@article {22898, title = {Flexible responses to visual and olfactory stimuli by foraging Manduca sexta: larval nutrition affects adult behaviour}, journal = {Proceedings of the Royal Society B}, volume = {276}, year = {2017}, pages = {2739-2745}, chapter = {2739}, abstract = {

\"Here, we show that the consequences of deficient micronutrient (\β-carotene) intake during larval stages of Manduca sexta are carried across metamorphosis, affecting adult behaviour. Our manipulation of larval diet allowed us to examine how developmental plasticity impacts the interplay between visual and olfactory inputs on adult foraging behaviour. Larvae of M. sexta were reared on natural (Nicotiana tabacum) and artificial laboratory diets containing different concentrations of \β-carotene (standard diet, low \β-carotene, high \β-carotene and cornmeal). This vitamin-A precursor has been shown to be crucial for photoreception sensitivity in the retina of M. sexta. After completing development, post-metamorphosis, starved adults were presented with artificial feeders that could be either scented or unscented. Regardless of their larval diet, adult moths fed with relatively high probabilities on scented feeders. When feeders were unscented, moths reared on tobacco were more responsive than moths reared on \β-carotene-deficient artificial diets. Strikingly, moths reared on artificial diets supplemented with increasing amounts of \β-carotene (low \β and high \β) showed increasing probabilities of response to scentless feeders. We discuss these results in relationship to the use of complex, multi-modal sensory information by foraging animals.\"

}, keywords = {BEHAVIOUR, CAROTENOID, ECOLOGY, LARVAL NUTRITION, MANDUCA, SPHINGIDAE, VISION}, doi = {10.1098/rspb.2009.0456}, author = {Goyret, J. and Kelber, A. and Pfaff, M. and Raguso, R.A.} } @article {22895, title = {Crepuscular and nocturnal illumination and its effects on color perception by the nocturnal hawkmoth Deilephila elpenor}, journal = {Journal of Experimental Biology}, volume = {209}, year = {2006}, month = {03/2006}, pages = {789-800}, chapter = {789}, abstract = {

\"Recent studies have shown that certain nocturnal insect and vertebrate species have true color vision under nocturnal illumination. Thus, their vision is potentially affected by changes in the spectral quality of twilight and nocturnal illumination, due to the presence or absence of the moon,artificial light pollution and other factors. We investigated this in the following manner. First we measured the spectral irradiance (from 300 to 700 nm) during the day, sunset, twilight, full moon, new moon, and in the presence of high levels of light pollution. The spectra were then converted to both human-based chromaticities and to relative quantum catches for the nocturnal hawkmoth Deilephila elpenor, which has color vision. The reflectance spectra of various flowers and leaves and the red hindwings of D. elpenor were also converted to chromaticities and relative quantum catches. Finally, the achromatic and chromatic contrasts (with and without von Kries color constancy) of the flowers and hindwings against a leaf background were determined under the various lighting environments. The twilight and nocturnal illuminants were substantially different from each other, resulting in significantly different contrasts. The addition of von Kries color constancy significantly reduced the effect of changing illuminants on chromatic contrast, suggesting that, even in this light-limited environment,the ability of color vision to provide reliable signals under changing illuminants may offset the concurrent threefold decrease in sensitivity and spatial resolution. Given this, color vision may be more common in crepuscular and nocturnal species than previously considered.\"

}, keywords = {COLOUR VISION, DEILEPHILA, OPTICS, SPHINGIDAE, VISION}, doi = {10.1242/jeb.02053}, author = {Johnsen, S. and Kelber, A. and Warrant, E. and Sweeney, A.M. and Widder, E.A. and Lee Jr, R.L. and Hernández-Andrés, J.} } @article {22894, title = {The relative importance of olfaction and vision in a diurnal and a nocturnal hawkmoth}, journal = {Journal of Comparative Physiology A}, volume = {192}, year = {2006}, month = {12/2006}, pages = {431-437}, chapter = {431}, abstract = {

\"Nectar-feeding animals can use vision and olfaction to find rewarding flowers and different species may give different weight to the two sensory modalities. We have studied how a diurnal or nocturnal lifestyle affects the weight given to vision and olfaction. We tested na\ïve hawkmoths of two species in a wind tunnel, presenting an odour source and a visual stimulus. Although the two species belong to the same subfamily of sphingids, the Macroglossinae, their behaviour was quite different. The nocturnal Deilephila elpenor responded preferably to the odour while the diurnal Macroglossum stellatarum strongly favoured the visual stimulus. Since a nocturnal lifestyle is ancestral for sphingids, the diurnal species, M. stellatarum, has evolved from nocturnal moths that primarily used olfaction. During bright daylight visual cues may have became more important than odour.\"

}, keywords = {DEILEPHILA, MACROGLOSSUM, OLFACTION, PHYSIOLOGY, SPHINGIDAE, VISION}, doi = {10.1007/s00359-005-0081-6}, author = {Balkenius, A. and Ros{\'e}n, W. and Kelber, A.} } @article {22893, title = {The effect of decoupling olfactory and visual stimuli on the foraging behavior of Manduca sexta}, journal = {Journal of Experimental Biology}, volume = {210}, year = {2007}, pages = {1398-1405}, chapter = {1398}, abstract = {

\"Within an appetitive context, Manduca sexta, a nectivorous nocturnal hawkmoth, can be attracted by a range of stimuli including floral volatiles and visual display, carbon dioxide and water vapor. Several studies on this and other flower-visiting insects have shown how olfactory and visual stimulation play (or do not play) a role in attraction and feeding. Nevertheless, these studies have consistently manipulated stimuli in a{\textquoteleft}presence\–absence\&$\#$39; manner. Here, we experimentally decoupled the presentation of both stimuli spatially and temporally in a wind tunnel, rather than entirely eliminating either one, and found that the decision-making process based on these stimuli is more flexible and complex than previously asserted. Manduca sexta was most responsive when both cues were present and emanated from the same source. When stimuli were spatially separated, responsiveness levels were comparable to those elicited by a single stimulus. However, transient olfactory stimulation either before or after visually guided approach (temporal decoupling) enhanced responsiveness to an odorless visual target. Additionally, searching times were increased by either a transient olfactory stimulation before take-off or by having the flower model spatially separated from the odor source tracked by the moths. Finally,in a dual-choice experiment, moths showed a strong bias for the visual display over the odor plume, suggesting the former to be the ultimate indicator of a nectar source. Our manipulation of floral cues shows that the feeding behavior of M. sexta, and probably of other nectivorous insects, is based not only on the sensory stimulation per se but also on the temporal and spatial pattern in which these stimuli are perceived.\"

}, keywords = {MANDUCA, OLFACTION, SPHINGIDAE, TEMPORAL DECOUPLING, VISION}, doi = {10.1242/jeb.02752}, author = {Goyret, J. and Markwell, P.M. and Raguso, R.A.} } @article {22892, title = {Interaction of visual and odour cues in the mushroom body of the hawkmoth Manduca sexta}, journal = {Journal of Experimental Biology}, volume = {212}, year = {2009}, pages = {535-541}, chapter = {535}, abstract = {

\"The responses to bimodal stimuli consisting of odour and colour were recorded using calcium-sensitive optical imaging in the mushroom bodies of the hawkmoth Manduca sexta. The results show that the activity in the mushroom bodies is influenced by both olfaction and vision. The interaction between the two modalities depends on the odour and the colour of the visual stimulus. A blue stimulus suppressed the response to a general flower scent (phenylacetaldehyde). By contrast, the response to a green leaf scent (1-octanol) was enhanced by the presence of the blue stimulus. A green colour had no influence on these odours but caused a marked increase in the response to an odour component (benzaldehyde) of the hawkmoth-pollinated Petunia axillaris.\"

}, keywords = {BRAIN, MANDUCA, MUSHROOM BODY, NEUROLOGY, OLFACTION, PETUNIA, SOLANACEAE, SPHINGIDAE, VISION}, doi = {10.1242/jeb.021220}, author = {Balkenius, A. and Bisch-Knaden, S. and Hansson, B.S.} } @article {22891, title = {Resolving the trade-off between visual sensitivity and spatial acuity{\textemdash}lessons from hawkmoths}, journal = {Integrative and Comparative Biology}, volume = {57}, year = {2017}, pages = {1093-1103}, chapter = {1093}, abstract = {

\"The visual systems of many animals, particularly those active during the day, are optimized for high spatial acuity. However, at night, when photons are sparse and the visual signal competes with increased noise levels, fine spatial resolution cannot be sustained and is traded-off for the greater sensitivity required to see in dim light. High spatial acuity demands detectors and successive visual processing units whose receptive fields each cover only a small area of visual space, in order to reassemble a finely sampled and well resolved image. However, the smaller the sampled area, the fewer the photons that can be collected, and thus the worse the visual sensitivity becomes\—leading to the classical trade-off between sensitivity and resolution. Nocturnal animals usually resolve this trade-off in favour of sensitivity, and thus have lower spatial acuity than their diurnal counterparts. Here we review results highlighting how hawkmoths, a highly visual group of insects with species active at different light intensities, resolve the trade-off between sensitivity and spatial resolution. We compare adaptations both in the optics and retina, as well as at higher levels of neural processing in a nocturnal and a diurnal hawkmoth species, and also give a perspective on the behavioral consequences. We broaden the scope of our review by drawing comparisons with the adaptive strategies used by other nocturnal and diurnal insects.\"

}, keywords = {DEILEPHILA, MACROGLOSSUM, OPTICS, SPATIAL ACUITY, SPHINGIDAE, TRADE-OFFS, VISION, VISUAL ACUITY, VISUAL PROCESSING}, doi = {10.1093/icb/icx058}, author = {St{\"o}ckl, A. and Smolka, J. and O{\textquoteright}Carroll, D. and Warrant, E.} } @article {22808, title = {The evolution and expression of the moth visual opsin family}, journal = {PLoS ONE}, volume = {8}, year = {2013}, month = {10/2013}, pages = {e78140}, chapter = {e78140}, keywords = {BAYESIAN INFERENCE, DIEL ACTIVITY, DIURNALISM, DNA, EVOLUTION, GENE EXPRESSION, LEPIDOPTERA, MACROGLOSSUM, MAXIMUM LIKELIHOOD, OPSIN, RNA, SPHINGIDAE, VISION}, issn = {1932-6203}, doi = {10.1371/journal.pone.0078140}, url = {http://dx.doi.org/10.1371/journal.pone.0078140}, author = {Xu, P. and Lu, B. and Xiao, H. and Fu, X. and Murphy, R.W. and Wu, K.} } @article {22734, title = {Parallel evolution of opsin visual pigments by tuning of spectral sensitivities during the transition from nocturnal to diurnal hawkmoths}, journal = {Journal of Experimental Biology}, volume = {XXX}, year = {2022}, month = {11/2022}, pages = {244541}, type = {Accepted manuscript}, chapter = {244541}, abstract = {

\"Light environments differ dramatically between day and night. The transition between diurnal and nocturnal visual ecology has happened repeatedly during evolution in many species. However, the molecular mechanism underlying the evolution of vision in recent diurnal\–nocturnal transition is poorly understood. Here, we focus on hawkmoths (Lepidoptera: Sphingidae) to address this question by investigating five nocturnal and five diurnal species. We performed RNA-sequencing analysis and identified opsin genes corresponding to the ultraviolet (UV), short-wavelength (SW), and long-wavelength (LW)-absorbing visual pigments. We found no significant differences in the expression patterns of opsin genes between the nocturnal and diurnal species. We then constructed the phylogenetic trees of hawkmoth species and opsins. The diurnal lineages had emerged at least three times from the nocturnal ancestors. The evolutionary rates of amino acid substitutions in the three opsins differed between the nocturnal and diurnal species. We found an excess number of parallel amino acid substitutions in the opsins in three independent diurnal lineages. The numbers were significantly more than those inferred from neutral evolution, suggesting that positive selection acted on these parallel substitutions. Moreover, we predicted the visual pigments\’ absorption spectra based on electrophysiologically-determined spectral sensitivity in two nocturnal and two diurnal species belonging to different clades. In the diurnal species, the LW pigments shift 10 nm toward the short wavelength, and the SW pigments shift 10 nm in the opposite direction. Taken together, our results suggest that parallel evolution of opsins may have enhanced the color discrimination properties of diurnal hawkmoths in ambient light.\"

}, keywords = {ABSORPTION SPECTRA, ADAPTATION, AMBULYX, AMPELOPHAGA, CEPHONODES, DAPHNIS, DIURNALISM, EVOLUTION, GENE EXPRESSION, HEMARIS, HISTOLOGY, MACROGLOSSUM, MARUMBA, MAXIMUM LIKELIHOOD, NEOGURELCA, OPSIN, PHYSIOLOGY, RNA, SPECTRAL SHIFT, SPHINGIDAE, THERETRA, ULTRAVIOLET VISION, VISION}, doi = {10.1242/jeb.244541}, url = { https://doi.org/10.1242/jeb.244541}, author = {Akiyama, T. and Uchiyama, H. and Yajima, S. and Arikawa, K. and Terai, Y.} } @article {22714, title = {Duplication and sub-functionalisation characterise diversification of opsin genes in the Lepidoptera}, year = {2022}, pages = {1-35}, type = {bioRxiv preprint}, abstract = {

\"Gene duplication is a vital process for evolutionary innovation. Functional diversification of duplicated genes is best explored in multicopy gene families such as histones, hemoglobin, and opsins. Rhodopsins are photo-sensitive proteins that respond to different wavelengths of light and contribute to diverse visual adaptations across insects. While there are several instances of gene duplications in opsin lineages, the functional diversification of duplicated copies and their ecological significance is properly characterised only in a few insect groups. We examined molecular and structural evolution that underlies diversification and sub-functionalisation of four opsin genes and their duplicated copies across 132 species of the diverse insect order\—Lepidoptera. Opsins have largely evolved under purifying selection with few residues showing signs of episodic and pervasive diversifying selection. Although these do not affect overall protein structures of opsins, substitutions in key amino acids in the chromophore-binding pocket of duplicated copies might cause spectral sensitivity shifts leading to sub-functionalisation or neofunctionalisation. Duplicated copies of opsins also exhibit developmental stage-specific expression in Papilio polytes, suggesting functional partitioning during development. Together, altered spectral sensitivities owing to key substitutions and differential expression of duplicated copies across developmental stages might enable enhanced colour perception and improved discrimination across wavelengths in this highly visual insect group.\"

}, keywords = {ANTHERAEA, BOMBYCIDAE, BOMBYX, EVOLUTION, GENE DUPLICATION, LEPIDOPTERA, MACROGLOSSUM, OPODIPHTHERA, OPSIN, PHYLOGENY, RHODOPSIN, SATURNIIDAE, SPHINGIDAE, VISION}, doi = {10.1101/2022.10.31.514481}, url = {https://doi.org/10.1101/2022.10.31.514481}, author = {Kuwalekar, M. and Deshmukh, R. and Baral, S. and Padvi, A. and Kunte, K.} } @article {22317, title = {Natural image statistics in the dorsal and ventral visual field match a switch in flight behaviour of a hawkmoth}, journal = {Current Biology}, volume = {31}, year = {2021}, month = {03/2021}, pages = {R280-R281}, chapter = {R280}, keywords = {BEHAVIOUR, MACROGLOSSUM, NATURAL IMAGE STATISTICS, OPTICS, SPHINGIDAE, VISION, VISUAL ACUITY}, author = {Bigge, R. and Pfefferle, M. and Pfeiffer, K. and St{\"o}ckl, A.} } @article {21488, title = {Visual position stabilization in the hummingbird hawk moth, Macroglossum stellatarum L. I. Behavioural analysis}, journal = {Journal of Comparative Physiology A}, volume = {182}, year = {1998}, pages = {225-237}, chapter = {225}, abstract = {

\"Optomotor responses of freely flying hawk moths, Macroglossum stellatarum, were characterized while the animals were hovering in front of and feeding on a dummy flower. Compensatory translational and rotational movements of the hawk moth were elicited by vertical grating patterns moving horizontally, mimicking imposed rotational and translational displacements of the animal in the horizontal plane. Oscillatory translational and rotational pattern motion leads to compensatory responses that peak in the frequency range between 2 Hz and 4\ Hz. The control systems mediating the translational and rotational components of the optomotor response do not seem to influence each other. The system mediating translational responses is more sensitive in the fronto-lateral part of the visual field than in the lateral part; the opposite is true for the rotational system. The sensitivity of the translational system does not change along the vertical, whereas the rotational system is much more sensitive to motion in the dorsal than in the ventral part of the visual field. These sensitivity gradients may reflect an adaptation to the specific requirements of position stabilization in front of flowers during feeding.\"

}, keywords = {HOVERING, MACROGLOSSUM, OPTOMOTOR BEHAVIOUR, POSITION STABILIZATION, SPHINGIDAE, VISION}, doi = {10.1007/s003590050173}, author = {Kern, R. and Varj{\'u}, D.} } @article {21486, title = {Fundamental differences in the optical structure of the eyes of nocturnal and diurnal mosquitoes}, journal = {Journal of Comparative Physiology A}, volume = {185}, year = {1999}, pages = {91-103}, chapter = {91}, abstract = {

\"We have studied the anatomy and optics of the eyes of a range of mosquito species from the wholly dark-active blood-feeding Anopheles gambiae to the diurnal plant-feeder Toxorhynchites brevipalpis. Consistent with studies by Sat\ô in the 1950s, we find that dark-active and crepuscular species have short fused rhabdoms with a conical construction. This maximises the amount of light the rhabdoms receive from the almost hemispherical wide-aperture lenses.\  Toxorhynchites, on the other hand, has long narrow rhabdomeres that are separated from each other over their entire length, and so resemble the open rhabdoms of advanced flies (Brachycera and Cyclorrhapha). These findings are confirmed by studies of the pseudopupil, whose form indicates the layout of the rhabdomere tips in the focal plane of each ommatidial lens. In anopheline species the pseudopupil is a single undivided ellipse, indicating a fused rhabdom structure, whereas in Toxorhynchites there is a ring of six outer elements surrounding a central one. This means that each rhabdomere views a separate direction in space, and our measurements indicate that, as in higher Diptera, adjacent rhabdomeres share their fields of view with one of the rhabdomeres in the immediately adjacent ommatidia. This in turn means that in the diurnal type of mosquito eye there is a basis for neural superposition, but the fused construction of anopheline rhabdoms precludes this. The Aedes species studied were similar to Anopheles but with lenses of less extreme aperture, and Sabethes cyaneus, a diurnal blood-feeder, was intermediate in structure, with fused conical rhabdoms in the centre of the eye and unfused
rhabdomeres around the periphery.\"

}, keywords = {AEDES, ANOPHELES, CULICIDAE, CULISETA, NEURAL SUPERPOSITION, OMMATIDIUM, OPTICS, PHYSIOLOGY, PSEUDOPUPIL, RHABDOMES, SABETHES, TOXORHYNCHITES, VISION}, author = {Land, M.F. and Gibson, G. and Horwood, J. and Zeil, J.} } @article {21484, title = {Higher-order neural processing tunes motion neurons to visual ecology in three species of hawkmoths}, journal = {Proceedings of the Royal Society B}, volume = {284}, year = {2017}, month = {06/2017}, pages = {20170880}, chapter = {20170880}, abstract = {

\"To sample information optimally, sensory systems must adapt to the ecological demands of each animal species. These adaptations can occur peripherally, in the anatomical structures of sensory organs and their receptors; and centrally, as higher-order neural processing in the brain. While a rich body of investigations has focused on peripheral adaptations, our understanding is sparse when it comes to central mechanisms. We quantified how peripheral adaptations in the eyes, and central adaptations in the wide-field motion vision system, set the trade-off between resolution and sensitivity in three species of hawkmoths active at very different light levels: nocturnal Deilephila elpenor, crepuscular Manduca sexta, and diurnal Macroglossum stellatarum. Using optical measurements and physiological recordings from the photoreceptors and wide-field motion neurons in the lobula complex, we demonstrate that all three species use spatial and temporal summation to improve visual performance in dim light. The diurnal Macroglossum relies least on summation, but can only see at brighter intensities. Manduca, with large sensitive eyes, relies less on neural summation than the smaller eyed Deilephila, but both species attain similar visual performance at nocturnal light levels. Our results reveal howthe visual systems of these three hawkmoth species are intimately matched to their visual ecologies.\"

}, keywords = {DEILEPHILA, ECOLOGY, MACROGLOSSUM, MANDUCA, NEURAL SUMMATION, PHYSIOLOGY, SPHINGIDAE, VISION, VISUAL PROCESSING}, url = {http://dx.doi.org/10.1098/rspb.2017.0880}, author = {St{\"o}ckl, A.L. and O{\textquoteright}Carroll, D.C. and Warrant, E.J.} } @article {21484, title = {Neural summation in the hawkmoth visual system extends the limits of vision in dim light}, journal = {Current Biology}, volume = {26}, year = {2016}, month = {03/2016}, pages = {821-826}, chapter = {821}, abstract = {

\"Most of the world\’s animals are active in dim light and depend on good vision for the tasks of daily life. Many have evolved visual adaptations that permit a performance superior to that of manmade imaging devices [1]. In insects, a major model visual system, nocturnal species show impressive visual abilities ranging from flight control [2, 3], to color discrimination [4, 5], to navigation using visual landmarks [6\–8] or dim celestial compass cues [9, 10]. In addition to optical adaptations that improve their sensitivity in dim light [11], neural summation of light in space and time\—which enhances the coarser and slower features of the scene at the expense of noisier finer and faster features\—has been suggested to improve sensitivity in theoretical [12\–14], anatomical [15\–17], and behavioral [18\–20] studies. How these summation strategies function neurally is, however, presently unknown. Here, we quantified spatial and temporal summation in the motion vision pathway of a nocturnal hawkmoth. We show that spatial and temporal summation combine supralinearly to substantially increase contrast sensitivity and visual information rate over four decades of light intensity, enabling hawkmoths to see at light levels 100 times dimmer than without summation. Our results reveal how visual motion is calculated neurally in dim light and how spatial and temporal summation improve sensitivity while simultaneously maximizing spatial and temporal resolution, thus extending models of insect motion vision derived predominantly from diurnal flies. Moreover, the summation strategies we have revealed may benefit manmade vision systems optimized for variable light levels [21].\"

}, keywords = {DEILEPHILA, NEURAL SUMMATION, PHYSIOLOGY, SPHINGIDAE, VISION, VISUAL PROCESSING}, url = {http://dx.doi.org/10.1016/j.cub.2016.01.030}, author = {St{\"o}ckl, A.L. and O{\textquoteright}Carroll, D.C. and Warrant, E.J.} } @article {21483, title = {Wide-field motion tuning in nocturnal hawkmoths}, journal = {Proceedings of the Royal Society B}, volume = {277}, year = {2010}, pages = {853-860}, chapter = {853}, abstract = {

\"Nocturnal hawkmoths are known for impressive visually guided behaviours in dim light, such as hovering while feeding from nectar-bearing flowers. This\  requires tight visual feedback to estimate and counter relative motion. Discrimination of low velocities, as required for stable hovering flight, is fundamentally limited by spatial resolution, yet in the evolution of eyes for nocturnal vision, maintenance of high spatial acuity compromises absolute sensitivity. To investigate these trade-offs, we compared responses of wide-field motion-sensitive neurons in three species of hawkmoth: Manduca sexta (a crepuscular hoverer), Deilephila elpenor (a fully nocturnal hoverer) and Acherontia atropos (a fully nocturnal hawkmoth that does not hover as it feeds uniquely from honey in bees\’\  nests). We show that despite smaller eyes, the motion pathway of D. elpenor is tuned to higher spatial frequencies and lower temporal frequencies than A. atropos, consistent with D. elpenor\’s need to detect low velocities for hovering. Acherontia atropos, however, presumably evolved low-light sensitivity without sacrificing temporal acuity. Manduca sexta, active at higher light levels, is tuned to the highest spatial frequencies of the three and temporal frequencies comparable with A. atropos. This yields similar tuning to low velocities as in D. elpenor, but with the advantage of shorter neural delays in processing motion.\"

}, keywords = {ACHERONTIA, DEILEPHILA, HOVERING, MANDUCA, NIGHT VISION, PHYSIOLOGY, SPHINGIDAE, VISION, VISUAL ACUITY, VISUAL PROCESSING}, doi = {10.1098/rspb.2009.1677}, author = {Theobald, J.C. and Warrant, E.J. and O{\textquoteright}Carroll, D.C.} } @article {21314, title = {Trichromatic colour vision in the hummingbird hawkmoth, Macroglossum stellatarum}, journal = {Journal of Comparative Physiology A}, volume = {184}, year = {1999}, pages = {535-541}, chapter = {535}, abstract = {

\"Hymenopterans have long been shown to choose colours by means of the spectral distribution and independently of the intensity (true colour vision). The same ability has only very recently been proven for two butterfly species. We present evidence for the existence of true colour vision in the European hummingbird hawkmoth, Macroglossum stellatarum. Moths were trained in dual-choice situations to spectral lights of a rewarding and an unrewarding wavelength. After training, unrewarded tests were performed during which the intensities of the lights were changed. The results confirm that the species has three spectral receptor types and uses true colour vision when learning the colour of a food source. If colour vision is not possible since only one receptor type is receiving input from both stimuli, the moths learn to associate some achromatic cue correlated to the receptor quantum catch, with the reward. The moths learn spectral cues rapidly and choose correctly after one to several rewarded visits even when trained to different colours in sequence.\"

}, keywords = {COLOUR VISION, MACROGLOSSUM, SPHINGIDAE, VISION}, doi = {10.1007/s003590050353}, author = {Kelber, A. and H{\'e}nique, U.} } @article {21313, title = {Wavelength discrimination and the role of ultraviolet vision in the feeding behaviour of hawkmoths}, journal = {Biotropica}, volume = {26}, year = {1994}, pages = {427-435}, chapter = {427}, abstract = {

\"Nocturnal Sphingidae (hawkmoths or sphinx moths) are important pollinators in tropical forests. Hawkmoth flowers are typically white to the human eye. As the retinas of hawkmoths contain ultraviolet-sensitive photoreceptors, flower patterns reflecting ultraviolet wavelengths (that are not visible to humans) might be significant to sphingid feeding behavior. The flowers of ten hawkmoth-pollinated species were examined with an ultraviolet sensitive video system in Monteverde, Costa Rica. All were found to lack ultraviolet reflectance. A common hawkmoth species, Manduca sexta, whose range extends to Costa Rica was then used in laboratory free choice experiments to determine which wavelengths elicited proboscis extension, probing and drinking of sugar water. When offered a choice between artificial flowers or backlighted filters, Manduca strongly preferred to feed at those reflecting or transmitting only wavelengths longer than 400 nm, avoiding those that also included ultraviolet wavelengths. That is, feeding behavior was best elicited by stimuli that mimicked the reflectance of typical hawkmoth flowers. Feeding behavior must be primarily activated by either the green- or violet-sensitive mechanisms (or both) of the hawkmoth visual system, while concurrent activation of the ultraviolet-sensitive mechanism interferes with it.\"

}, keywords = {COSTA RICA, MANDUCA, SPHINGIDAE, ULTRAVIOLET VISION, VISION}, url = {http://www.jstor.org/stable/2389237 }, author = {White, R.H. and Stevenson, R.D. and Bennett, R.R. and Cutler, D.E. and Haber, W.A.} } @article {21311, title = {Three visual pigments in Deilephila elpenor (Lepidoptera, Sphingidae)}, journal = {Journal of Comparative Physiology A}, volume = {86}, year = {1973}, pages = {215-229}, chapter = {215}, abstract = {

\"Selective irradiation of digitonin extracts (pH 6.1) from dark-adapted eyes of the Sphingid moth Deilephila elpenor revealed three visual pigments with \λmax at about 345, 440 and 520 nm. From the reaction of the chromophoric group with hydroxylamine it is concluded that these pigments are based on retinaldehyde, and therefore can be classed with rhodopsins. At \−15 \°C, each of the rhodopsins can completely (P 520) or partially (P 440 and P 345) be converted by light to acid metarhodopsin absorbing maximally at 480 \± 10 nm, which decay at room temperature to retinaldehyde and protein moiety. Spectrophotometric measurements on isolated retinae (pH 7.4) indicate that the metarhodopsins, in contrast to the findings on extracts, are thermostable in the photoreceptor membrane at room temperature. The results are compared with electrophysiological and microspectrophotometrical data. The different properties of the metarhodopsins in detergent solution and in the isolated retina are discussed.\"

}, keywords = {COLOUR VISION, DEILEPHILA, RHODOPSIN, SPHINGIDAE, VISION}, doi = {10.1007/BF00696341}, author = {Schwemer, J. and Paulsen, R.} } @article {18705, title = {Scotopic colour vision in nocturnal hawkmoths}, journal = {Nature}, volume = {419}, year = {2002}, month = {10/2002}, pages = {922-925}, chapter = {922}, abstract = {

\"Humans are colour-blind at night, and it has been assumed that this is true of all animals. But colour vision is as useful for discriminating objects at night as it is during the day. Here we show, through behavioural experiments, that the nocturnal hawkmoth Deilephila elpenor uses colour vision to discriminate coloured stimuli at intensities corresponding to dim starlight (0.0001 cd m-2). It can do this even if the illumination colour changes, thereby showing colour constancy\—a property of true colour vision systems. In identical conditions humans are completely colour-blind. Our calculations show that the possession of three photoreceptor classes reduces the absolute sensitivity of the eye, which indicates that colour vision has a high ecological relevance in nocturnal moths. In addition, the photoreceptors of a single ommatidium absorb too few photons for reliable discrimination, indicating that spatial and/or temporal summation must occur for colour vision to be possible. Taken together, our results show that colour vision occurs at nocturnal intensities in a biologically relevant context.\"

}, keywords = {COLOUR VISION, DEILEPHILA, HYLES, PHYSIOLOGY, SPHINGIDAE, VISION}, doi = {10.1038/nature01065}, author = {Kelber, A. and Balkenius, A. and Warrant, E.J.} } @article {21298, title = {Conservation, duplication, and divergence of five opsin genes in insect evolution}, journal = {Genome Biology and Evolution}, volume = {8}, year = {2016}, month = {02/2016}, pages = {579-587}, chapter = {579}, abstract = {

\"Opsin proteins covalently bind to small molecular chromophores and each protein-chromophore complex is sensitive to particular wavelengths of light. Multiple opsins with different wavelength absorbance peaks are required for color vision. Comparing opsin responses is challenging at low light levels, explaining why color vision is often lost in nocturnal species. Here, we investigated opsin evolution in 27 phylogenetically diverse insect species including several transitions between photic niches (nocturnal, diurnal, and crepuscular).We find widespread conservation of five distinct opsin genes, more than commonly considered.\  These comprise one c-opsin plus four r-opsins (long wavelength sensitive or LWS, blue sensitive, ultra violet [UV] sensitive and the often overlooked Rh7 gene). Several recent opsin gene duplications are also detected. The diversity of opsin genes is consistent with color vision in diurnal, crepuscular, and nocturnal insects. Tests for positive selection in relation to photic niche reveal evidence for adaptive evolution in UVsensitive opsins in day-flying insects in general, and in LWS opsins of day-flying Lepidoptera specifically.\"

}, keywords = {BOMBYCIDAE, BOMBYX, C-OPSINS, COLOUR VISION, DAY/NIGHT TRANSITIONS, EREBIDAE, EVOLUTION, GENE DUPLICATION, INSECTA, LWS-OPSIN, MANDUCA, OPSIN, PHYLOGENY, R-OPSINS, SPHINGIDAE, VISION}, doi = {10.1093/gbe/evw015}, author = {Feuda, R. and Marl{\'e}taz, F. and Bentley, M.A. and Holland, P.W.H.} } @article {20942, title = {Adaptations for nocturnal and diurnal vision in the hawkmoth lamina}, journal = {Journal of Comparative Neurology}, volume = {524}, year = {2016}, pages = {160-175}, chapter = {160}, keywords = {ADAPTATION, DEILEPHILA, LAMINA, MACROGLOSSUM, MANDUCA, NEUROLOGY, PHYSIOLOGY, SPATIAL SUMMATION, SPHINGIDAE, VISION, VISUAL PROCESSING}, doi = {10.1002/cne.23832}, author = {St{\"o}ckl, A.L. and Ribi, W.A. and Warrant, E.J.} } @article {20491, title = {Physiological optics in the hummingbird hawkmoth: a compound eye without ommatidia}, journal = {Journal of Experimental Biology}, volume = {202}, year = {1999}, month = {1999}, pages = {497-511}, chapter = {497}, keywords = {MACROGLOSSUM, OPTICS, PHYSIOLOGY, SPHINGIDAE, VISION}, url = {https://journals.biologists.com/jeb/article/202/5/497/8082/Physiological-optics-in-the-hummingbird-hawkmoth-a}, author = {Warrant, E. and Bartsch, K. and G{\"u}nther, C.} } @article {19656, title = {Sensory flexibility in hawkmoth foraging behavior: lessons from {IManduca sexta} and other species}, journal = {Chemical Senses}, volume = {30}, year = {2005}, month = {2005}, pages = {i295-i296}, keywords = {BEHAVIOUR, FORAGING, MANDUCA, PHYSIOLOGY, SPHINGIDAE, VISION}, author = {Raguso, R. A. and LeClere, A. R. and Schlumpberger, R. O.} } @article {19658, title = {Synergy between visual and olfactory cues in nectar feeding by na{\"\i}ve hawkmoths, Manduca sexta}, journal = {Animal Behaviour}, volume = {64}, year = {2002}, month = {2002}, pages = {685-695}, chapter = {685}, keywords = {BEHAVIOUR, FLORAL SCENT, MANDUCA, OLFACTION, POLLINATION, SPHINGIDAE, VISION}, doi = {10.1006/anbe.2002.4010}, author = {Raguso, R.A. and Willis, M.A.} } @article {18704, title = {Pattern discrimination in a hawkmoth: innate preferences, learning performance and ecology}, journal = {Proceedings of the Royal Society of London}, volume = {269}, year = {2002}, note = {B}, month = {11/2002}, pages = {2573-2577}, chapter = {2573}, keywords = {BEHAVIOUR, LEARNING, MACROGLOSSUM, PATTERN DISCRIMINATION, SPHINGIDAE, VISION}, doi = {10.1098/rspb.2002.2201}, author = {Kelber, A.} } @article {18705, title = {Colour vision in diurnal and nocturnal hawkmoths}, journal = {Integrative and Comparative Biology}, volume = {43}, year = {2003}, month = {2003}, pages = {571-579}, chapter = {571}, keywords = {COLOUR VISION, DEILEPHILA, HYLES, MACROGLOSSUM, PHYSIOLOGY, SPHINGIDAE, VISION}, url = {https://academic.oup.com/icb/article/43/4/571/617782}, author = {Kelber, A. and Balkenius, A. and Warrant, E.J.} } @article {17974, title = {Morphology and structure of the compound eye of a sphingid moth, {IHerse convolvuli} L}, journal = {Acta Entomologica Sinica}, volume = {29}, year = {1986}, month = {1986}, pages = {267-271}, keywords = {ANATOMY, SPHINGIDAE, VISION}, author = {Gao, W. Z.} } @article {17378, title = {Feeding behavior in the nocturnal moth {IManduca sexta} is mediated mainly by blue receptors, but where are they located in the retina?}, journal = {Journal of Experimental Biology}, volume = {198}, year = {1995}, month = {1995}, pages = {1909-1917}, keywords = {PHYSIOLOGY, SPHINGIDAE, VISION}, author = {Cutler, D. E. and Bennett, R. R. and Stevenson, R. D. and White, R. H.} } @article {16735, title = {Color constancy in diurnal and nocturnal hawkmoths}, journal = {Journal of Experimental Biology}, volume = {207}, year = {2004}, month = {2004}, pages = {3307-3316}, keywords = {COLOUR VISION, DEILEPHILA, PHYSIOLOGY, SPHINGIDAE, VISION}, author = {Balkenius, A.} }