Many insects exploit the partial plane polarization of skylight for visual compass orientation and/or navigation. In the present study, using a tethering system, we investigated how flying bees ...respond to polarized light stimuli. The behavioral responses of honeybees (
) to a zenithal polarized light stimulus were observed using a tethered animal in a flight simulator. Flight direction of the bee was recorded by monitoring the horizontal movement of its abdomen, which was strongly anti-correlated with its torque. When the e-vector orientation of the polarized light was rotated clockwise or counterclockwise, the bee responded with periodic right-and-left abdominal movements; however, the bee did not show any clear periodic movement under the static e-vector or depolarized stimulus. The steering frequency of the bee was well coordinated with the e-vector rotation frequency of the stimulus, indicating that the flying bee oriented itself to a certain e-vector orientation, i.e. exhibited polarotaxis. The percentage of bees exhibiting clear polarotaxis was much smaller under the fast stimulus (3.6 deg s
) compared with that under a slow stimulus (0.9 or 1.8 deg s
). Bees did not demonstrate any polarotactic behavior after the dorsal rim area of the eyes, which mediates insect polarization vision in general, was bilaterally covered with black paint. Preferred e-vector orientations under the clockwise stimulus varied among individuals and distributed throughout -90 to 90 deg. Some bees showed similar preferred e-vector orientations for clockwise and counterclockwise stimuli whereas others did not. Our results strongly suggest that flying honeybees utilize the e-vector information from the skylight to deduce their heading orientation for navigation.
Many insects utilise the polarisation pattern of the sky to adjust their travelling directions. The extraction of directional information from this sky-wide cue is mediated by specialised ...photoreceptors located in the dorsal rim area (DRA). While this part of the eye is known to be sensitive to the ultraviolet, blue or green component of skylight, the latter has only been observed in insects active in dim light. To address the functional significance of green polarisation sensitivity, we define the spectral and morphological adaptations of the DRA in a nocturnal ball-rolling dung beetle-the only family of insects demonstrated to orient to the dim polarisation pattern in the night sky. Intracellular recordings revealed polarisation-sensitive green photoreceptors in the DRA of Escarabaeus satyrus. Behavioural experiments verified the navigational relevance of this finding. To quantify the adaptive value of green sensitivity for celestial orientation at night, we also obtained the polarisation properties of the night sky in the natural habitat of the beetle. Calculations of relative photon catch revealed that under a moonlit sky the green-sensitive DRA photoreceptors can be expected to catch an order of magnitude more photons compared with the UV-sensitive photoreceptors in the main retina. The green-sensitive photoreceptors - which also show a range of morphological adaptations for enhanced sensitivity - provide E. satyrus with a highly sensitive system for the extraction of directional information from the night sky.
For compass orientation many insects rely on the pattern of sky polarization, but some species also exploit the sky chromatic contrast. Desert locusts, Schistocerca gregaria, detect polarized light ...through a specialized dorsal rim area (DRA) in their compound eye. To better understand retinal mechanisms underlying visual navigation, we compared opsin expression, spectral and polarization sensitivities and response-stimulus intensity functions in the DRA and main retina of the locust. In addition to previously characterized opsins of long-wavelength-absorbing (Lo1) and blue-absorbing visual pigments (Lo2), we identified an opsin of an ultraviolet-absorbing visual pigment (LoUV). DRA photoreceptors exclusively expressed Lo2, had peak spectral sensitivities at 441 nm and showed high polarization sensitivity (PS 1.3-31.7). In contrast, ommatidia in the main eye co-expressed Lo1 and Lo2 in five photoreceptors, expressed Lo1 in two proximal photoreceptors, and Lo2 or LoUV in one distal photoreceptor. Correspondingly, we found broadband blue- and green-peaking spectral sensitivities in the main eye and one narrowly tuned UV peaking receptor. Polarization sensitivity in the main retina was low (PS 1.3-3.8). V-log I functions in the DRA were steeper than in the main retina, supporting a role in polarization vision. Desert locusts occur as two morphs, a day-active gregarious and a night-active solitarious form. In solitarious locusts, sensitivities in the main retina were generally shifted to longer wavelengths, particularly in ventral eye regions, supporting a nocturnal lifestyle at low light levels. The data support the role of the DRA in polarization vision and suggest trichromatic colour vision in the desert locust.
Polarized light-based navigation in insects is facilitated by a polarization-sensitive part of the eye, the dorsal rim area (DRA). Existing methods to study the anatomy of the DRA are destructive and ...time-consuming. We presented a novel method for DRA localization, dissection, and measurement using 3D volumetric images from X-ray micro-computed tomography in combination with 2D photographs. Applying the method on size-polymorphic buff-tailed bumblebees, Bombus terrestris, we found that the DRA was easily obtainable from photographs of the dorsal eye region. Allometric analysis of the DRA in relation to body size in B. terrestris showed that it increased with the body size but not at the same rate. By localizing the DRA of individual bumblebees, we could also perform individual-level descriptions and inter-individual comparisons between the ommatidial structures (lens, crystalline cones, rhabdoms) of three different eye regions (DRA, non-DRA, proximate to DRA). One feature distinct to the bumblebee DRA was the smaller dimension of the crystalline cones in comparison to other regions of the eye. Using our novel methodology, we provide the first individual-level description of DRA ommatidial features and a comparison of how the DRA varies with body size in bumblebees.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, UILJ, UKNU, UL, UM, UPUK
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 for understanding 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.
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.
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.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Insects are well known to orient using celestial cues. The pattern of polarised skylight is the dominant celestial compass information that insects use, which they detect using a specialised set of ...ommatidia. The number of ommatidia decreases with body size, and it is unknown how this reduction in the number of ommatidia affects the precision of orienting using celestial cues. We investigated this in eight different ant species that had varying numbers of ommatidia. We captured ants returning home, displaced them to an unfamiliar location and measured their precision in determining heading direction using celestial cues. The heading direction of the ants measured at a fixed distance from the release and also at a distance scaled to their body size was not correlated with the number of ommatidia. However, both the path straightness and walking speed were lower in smaller ants indicating the ability to orient at a finer scale was affected by miniaturisation.
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EMUNI, FIS, FZAB, GEOZS, GIS, IJS, IMTLJ, KILJ, KISLJ, MFDPS, NLZOH, NUK, OBVAL, OILJ, PNG, SAZU, SBCE, SBJE, SBMB, SBNM, UKNU, UL, UM, UPUK, VKSCE, ZAGLJ
In many insect species, photoreceptors of a small dorsal rim area of the eye are specialized for sensitivity to the oscillation plane of polarized skylight and, thus, serve a role in sky compass ...orientation. To further understand peripheral mechanisms of polarized-light processing in the optic lobe, we have studied the projections of photoreceptors and their receptive fields in the main eye and dorsal rim area of the desert locust, a model system for polarization vision analysis. In both eye regions, one photoreceptor per ommatidium, R7, has a long visual fiber projecting through the lamina to the medulla. Axonal fibers from R7 receptors of the dorsal rim area have short side branches throughout the depth of the dorsal lamina and maintain retinotopic projections to the dorsal medulla following the first optic chiasma. Receptive fields of dorsal rim photoreceptors are considerably larger (average acceptance angle 33°) than those of the main eye (average acceptance angle 2.04°) and, taken together, cover almost the entire sky. The data challenge previous reports of two long visual fibers per ommatidium in the main eye of the locust and provide data for future analysis of peripheral networks underlying polarization opponency in the locust brain.
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EMUNI, FIS, FZAB, GEOZS, GIS, IJS, IMTLJ, KILJ, KISLJ, MFDPS, NLZOH, NUK, OILJ, PNG, SAZU, SBCE, SBJE, SBMB, SBNM, UKNU, UL, UM, UPUK, VKSCE, ZAGLJ
本研究では，チャバネアオカメムシPlautia crossota ...staliの複眼の外部および内部構造を詳細に観察し，視覚定位行動において機能的な役割を果たす複眼の構造的部域差について検討した。チャバネアオカメムシの赤道面前方部域の個眼は，背側や腹側の領域に比べて個眼間密度が高く，角膜レンズや円錐細胞などの光学系構造が大きいことや，長い感桿構造をもつことから，空間分解能と感度が共に高い部域である複眼のフォベアであることが強く示唆された。本種は光に対して強い走光性を示すが，本種が上方光源に対して視覚定位する際，姿勢を変えて複眼の赤道面前方部付近のフォベアで光をとらえていることが明らかになった。また，複眼の背側領域の一部には，中心視細胞であるR7とR8の感桿の微絨毛の配向は直交配向で偏光アナライザー機能をもち，微絨毛の直交配向は途中で変化せず，光軸は天頂方向へ向き，感桿の長さが短く，角膜レンズは他の部域に比べて平らで円錐細胞も短い，といった天空の偏光パターンの弁別に関わる複眼背縁部(dorsal rim area)の構造特徴を示す個眼領域が確認された。
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