Dual Sensitivities of Cells in Wolf Spider Eyes at Ultraviolet and Visible Wavelengths of Light

Intracellular recordings have been made from visual cells in principal and secondary eyes of in vitro wolf spider preparations. The responses of all cells to all wavelengths of light were graded depolarizations; no hyperpolarizations or nerve discharges were seen. Cells in a secondary eye, the anterior lateral eye, had a maximum sensitivity in the visible at 510 nm and a secondary maximum, or shoulder, of sensitivity in the near ultraviolet at 380 nm. Cells in principal eyes, the anterior median eyes, all responded maximally both in the visible at 510 nm and in the ultraviolet at 360–370 nm or less. However, there was no typical ratio of ultraviolet to visible sensitivities; the differences in log sensitivities (log UV/VIS) varied from 3.3 to -0.5. Each principal eye had a population of cells with different ratios. These populations varied with the time of the year, possibly due to changes in light upon the animals. Chromatic adaptations of cells in anterior median (but not anterior lateral) eyes resulted in small, selective changes in spectral sensitivities, and there was some facilitation of responses from cells repeatedly stimulated. It is concluded that cells of secondary eyes contain only a visual pigment absorbing maximally in the visible, while cells of principal eyes probably contain variable amounts of both this pigment and one absorbing in the ultraviolet as well.     

Ultraviolet-Induced Sensitivity to Visible Light in Ultraviolet Receptors of Limulus

In the UV-sensitive photoreceptors of the median ocellus (UV cells), prolonged depolarizing afterpotentials are seen following a bright UV stimulus. These afterpotentials are abolished by long-wavelength light. During a bright UV stimulus, long-wavelength light elicits a sustained negative-going response. These responses to long-wavelength light are called repolarizing responses. The spectral sensitivity curve for the repolarizing responses peaks at 480 nm; it is the only spectral sensitivity curve for a median ocellus electrical response known to peak at 480 nm. The reversal potentials of the repolarizing response and the depolarizing receptor potential are the same, and change in the same way when the external sodium ion concentration is reduced. We propose that the generation of repolarizing responses involves a thermally stable intermediate of the UV-sensitive photopigment of UV cells.     

The Spectral Sensitivities of Single Receptor Cells in the Lateral, Median, and Ventral Eyes of Normal and White-Eyed Limulus

Spectral sensitivity curves can be distorted by screening pigments. We have determined whether this is true for Limulus polyphemus by determining, from receptor potentials recorded using intracellular microelectrodes, spectral sensitivity curves for normal animals and for white-eyed animals (which lack screening pigment). Our results show: (a) In median ocelli, the curve for UV-sensitive receptor cells peaks at 360 nm and does not depend on the presence of screening pigment, (b) The curve for ventral eye photoreceptors is identical to that for retinular cells from the lateral eyes of white-eyed animals and peaks at 520–525 nm. (c) In normal lateral eyes, when the stimulating light passes through screening pigment, the curve indicates relatively more sensitivity in the red region of the spectrum than does the curve for white-eyed animals. Therefore, the screening pigment is probably red-transmitting, (d) In median ocelli, the curve for visible-sensitive cells peaks at 525 nm and is approximately the same whether the ocelli are from normal or white-eyed animals. However, the curve is significantly broader than that for ventral eyes and for lateral eyes from white-eyed animals.     

Electroretinogram Characteristics and the Spectral Mechanisms of the Median Ocellus and the Lateral Eye in Limulus polyphemus

Electrical responses (ERG) to light flashes of various wavelengths and energies were obtained from the dorsal median ocellus and lateral compound eye of Limulus under dark and chromatic light adaptation. Spectral mechanisms were studied by analyzing (a) response waveforms, e.g. response area, rise, and fall times as functions of amplitude, (b) slopes of amplitude-energy functions, and (c) spectral sensitivity functions obtained by the criterion amplitude method. The data for a single spectral mechanism in the lateral eye are (a) response waveforms independent of wavelength, (b) same slope for response-energy functions at all wavelengths, (c) a spectral sensitivity function with a single maximum near 520 mµ, and (d) spectral sensitivity invariance in chromatic adaptation experiments. The data for two spectral mechanisms in the median ocellus are (a) two waveform characteristics depending on wavelength, (b) slopes of response-energy functions steeper for short than for long wavelengths, (c) two spectral sensitivity peaks (360 and 530–535 mµ) when dark-adapted, and (d) selective depression of either spectral sensitivity peak by appropriate chromatic adaptation. The ocellus is 200–320 times more sensitive to UV than to visible light. Both UV and green spectral sensitivity curves agree with Dartnall's nomogram. The hypothesis is favored that the ocellus contains two visual pigments each in a different type of receptor, rather than (a) various absorption bands of a single visual pigment, (b) single visual pigment and a chromatic mask, or (c) fluorescence. With long duration light stimuli a steady-state level followed the transient peak in the ERG from both types of eyes.     

Retinal receptor types inAglais urticae andPieris brassicae (Lepidoptera), revealed by analysis of the electroretinogram obtained with Fourier interferometric stimulation (FIS)

Summary Electroretinograms obtained in the butterfliesAglais urticae andPieris brassicae by the procedure of Fourier interferometric stimulation (FIS) were used to construct spectral sensitivity curves. These curves, representing the combined responses of several receptor types, were approximated by summation of spectral sensitivity curves for individual pigments, and the presence of these pigments was corroborated by chromatic adaptation experiments. The results show that the retina in the compound eye ofAglais urticae contains 3 photopigments, with maximal absorption at ca. 360 nm, 460 nm and 530 nm, respectively (Fig. 5). The retina in the compound eye ofPieris brassicae has two subdivisions. In the dorsal region of the eye 3 photopigments were found, with maxima at ca. 360 nm, 450 nm and 560 nm (Fig. 8). In the medioventral region pigments with essentially the same maxima are present together with an additional, fourth long-wavelength component with effective maximal absorption at ca. 620 nm (Fig. 11). Its absorption curve is considerably narrower than would be expected for a rhodopsin with the same absorption maximum, and presumably results from the spectral combination of a photopigment and a photostable screening pigment.Abbreviations FIS Fourier interferometric stimulation - WLP White-light position - ERG Electroretinogram     

Spectral Sensitivities of Wolf Spider Eyes

ERG's to spectral lights were recorded from all eyes of intact wolf spiders. Secondary eyes have maximum relative sensitivities at 505–510 nm which are unchanged by chromatic adaptations. Principal eyes have ultraviolet sensitivities which are 10 to 100 times greater at 380 nm than at 505 nm. However, two animals' eyes initially had greater blue-green sensitivities, then in 7 to 10 wk dropped 4 to 6 log units in absolute sensitivity in the visible, less in the ultraviolet. Chromatic adaptations of both types of principal eyes hardly changed relative spectral sensitivities. Small decreases in relative sensitivity in the visible with orange adaptations were possibly retinomotor in origin. Second peaks in ERG waveforms were elicited from ultraviolet-adapted principal eyes by wavelengths 400 nm and longer, and from blue-, yellow-, and orange-adapted secondary eyes by wavelengths 580 nm and longer. The second peaks in waveforms were most likely responses of unilluminated eyes to scattered light. It is concluded that both principal and secondary eyes contain cells with a visual pigment absorbing maximally at 505–510 nm. The variable absolute and ultraviolet sensitivities of principal eyes may be due to a second pigment in the same cells or to an ultraviolet-absorbing accessory pigment which excites the 505 nm absorbing visual pigment by radiationless energy transfer.     

Ultraviolet vision in birds: the importance of transparent eye media

Ultraviolet (UV)-sensitive visual pigments are widespread in the animal kingdom but many animals, for example primates, block UV light from reaching their retina by pigmented lenses. Birds have UV-sensitive (UVS) visual pigments with sensitivity maxima around 360–373 nm (UVS) or 402–426 nm (violet-sensitive, VS). We describe how these pigments are matched by the ocular media transmittance in 38 bird species. Birds with UVS pigments have ocular media that transmit more UV light (wavelength of 50% transmittance, λ_T0.5, 323 nm) than birds with VS pigments (λ_T0.5, 358 nm). Yet, visual models predict that colour discrimination in bright light is mostly dependent on the visual pigment (UVS or VS) and little on the ocular media. We hypothesize that the precise spectral tuning of the ocular media is mostly relevant for detecting weak UV signals, e.g. in dim hollow-nests of passerines and parrots. The correlation between eye size and UV transparency of the ocular media suggests little or no lens pigmentation. Therefore, only small birds gain the full advantage from shifting pigment sensitivity from VS to UVS. On the other hand, some birds with VS pigments have unexpectedly low UV transmission of the ocular media, probably because of UV blocking lens pigmentation.     

Photoreceptor Spectral Sensitivity in the Bumblebee,Bombus impatiens (Hymenoptera: Apidae)

The bumblebee Bombus impatiens is increasingly used as a model in comparative studies of colour vision, or in behavioural studies relying on perceptual discrimination of colour. However, full spectral sensitivity data on the photoreceptor inputs underlying colour vision are not available for B. impatiens. Since most known bee species are trichromatic, with photoreceptor spectral sensitivity peaks in the UV, blue and green regions of the spectrum, data from a related species, where spectral sensitivity measurements have been made, are often applied to B impatiens. Nevertheless, species differences in spectral tuning of equivalent photoreceptor classes may result in peaks that differ by several nm, which may have small but significant effects on colour discrimination ability. We therefore used intracellular recording to measure photoreceptor spectral sensitivity in B. impatiens. Spectral peaks were estimated at 347, 424 and 539 nm for UV, blue and green receptors, respectively, suggesting that this species is a UV-blue-green trichromat. Photoreceptor spectral sensitivity peaks are similar to previous measurements from Bombus terrestris, although there is a significant difference in the peak sensitivity of the blue receptor, which is shifted in the short wave direction by 12–13 nm in B. impatiens compared to B. terrestris.     

Spectral sensitivity of photoreceptors mediating phase-shifts of circadian rhythms in retinally degenerate CBA/J (rd/rd) and normal CBA/N (+/+) mice

Light-dark cycles are the most important time cue for the circadian system to entrain the endogenous circadian clock to the environmental 24 h cycle. Although photic entrainment of circadian rhythms is mediated by the eye in mammals, photoreceptors implicated in circadian photoreception remain unknown. In our previous study, retinally degenerate CBA/J (rd/rd) mice were found to have lower circadian photo-sensitivity for phase-shifting the locomotor activity rhythms than normal CBA/N(+/+) mice. In the present study, the spectral sensitivity for phase-shifting the rhythms was examined in order to characterize the photopigments involved in circadian photoreception of these mice. The spectral sensitivity of CBA/J-rd/rd mice clearly fitted to the Dartnall nomogram for a retinal₁-based pigment with a maximum at 480 nm, while the best fitted nomogram had a maximum at 500 nm in CBA/N- +/+ mice. These results suggest that circadian photopigments involved in CBA/J-rd/rd and CBA/N- +/+ mice may be different.     

Spectral sensitivities including the ultraviolet of the passeriform bird Leiothrix lutea

Spectral sensitivity functions of a passeriform bird, the Red-billed Leiothrix Leiothrix lutea (Timalidae) were determined in a behavioural test under different background illuminations.

Six spectral sensitivity classes in crab visual interneurons

Summary Electrophysiological determination of the spectral sensitivity of units from the retina, optic lobes and brain of the crabParagrapsus gaimardii revealed six major colour types. Single peaks occurred in the UV, blue-green, green and yellow, and further units showed either a broad-band sensitivity or double peaks in the violet and yellow.Many of the curves had broad shoulders and offset peaks, markedly different in shape to curves suggested by Dartnall for rhodopsins with the same peak values, but similar to those previously recorded for other crabs. This finding, allied with limited evidence that a number of the colour types changed one from the other depending on illumination level, suggests that at least some of the colour types may result from a filtering of the light reaching a single retinal type.The UV sensitivity, recorded from brain interneurons, is the highest so far found in crabs.The range of colour types recorded provides a good potential substrate for colour vision. It is therefore of interest that a survey of a total of 105 units provided no evidence for colour opponency.Queen Elizabeth II Fellow in Marine Science, 1978–1979     

Colour choice behaviour in the pollen beetle Meligethes aeneus (Coleoptera: Nitidulidae)

The pollen beetle Meligethes aeneus Fabricius (Coleoptera, Nitidulidae), a pest of oilseed rape (Brassica napus), is known to respond to coloured stimuli; however, current understanding of the underlying mechanisms of colour choice in this species is limited. In the present study, physiological and behavioural experiments are conducted to determine the response of the pollen beetle to colours in the field. Spectral sensitivity is measured in 10 animals using the electroretinogram technique. Light flashes (100 ms) at varied wavelengths (340–650 nm, 10‐nm steps) and at different light intensities are applied to the eye after dark adaptation. In behavioural experiments in the field, 100 water traps of varying colours (from yellow to green to blue with varying amounts of white and black added, and with known spectral reflectance) are set out on a bare soil field in May 2008. The mean spectral sensitivity curve of M. aeneus peaks at 520 nm; however, a model template fitted to the long wavelength tail of the observed curve reveals a peak at approximately 540 nm (green). A secondary sensitivity peak is observed in the ultraviolet (UV) range (370 nm). A total of 2482 pollen beetles are captured in the coloured traps. The results show that the pollen beetles' preference for yellow over other colours can be modelled as a colour opponent mechanism (green versus blue); however, further experiments are needed to specify responses to colours with higher UV reflectance. These findings may be used to optimize trap colours for monitoring to help develop integrated pest management strategies for pollen beetle control.     

An expanded set of photoreceptors in the Eastern Pale Clouded Yellow butterfly, Colias erate

We studied the spectral and polarisation sensitivities of photoreceptors of the butterfly Colias erate by using intracellular electrophysiological recordings and stimulation with light pulses. We developed a method of response waveform comparison (RWC) for evaluating the effective intensity of the light pulses. We identified one UV, four violet-blue, two green and two red photoreceptor classes. We estimated the peak wavelengths of four rhodopsins to be at about 360, 420, 460 and 560 nm. The four violet-blue classes are presumably based on combinations of two rhodopsins and a violet-absorbing screening pigment. The green classes have reduced sensitivity in the ultraviolet range. The two red classes have primary peaks at about 650 and 665 nm, respectively, and secondary peaks at about 480 nm. The shift of the main peak, so far the largest amongst insects, is presumably achieved by tuning the effective thickness of the red perirhabdomal screening pigment. Polarisation sensitivity of green and red photoreceptors is higher at the secondary than at the main peak. We found a 20-fold variation of sensitivity within the cells of one green class, implying possible photoreceptor subfunctionalisation. We propose an allocation scheme of the receptor classes into the three ventral ommatidial types.     

Optimal mechanisms for finding and selecting mates: how threespine stickleback (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>) should encode male throat colors

Male threespine stickleback (Gasterosteus aculeatus) use nuptial colors to attract mates and intimidate rivals. We quantified stickleback color and environmental lighting using methods independent of human perception to evaluate the information transmitted by male signals in a habitat where these signals are displayed. We also developed models of chromatic processing based on four cone photopigments (peak absorptions at 360, 445, 530, and 605 nm) characterized microspectrophotometrically in G. aculeatus and three other stickleback species. We show that a simple opponent mechanism receiving equally weighted inputs from cones with peak absorptions at 445 nm and 605 nm efficiently encodes variation in male throat colors. An orthogonal opponent mechanism—the difference between outputs of 530-nm cones and mean of outputs of 445- and 605-nm cones—produces a neural signal that could be used for species recognition and would be largely insensitive to variation in male throat color. We also show that threespine stickleback throats/photopigments are optimized for this coding scheme. These and other findings lead to testable hypotheses about the spectral processing mechanisms present in the threespine stickleback visual systems and the evolutionary interactions that have shaped this signal/receiver system.Abbreviations LWS long-wave sensitive - MWS middle-wave sensitive - SWS short-wave sensitive - UVS ultra-violet sensitive     

Evidence for trichromacy in the green peach aphid, Myzus persicae (Sulz.) (Hemiptera: Aphididae)

The green peach aphid, Myzus persicae (Hemiptera: Aphididae) is an important phytophagous pest of greenhouse and field crops. In the host finding process visual cues are of paramount importance. In order to contribute to the understanding of the perception of visual stimuli in this species, we measured the electroretinogram of alate female summer migrants of M. persicae. The spectral sensitivity was measured in 10nm steps under both dark and light adaptation from 320 to 640 nm. The dark adapted spectral sensitivity curve showed one maximum in the green region around 530 nm and a distinct shoulder between 500 and 510 nm. In presence of adapting light, a secondary blue-green peak (490 nm) and a third peak in the near UV (330-340 nm) were observed. From these results we conclude that M. persicae has three spectral types of photoreceptors.     

The inhibition of mitosis in tissue culture cells by blue light

Blue light (wavelength 350-480 nm) irradiation of the early mitotic (prophase and prometaphase) tissue culture cells at the dose of 50-3000 J/cm2 delay mitosis or completely block it at the metaphase. Cell sensitivity to the near UV light (wavelength 360 nm) was few times more as compared with the sensitivity to the visible light (wavelength 400-480 nm). Mitotic cells irradiated with the green light (wavelength more than 500 nm; dose up to 7500 J/cm2) completed division normally. The effect of the blue light did not depend on the presence of phenol red in tissue culture medium. Rhodamin 123 staining did not show any changes in the mitochondrial system in the irradiated mitotic cells. Blue light irradiation with the dose enough for the induction of mitotic delay appears to be insufficient to affect the proliferation of interphase cells.     

Spectral Sensitivity of the Barnacle, Balanus amphitrite

The extracellular ocellar potential was used to evaluate the spectral sensitivity of the ocellus of the barnacle, Balanus amphitrite. Maximum relative sensitivity was at 530–540 nm. Studies with chromatic adapting lights suggest that the receptors contain a single photopigment. The spectra were relatively broader in the dark as compared to the light-adapted state. This effect was shown to be due to an increase in the slope of the amplitude-intensity function, caused by light adaptation. Studies of tapetal fluorescence and corneal transmission indicate little effect of the ocellar media on the determination of sensitivity.     

A look into the invisible: ultraviolet-B sensitivity in an insect (Caliothrips phaseoli) revealed through a behavioural action spectrum

Caliothrips phaseoli, a phytophagous insect, detects and responds to solar ultraviolet-B radiation (UV-B; λ ≤ 315 nm) under field conditions. A highly specific mechanism must be present in the thrips visual system in order to detect this narrow band of solar radiation, which is at least 30 times less abundant than the UV-A (315–400 nm), to which many insects are sensitive. We constructed an action spectrum of thrips responses to light by studying their behavioural reactions to monochromatic irradiation under confinement conditions. Thrips were maximally sensitive to wavelengths between 290 and 330 nm; human-visible wavelengths (λ ≥ 400 nm) failed to elicit any response. All but six ommatidia of the thrips compound eye were highly fluorescent when exposed to UV-A of wavelengths longer than 330 nm. We hypothesized that the fluorescent compound acts as an internal filter, preventing radiation with λ > 330 nm from reaching the photoreceptor cells. Calculations based on the putative filter transmittance and a visual pigment template of λ_max = 360 nm produced a sensitivity spectrum that was strikingly similar to the action spectrum of UV-induced behavioural response. These results suggest that specific UV-B vision in thrips is achieved by a standard UV-A photoreceptor and a sharp cut-off internal filter that blocks longer UV wavelengths in the majority of the ommatidia.     

Photoreceptors and photopigments in a subterranean rodent,the pocket gopher (<Emphasis Type="Italic">Thomomys bottae</Emphasis>)

Pocket gophers (Thomomys bottae) are rodents that spend much of their lives in near-lightless subterranean burrows. The visual adaptations associated with this extreme environment were investigated by making anatomical observations of retinal organization and by recording retinal responses to photic stimulation. The size of the eye is within the normal range for rodents, the lens transmits light well down into the ultraviolet, and the retina conforms to the normal mammalian plan. Electroretinogram recording revealed the presence of three types of photopigments, a rod pigment with a spectral peak of about 495 nm and two types of cone pigment with respective peak values of about 367 nm (UV) and 505 nm (medium-wavelength sensitive). Both in terms of responsivity to lights varying in temporal frequency and in response recovery following intense light adaptation, the cone responses of the pocket gopher are similar to those of other rodents. Labeling experiments indicate an abundance of cones that reach densities in excess of 30,000 mm^–2. Cones containing UV opsin are found throughout the retina, but those containing medium-wavelength sensitive opsin are mostly restricted to the dorsal retina where coexpression of the two photopigments is apparently the rule. Rod densities are lower than those typical for nocturnal mammals.     

Photosensitivity spectrum of crayfish rhodopsin measured using fluorescence of metarhodopsin

Discrepancies exist among spectral measurements of sensitivity of crayfish photoreceptors, their absorption in situ, and the number and absorption spectra of crayfish photopigments that are extracted by digitonin solutions. We have determined the photosensitivity spectrum of crayfish rhodopsin in isolated rhabdoms using long wavelength fluorescence emission from crayfish metarhodopsin as an intrinsic probe. There is no measurable metarhodopsin in the dark-adapted receptor, so changes in the emission level are directly proportional to metarhodopsin concentration. We therefore used changes in metarhodopsin fluorescence to construct relaxation and saturation ("photoequilibrium") spectra, from which the photosensitivity spectrum of crayfish rhodopsin was calculated. This spectrum peaks at or approximately 530 nm and closely resembles the previously measured difference spectrum for total bleaches of dark-adapted rhabdoms. Measurements of the kinetics of changes in rhabdom fluorescence and in transmittance at 580 nm were compared with predictions derived from several model systems containing one or two photopigments. The comparison shows that only a single rhodopsin and its metarhodopsin are present in the main rhabdom of crayfish, and that other explanations must be sought for the multiple pigments seen in digitonin solution. The same analysis shows that there is no detectable formation of isorhodopsin in the rhabdom.