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     

Visual pigment absorbance and spectral sensitivity of the <Emphasis Type="Italic">Mysis relicta</Emphasis> species group (Crustacea,Mysida) in different light environments

Visual-pigment absorbance spectra and eye spectral sensitivities were examined in eight populations of opossum shrimp from different light environments. Four Finnish populations, two from the Baltic Sea and two from freshwater lakes, represent Mysis relicta, sensu stricto. The sibling species M. salemaai and M. diluviana are represented by, respectively, two Baltic Sea populations and two populations from freshwater lakes in Idaho, USA. In M. relicta, the visual pigments of the two lake populations were similar (λ_max=554.3±0.8 nm and 556.4±0.4 nm), but significantly red-shifted compared with the sea populations (at 529 and 535 nm) and with M. salemaai (at 521 and 525 nm). All these pigments had only A2 chromophore and the lake/sea difference indicates adaptive evolution of the opsin. In M. diluviana, λ_max varied in the range 505–529 nm and the shapes of spectra suggested varying A1/A2 chromophore proportions, with pure A1 in the 505 nm animals. Eye sensitivity spectra were flatter and peaked at longer wavelengths than the relevant visual-pigment templates, but declined with the same slope beyond ca. 700 nm. The deviations from visual-pigment spectra can be explained by ocular light filters based on three types of identified screening pigments.     

Properties of pupil mechanisms in owl-fly Ascalaphus macaronius (Neuroptera)

Regulation of light flux by pupil mechanisms in the UV-sensitive superposition eye of owl-fly Ascalaphus macaronius (Neuroptera) was studied with a fast reflection microspectrophotometric technique. The spectral sensitivity of pupil reaction, which was calculated on the basis of changes of transient amplitude reflection, was almost identical with the one of Deilephila eye. This indicates that in spite of different life styles and spectral sensitivities of photoreceptors, pupil closing is triggered by the same photosensitive structure in both eyes. By measuring the spectra of reflected light from the Ascalaphus eye between 400 and 700 nm after different dark periods following light stimulation, it was established that the restoration of reflection was much faster in the red than in the blue spectral range. Based on this, we propose that two different pupil mechanisms with different spectral absorption characteristics are involved in light-flux regulation. Fast-reacting pupil is probably represented by screening pigment migration in the secondary pigment cells and a slow blue-absorbing system by the activity in primary pigment cells. The importance of two different pupils for the photoregeneration of visual pigment is discussed. Accepted: 1 October 1998     

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.     

Tuning of photoreceptor spectral sensitivity in fireflies (Coleoptera: Lampyridae)

Sexual communication between male and female fireflies involves the visual detection of species-specific bioluminescent signals. Firefly species vary spectrally in both their emitted light and in the sensitivity of the eye, depending on the time when each is active. Tuning of spectral sensitivity in three firefly species that occupy different photic niches was investigated using light and electron microscopy, microspectrophotometry, and intracellular recording to characterize the location and spectral absorption of the screening pigments that filter incoming light, the visual pigments that receive this filtered light, and the visual spectral sensitivity. Twilight-active species had similar pink screening pigments, but the visual pigment of Photinus pyralis peaked near 545 nm, while that of P. scintillans had a λ_max near 557 nm. The night-active Photuris versicolor had a yellow screening pigment that was uniquely localized, while its visual pigment was similar to that of P. pyralis. These results show that both screening and visual pigments vary among species. Modeling of spectral tuning indicates that the combination of screening and visual pigments found in the retina of each species provides the best possible match of sensitivity to bioluminescent emission. This combination also produced model sensitivity spectra that closely resemble sensitivities measured either with electroretinographic or intracellular techniques. Vision in both species of Photinus appears to be evolutionarily tuned for maximum discrimination of conspecific signals from spectrally broader backgrounds. Ph. versicolor, on the other hand, appears to have a visual system that offers a compromise between maximum sensitivity to, and maximum discrimination of, their signals. Accepted: 29 September 1999     

Colour discrimination in dim light: An analysis of the photoreceptor arrangement in the mothDeilephila

Summary Visual pigment absorption and spectral sensitivity are calculated for a model rhabdom based on theDeilephila rhabdom. The effect of different sky light intensity spectra on absorption and spectral sensitivity is examined, and the importance of the receptor arrangement for colour vision discussed. The quality of colour perception which can be expected for such an eye is estimated. The calculations reveal, firstly, a balance between the spectral bandwidths of rhodopsin absorption spectra and the distances between their maxima, which is of great significance with respect to colour vision. Secondly, they show that the quality of colour discrimination for dim light, at luminance levels between 0.1 and 10 cd/m², is comparable to the performance of the human eye at much higher levels of luminance.     

Colour in the eyes of insects

Many insect species have darkly coloured eyes, but distinct colours or patterns are frequently featured. A number of exemplary cases of flies and butterflies are discussed to illustrate our present knowledge of the physical basis of eye colours, their functional background, and the implications for insect colour vision. The screening pigments in the pigment cells commonly determine the eye colour. The red screening pigments of fly eyes and the dorsal eye regions of dragonflies allow stray light to photochemically restore photoconverted visual pigments. A similar role is played by yellow pigment granules inside the photoreceptor cells which function as a light-controlling pupil. Most insect eyes contain black screening pigments which prevent stray light to produce background noise in the photoreceptors. The eyes of tabanid flies are marked by strong metallic colours, due to multilayers in the corneal facet lenses. The corneal multilayers in the gold-green eyes of the deer fly Chrysops relictus reduce the lens transmission in the orange-green, thus narrowing the sensitivity spectrum of photoreceptors having a green absorbing rhodopsin. The tapetum in the eyes of butterflies probably enhances the spectral sensitivity of proximal long-wavelength photoreceptors. Pigment granules lining the rhabdom fine-tune the sensitivity spectra.     

Deriving room temperature excitation spectra for photosystem I and photosystem II fluorescence in intact leaves from the dependence of <Emphasis Type="Italic">F</Emphasis><Subscript>V</Subscript>/<Emphasis Type="Italic">F</Emphasis><Subscript>M</Subscript> on excitation wavelength

The F ₀ and F _M level fluorescence from a wild-type barley, a Chl b-less mutant barley, and a maize leaf was determined from 430 to 685 nm at 10 nm intervals using pulse amplitude-modulated (PAM) fluorimetry. Variable wavelengths of the pulsed excitation light were achieved by passing the broadband emission of a Xe flash lamp through a birefringent tunable optical filter. For the three leaf types, spectra of F _V/F _M (=(F _M − F ₀)/F _M) have been derived: within each of the three spectra of F _V/F _M, statistically meaningful variations were detected. Also, at distinct wavelength regions, the F _V/F _M differed significantly between leaf types. From spectra of F _V/F _M, excitation spectra of PS I and PS II fluorescence were calculated using a model that considers PS I fluorescence to be constant but variable PS II fluorescence. The photosystem spectra suggest that LHC II absorption results in high values of F _V/F _M between 470 and 490 nm in the two wild-type leaves but the absence of LHC II in the Chl b-less mutant barley leaf decreases the F _V/F _M at these wavelengths. All three leaves exhibited low values of F _V/F _M around 520 nm which was tentatively ascribed to light absorption by PS I-associated carotenoids. In the 550–650 nm region, the F _V/F _M in the maize leaf was lower than in the barley wild-type leaf which is explained with higher light absorption by PS I in maize, which is a NADP-ME C₄ species, than in barley, a C₃ species. Finally, low values of F _V/F _M at 685 in maize leaf and in the Chl b-less mutant barley leaf are in agreement with preferential PS I absorption at this wavelength. The potential use of spectra of the F _V/F _M ratio to derive information on spectral absorption properties of PS I and PS II is discussed.     

Unconventional ultraviolet sensitivity spectra of Ascalaphus (Insecta,Neuroptera)

The spectral sensitivity of 21 eye preparations of Ascalaphus (Libelluloides) macaronius (Insecta, Neuroptera) has been re-measured using an up-to-date spectral scan method. 1. Dorso-frontal and ventro-lateral eyes have different spectral characteristics with peaks of sensitivity at 329 ± 8 nm (n = 15) and 343 ± 4 nm (n = 5) (P = 0.002), respectively. 2. The absorbance of the visual pigment layer, K, determined from the shape of the spectral sensitivity curves is 1.3 ± 1.8(n = 15) for dorso-frontal eyes and – 1.0 ± 0.3(n = 5) for ventrolateral eyes, thus implying higher selfscreening in the dorso-frontal eyes and narrowing of the spectral sensitivity curves as regards to a template visual pigment in ventro-lateral eyes. 3. Plotting K versus spectral sensitivity peak wavelength _max revealed an inverse correlation between these variables with K = 42.5 – 0.126 _max at r = 0.88(n = 19). 4. Extracts of ommochromes and carotenoids (Figs. 4 to 6) do not allow to account for the above diversity of optical properties of the Ascalaphus eye (Fig. 7).Abbreviations SSC spectral sensitivity curve - DF dorso-frontal eye - UV ultraviolet - VL ventro-lateral eye     

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.     

An evaluation of the diet of <Emphasis Type="Italic">Mysis relicta</Emphasis> using gut contents and fatty acid profiles in lakes with and without the invader <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis> (<Emphasis Type="Italic">Onychopoda,Cercopagidae</Emphasis>)

Diets of Mysis relicta from four lakes in central Ontario that had been invaded by Bythotrephes longimanus and three lakes that had not been invaded were investigated using gut content analysis and fatty acid (FA) composition. Gut content analysis of M. relicta revealed a high incidence of cannibalism in all lakes, and consumption of B. longimanus and native zooplanktivorous midges in the genus Chaoborus in lakes where these were present. Cladocera other than B. longimanus were present in the guts of all M. relicta examined except those from Bernard Lake, the lake with the most B. longimanus. In that lake, B. longimanus was the most frequent diet item. Copepod remains were found in 60–100% of M. relicta guts with the lowest frequency occurring in Bernard Lake. Fatty acids (FA) that contributed strongly to the variation in FA composition in M. relicta, as revealed by a principal component analysis, were C16:0 (palmitic acid), C16:1n7 (palmitoleic acid), C18:1n9c (oleic acid), C20:4n6 (arachidonic acid), C20:5n3 (eicosapentaenoic acid), and C22:6n3 (docosahexaenoic acid). Significant differences in FA amount and composition of M. relicta were found between invaded and non-invaded lakes, and among lakes within these groups. Generally, M. relicta in non-invaded lakes had higher concentrations of C16:0, C18:1n9c, C18:2n6c (linoleic acid), C18:3n3 (α-linolenic acid) and C20:4n6, while M. relicta in invaded lakes had higher concentrations of C22:6n3. Two of the non-invaded lakes had lower water transparency, as measured by Secchi depth, which may be the reason why mysids and abundant populations of Chaoborus spp. could be found in the water column during the day. However, differences in FA profiles and gut contents of M. relicta between invaded and non-invaded lakes are consistent with competition for Cladocera in the presence of the invader rather than pre-existing differences among lakes. We conclude that the diet of M. relicta is affected by the invasion of B. longimanus.     

Spectral absorption by screening pigment granules in the compound eyes of a moth and a wasp

Summary The spectral absorption by single ommin containing pigment granules or clusters of granules from compound eyes was measured spectrophotometrically between 300 and 700 nm. The measurements were made on fresh and fixed slices from compound eyes of Celerio euphorbiae and Vespa spec. In the visible part of the spectrum there is an absorption maximum between 540 and 550 nm, situated nearly 30 nm more towards the red than that of pure ommin in solution. A frequently found side maximum of variable height at about 450 nm is probably caused by oxidized xanthommatin occurring additionally within the granules. The absorption increases from 350 nm towards shorter wavelengths, and gradually declines between 550 and 700 nm.This work was supported partly by the Swedish Medical Research Council, Stiftelsen Gustaf och Tyra Svenssons Minne, Reservationsanslaget, and partly by the Air Force Office of Scientific Research through the European Office of Aerospace Research, OAR, United States Air Force under grant number EOOAR-68-0036.     

Photoreceptor spectral sensitivities of the Small White butterfly <Emphasis Type="Italic">Pieris rapae crucivora</Emphasis> interpreted with optical modeling

The compound eye of the Small White butterfly, Pieris rapae crucivora, has four classes of visual pigments, with peak absorption in the ultraviolet, violet, blue and green, but electrophysiological recordings yielded eight photoreceptors classes: an ultraviolet, violet, blue, double-peaked blue, green, blue-suppressed-green, pale-red and deep-red class. These photoreceptor classes were identified in three types of ommatidia, distinguishable by the different eye shine spectra and fluorescence; the latter only being present in the eyes of males. We present here two slightly different optical models that incorporate the various visual pigments, the light-filtering actions of the fluorescent, pale-red and deep-red screening pigment, located inside or adjacent to the rhabdom, and the reflectance spectrum of the tapetum that abuts the rhabdom proximally. The models serve to explain the photoreceptor spectral sensitivities as well as the eye shine.     

The lens eyes of the box jellyfish <Emphasis Type="Italic">Tripedalia cystophora</Emphasis> and <Emphasis Type="Italic">Chiropsalmus sp.</Emphasis> are slow and color-blind

Box jellyfish, or cubomedusae, possess an impressive total of 24 eyes of four morphologically different types. Compared to other cnidarians they also have an elaborate behavioral repertoire, which for a large part seems to be visually guided. Two of the four types of cubomedusean eyes, called the upper and the lower lens eye, are camera type eyes with spherical fish-like lenses. Here we explore the electroretinograms of the lens eyes of the Caribbean species, Tripedalia cystophora, and the Australian species, Chiropsalmus sp. using suction electrodes. We show that the photoreceptors of the lens eyes of both species have dynamic ranges of about 3 log units and slow responses. The spectral sensitivity curves for all eyes peak in the blue-green region, but the lower lens eye of T. cystophora has a small additional peak in the near UV range. All spectral sensitivity curves agree well with the theoretical absorbance curve of a single opsin, strongly suggesting color-blind vision in box jellyfish with a single receptor type. A single opsin is supported by selective adaptation experiments.     

Base composition of intact nucleic acid oligomers

The base composition of intact, purified, oligonucleotides has been determined by comparing the absorption spectrum of each oligomer with calculated curves. The spectral curve of each oligomer was measured in 7M urea at three pH values and digitized at 1-mμ intervals. The calculated curves consisted of the sums of the absorption spectra of mononucleotides in 7M urea at the same pH's. A computer was used to make the comparisons and establish the best fit. Results are presented for nine of the ten possible dimer composition isomers; trimers from pancreatic and T₁ ribonuclease hydrolyzates of TMV-RNA; poly-A, poly-C, and poly-U; and intact tobacco mosaic virus-RNA.     

WAVELENGTH DEPENDENCY OF THE MAXIMUM QUANTUM YIELD OF CARBON FIXATION FOR TWO RED TIDE DINOFLAGELLATES,HETEROCAPSA PYGMAEA AND PROROCENTRUM MINIMUM (PYRROPHYTA): IMPLICATIONS FOR MEASURING PHOTOSYNTHETIC RATES1

The influence of photoadaptive state on the spectral dependency of the maximum quantum yield for carbon fixation was determined for two red tide dinoflagellates, Heterocapsa pygmaea Loeblich, Schmidt, et Sherley and Prorocentrum minimum Pavillard. Cultures were acclimated to green, blue, red, and white light. The spectral dependency in the light-limited slope of the photosynthesis–irradiance curves (α) was measured with carbon action spectra that, when divided by the spectrally weighted absorption coefficient, provided estimates of the maximum quantum yield (φ_max) for carbon fixation. Values of φ_max varied with wavelength within each culture condition as well as between different culture conditions. The degree to which the spectral dependency in φ_max was influenced by the presence of photoprotective carotenoids and/or energy imbalances between photosystems I and II was assessed for both dinoflagellates. The impact of photoprotective pigmentation on the spectral dependency of φ_max was most significant for cells grown under high light conditions reflecting the enrichment of diadinoxanthin. Energy imbalances between the photosystems was assessed by quantifying enhancement effects on spectral φ_max in the presence of background illumination. Under our experimental conditions, enhancement effects on carbon action spectra were evident for H. pygmaea under nearly all growth conditions but were not detectable for P. minimum under any growth condition. We hypothesize that sensitivity to enhancement effects reflected differences in the structure of the photosynthetic machinery of these two peridinin-containing dinoflagellates. While measurements of φ_max are sensitive to the color of the light within an incubator, the relative impact on the spectral dependency of a was less than the wavelength dependency associated with the cellular absorption properties. Finally we used our data to validate an approach proposed by others to aid in the correction of photosynthetic measurements where the in situ spectral light field cannot be easily mimicked. The average error using this approach was 8%, which was significantly less than the error associated with ignoring the spectral dependency in α.     

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.     

Spectral Sensitivity of Larval Mosquito Ocelli

The spectral sensitivity of lateral ocelli in both wild-type and white-eyed larvae of the yellow fever mosquito Aedes aegypti L. (reared in darkness) was measured by means of the electroretinogram. The spectral sensitivity is maximal at about 520 nm, with a small secondary peak near 370 nm. When allowance is made for some screening and filtering by the eye tissues, the spectral sensitivity is in reasonable agreement with the absorption spectrum of ocellar rhodopsin (λ_max = 515 nm).     

Comparative energetics and life cycle of the opossum shrimp ( Mysis relicta ) in native and non-native environments

1. Life history, fecundity and energy ingestion were compared for non-native Mysis relicta in Flathead Lake, Montana, and in Waterton Lake, Montana/Alberta, where the mysid is native. 2. Based on estimates from stomach contents, M. relicta in Flathead Lake consumed three to four times more energy per hour while foraging than M. relicta from Waterton Lake. The primary prey of M. relicta in Flathead Lake was Daphnia thorata. Diaptomus sicilis was the primary prey for M. relicta in Waterton Lake. 3. Mysis relicta in Flathead Lake had a 1-year life history. It took 2 years for M. relicta in Waterton Lake to complete a generation. Mysis relicta from Flathead Lake produced significantly (P < 0.05) more eggs per female (19.9 ± 4.7) compared to M. relicta in Waterton Lake (13.7 ± 3.6). The lipid content in eggs from M. relicta in Flathead Lake was also significantly (P < 0.05) higher than eggs from M. relicta in Waterton Lake. 4. In Flathead Lake, M. relicta is able effectively to exploit Daphnia thorata, consuming greater biomass than M. relicta in Waterton Lake. Mysis relicta efficiently assimilates the greater biomass, which reduces generation time, increases fecundity, and increases lipid reserves of offspring. These results provide evidence of energetic mechanisms that enable M. relicta to colonize and affect foodwebs when introduced into lakes.