The photoactivation energy of the visual pigment in two spectrally different populations of <Emphasis Type="Italic">Mysis relicta</Emphasis> (Crustacea,Mysida)

We report the first study of the relation between the wavelength of maximum absorbance (λ_max) and the photoactivation energy (E _a) in invertebrate visual pigments. Two populations of the opossum shrimp Mysis relicta were compared. The two have been separated for 9,000 years and have adapted to different spectral environments (“Sea” and “Lake”) with porphyropsins peaking at λ_max=529 nm and 554 nm, respectively. The estimation of E _a was based on measurement of temperature effects on the spectral sensitivity of the eye. In accordance with theory (Stiles in Transactions of the optical convention of the worshipful company of spectacle makers. Spectacle Makers’ Co., London, 1948), relative sensitivity to long wavelengths increased with rising temperature. The estimates calculated from this effect are E _a,529=47.8±1.8 kcal/mol and E _a,554=41.5±0.7 kcal/mol (different at P<0.01). Thus the red-shift of λ_max in the “Lake” population, correlating with the long-wavelength dominated light environment, is achieved by changes in the opsin that decrease the energy gap between the ground state and the first excited state of the chromophore. We propose that this will carry a cost in terms of increased thermal noise, and that evolutionary adaptation of the visual pigment to the light environment is directed towards maximizing the signal-to-noise ratio rather than the quantum catch.     

Spectral sensitivity differences in two Mysis sibling species (Crustacea, Mysida): Adaptation or phylogenetic constraints?

The variation in eye spectral sensitivities of the closely related mysid species Mysis relicta Lovén, 1862 and Mysis salemaai Audzijonyt? and Väinölä, 2005 was studied in sympatric and allopatric populations from the brackish Baltic Sea and from two lakes representing different light environments. In the Baltic Sea the maximum spectral sensitivity of M. relicta, measured by the electroretinogram (ERG) technique, was shifted by ca 20 nm to longer wavelengths than in M. salemaai (564 and 545 nm, respectively). The spectral sensitivity of M. salemaai was closer to that of marine mysid species, which is consistent with its broader euryhalinity and the presumed longer brackish-water history. The species-specific sensitivities in the Baltic Sea were not affected by regional differences in light environments. In two lake populations of M. relicta, the spectral sensitivity was further shifted by ca 28 nm towards the longer wavelengths compared with the conspecific Baltic Sea populations. The spectral sensitivities in the four M. relicta populations were not correlated to the current light conditions, but rather to the phylogeographic histories and fresh- vs. brackish-water environments. A framework to further explore factors affecting spectral sensitivities in Mysis is suggested.     

Visual pigments of marine carnivores: pinnipeds, polar bear, and sea otter

Rod and cone visual pigments of 11 marine carnivores were evaluated. Rod, middle/long-wavelength sensitive (M/L) cone, and short-wavelength sensitive (S) cone opsin (if present) sequences were obtained from retinal mRNA. Spectral sensitivity was inferred through evaluation of known spectral tuning residues. The rod pigments of all but one of the pinnipeds were similar to those of the sea otter, polar bear, and most other terrestrial carnivores with spectral peak sensitivities (λ_max) of 499 or 501 nm. Similarly, the M/L cone pigments of the pinnipeds, polar bear, and otter had inferred λ_max of 545 to 560 nm. Only the rod opsin sequence of the elephant seal had sensitivity characteristic of adaptation for vision in the marine environment, with an inferred λ_max of 487 nm. No evidence of S cones was found for any of the pinnipeds. The polar bear and otter had S cones with inferred λ_max of ∼440 nm. Flicker-photometric ERG was additionally used to examine the in situ sensitivities of three species of pinniped. Despite the use of conditions previously shown to evoke cone responses in other mammals, no cone responses could be elicited from any of these pinnipeds. Rod photoreceptor responses for all three species were as predicted by the genetic data.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Interspecific variation in the visual pigments of deep-sea fishes

Summary Visual pigments in the rods of 38 species of deep-sea fish were examined by microspectrophotometry. 33 species were found to have a single rhodopsin with a wavelength of maximum absorbance ( _max) in the range 470–495 nm. Such visual pigments have absorbance maxima close to the wavelengths of maximum spectral transmission of oceanic water. 5 species, however, did not conform to this pattern and visual pigments were found with _max values ranging from 451 nm to 539 nm. In 4 of these species two visual pigments were found located in two types of rod. Some 2-pigment species which have unusual red sensitivity, also have red-emitting photophores. These species have both rhodopsin and porphyropsin pigments in their retinae, which was confirmed by HPLC, and the two pigments are apparently located in separate rods in the same retinal area. In deep-sea fishes the occurrence of unusual visual pigments seems to be correlated with aspects of the species' depth ranges. In addition to ecological influences we present evidence, in the form of _max spectral clustering, that indicates the degree of molecular constraint imposed on the evolution of visual pigments in the deep-sea.     

On the three visual pigments in the retina of the firefly squid,Watasenia scintillans

Summary The deep-sea bioluminescent squid, Watasenia scintillans, has three visual pigments: The major one (A1 pigment) is based on retinal and has _max = 484 nm, the second one (A2 pigment) is based on 3-dehydroretinal and has _max = 500 nm, and the third one (A4 pigment) is based on 4-hydroxyretinal and has _max = 470 nm. The distribution of these 3 visual pigments in the retina was studied by HPLC analysis of the retinals in retina slices obtained by microdissection. It was found that A1 pigment was not located in the specific region of the ventral retina receiving the down-welling light which contains very long photoreceptor cells, forming two strata. A2 and A4 pigment were found exclusively in the proximal pinkish stratum and in the distal yellowish stratum. The role of these pigments in the retina is hypothesized to involve spectral discrimination. The extraction and analysis of retinoids to determine the origin of 3-dehydroretinal and 4-hydroxyretinal in the mature squid showed only a trace amount of 4-hydroxyretinol in the eggs. Similar analysis of other cephalopods collected near Japan showed the absence of A2 or A4 pigment in their eyes.Abbreviations HPLC high-performance liquid chromatography - IS inner segment - OS outer segment     

A charge transfer process in the visual pigments

A physical model that incorporates all the experimental information on the formation of the visual pigment rhodopsin is presented. The visual pigments consist of a chromophore bound to an appropriate protein. Thus rhodopsin (λ_m 505 mμ) is formed by a Schiff’s base linkage C₁₉H₂₇CH=NH⁺-opsin (λ_m 440 mμ) between 11-cis retinal (λ_m 380 mμ) and the protein opsin (λ_m 280 mμ). It is found that there exists a red shift in the spectrum of rhodopsin from the Schiff’s base. The model brings an explanation for this red shift. It is shown that such a shift may be due to a charge transfer process (R. S. Mulliken,J. Am. Chem. Soc.,74, 811–824, 1952) between an electron at the double bond of carbons C₁₁−C₁₂ and an atomic orbital of the sulphur present in cysteine. This provides an explanation of the presence of SH-groups in the protein after the absorption of light. A one-electron approximation is used and the dipole momentμ _NV ; hence, the oscillator strengthf of the transitionNV is estimated and compared with the experimentally determined extinction coefficient ∈_m by mixing 3.5×10⁻³ M of 11-cis retinal with 8.3×10⁻⁵ M of cysteine at pH ranges 6 through 8. Reasonable agreement is found. Solvent, concentration and temperature dependence are shown also.     

Visual pigments and spectral sensitivity of the diurnal gecko Gonatodes albogularis

The visual pigments and oil droplets in the retina of the diurnal gecko Gonatodes albogularis were examined microspectrophotometrically, and the spectral sensitivity under various adapting conditions was recorded using electrophysiological responses. Three classes of visual pigments were identified, with _max at about 542, 475, and 362 nm. Spectral sensitivity functions revealed a broad range of sensitivity, with a peak at approximately 530–540 nm. The cornea and oil droplets were found to be transparent across a range from 350–700 nm, but the lens absorbed short wavelength light below 450 nm. Despite the filtering effect of the lens, a secondary peak in spectral sensitivity to ultraviolet wavelengths was found. These results suggest that G. albogularis does possess the visual mechanisms for discrimination of the color pattern of conspecifics based on either hue or brightness. These findings are discussed in terms of the variation in coloration and social behavior of Gonatodes.Abbreviations ERG electroretinogram - MSP microspectrophotometry - UV ultraviolet - _max wavelength of maximum absorbance     

Comparative study of spectral and antioxidant properties of pigments from the eyes of two Mysis relicta (Crustacea, Mysidacea) populations, with different light damage resistance

Retinal visual and screening pigments of two populations (one marine and the other freshwater) of the opossum shrimp Mysis relicta Lovén (Crustacea, Mysidacea), which have different ocular tolerance to light, was investigated. Visual pigments were extracted by detergent and their bleaching difference spectra were determined. The difference between the visual pigment absorption maximum of the two populations correlated with their difference in spectral sensitivity. Using buffer or neutral methanol, a yellow pigment was extracted which had absorption maxima at 440 nm and 325 nm and bright blue fluorescence (λ_max 415 nm). A screening pigment (ommochrome) with maximum at 525 nm was extracted by acid methanol, and was probably related to the group of ommines. The eyes of the lake population had 1.8–2.7 times less of this pigment than the eyes of the sea population. The sea population is more resistant to photo-induced accumulation of thiobarbituric acid-reactive substances in eye tissues. This resistance may be due to the higher ommochrome content. Accepted: 8 December 1998     

Photoregeneration of visual pigments in a moth

Summary The spectral absorbance by the visual pigments in the compound eye of the mothDeilephila elpenor was determined by microphotometry. Two visual pigments and their photoproducts were demonstrated. The photoproducts are thermostable and are reconverted to the visual pigments by light. The concentrations of the visual pigments and the photoproducts at each wavelength are determined by their absorbance coefficients at this wavelength. P 525: The experimental recordings (difference spectra and spectral absorbance changes after exposure to monochromatic lights) were completely reproduced by calculations using nomograms for vertebrate rhodopsin. The identity between experimental recordings and calculations show: One visual pigment absorbs maximally at 525 nm (P 525). The resonance spectrum of the visual pigment is identical to that for a vertebrate rhodopsin (_max at 525 nm). The photoproduct of this pigment absorbs maximally at 480 nm (M 480). It is similar to the acid metarhodopsin in cephalopods. The relative absorbance of P 525 to that of M 480 is 11.75. The quantum efficiency for photoconversion of P 525 to M 480 is nearly equal to that for reconversion of M 480 to P 525. Wavelengths exceeding about 570 nm are absorbed only by P 525, i. e. P 525 is completely converted to M 480. Shorter wavelengths are absorbed both by P 525 and M 480. At these wavelengths a photoequilibrium between the two pigments is formed. Maximal concentration of P 525 is obtained at about 450 nm. P 350: A second visual pigment absorbs maximally at about 350 nm (P 350), and its photoproduct at 450 to 460 nm. In the region of spectral overlap a photoequilibrium between the two pigments is formed.The visual pigment and the photoproduct are similar to those in the neuropteran insectAscalaphus.The work reported in this article was supported by Deutsche Forschungsgemeinschaft, Schwerpunktsprogramm Rezeptorphysiologie Ha 258-10, and SFB 114, by the Swedish Medical Research Council (grant no B 73-04X-104-02B), by Karolinska Institutet, and by a grant (to G. Höglund) from Deutscher Akademischer Austauschdienst.     

Temporal shifts in visual pigment absorbance in the retina of Pacific salmon

The visual pigments and photoreceptor types in the retinas of three species of Pacific salmon (coho, chum, and chinook) were examined using microspectrophotometry and histological sections for light microscopy. All three species had four cone visual pigments with maximum absorbance in the UV (_max: 357–382 nm), blue (_max: 431–446 nm), green (_max: 490–553 nm) and red (_max: 548–607 nm) parts of the spectrum, and a rod visual pigment with _max: 504–531 nm. The youngest fish (yolk-sac alevins) did not have blue visual pigment, but only UV pigment in the single cones. Older juveniles (smolts) had predominantly single cones with blue visual pigment. Coho and chinook smolts (>1 year old) switched from a vitamin A₁- to a vitamin A₂-dominated retina during the spring, while the retina of chum smolts and that of the younger alevin-to-parr coho did not. Adult spawners caught during the Fall had vitamin A₂-dominated retinas. The central retina of all species had three types of double cones (large, medium and small). The small double cones were situated toward the ventral retina and had lower red visual pigment _max than that of medium and large double cones, which were found more dorsally. Temperature affected visual pigment _max during smoltification.     

Spectral sensitivity of visual cells of the compound eye ofBlatta orientalis

Responses of single visual cells of the anterior part of the compound eye of the oriental cockroachBlatta orientalis were recorded intracellularly. Two spectral types of cells were discovered: ultraviolet receptors with _max 361 nm and green-sensitive receptors with _max 503 nm. The spectral curve of the whole eye, measured by the electroretinogram, included two peaks (=350–370 and 500 nm) and a minimum between 400 and 430 nm. This last fact is interpreted as additional evidence of the dichromatic vision of the cockroach.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 57–61, January–February, 1985.     

A new pathway in the photoreaction cycle of trans-bacteriorhodopsin and the absorption spectra of its intermediates

The intermediates of trans-bacteriorhodopsin (trans-bR) in the photoreaction cycle were investigated under two different conditions. In a low salt and neutral pH medium (10 mM phosphate buffer, pH 6.6), trans-bR was irradiated with 500 nm light at –190 C, resulting in formation of batho-trans-bR (batho-bR^t). On warming in the dark, batho-bR^t converted to lumi-trans-bR (lumi-bR^t), meta-trans-bR (meta-bR^t) and finally to trans-bR. The intermediates N and O, which had been detected by others by flash photolysis, were not observed. The thermal decay of lumi-bR^t in a high salt and high pH medium (10 mM borate buffer with l M NaCl, pH 10.0) proceeded simultaneously through two pathways; one to meta-bR^t and another to trans-bR. About 72% of lumi-bR^t converted to trans-bR directly and the residue converted to meta-bR^t. By use of this value, the absorption spectra of batho-bR^t (_max: 626 nm), lumi-bR^t (_max: 543 nm) and meta-bR^t (_max: 418 nm) were calculated. A photoreaction cycle of bacteriorhodopsin was proposed on the basis of the above findings.     

Microspectrophotometric characterization and localization of three visual pigments in the compound eye ofNotonecta glauca L. (Heteroptera)

Summary The absorption maxima ( _max) of the visual pigments in the ommatidia ofNotonecta glauca were found by measuring the difference spectra of single rhabdomeres after alternating illumination with two different adaptation wavelengths. All the peripheral rhabdomeres contain a pigment with an extinction maximum at 560 nm. This pigment is sensitive to red light up to wavelengths > 700 nm. In a given ommatidium in the dorsal region of the eye, the two central rhabdomeres both contain one of two pigments, either a pigment with an absorption maximum in the UV, at 345 nm, or — in neighboring rhabdoms — a pigment with an absorption maximum at 445 nm. In the ventral part of the eye only the pigment absorbing maximally in the UV was found in the central rhabdomeres. The spectral absorption properties of various types of screening-pigment granules were measured.     

Pityrialactone- a new fluorochrome from the tryptophan metabolism of M. alassezia furfur

As the main nitrogen source in Malassezia (M.) furfur, tryptophan induces the formation of fluorochromes and pigments, which makes the yeast less sensitive towards UV light. For the investigation of the fluorochromes, M. furfur (CBS1878) was incubated at 32 °C for 14 days on a pigment-inducing medium, and the agar extract was purified by column chromatography, preparative TLC and HPLC. The structures of the pure metabolites were determined by mass spectrometry and NMR spectroscopy. A pale yellow compound eluting from the column with 22% acetonitrile was found to exhibit a strong green-yellow fluorescence. The fluorochrome is a new bisindolyl compound (C₂₀H₁₂N₂O₃, MW 328.33) named pityrialactone because of its furan-2,3-dione structure. The UV protective properties (λ_max 352, 292, 276, 224 nm) of this metabolite were confirmed in a yeast model. As shown by the fluorescence spectrum, pityrialactone appears to be responsible for the green-yellow fluorescence of pityriasis versicolor lesions under Wood light. Pityrialactone is accompanied by the isomeric bisindolylmaleic anhydride (pityriaanhydride), which has not yet been described as a natural product but is a known intermediate in the total synthesis of bisindolylmaleimides. This revised version was published online in August 2006 with corrections to the Cover Date.     

Non-enzymatic glycosylation induced changesin vitro in some molecular parameters of collagen

Non-enzymatic glycosylation of rat tail tendon collagen was examined by incubation with D-glucosein vitro. The changes in molecular parameters such as viscosity, thermal stability, electrophoretic mobility and solubility were determined on nonenzymatically glycosylated collagenin vitro. Tendons incubated with 8 and 24 mg glucose/ml showed an increase in dissolution temperature and a l.6-3-fold increase in thermal isometric tension respectively when compared to tendons incubated in the absence of glucose, indicating the formation of new intermolecular bonds. This conclusion was further supported by the decreased solubility of glycosylated collagen in 0.5 N acetic acid and the change in sub-unit composition as measured from the sodium dodecyl sulphate Polyacrylamide gel electrophoresis pattern. Glycosylated collagen gave a characteristic absorption spectra λ_max 248 nm) as distinct from that of control (λ_max 242 nm). Denaturation temperature of glycosylated collagen, as determined from temperature dependent viscosity measurements, was reduced. These studies indicate that glycosylation affects the molecular interactions as well as the crosslinking of collagen.     

Light-inducedtrans-cis isomerization of retinal by a protein from honeybee retina

Summary Two retinal-binding proteins (RBP-A and RBP-B) isolated from the honeybee retina were further purified by ion-exchange chromatography. Whereas RBP-A seems to be denatured by this procedure, RBP-B remains intact with respect to its photochemical characteristics (Fig. 3a). Analysis of the geometric isomers of retinal bound to RBP-B by high performance liquid chromatography demonstrated that all-trans retinal was the chromophore of the non-irradiated RBP-B. Irradiation converted RBP-B (_max 440 nm) into a photoproduct (_max 370 nm) the chromophore of which was 11-cis retinal, i.e., light isomerized all-trans retinal almost exclusively to the 11-cis form (Fig. 3b). Irradiation of a solution of RBP-B in the presence of excess all-trans retinal also led to the formation of 11-cis retinal indicating that RBP catalyzes the photoisomerization of all-trans retinal. The physiological significance of RBP-B is discussed with respect to the renewal of rhodopsin.Abbreviations RBP retinal-binding protein - HPLC high performance liquid chromatography     

Physiological characterization of the compound eye in monarch butterflies with focus on the dorsal rim area

The spectral, angular and polarization sensitivities of photoreceptors in the compound eye of the monarch butterfly (Danaus plexippus) are examined using electrophysiological methods. Intracellular recordings reveal a spectrally homogenous population of UV receptors with optical axes directed upwards and ≥10° to the contralateral side. Based on optical considerations and on the opsin expression pattern (Sauman et al. 2005), we conclude that these UV receptors belong to the anatomically specialized dorsal rim area (DRA) of the eye. Photoreceptors in the main retina with optical axes <10° contralateral or ipsilateral have maximal sensitivities in the UV (λ_max≤340 nm), the blue (λ_max=435 nm) or in the long-wave range (green, λ_max=540 nm). The polarization sensitivity (PS) of the UV receptors in the DRA is much higher (PS=9.4) than in the UV cells (PS=2.9) or green cells (PS=2.8) of the main retina. The physiological properties of the photoreceptors in the DRA and in the main retina fit closely with the anatomy and the opsin expression patterns described in these eye regions. The data are discussed in the light of present knowledge about polarized skylight navigation in Lepidopterans.     

Visual pigments in the individual rods of deep-sea fishes

Summary The visual pigments in the rods of 15 species of deep-sea fish were examined by microspectrophotometry. In 13 species a single visual pigment was found. The max of these pigments, which ranged from 475 nm to 488 nm, suggest they give the fish maximum sensitivity to the ambient light in the deep, blue ocean waters where they live. In two species two visual pigments were found in separate rods.Bathylagus bericoides had rhodopsins of max 466 nm and 500 nm andMalacocephalus laevis had two rhodopsins of max 478 nm and 485 nm. It is noted that the species with two visual pigments tend to be dark in colour and live in deeper, darker, water.