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     

Visual pigments, oil droplets, ocular media and cone photoreceptor distribution in two species of passerine bird: the blue tit (Parus caeruleus L.) and the blackbird (Turdus merula L.)

The spectral absorption characteristics of the retinal photoreceptors of the blue tit (Parus caeruleus) and blackbird (Turdus merula) were investigated using microspectrophotometry. The retinae of both species contained rods, double cones and four spectrally distinct types of single cone. Whilst the visual pigments and cone oil droplets in the other receptor types are very similar in both species, the wavelength of maximum sensitivity (λ_max) of long-wavelength-sensitive single and double cone visual pigment occurs at a shorter wavelength (557 nm) in the blackbird than in the blue tit (563 nm). Oil droplets located in the long-wavelength-sensitivesingle cones of both species cut off wavelengths below 570–573 nm, theoretically shifting cone peak spectral sensitivity some 40 nm towards the long-wavelength end of the spectrum. This raises the possibility that the precise λ_max of the long-wavelength-sensitive visual pigment is optimised for the visual function of the double cones. The distribution of cone photoreceptors across the retina, determined using conventional light and fluorescence microscopy, also varies between the two species and may reflect differences in their visual ecology. Accepted: 8 January 2000     

Correlation between auditory sensitivity and vocalization in anabantoid fishes

Several anabantoid species produce broad-band sounds with high-pitched dominant frequencies (0.8–2.5 kHz), which contrast with generally low-frequency hearing abilities in (perciform) fishes. Utilizing a recently developed auditory brainstem response recording-technique, auditory sensitivities of the gouramis Trichopsis vittata, T. pumila, Colisa lalia, Macropodus opercularis and Trichogaster trichopterus were investigated and compared with the sound characteristics of the respective species. All five species exhibited enhanced sound-detecting abilities and perceived tone bursts up to 5 kHz, which qualifies this group as hearing specialists. All fishes possessed a high-frequency sensitivity maximum between 800 Hz and 1500 Hz. Lowest hearing thresholds were found in T. trichopterus (76 dB re 1 μPa at 800 Hz). Dominant frequencies of sounds correspond with the best hearing bandwidth in T. vittata (1–2 kHz) and C. lalia (0.8–1 kHz). In the smallest species, T. pumila, dominant frequencies of acoustic signals (1.5–2.5 kHz) do not match lowest thresholds, which were below 1.5 kHz. However, of all species studied, T. pumila had best hearing sensitivity at frequencies above 2 kHz. The association between high-pitched sounds and hearing may be caused by the suprabranchial air-breathing chamber, which, lying close to the hearing and sonic organs, enhances both sound perception and emission at its resonant frequency. Accepted: 26 November 1997     

Phototaxis in water fleas (Daphnia magna) is differently influenced by visible and UV light

The effects of vertical illumination with monochromatic lights on phototaxis of Daphnia magna in a test chamber were determined at five levels of equal quantal flux density (between 188 and 6.42 · 10⁻⁵ nEinstein). Visible adaptation light (500 nm) and subsequent spectral test light had the same quantal flux density. The animals reacted to ultraviolet light (260–380 nm) with negative phototaxis, whereas visible light (420–600 nm) caused positive phototaxis. Action spectra were determined, based on the evaluation of different parameters of phototactic behavior. The maximum spectral sensitivity in the ultraviolet was found at 340 nm. The maximum spectral efficiency in the visible varied in dependence on light intensity. Ecological consequences of the results are discussed. Accepted: 3 August 1998     

A comparison of electrophysiologically determined spectral responses in 35 species of Lepidoptera

Spectral responses from the compound eyes of 35 lepidopteran species representing 14 families were investigated electrophysiologically using ERG recordings. The light-stimuli used overed the range of 383–700 nm wavelengths. All species show three or four maxima in their spectral sensitivity curves. Two of these peaks were usually associated with ultraviolet and blue light (383 and 460 nm, respectively). The other maxima occurred in the 500–620 nm region. In Nymphalidae the highest peak was found in response to 560–580 nm stimuli. Of all wavelengths tested, these are the longest wavelengths to produce principal peak sensitivities.Pieridae and Lycaenidae have maxima in the UV region which represent significantly higher sensitivities than the secondary peaks to stimuli of longer wavelengths.Satyridae, Danaidae, Hesperiidae and diurnal moths except Epicopeia (Epicopeidae) generally have similar sensitivity curves with principal peaks between 500 and 520 nm.In Papilionid species except Graphium (max = 560 nm) high maxima occur in the UV and blue (460 nm) region.Noctural Sphingid moths possess the highest peak sensitivity at 540 nm. All other noctural moths tested have three or four maxima.     

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     

Dichromatic colour vision in an Australian marsupial, the tammar wallaby

Despite earlier assertions that most mammals are colour blind, colour vision has in recent years been demonstrated in a variety of eutherian mammals from a wide range of different orders. This paper presents the first behavioural evidence from colour discrimination experiments, that an Australian marsupial, the tammar wallaby (Macropus eugenii), has dichromatic colour vision. In addition, the experiments show that the wallabies readily learn the relationship between the presented colours rather than the absolute hues. This provides a sensitive method to measure the location of the neutral-point, which is the wavelength of monochromatic light that is indistinguishable from white. This point is a diagnostic feature for dichromats. The spectral sensitivity of the wallabies' middle-wavelength-sensitive photoreceptor is known (peak: 539 nm) and the behavioural results imply that the sensitivity of the short-wavelength-sensitive receptor must be near 420 nm. These spectral sensitivities are similar to those found in eutherian mammals, supporting the view that the earliest mammals had dichromatic colour vision. Accepted: 18 July 1999     

Isolation and characterization of photoactive complexes of NADPH:protochlorophyllide oxidoreductase from wheat

A photoactive substrate-enzyme complex of the NADPH:protochlorophyllide oxidoreductase (POR; EC 1. 3. 1. 33) was purified from etiolated Triticum aestivum L. by gel chromatography after solubilization of prolamellar bodies by dodecyl-maltoside. Irradiation by a 1-ms flash induced the phototransformation of protocholorophyllide a (Pchlide) with −196 °C absorbance and emission maxima at 640 and 643 nm, respectively. The apparent molecular weight of this complex was 112 ± 24 kDa, which indicates aggregation of enzyme subunits. By lowering the detergent concentration in the elution buffer, a 1080 ± 250-kDa particle was obtained which displayed the spectral properties of the predominant form of photoactive Pchlide in vivo (−196 °C absorbance and fluorescence maxima at 650 and 653 nm). In this complex, POR was the dominant polypeptide. Gel chromatography in the same conditions of an irradiated sample of solubilized prolamellar bodies indicated rapid disaggregation of the complex after Pchlide phototransformation. High performance liquid chromatographic analysis of the POR complexes obtained using two detergent concentrations indicates a possible association of zeaxanthin and violaxanthin with the photoactive complex. Received: 25 February 1998 / Accepted: 8 June 1998     

The photoreceptors and visual pigments of two species of Acipenseriformes, the shovelnose sturgeon (Scaphirhynchus platorynchus ) and the paddlefish (Polyodon spathula )

Scanning electron microscopy, microspectrophotometry, and spectrophotometry of digitonin extracts were employed to characterize the photoreceptors and visual pigments of two freshwater Acipenseriformes. The retinas of the shovelnose sturgeon, Scaphirhynchus platorynchus (Acipenseridae), and the paddlefish, Polyodon spathula (Polyodontidae) are dominated by large rods with long, broad outer segments. A second rod, rare and much narrower than the dominant rod, is present in Scaphirhynchus but not seen in Polyodon. The absorbance maximum of the visual pigment in the rods of Polyodon is near 540 nm; that of Scaphirhynchus near 534 nm. The retinas of both species contain substantial numbers of large, single cones, about 33% of the photoreceptors in Scaphirhynchus; 37% in Polyodon. Scaphirhynchus cone pigments have absorbance maxima near 610 nm, 521 nm and 470 nm, respectively. Polyodon cone pigments absorb maximally near 607 nm and 535 nm, respectively. All visual pigments are based on vitamin A₂. The data are compared to those from other Acipenseriformes and are discussed in terms of lifestyle and behavior. Accepted: 7 October 1998     

Photon Hunting in the Twilight Zone: Visual Features of Mesopelagic Bioluminescent Sharks

The mesopelagic zone is a visual scene continuum in which organisms have developed various strategies to optimize photon capture. Here, we used light microscopy, stereology-assisted retinal topographic mapping, spectrophotometry and microspectrophotometry to investigate the visual ecology of deep-sea bioluminescent sharks [four etmopterid species (Etmopterus lucifer, E. splendidus, E. spinax and Trigonognathus kabeyai) and one dalatiid species (Squaliolus aliae)]. We highlighted a novel structure, a translucent area present in the upper eye orbit of Etmopteridae, which might be part of a reference system for counterillumination adjustment or acts as a spectral filter for camouflage breaking, as well as several ocular specialisations such as aphakic gaps and semicircular tapeta previously unknown in elasmobranchs. All species showed pure rod hexagonal mosaics with a high topographic diversity. Retinal specialisations, formed by shallow cell density gradients, may aid in prey detection and reflect lifestyle differences; pelagic species display areae centrales while benthopelagic and benthic species display wide and narrow horizontal streaks, respectively. One species (E. lucifer) displays two areae within its horizontal streak that likely allows detection of conspecifics'' elongated bioluminescent flank markings. Ganglion cell topography reveals less variation with all species showing a temporal area for acute frontal binocular vision. This area is dorsally extended in T. kabeyai, allowing this species to adjust the strike of its peculiar jaws in the ventro-frontal visual field. Etmopterus lucifer showed an additional nasal area matching a high rod density area. Peak spectral sensitivities of the rod visual pigments (λ_max) fall within the range 484–491 nm, allowing these sharks to detect a high proportion of photons present in their habitat. Comparisons with previously published data reveal ocular differences between bioluminescent and non-bioluminescent deep-sea sharks. In particular, bioluminescent sharks possess higher rod densities, which might provide them with improved temporal resolution particularly useful for bioluminescent communication during social interactions.     

Four spectral classes of cone in the retinas of birds

Summary The spectral sensitivity of 15 species of birds has been measured by recording transretinal voltages from opened eyecups. With suitable combinations of colored adapting lights, we find that a variety of passerines have four peaks of photopic sensitivity, with maxima at 370, 450, 480, and 570 nm. Additional sensitivity maxima at 510 nm are found in some species. The spectral sensitivity functions are not altered by bathing the retinas in 50 mM sodium aspartate, suggesting that they reflect the properties of cones and do not result from inhibitory interactions between retinal interneurons.Comparison of the results with a general mathematical model that describes spectral sensitivity functions recorded extracellularly from populations of receptors in different states of adaptation (Goldsmith 1986) shows that the retinal spectral sensitivity functions are consistent with the presence of (at least) four types of cone, but indicate as well that many of the cones that are maximally sensitive in the blue and violet likely contain oil droplets that attenuate the deep violet and near uv.     

Neuronal basis for spectral song discrimination in the bushcricket Tettigonia cantans

Previous experiments demonstrated that in choice situations female Tettigonia cantans prefer signals with the conspecific spectrum over signals with the spectrum of the sibling species T. viridissima, whose signals lack the T. cantans-typical 8-kHz component but are otherwise identical in spectral composition. The underlying neuronal mechanisms were investigated using whole-nerve recordings of the tympanal nerve. Hearing thresholds did not differ significantly between the species in the range between 7 and 25 kHz. The responses of the tympanic nerve at 30 dB above threshold did, however, differ significantly between the species. While in T. viridissima response amplitudes did not differ between 7 and 12 kHz, in T. cantans responses at 8 kHz were significantly larger than at 10 and 12 kHz. Since possible influences of axonal diameters on response magnitudes were excluded, these results indicate that in T. cantans more receptor cells are tuned to 8 kHz than to 10–12 kHz, while in T. viridissima tuning of receptor cells is more evenly distributed. The higher response magnitude in T. cantans at 8 kHz than at 10–12 kHz is equivalent to an amplitude difference of 3.8–7 dB, which agrees well with previous behavioural estimates. Accepted: 13 January 1999     

The visual pigments of four deep-sea crustacean species

Summary The visual pigments of four mesopelagic crustacean species were studied at sea by means of microspectrophotometry. The absorbance maxima obtained for the visual pigments and their metarhodopsins, respectively, were: 493 nm and 481 nm (Systellaspis debilis), 485 nm and 480 nm (Acanthephyra curtirostris), 491 nm and 482 nm (A. smithi), and 495 nm and 487 nm (Sergestes tenuiremis). The spectral characteristics of the rhodopsins and metarhodopsins permit high photosensitivity and facilitate photoregeneration in a nearly monochromatic environment. Photic regeneration of rhodopsins from the deep-sea environment was demonstrated, and data were obtained which are consistent with the occurrence of dark regeneration. Specific optical density of the observed visual pigments was calculated for two species.     

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.     

Day and night mesopelagic fish assemblages off the Kerguelen Islands (Southern Ocean)

Mesopelagic fish assemblages were investigated in the Polar Frontal Zone off the Kerguelen Islands during summer 1995, in parallel with a king penguin tracking study. During the day, the upper offshore water layers (0–200 m) have low potential prey diversity and abundance with only three fish species: a lanternfish, Krefftichthys anderssoni, a member of the Muraenolepididae, Muraenolepis marmoratus, and the early stages of the nototheniid, Lepidonotothen squamifrons. The mesopelagic fish community, including the typical myctophids, first appears in the deeper layer (300 m). At night, the surface layer (50 m) is invaded by the mesopelagic Myctophidae Electrona antarctica, Gymnoscopelus braueri, G. piabilis, G. fraseri, G. nicholsi, Protomyctophum bolini and P. tenisoni. Deeper (>100 m), a cline of species assemblages from the coast to offshore is observed. Accepted: 4 August 1999     

Spectral sensitivity of the compound eye in butterflies (Heliconius)

Summary The spectral sensitivity of the compound eye in three butterfly species (Heliconius erato, H. numata, H, sara) was tested electrophysiologically in the wavelength region 310 to 650 nm. Sensitivity maxima were found at 370 to 390 nm, 450 to 470 nm, and 550 to 570 nm, for all species. The three sensitivity maxima are suggested to be due to different photoreceptor types effecting wave-length discrimination. An interspecies difference in spectral sensitivity was also found. The difference is suggested to be due to the relative number of photoreceptors of each type. In some of the present experiments a small discontinuity in sensitivity was found at 610 or 630 nm. It is probably caused by a selective reflection of these wavelengths from a tapetum.     

Breaking the ice: large-scale distribution of mesozooplankton after a decade of Arctic and transpolar cruises

Mesozooplankton collected during five summer expeditions to the Arctic Ocean between 1987 and 1991 was analysed for regional patterns in biomass and species distribution, distinguishing between an epipelagic (0–100 m) and a deeper (0–500 m) layer. A total of 58 stations was sampled mainly in the Nansen, Amundsen and Makarov Basins of the central Arctic Ocean and in areas of the Greenland Sea, West Spitsbergen Current and Barents Sea. Results from the different expeditions were combined to create a transect extending from the Fram Strait across the Eurasian Basin into the Makarov Basin. Mesozooplankton dry mass in the upper 500 m decreased from 8.4 g m⁻² in the West Spitsbergen Current to less than 2 g m⁻² in the high-Arctic deep-sea basins. In the central Arctic Ocean, biomass was concentrated in the upper 100 m and was dominated by the large copepods Calanus hyperboreus and C. glacialis. In contrast, the mesozooplankton in the West Spitsbergen Current was more evenly distributed throughout the upper 500 m, with C. finmarchicus as the prevailing species. The distribution of abundant mesopelagic species reflected the hydrographic regime: the calanoid copepod Gaetanus tenuispinus and the hyperiid amphipod Themisto abyssorum were most abundant in the Atlantic inflow, while Scaphocalanus magnus was a typical component of the high-Arctic fauna. The relatively high mesozooplankton biomass and the occurrence of boreal-Atlantic species in the central Arctic Ocean are indicators for the import of organic material from allochthonous sources, especially from the northern North Atlantic. Hence, in spite of its enclosure by land masses, the Arctic Ocean is characterized by an exchange of water masses and organisms with the North Atlantic, and advection processes strongly influence the distribution of plankton species in this high-latitude ecosystem. Received: 18 December 1997 / Accepted: 11 April 1998     

The spectral sensitivity of eye movements in response to light flashes inDaphnia magna

Summary The crustaceanDaphnia magna responds to a flash of light with a ventral rotation of its compound eye; this behavior is termed eye flick. We determined the spectral sensitivity for the threshold of eye flick in response to light flashes having three different spatial characteristics: (1) full-field, extending 180° from dorsal to ventral in the animal's field of view; (2) dorsal, 30° wide and located in the dorsal region of the visual field; (3) ventral, same as dorsal but located ventrally. All three stimuli extended 30° to the right and to the left of the animal's midplane. We found that spectral sensitivity varies with the spatial characteristics of the stimulus. For full-field illumination, the relative sensitivity was maximal at 527 nm and between 365 nm and 400 nm, with a significant local minimum at 420 nm. For the dorsal stimulus, the relative sensitivity was greatest at 400 nm, but also showed local maxima at 440 nm and 517 nm. For the ventral stimulus, the relative sensitivity maxima occurred at the same wavelengths as those for the full-field stimulus. At wavelengths of 570 nm and longer, the responses to both dorsal and ventral stimuli showed lower relative sensitivity than the full-field stimulus. No circadian or other periodic changes in threshold spectral sensitivity were observed under our experimental conditions. Animals which had their nauplius eyes removed by means of laser microsurgery had the same spectral sensitivity to full-field illumination as normal animals. Our results are discussed in terms of our current knowledge of the spectral classes of photoreceptors found in theDaphnia compound eye.     

Spectral discrimination of coral reef benthic communities

Effective identification and mapping of coral reef benthic communities using high-spatial and -spectral resolution digital imaging spectrometry requires that the different communities are distinguishable by their spectral reflectance characteristics. In Kaneohe Bay, Oahu, Hawaii, USA, we collected in situ a total of 247 spectral reflectances of three coral species (Montipora capitata, Porites compressa, Porites lobata), five algal species (Dictyosphaeria cavernosa, Gracilaria salicornia, Halimeda sp., Porolithon sp., Sargassum echinocarpum) and three sand benthic communities (fine-grained carbonate sand, sand mixed with coral rubble, coral rubble). Major reflectance features were identified by peaks in fourth derivative reflectance spectra of coral (at 573, 604, 652, 675 nm), algae (at 556, 601, 649 nm) and sand (at 416, 448, 585, 652, 696 nm). Stepwise wavelength selection and linear discriminant function analysis revealed that spectral separation of the communities is possible with as few as four non-contiguous wavebands. These linear discriminant functions were applied to an airborne hyperspectral image of a patch reef in Kaneohe Bay. The results demonstrate the ability of spectral reflectance characteristics, determined in situ, to discriminate the three basic benthic community types: coral, algae and sand. Accepted: 12 January 2000     

Spectral sensitivity of ground squirrel cones measured with ERG flicker photometry

Summary Ground squirrels have dichromatic color vision. The spectral sensitivities of the two classes of cones found in the retinas of two species of ground squirrel were measured using ERG flicker photometry. The spectral sensitivity curves for these cone classes were closely fit by curves from wavelength-dependent visual pigment nomograms. One cone type had an average peak sensitivity of 518.9 nm (California ground squirrels,Spermophilus beecheyi) or 517.0 nm (thirteen-lined ground squirrels,Spermophilus tridecemlineatus). The second type of cone found in these ground squirrels had an average peak sensitivity of 436.7 nm. An examination of the variation in spectral sensitivity among individual animals suggests that the sensitivity peaks for the middle-wavelength cone cover a range of not greater than 4 nm.