Photoreceptor processing speed and input resistance changes during light adaptation correlate with spectral class in the bumblebee, Bombus impatiens

Colour vision depends on comparison of signals from photoreceptors with different spectral sensitivities. However, response properties of photoreceptor cells may differ in ways other than spectral tuning. In insects, for example, broadband photoreceptors, with a major sensitivity peak in the green region of the spectrum (>500 nm), drive fast visual processes, which are largely blind to chromatic signals from more narrowly-tuned photoreceptors with peak sensitivities in the blue and UV regions of the spectrum. In addition, electrophysiological properties of the photoreceptor membrane may result in differences in response dynamics of photoreceptors of similar spectral class between species, and different spectral classes within a species. We used intracellular electrophysiological techniques to investigate response dynamics of the three spectral classes of photoreceptor underlying trichromatic colour vision in the bumblebee, Bombus impatiens, and we compare these with previously published data from a related species, Bombus terrestris. In both species, we found significantly faster responses in green, compared with blue- or UV-sensitive photoreceptors, although all 3 photoreceptor types are slower in B. impatiens than in B. terrestris. Integration times for light-adapted B. impatiens photoreceptors (estimated from impulse response half-width) were 11.3 ± 1.6 ms for green photoreceptors compared with 18.6 ± 4.4 ms and 15.6 ± 4.4 for blue and UV, respectively. We also measured photoreceptor input resistance in dark- and light-adapted conditions. All photoreceptors showed a decrease in input resistance during light adaptation, but this decrease was considerably larger (declining to about 22% of the dark value) in green photoreceptors, compared to blue and UV (41% and 49%, respectively). Our results suggest that the conductances associated with light adaptation are largest in green photoreceptors, contributing to their greater temporal processing speed. We suggest that the faster temporal processing of green photoreceptors is related to their role in driving fast achromatic visual processes.     

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.     

Spectral correspondence between visual spectral sensitivity and bioluminescence emission spectra in the click beetle Pyrophorus punctatissimus (Coleoptera: Elateridae)

The presence of two spectral mechanisms, near-ultraviolet and green (lambda(max)=545nm), is strongly suggested by electroretinographic visual spectral sensitivity curves obtained under dark and red chromatic adaptation conditions in the compound eyes of the click beetle Pyrophorus punctatissimus. The bioluminescence emission of the dorsal prothoracic lanterns is deep green (lambda(max)=543nm) and that of the ventral abdominal lantern is lime green (lambda(max)=556nm) in colour in P. punctatissimus. A broad green visual receptor would detect both deep green and lime green bioluminescent optical signals.     

Separation of Myzus (Nectarosiphon) antirrhinii (Macchiati) from Myzus (N.) persicae (Sulzer) and related species in Europe (Homoptera: Aphididae)

Abstract. Multivariate morphometric analysis (canonical variates) was used to discriminate between closely related taxa within the Myzus persicae group. It is demonstrated that dark green, anholocyclic populations with a 2n=13 or 2n=14 karyotype in Europe, hitherto treated as a form or biotype of M.persicae, all conform to a discrete morphometric grouping and should therefore be treated as a separate taxonomic entity, permanently isolated from the M.persicae gene pool. It is suggested that this taxon was first described as Siphonophora antirrhinii by Macchiati in 1883. A discriminant function is provided to separate most individual apterae of antirrhinii from those of persicae. Myzus icelandicus Blackman sp.n., on Caryophyllaceae and other plants in Iceland, is distinguished from M.polaris H.R.L. and M.certus (Walker), and a key is given to apterous virginoparae of the species of the M.persicae group in Europe.     

Light emitting diode(LED)‐based trapping of the greenhouse whitefly (Trialeurodes vaporariorum)

Visual traps like yellow sticky card traps are used for monitoring and control of the greenhouse whitefly (Trialeurodes vaporariorum). However, reflected intensity (brightness) and hence, attractiveness depend on the ambient light conditions, and the colour (wavelength) might not fit with the sensitivity of whitefly photoreceptors. The use of light emitting diodes (LEDs) is a promising approach to increase the attractiveness, specificity and adaptability of visual traps. We constructed LED‐based visual traps equipped with blue and green high‐power LEDs and ultraviolet (UV) standard LEDs according to the putative spectral sensitivities of the insects' photoreceptors. In a series of small‐scale choice and no‐choice recapture experiments, the factors time of day as well as light intensity and light quality (colour) of LED traps were studied in terms of attractiveness compared to yellow traps without LEDs. Green LED traps (517 nm peak wavelength) were comparably attractive in no‐choice experiments but clearly preferred over yellow traps in all choice experiments. The time of day had a clear effect on the flight activity of the whiteflies and thereby on the trapping success. Blue LEDs (474 nm) suppressed the attractiveness of the light traps when combined with green LEDs suggesting that a yet undetected photoreceptor, sensitive for blue light, and an inhibiting interaction with the green receptor, might exist in T. vaporariorum. In choice experiments between LED traps emitting green light only or in combination with UV (368 nm), the green‐UV combination was preferred. In no‐choice night‐time experiments, UV LEDs considerably increased whitefly flight activity and efficacy of trapping. Most likely, the reason for the modifying effect of UV is the stimulating influence on flight activity. In conclusion, it seems that the use of green LEDs alone or in combination with UV LEDs could be an innovative option for improving attractiveness of visual traps.     

Physiological basis of phototaxis to near-infrared light in Nephotettix cincticeps

In a previous study of the phototaxis of green rice leafhoppers, Nephotettix cincticeps (Hemiptera, Cicadellidae), we found positive responses to 735 nm light. Here, we investigated the mechanism underlying this sensitivity to near-infrared light. We first measured the action spectrum using a Y-maze with monochromatic lights from 480 to 740 nm. We thus found that the action spectrum peaks at 520 nm in the tested wavelength range, but that a significant effect is still observed at 740 nm, albeit with a sensitivity 5 log units lower than the peak. Second, we measured the spectral sensitivity of the eye, and found that the sensitivity in the long-wavelength region parallels the behaviorally determined action spectrum. We further identified mRNAs encoding opsins of ultraviolet, blue, and green-absorbing visual pigments, and localized the mRNAs in the ommatidia by in situ hybridization. The electrophysiology, molecular biology and the anatomy of the eye together indicate that the eyes of N. cincticeps do not contain true “red” receptors, but rather that the behavioral response to near-infrared light is mediated by the tail sensitivity of the green receptors in the long-wavelength region of the spectrum.     

Spectral sensitivity of the cod,Gadus morhua L.

The spectral sensitivity of the cod was determined under both dark adapted and light adapted conditions in the laboratory. Cod were trained by cardiac conditioning to detect a difference in radiance between an image of spots and the background radiance of a screen. Thresholds for this response were measured for a range of different wavelengths, and expressed as quantum adjusted values. Electroretino‐graphic studies were also performed on the eyes of cod, and spectral sensitivity curves prepared. Under dark adapted conditions both the behavioural and e.r.g. derived curves showed greatest sensitivity in the blue/green at 490 nm, matching the absorption curve for rhodopsin. A secondary peak in the behaviourally derived curve in the green/yellow at 550 nm indicated that a population of yellow cones may be implicated with the rods in scotopic vision. Under light adapted conditions the behavioural curves showed a shift to the blue, perhaps indicating an adaption to the high red content of the illuminating source. The e.r.g. curve showed greatest sensitivity to blue/green, as in the scotopic experiments but with an enhanced response at 550 nm, indicating greater cone activity. It is suggested that there is complex interaction between rods and cones in the cod retina, both types of receptor being active over a wide range of light intensities.     

Molecular basis of spectral tuning in the red- and green-sensitive (M/LWS) pigments in vertebrates

Vertebrate vision is mediated by five groups of visual pigments, each absorbing a specific wavelength of light between ultraviolet and red. Despite extensive mutagenesis analyses, the mechanisms by which contemporary pigments absorb variable wavelengths of light are poorly understood. We show that the molecular basis of the spectral tuning of contemporary visual pigments can be illuminated only by mutagenesis analyses using ancestral pigments. Following this new principle, we derive the "five-sites" rule that explains the absorption spectra of red and green (M/LWS) pigments that range from 510 to 560 nm. Our findings demonstrate that the evolutionary method should be used in elucidating the mechanisms of spectral tuning of four other pigment groups and, for that matter, functional differentiations of any other proteins.     

Out of the blue: the spectral sensitivity of hummingbird hawkmoths

The European hummingbird hawkmoth Macroglossum stellatarum is a diurnal nectar forager like the honeybee, and we expect similarities in their sensory ecology. Using behavioural tests and electroretinograms (ERGs), we studied the spectral sensitivity of M. stellatarum. By measuring ERGs in the dark-adapted eye and after adaptation to green light, we determined that M. stellatarum has ultraviolet (UV), blue and green receptors maximally sensitive at 349, 440 and 521 nm, and confirmed that green receptors are most frequent in the retina. To determine the behavioural spectral sensitivity (action spectrum) of foraging moths, we trained animals to associate a disk illuminated with spectral light, with a food reward, and a dark disk with no reward. While the spectral positions of sensitivity maxima found in behavioural tests agree with model predictions based on the ERG data, the sensitivity to blue light was 30 times higher than expected. This is different from the honeybee but similar to earlier findings in the crepuscular hawkmoth Manduca sexta. It may indicate that the action spectrum of foraging hawkmoths does not represent their general sensory capacity. We suggest that the elevated sensitivity to blue light is related to the innate preference of hawkmoths for blue flowers.     

Spectral sensitivity of the compound eyes in two day-active fireflies (Coleoptera: Lampyridae: Lucidota)

Summary The electroretinographic visual spectral sensitivity functions in day-active fireflies Lucidota luteicollis and Lucidota atra show a broad green sensitivity and a shoulder in the near-ultraviolet region of the spectrum (Figs. 1, 2) as is commonly found among day-active insects. The nomogram for P530 visual pigment matches the spectral sensitivity curves in the green. The adult L. luteicollis retains its larval bioluminescent light organ which has a peak emission at 562 nm. The _max of the ERG spectral sensitivity does not match the bioluminescent peak (Fig. 1B) as it does in twilight- and dark-active fireflies. Some relevant behavioural observations with respect to mating are presented.     

Spectral sensitivity of vision and bioluminescence in the midwater shrimp Sergestes similis

In the oceanic midwater environment, many fish, squid, and shrimp use luminescent countershading to remain cryptic to silhouette-scanning predators. The mid-water penaeid shrimp, Sergestes similis Hansen, responds to downward-directed light with a dim bioluminescence that dynamically matches the spectral radiance of oceanic down-welling light at depth. Although the sensory basis of luminescent countershading behavior is visual, the relationship between visual and behavioral sensitivity is poorly understood. In this study, visual spectral sensitivity, based on microspectrophotometry and electrophysiological measurements of photoreceptor response, is directly compared to the behavioral spectral efficiency of luminescent countershading. Microspectrophotometric measurements on single photoreceptors revealed only a single visual pigment with peak absorbance at 495 nm in the blue-green region of the spectrum. The peak electrophysiological spectral sensitivity of dark-adapted eyes was centered at about 500 nm. The spectral efficiency of luminescent countershading showed a broad peak from 480 to 520 nm. Both electrophysiological and behavioral data closely matched the normalized spectral absorptance curve of a rhodopsin with lambda(max) = 495 nm, when rhabdom length and photopigment specific absorbance were considered. The close coupling between visual spectral sensitivity and the spectral efficiency of luminescent countershading attests to the importance of bioluminescence as a camouflage strategy in this species.     

Phototactic Behavior of <Emphasis Type="Italic">Scleroderma guani</Emphasis> (Hymenoptera: Bethylidae) - Parasitoid of <Emphasis Type="Italic">Pissodes punctatus</Emphasis> (Coleoptera: Curculionidae)

Although light trap can be used to control pest populations, they can also kill the natural enemies of pests. Scleroderma guani (Hymenoptera: Bethylidae) is a parasitoid of a bark-weevil Pissodes punctatus (Coleoptera: Curculionidae). To understand the phototactic behavior of S. guani, we investigated its diurnal and nocturnal behavior, then examined its phototactic response to nine monochromatic lights and to five intensities of the two most attractive lights. Our results showed that S. guani is most active during the day, while remain still in a dark room or at night. S. guani showed a positive response both to a broad spectrum of monochromatic light and total light (natural light), which implies a broad sensitivity to the light spectrum. S. guani was most sensitive to blue (450 nm) and green (549 nm) lights, suggesting its visual system composed of blue and green receptors. S. guani was least sensitive to ultraviolet (340 nm) light, which may be caused by long-term mass rearing and propagating under artificial conditions. Furthermore, low intensities elicited a positive phototactic response, while high intensities showed a decreased trend under both blue and green lights. Thus, S. guani is a phototactic insect which shows preferences for light in both color and intensity. This study suggests that light trap can only be utilized to control the adult P. punctatus during and after its peak emergence, due to the overlap in the spectral sensitivity of both pest and parasitoid adults.     

SPECTRAL SENSITIVITY IN TWO SPECIES OF PINNIPEDS (PHOCA VITULINA AND OTARIA FLAVESCENS)

Spectral sensitivity was measured in air in the light adapted state in two harbor seals and a South American sea lion using a behavioral training technique. Increment thresholds were determined in a spectral range from 390 nm to 670 nm in a simultaneous two‐choice discrimination task. The spectral sensitivity curves show two maxima in sensitivity, one main peak with a maximum around 500 nm in the harbor seal and around 550 nm in the South American sea lion, and a second, smaller peak with a maximum in the range of 410 nm in both species. The broad shape and the position of the maximum of the spectral sensitivity curve of the harbor seals suggests that even under photopic conditions both rods and cones are contributing to the measurements since harbor seals possess only one cone type. The maximum sensitivity in the green part of the spectrum may indicate an adaptation to a specific underwater environment.     

Spectral preferences of Lycorma delicatula (Hemiptera: Fulgoridae)

Many insects prefer lights with certain spectral properties, and such preferences may be associated with behavioral contexts such as mating, host finding or dispersal. Lycorma delicatula (Hemiptera: Fulgoridae) is a newly invasive species in Korea and is spreading rapidly. It is diurnal and may rely on visual cues for orientation. We conducted a series of three phototaxis experiments to understand spectral preferences in L. delicatula: light/dark choice, UV/white light choice, and color preference experiments. Nymphs of the third and final stages as well as adults were used for these experiments. In the light/dark choice, the orientation of L. delicatula was bimodal between the white light and darkness, for all stages and both sexes. In a choice of UV (395–410 nm wavelengths) vs. white light, L. delicatula of both sexes and all stages preferred the UV light. In the color preference experiment where insects had a choice of four colors in a circular arena, L. delicatula stayed significantly longer in the blue light than in white, yellow or green lights. Overall, nymphs and adults of L. delicatula oriented toward lights with shorter wavelengths, and this orientation was consistent throughout all stages, regardless of sex. It is necessary to investigate the behavioral contexts under which L. delicatula prefers the UV and blue lights.     

Vision in the fireflyPhoturis lucicrescens (Coleoptera: Lampyridae): Spectral sensitivity and selective adaptation in the compound eye

Summary The waveform of the electroretinograms (ERGs) recorded from the compound eyes in the dark-active fireflyPhoturis lucicrescens was different in the short (near-UV and violet) and long (green-yellow) wavelengths (Fig. 1). The spectral sensitivity curves in the dark and chromatic adaptation conditions suggested the presence of receptor types in the short (near-UV, Fig. 4, and violet, Fig. 5) and long wavelength (green; max 550 nm, Figs. 3–5) regions of the spectrum. The green peak is in correspondence with the species' bioluminescence emission peak at 554 nm (Fig. 3c).Abbreviations DA dark-adapted - ERG electroretinogram - VP visual pigment Contribution No. 1112 of the McCollum-Pratt Institute and Department of Biology, The Johns Hopkins University     

Short-wavelength light sensitivity of circadian, pupillary, and visual awareness in humans lacking an outer retina

As the ear has dual functions for audition and balance, the eye has a dual role in detecting light for a wide range of behavioral and physiological functions separate from sight. These responses are driven primarily by stimulation of photosensitive retinal ganglion cells (pRGCs) that are most sensitive to short-wavelength ( approximately 480 nm) blue light and remain functional in the absence of rods and cones. We examined the spectral sensitivity of non-image-forming responses in two profoundly blind subjects lacking functional rods and cones (one male, 56 yr old; one female, 87 yr old). In the male subject, we found that short-wavelength light preferentially suppressed melatonin, reset the circadian pacemaker, and directly enhanced alertness compared to 555 nm exposure, which is the peak sensitivity of the photopic visual system. In an action spectrum for pupillary constriction, the female subject exhibited a peak spectral sensitivity (lambda(max)) of 480 nm, matching that of the pRGCs but not that of the rods and cones. This subject was also able to correctly report a threshold short-wavelength stimulus ( approximately 480 nm) but not other wavelengths. Collectively these data show that pRGCs contribute to both circadian physiology and rudimentary visual awareness in humans and challenge the assumption that rod- and cone-based photoreception mediate all "visual" responses to light.     

蝗虫复眼光谱敏感性的比较研究——Ⅱ.视网膜电图的光谱敏感曲线

(1)用视网膜电图(ERG)方法测定了9种蝗虫在黑暗、蓝光和橙光适应下的光谱敏感性。(2)9种蝗虫的碚适应光谱敏感曲线峰值均在520—546nm 之间。(3)橙光或蓝光明适应导致不同程度的峰值位移,蓝区的相对敏感性提高,这与光引起屏蔽色素移动效应有关。(4)黑背蝗和稻蝗复眼表面均没有黑白间,橙光适应时出现第二个峰值在蓝区,而蓝光适应则压抑蓝区的敏感性。可能这两种蝗虫还具有蓝敏视色素。(5)佛蝗和黄脊蝗复眼表面均有明显的黑白相间的区域,在有色光适应下这两种蝗虫的光谱敏感性变化最小,没有证据说明多于一种光敏色素。     

Laser-induced fluorescence (LIF) spectra of herbaceous and woody pre- and post-digested plant material

Filtrate from pre- and post-digested plant material was exposed to 355-nm pulsed laser light and the subsequent laser-induced fluorescence (LIF) was recorded. Similarities and differences among spectra from 20 materials are discussed. Each material was replicated once, dried, ground, and exposed to chloroform (CHCl₃) for 24 h. The material represented aged (1 to 18 years old) plants from different herbaceous (grasses and forbs) and woody plant life forms. Mean peak fluorescence recorded among materials differed (P < 0.0001) in both wavelength and peak amplitude (counts) across the spectral range (387 to 788 nm). Peak fluorescence was evaluated within each of three arbitrary color categories, blue near 455 nm and red near 674 nm, while only 16 of the materials produced a green peak near 528 nm. In general, the blue and green fluorescence peaks were broad while the red peak was narrow. Mean peak counts were largest in the red range. Varying amounts of laser beam absorption occurred among the materials evaluated due to different concentrations of filtrate and different absorption efficiencies; therefore, amplitude data (counts) were not used to determine statistical differences among materials. To overcome difficulties attributed to the raw count data, red/blue, red/green and blue/green count ratios within replicates were calculated. Using all three count ratios in a multivariate analysis of variance, the 16 materials could be separated into nine different (P < 0.05) material groupings. The LIF technique may provide a reliable means to separate ground pre- and post-digested plant materials following further research into determining what fluorophores are producing the spectral signatures and how sample preparation affect peak wavelengths.     

Opsin evolution and expression in Arthropod compound Eyes and Ocelli: Insights from the cricket Gryllus bimaculatus

ABSTRACT: BACKGROUND: Opsins are key proteins in animal photoreception. Together with a light-sensitive group, the chromophore, they form visual pigments which initiate the visual transduction cascade when photoactivated. The spectral absorption properties of visual pigments are mainly determined by their opsins, and thus opsins are crucial to understand the adaptations of animal eyes. Studies on the phylogeny and expression pattern of opsins have received considerable attention, but our knowledge about insect visual opsins is still limited. Up to now, researchers have focused on holometabolous insects, while general conclusions require sampling from a broader range of taxa. We have therefore investigated visual opsins in the ocelli and compound eyes of the two-spotted cricket Gryllus bimaculatus, a hemimetabolous insect. RESULTS: Phylogenetic analyses place all identified cricket sequences within the three main visual opsin clades of insects. We assign three of these opsins to visual pigments found in the compound eyes with peak absorbances in the green (515 nm), blue (445 nm) and UV (332 nm) spectral range. Their expression pattern divides the retina into distinct regions: (1) the polarization-sensitive dorsal rim area with blue- and UV-opsin, (2) a newly-discovered ventral band of ommatidia with blue- and green-opsin and (3) the remainder of the compound eye with UV- and green-opsin. In addition, we provide evidence for two ocellar photopigments with peak absorbances in the green (511 nm) and UV (350 nm) spectral range, and with opsins that differ from those expressed in the compound eyes. CONCLUSIONS: Our data show that cricket eyes are spectrally more specialized than has previously been assumed, suggesting that similar adaptations in other insect species might have been overlooked. The arrangement of spectral receptor types within some ommatidia of the cricket compound eyes differs from the generally accepted pattern found in holometabolous insect taxa and awaits a functional explanation. From the opsin phylogeny, we conclude that gene duplications, which permitted differential opsin expression in insect ocelli and compound eyes, occurred independently in several insect lineages and are recent compared to the origin of the eyes themselves.     

桃蚜对不同单色光趋性反应的测定

为了探讨蚜虫对不同色光选择反应的定量指标, 采用滤光片技术测定了有翅和无翅桃蚜Myzus persicae对不同波长单色光的趋性反应。结果表明: 有翅蚜对490~550 nm范围内的单色光表现出明显趋性, 其中对538.9和549.9 nm的绿偏黄色光趋性最强, 平均位移分别达25.29和22.97 cm, 其次为491.5 nm的蓝绿色光, 而对于波长576.0 nm的黄色光并没有表现出明显趋性。无翅蚜对不同单色光的趋性反应则没有明显的峰值, 最高相对平均位移仅1.41。行为测定结果与前人电生理测定的结果基本一致, 说明以位移作为小体昆虫趋光性强弱的指标是可靠的。