq and Go alpha subunits. We then determined the spectral sensitivity of the four visual opsins using biochemical and spectroscopic methods. Gq-coupled rhodopsin and xenopsin exhibit maximum sensitivity at ~ 456 and 475 nm, respectively, and Opn5A and Opn5B exhibit maximum sensitivity at ~ 500 and 470 nm, respectively, with significant UV sensitivity. Notably, in vitro experiments revealed that Go alpha was activated by all four visual opsins, in contrast to the specific activation of Gq alpha by Gq-coupled rhodopsin, suggesting that the eye photoreceptor of Limax uses complex G protein signaling pathways.ConclusionsThe eye photoreceptor in Limax expresses as many as four different visual opsin species belonging to three distinct classes. The combination of opsins with different spectral sensitivities and G protein selectivities may underlie physiological properties of the ocular photoreception, such as a shift in spectral sensitivity between dark- and light-adapted states. This may be allowed by adjustment of the relative contribution of the four opsins without neural networks, enabling a simple strategy for fine-tuning of vision.

Enzymatic vitamin A2 production enables red-shifted optogenetics

Optogenetics is a technology using light-sensitive proteins to control signaling pathways and physiological processes in cells and organs and has been applied in neuroscience, cardiovascular sciences, and many other research fields. Most commonly used optogenetic actuators are sensitive to blue and green light, but red-light activation would allow better tissue penetration and less phototoxicity. Cyp27c1 is a recently deorphanized cytochrome P450 enzyme that converts vitamin A1 to vitamin A2, thereby red-shifting the spectral sensitivity of visual pigments and enabling near-infrared vision in some aquatic species.Here, we investigated the ability of Cyp27c1-generated vitamin A2 to induce a shift in spectral sensitivity of the light-gated ion channel Channelrhodopsin-2 (ChR2) and its red-shifted homolog ReaChR. We used patch clamp to measure photocurrents at specific wavelengths in HEK 293 cells expressing ChR2 or ReaChR. Vitamin A2 incubation red-shifted the wavelength for half-maximal currents (λ50%) by 6.8 nm for ChR2 and 12.4 nm for ReaChR. Overexpression of Cyp27c1 in HEK 293 cells showed mitochondrial localization, and HPLC analysis showed conversion of vitamin A1 to vitamin A2. Notably, the λ50% of ChR2 photocurrents was red-shifted by 10.5 nm, and normalized photocurrents at 550 nm were about twofold larger with Cyp27c1 expression. Similarly, Cyp27c1 shifted the λ50% of ReaChR photocurrents by 14.3 nm and increased normalized photocurrents at 650 nm almost threefold.Since vitamin A2 incubation is not a realistic option for in vivo applications and expression of Cyp27c1 leads to a greater red-shift in spectral sensitivity, we propose co-expression of this enzyme as a novel strategy for red-shifted optogenetics.

Transcriptomic evidence for visual adaptation during the aquatic to terrestrial metamorphosis in leopard frogs

BackgroundDifferences 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.ResultsWe 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.ConclusionsOverall, 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.

Evolutionary ecology of the visual opsin gene sequence and its expression in turbot (Scophthalmus maximus)

BackgroundAs flatfish, turbot undergo metamorphosis as part of their life cycle. In the larval stage, turbot live at the ocean surface, but after metamorphosis they move to deeper water and turn to benthic life. Thus, the light environment differs greatly between life stages. The visual system plays a great role in organic evolution, but reports of the relationship between the visual system and benthic life are rare. In this study, we reported the molecular and evolutionary analysis of opsin genes in turbot, and the heterochronic shifts in opsin expression during development.ResultsOur gene synteny analysis showed that subtype RH2C was not on the same gene cluster as the other four green-sensitive opsin genes (RH2) in turbot. It was translocated to chromosome 8 from chromosome 6. Based on branch-site test and spectral tuning sites analyses, E122Q and M207L substitutions in RH2C, which were found to be under positive selection, are closely related to the blue shift of optimum light sensitivities. And real-time PCR results indicated the dominant opsin gene shifted from red-sensitive (LWS) to RH2B1 during turbot development, which may lead to spectral sensitivity shifts to shorter wavelengths.ConclusionsThis is the first report that RH2C may be an important subtype of green opsin gene that was retained by turbot and possibly other flatfish species during evolution. Moreover, E122Q and M207L substitutions in RH2C may contribute to the survival of turbot in the bluish colored ocean. And heterochronic shifts in opsin expression may be an important strategy for turbot to adapt to benthic life.

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.

Spectral Diversification and Trans-Species Allelic Polymorphism during the Land-to-Sea Transition in Snakes

Snakes are descended from highly visual lizards [1] but have limited (probably dichromatic) color vision attributed to a dim-light lifestyle of early snakes [2-4]. The living species of front-fanged elapids, however, are ecologically very diverse, with ∼300 terrestrial species (cobras, taipans, etc.) and ∼60 fully marine sea snakes, plus eight independently marine, amphibious sea kraits [1]. Here, we investigate the evolution of spectral sensitivity in elapids by analyzing their opsin genes (which are responsible for sensitivity to UV and visible light), retinal photoreceptors, and ocular lenses. We found that sea snakes underwent rapid adaptive diversification of their visual pigments when compared with their terrestrial and amphibious relatives. The three opsins present in snakes (SWS1, LWS, and RH1) have evolved under positive selection in elapids, and in sea snakes they have undergone multiple shifts in spectral sensitivity toward the longer wavelengths that dominate below the sea surface. Several relatively distantly related Hydrophis sea snakes are polymorphic for shortwave sensitive visual pigment encoded by alleles of SWS1. This spectral site polymorphism is expected to confer expanded "UV-blue" spectral sensitivity and is estimated to have persisted twice as long as the predicted survival time for selectively neutral nuclear alleles. We suggest that this polymorphism is adaptively maintained across Hydrophis species via balancing selection, similarly to the LWS polymorphism that confers allelic trichromacy in some primates. Diving sea snakes thus appear to share parallel mechanisms of color vision diversification with fruit-eating primates.

Photoreceptors and diurnal variation in spectral sensitivity in the fiddler crab Gelasimus dampieri.

Colour signals, and the ability to detect them, are important for many animals and can be vital to their survival and fitness. Fiddler crabs use colour information to detect and recognise conspecifics, but their colour vision capabilities remain unclear. Many studies have attempted to measure their spectral sensitivity and identify contributing retinular cells, but the existing evidence is inconclusive. We used electroretinogram (ERG) measurements and intracellular recordings from retinular cells to estimate the spectral sensitivity of Gelasimus dampieri and to track diurnal changes in spectral sensitivity. G. dampieri has a broad spectral sensitivity and is most sensitive to wavelengths between 420 and 460 nm. Selective adaptation experiments uncovered an ultraviolet (UV) retinular cell with a peak sensitivity shorter than 360 nm. The species' spectral sensitivity above 400 nm is too broad to be fitted by a single visual pigment and using optical modelling, we provide evidence that at least two medium-wavelength sensitive (MWS) visual pigments are contained within a second blue-green sensitive retinular cell. We also found a ∼25 nm diurnal shift in spectral sensitivity towards longer wavelengths in the evening in both ERG and intracellular recordings. Whether the shift is caused by screening pigment migration or changes in opsin expression remains unclear, but the observation shows the diel dynamism of colour vision in this species. Together, these findings support the notion that G. dampieri possesses the minimum requirement for colour vision, with UV and blue/green receptors, and help to explain some of the inconsistent results of previous research.

A novel cellular structure in the retina of insectivorous birds

The keen visual systems of birds have been relatively well-studied. The foundations of avian vision rest on their cone and rod photoreceptors. Most birds use four cone photoreceptor types for color vision, a fifth cone for achromatic tasks, and a rod for dim-light vision. The cones, along with their oil droplets, and rods are conserved across birds – with the exception of a few shifts in spectral sensitivity – despite taxonomic, behavioral and ecological differences. Here, however, we describe a novel photoreceptor organelle in a group of New World flycatchers (Empidonax spp.) in which the traditional oil droplet is replaced with a complex of electron-dense megamitochondria surrounded by hundreds of small, orange oil droplets. The photoreceptors with this organelle were unevenly distributed across the retina, being present in the central region (including in the fovea), but absent from the retinal periphery and the area temporalis of these insectivorous birds. Of the many bird species with their photoreceptors characterized, only the two flycatchers described here (E. virescens and E. minimus) possess this unusual retinal structure. We discuss the potential functional significance of this unique sub-cellular structure, which might provide an additional visual channel for these small predatory songbirds.

Archelosaurian Color Vision, Parietal Eye Loss, and the Crocodylian Nocturnal Bottleneck.

Vertebrate color vision has evolved partly through the modification of five ancestral visual opsin proteins via gene duplication, loss, and shifts in spectral sensitivity. While many vertebrates, particularly mammals, birds, and fishes, have had their visual opsin repertoires studied in great detail, testudines (turtles) and crocodylians have largely been neglected. Here I examine the genomic basis for color vision in four species of turtles and four species of crocodylians, and demonstrate that while turtles appear to vary in their number of visual opsins, crocodylians experienced a reduction in their color discrimination capacity after their divergence from Aves. Based on the opsin sequences present in their genomes and previous measurements of crocodylian cones, I provide evidence that crocodylians have co-opted the rod opsin (RH1) for cone function. This suggests that some crocodylians might have reinvented trichromatic color vision in a novel way, analogous to several primate lineages. The loss of visual opsins in crocodylians paralleled the loss of various anatomical features associated with photoreception, attributed to a "nocturnal bottleneck" similar to that hypothesized for Mesozoic mammals. I further queried crocodylian genomes for nonvisual opsins and genes associated with protection from ultraviolet light, and found evidence for gene inactivation or loss for several of these genes. Two genes, encoding parietopsin and parapinopsin, were additionally inactivated in birds and turtles, likely co-occurring with the loss of the parietal eye in these lineages.

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

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

Rapid adaptive evolution of colour vision in the threespine stickleback radiation.

Vision is a sensory modality of fundamental importance for many animals, aiding in foraging, detection of predators and mate choice. Adaptation to local ambient light conditions is thought to be commonplace, and a match between spectral sensitivity and light spectrum is predicted. We use opsin gene expression to test for local adaptation and matching of spectral sensitivity in multiple independent lake populations of threespine stickleback populations derived since the last ice age from an ancestral marine form. We show that sensitivity across the visual spectrum is shifted repeatedly towards longer wavelengths in freshwater compared with the ancestral marine form. Laboratory rearing suggests that this shift is largely genetically based. Using a new metric, we found that the magnitude of shift in spectral sensitivity in each population corresponds strongly to the transition in the availability of different wavelengths of light between the marine and lake environments. We also found evidence of local adaptation by sympatric benthic and limnetic ecotypes to different light environments within lakes. Our findings indicate rapid parallel evolution of the visual system to altered light conditions. The changes have not, however, yielded a close matching of spectrum-wide sensitivity to wavelength availability, for reasons we discuss.

Functional characterization of spectral tuning mechanisms in the great bowerbird short-wavelength sensitive visual pigment (SWS1), and the origins of UV/violet vision in passerines and parrots

BackgroundOne of the most striking features of avian vision is the variation in spectral sensitivity of the short wavelength sensitive (SWS1) opsins, which can be divided into two sub-types: violet- and UV- sensitive (VS & UVS). In birds, UVS has been found in both passerines and parrots, groups that were recently shown to be sister orders. While all parrots are thought to be UVS, recent evidence suggests some passerine lineages may also be VS. The great bowerbird (Chlamydera nuchalis) is a passerine notable for its courtship behaviours in which males build and decorate elaborate bower structures.ResultsThe great bowerbird SWS1 sequence possesses an unusual residue combination at known spectral tuning sites that has not been previously investigated in mutagenesis experiments. In this study, the SWS1 opsin of C. nuchalis was expressed along with a series of spectral tuning mutants and ancestral passerine SWS1 pigments, allowing us to investigate spectral tuning mechanisms and explore the evolution of UV/violet sensitivity in early passerines and parrots. The expressed C. nuchalis SWS1 opsin was found to be a VS pigment, with a λmax of 403 nm. Bowerbird SWS1 mutants C86F, S90C, and C86S/S90C all shifted λmax into the UV, whereas C86S had no effect. Experimentally recreated ancestral passerine and parrot/passerine SWS1 pigments were both found to be VS, indicating that UV sensitivity evolved independently in passerines and parrots from a VS ancestor.ConclusionsOur mutagenesis studies indicate that spectral tuning in C. nuchalis is mediated by mechanisms similar to those of other birds. Interestingly, our ancestral sequence reconstructions of SWS1 in landbird evolution suggest multiple transitions from VS to UVS, but no instances of the reverse. Our results not only provide a more precise prediction of where these spectral sensitivity shifts occurred, but also confirm the hypothesis that birds are an unusual exception among vertebrates where some descendants re-evolved UVS from a violet type ancestor. The re-evolution of UVS from a VS type pigment has not previously been predicted elsewhere in the vertebrate phylogeny.

Amorphous Photodiode with an Intermediate Contact for Improved Color Separation

ABSTRACTMost of actual photonic devices being sensitive in the visible (VIS) spectrum are based on crystalline silicon (x-Si). The production of x-Si requires an expensive high temperature process. The color reproduction with x-Si diodes additionally requires an integration of color filters [1] to realize a shift in spectral sensitivity. This work presents an amorphous silicon (a-Si:H) photodiode with an intermediate contact for color separation without color filters. Such detectors can be produced in a low cost and low temperature PECVD process, which allows their direct deposition on a custom specific ASIC [2]. Another advantage of a-Si:H is the up to 10 times higher light absorption compared to that of x-Si in the VIS spectrum [3]. The device consists of a metal (Cr) cathode, an amorphous NIP diode structure and a TCO (Al doped ZnO) anode. The I-layer includes an interior TCO contact buried between two P-layers. The thickness of this TCO layer is about 200 nm; the P-layers have a thickness of about 10 nm. The chromium cathode is sputtered on a glass substrate in a PVD process. The amorphous layers are deposited in a multi-chamber PECVD line; the buried and top TCO contacts are sputtered in the same line continuously under high-vacuum conditions. In a first photolithography step the top anode is patterned while the buried anode is uncovered. Afterwards, the diode must be patterned again, resulting in a final Cr, NIP-a-Si:H, TCO, PIP-a-Si:H and TCO multi-layer stack.The spectral sensitivity of a common NIP diode can be shifted by external bias voltages. The spectral sensitivity at higher negative voltages overlays those of that at lower voltages and additionally shifts to longer wavelengths. The color reproduction is difficult; it can be improved by reducing the overlap of the spectral sensitivity [4]. The spectral response of the diodes presented in this work also can be shifted by the bias voltage. Furthermore, it can be split by substituting the disclosed anodes. The spectral response, using the cathode and top anode has a maximum at short wavelengths. If the diode between the interior anode and the cathode is used, the spectral sensitivity for longer wavelengths increases. Shorter wavelengths are blocked by the top part of the diode; it works like a filter. The presented device structure offers good prospects to improve color separation compared to currently existing detectors by using an additional intermediate contact.

Photometry in the dark: time dependent visibility of low intensity light sources

This paper aims at describing the perceived brightness of persistent luminescent materials for emergency signage. In case of emergency, typically, a fully light adapted person is left in the dark, except for the emergency sign. The available photometric models cannot describe visibility of such light source, as they do not consider the slow dark adaptation of the human eye. The model proposed here fully takes into account the shift from photopic to scotopic vision, the related shift in spectral sensitivity and the dark adaptation. The resulting metric is a 'visibility index' and preliminary tests show that it more realistically describes the perceived brightness of persistent luminescent materials than the common photometric standards.

The Green-absorbing Drosophila Rh6 Visual Pigment Contains a Blue-shifting Amino Acid Substitution That Is Conserved in Vertebrates

The molecular mechanisms that regulate invertebrate visual pigment absorption are poorly understood. Through sequence analysis and functional investigation of vertebrate visual pigments, numerous amino acid substitutions important for this adaptive process have been identified. Here we describe a serine/alanine (S/A) substitution in long wavelength-absorbing Drosophila visual pigments that occurs at a site corresponding to Ala-292 in bovine rhodopsin. This S/A substitution accounts for a 10-17-nm absorption shift in visual pigments of this class. Additionally, we demonstrate that substitution of a cysteine at the same site, as occurs in the blue-absorbing Rh5 pigment, accounts for a 4-nm shift. Substitutions at this site are the first spectrally significant amino acid changes to be identified for invertebrate pigments sensitive to visible light and are the first evidence of a conserved tuning mechanism in vertebrate and invertebrate pigments of this class.

Seeing red: behavioral evidence of trichromatic color vision in strepsirrhine primates

Among primates, catarrhines (Old World monkeys and apes) and certain platyrrhines (New World monkeys) possess trichromatic color vision, which might confer important evolutionary advantages, particularly during foraging. Recently, a polymorphism has been shown to shift the spectral sensitivity of the X-linked opsin protein in certain strepsirrhines (e.g., Malagasy lemurs); however, its behavioral significance remains unknown. We assign genotypes at the X-linked variant to 45 lemurs, representing 4 species, and test if the genetic capacity for trichromacy impacts foraging performance, particularly under green camouflage conditions in which red detection can be advantageous. We confirm polymorphism at the critical site in sifakas and ruffed lemurs and fail to find this polymorphism in collared lemurs and ring-tailed lemurs. We show that this polymorphism may be linked to ‘‘behavioral trichromacy’’ in heterozygous ruffed lemurs but find no comparable evidence in a single heterozygous sifaka. Despite their putative dichromatic vision, female collared lemurs were surprisingly efficient at retrieving both red and green food items under camouflage conditions. Thus, species-specific feeding ecologies may be as important as trichromacy in influencing foraging behavior. Although the lemur opsin polymorphism produced measurable behavioral effects in at least one species, the ruffed lemur, these effects were modest, consistent with the modest shift in spectral sensitivity. Additionally, the magnitude of these effects varied across individuals of the same genotype, emphasizing the need for combined genetic and behavioral studies of trichromatic vision. We conclude that trichromacy may be only one of several routes toward increased foraging efficiency in visually complex environments. Key words: dichromacy, feeding ecology, lemurs, opsin gene polymorphism, trichromacy. [Behav Ecol]

Paradoxical shifts in human color sensitivity caused by constructive and destructive interference between signals from the same cone class

Paradoxical shifts in human color (spectral) sensitivity occur on deep-red (658 nm) background fields. As the radiance of the deep-red background is increased from low to moderate levels, the spectral sensitivity for detecting 15-Hz flicker shifts toward shorter wavelengths, although by more than is predicted by selective chromatic adaptation (e.g., Eisner & MacLeod, 1981; Stromeyer et al., 1987; Stockman et al., 1993). Remarkably, though, at higher background radiances, the spectral sensitivity then shifts precipitously back towards longer wavelengths. Here, we show that both effects are due in large part to destructive and constructive interference between signals generated by the same cone type. Contrary to the conventional model of the human visual system, the M- and L-cone types contribute not just the customary fast signals to the achromatic or luminance pathway, but also slower signals of the same or opposite sign. The predominant signs of the slow M- and L-cone signals change with background radiance, but always remain spectrally opposed (M-L or L-M). Consequently, when the slow and fast signals from one cone type destructively interfere, as they do near 15 Hz, those from the other cone type constructively interfere, causing the paradoxical shifts in spectral sensitivity. The shift in spectral sensitivity towards longer wavelengths is accentuated at higher temporal frequencies by a suppression of fast M-cone signals by deep-red fields.

Large area single and stacked p-i-n photodiodes as a color image sensors

AbstractSingle and stacked p-i-n sensing elements for image recognition and color extraction applications are presented. The aim of this work is to optimize the performance of the a-SiC:H thin films layers in order to enhance its performance when making part of the structure of large area image and color sensors. The efforts are focused mainly on doped n- and p-type layers at high and low doping levels with and without carbon. The structural and optoelectronic properties of the single layers were determined through infrared and visible spectroscopy, temperature-dependent conductivity, and were complemented by CPM measurements. Junction properties, carrier transport, photogeneration and collection efficiency are investigated from dark and illuminated current-voltage characteristics and spectral response measurements, with and without additional background illumination and under different light bias conditions. The spectral response dependence on the applied voltage and on optical bias was also studied. Results show that the spectral sensitivity is strongly dependent on the applied voltage, namely the maximum spectral sensitivity shifts with the voltage, and at certain wavelengths the spectral response goes down to zero, which allows a different selectivity, and enables color recognition.

Spectral sensitivity of larval and juvenile coral reef fishes: implications for feeding in a variable light environment

The spectral sensitivity of larval and juvenile stages of 3 species of coral reef fishes, Apogon compressus (Apogonidae), Pomacentrus amboinensis (Pomacentridae) and Premnas biaculeatus (Pomacentridae) has been investigated using feeding behaviour. Ontogenetic and taxonomic differences in spectral sensitivity were determined by establishing the minimum light intensity at which larvae and juveniles could strike prey at 12 restricted wavelength bands between 355 and 650 nm. Following construction of chromatic action spectra, the wavelength of maximum sensitivity (λmax) and the median wavelength (λP50) of the 3 species were found to be located close to 500 nm. All 3 species increased in sensitivity during growth, with A. compressus becoming the most sensitive prior to settlement. Ontogenetic shifts in spectral sensitivity towards longer wavelengths occurred in P. amboinensis and P. biaculeatus, but not in A. compressus. Spectral efficiency (wavelength-dependent efficiency of photon capture) was modelled for eutrophic and oligotrophic coral reef waters (Jerlov types Oceanic IA, Oceanic III and Coastal 1) at 2 different optical depths. Spectral efficiency was highest in the intermediate coral reef water type (JOIII) in all 3 larval fish taxa throughout early ontogeny, regardless of water depth. The results imply that the larvae would be able to feed across a broad spectrum of coral reef water types and depths.

The operational principle of a new amorphous silicon based p-i-i-n color detector

The operational principle of a new type p-i-i-n color sensor is described with the aid of numerical modeling. The modeling results account for the color detection mechanism recently presented that this kind of structure exhibits [T. Neidlinger, M. B. Schubert, G. Schmid, and H. Brummack, in Amorphous Silicon Technology—1996, edited by E. A. Schiff et al. (Materials Research Society, Pittsburgh, 1996), p. 147]. By band gap engineering the experimental red response is maximized at larger reverse bias voltage whereas the green response has its maximum at low reverse bias voltage. The numerical modeling qualitatively reproduces the characteristic shape of the steady-state current-voltage curves at different illumination wavelengths. At low and at high reverse bias voltages the influence of the internal variables and parameters is identified and leads to the experimentally observed response. The potential profile of the p-i-i-n structure is of crucial importance to the color detection mechanism. At larger wavelengths the large potential drop across the two highly defective front layers assists recombination in the back part of the device, which thus leads to the drop in the red response at low reverse voltage. For the voltage-dependent shift in spectral sensitivity it is important that photogenerated carriers under green bias illumination are lost by recombination in the front part of the device.