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Fruit-feeding pteropodid bats roost under varying light conditions. Some roost in trees with high exposure to daylight (> 1000 lx), while others roost in dark caves (< 0.1 lx). To understand the effect of ambient light intensity and moon phase on flight activity, we examined flight times across five lunar cycles in three pteropodid species whose roosts differ in daylight exposure. We found significant interspecific differences in flight emergence and termination times. All species initiated flights after sunset but Rousettus leschenaultii, which typically roosts in caves, delayed emergence (40 ± 11 min) more than the two tree-roosting species Pteropus giganteus (16 ± 6 min) and Cynopterus sphinx (19 ± 7 min). R. leschenaultii terminated flights earlier (30 ± 7 min before sunrise) than P. giganteus (11 ± 11 min) and C. sphinx (16 ± 10 min). All individuals from P. giganteus and C. sphinx roosts emerged within less than an hour, while emergence times were more spread out in the R. leschenaultii colony. Peak emergence times differed across moon phases in the cave-roosting R. leschenaultii but not in the other species. Flight activity in R. leschenaultii is restricted to comparatively lower light levels than the tree-roosting species. The observed interspecific differences suggest that bat species, sharing same landscapes may respond differently to light pollution. [ABSTRACT FROM AUTHOR]

Hawkmoths (Lepidoptera, Sphingidae) comprise around 1500 species, most of which forage on nectar from flowers in their adult stage, usually while hovering in front of the flower. The majority of species have a nocturnal lifestyle and are important nocturnal pollinators, but some species have turned to a diurnal lifestyle. Hawkmoths use visual and olfactory cues including CO2 and humidity to detect and recognise rewarding flowers; they find the nectary in the flowers by means of mechanoreceptors on the proboscis and vision, evaluate it with gustatory receptors on the proboscis, and control their hovering flight position using antennal mechanoreception and vision. Here, we review what is presently known about the sensory organs and sensory-guided behaviour that control feeding behaviour of this fascinating pollinator taxon. We also suggest that more experiments on hawkmoth behaviour in natural settings are needed to fully appreciate their sensory capabilities. [ABSTRACT FROM AUTHOR]

Many insects rely on vision to find food, to return to their nest and to carefully control their flight between these two locations. The amount of information available to support these tasks is, in part, dictated by the spatial resolution and contrast sensitivity of their visual systems. Here, we investigate the absolute limits of these visual properties for visually guided position and speed control in Bombus terrestris. Our results indicate that the limit of spatial vision in the translational motion detection system of B. terrestris lies at 0.21 cycles deg with a peak contrast sensitivity of at least 33. In the perspective of earlier findings, these results indicate that bumblebees have higher contrast sensitivity in the motion detection system underlying position control than in their object discrimination system. This suggests that bumblebees, and most likely also other insects, have different visual thresholds depending on the behavioral context. [ABSTRACT FROM AUTHOR]

A ripe strawberry looks red to our eyes in sunlight and in the green light of a forest, although the spectrum of light reflected from its surface differs dramatically. This is caused by two effects: colour constancy and our ability to learn relative colour cues - the ripe strawberry remains relatively 'redder' than an unripe green strawberry. While colour constancy - the ability to recognize colours in shifted illumination - has been studied in many animals, the use of relative colour cues is investigated more rarely. In a previous study on chickens, we measured how large a shift in illumination their colour constancy mechanisms tolerate without reliable relative colour cues. Here, we show that chickens remain colour constant over larger illumination shifts, if they can use such relative colour cues. As relative colour cues are readily available in natural environments, we suggest that their use contributes strongly to colour constancy performance in nature. [ABSTRACT FROM AUTHOR]

Colour discrimination is based on opponent photoreceptor interactions, and limited by receptor noise. In dim light, photon shot noise impairs colour vision, and in vertebrates, the absolute threshold of colour vision is set by dark noise in cones. Nocturnal insects (e.g. moths and nocturnal bees) and vertebrates lacking rods (geckos) have adaptations to reduce receptor noise and use chromatic vision even in very dim light. In contrast, vertebrates with duplex retinae use colour-blind rod vision when noisy cone signals become unreliable, and their transition from cone- to rod-based vision is marked by the Purkinje shift. Rod–cone interactions have not been shown to improve colour vision in dim light, but may contribute to colour vision in mesopic light intensities. Frogs and toads that have two types of rods use opponent signals from these rods to control phototaxis even at their visual threshold. However, for tasks such as prey or mate choice, their colour discrimination abilities fail at brighter light intensities, similar to other vertebrates, probably limited by the dark noise in cones. [ABSTRACT FROM AUTHOR]

ABSTRACT The recognition that animals sense the world in a different way than we do has unlocked important lines of research in ecology and evolutionary biology. In practice, the subjective study of natural stimuli has been permitted by perceptual spaces, which are graphical models of how stimuli are perceived by a given animal. Because colour vision is arguably the best-known sensory modality in most animals, a diversity of colour spaces are now available to visual ecologists, ranging from generalist and basic models allowing rough but robust predictions on colour perception, to species-specific, more complex models giving accurate but context-dependent predictions. Selecting among these models is most often influenced by historical contingencies that have associated models to specific questions and organisms; however, these associations are not always optimal. The aim of this review is to provide visual ecologists with a critical perspective on how models of colour space are built, how well they perform and where their main limitations are with regard to their most frequent uses in ecology and evolutionary biology. We propose a classification of models based on their complexity, defined as whether and how they model the mechanisms of chromatic adaptation and receptor opponency, the nonlinear association between the stimulus and its perception, and whether or not models have been fitted to experimental data. Then, we review the effect of modelling these mechanisms on predictions of colour detection and discrimination, colour conspicuousness, colour diversity and diversification, and for comparing the perception of colour traits between distinct perceivers. While a few rules emerge (e.g. opponent log-linear models should be preferred when analysing very distinct colours), in general model parameters still have poorly known effects. Colour spaces have nonetheless permitted significant advances in ecology and evolutionary biology, and more progress is expected if ecologists compare results between models and perform behavioural experiments more routinely. Such an approach would further contribute to a better understanding of colour vision and its links to the behavioural ecology of animals. While visual ecology is essentially a transfer of knowledge from visual sciences to evolutionary ecology, we hope that the discipline will benefit both fields more evenly in the future. [ABSTRACT FROM AUTHOR]

Colour vision — the ability to discriminate spectral differences irrespective of variations in intensity — has two basic requirements: (1) photoreceptors with different spectral sensitivities, and (2) neural comparison of signals from these photoreceptors. Major progress has been made understanding the evolution of the basic stages of colour vision–opsin pigments, screening pigments, and the first neurons coding chromatic opponency, and similarities between mammals and insects point to general mechanisms. However, much work is still needed to unravel full colour pathways in various animals. While primates may have brain regions entirely dedicated to colour coding, animals with small brains, such as insects, likely combine colour information directly in parallel multisensory pathways controlling various behaviours. [ABSTRACT FROM AUTHOR]

Despite the strong relationship between insect vision and the spectral properties of flowers, the visual system has been studied in detail in only a few insect pollinator species. For instance, wavelength discrimination thresholds have been determined in two species only: the honeybee (Apis mellifera) and the butterfly Papilio xuthus. Here, we present the wavelength discrimination thresholds (Δλ) for the hawkmoth Macroglossum stellatarum. We compared the data with those found for the honeybee, the butterfly P. xuthus and the predictions of a colour discrimination model. After training moths to feed from a rewarded disc illuminated with a monochromatic light, we tested them in a dual-choice situation, in which they had to choose between light of the training wavelength and a novel unrewarded wavelength. To characterise the Δλ function, we decreased the difference between wavelengths in subsequent tests. We also varied the light intensity to test its effect on the discrimination capacity. In agreement with the predictions of the model, we found two expected minima of discrimination where photoreceptor sensitivities overlap, as well as a third, minor, unpredicted minimum around the peak of the blue photoreceptor. Macroglossum stellatarum is capable of discriminating between lights with a wavelength difference of 1-2 nm. These discrimination minima are similar to those found for the tetrachromatic P. xuthus, and are better than those of the honeybee. The moth is also capable of using achromatic information to discriminate between lights of long wavelengths. [ABSTRACT FROM AUTHOR]

All seals and cetaceans have lost at least one of two ancestral cone classes and should therefore be colour-blind. Nevertheless, earlier studies showed that these marine mammals can discriminate colours and a colour vision mechanism has been proposed which contrasts signals from cones and rods. However, these earlier studies underestimated the brightness discrimination abilities of these animals, so that they could have discriminated colours using brightness only. Using a psychophysical discrimination experiment, we showed that a harbour seal can solve a colour discrimination task by means of brightness discrimination alone. Performing a series of experiments in which two harbour seals had to discriminate the brightness of colours, we also found strong evidence for purely scotopic (rod-based) vision at light levels that lead to mesopic (rod-cone-based) vision in other mammals. This finding speaks against rod-cone-based colour vision in harbour seals. To test for colour-blindness, we used a cognitive approach involving a harbour seal trained to use a concept of same and different. We tested this seal with pairs of isoluminant stimuli that were either same or different in colour. If the seal had perceived colour, it would have responded to colour differences between stimuli. However, the seal responded with 'same', providing strong evidence for colour-blindness. [ABSTRACT FROM AUTHOR]

Many birds move fast into dark nest cavities forcing the visual system to adapt to low light intensities. Their visual system takes between 15 and 60 min for complete dark adaptation, but little is known about the visual performance of birds during the first seconds in low light intensities. In a forced two-choice behavioural experiment we studied how well budgerigars can discriminate stimuli of different luminance directly after entering a darker environment. The birds made their choices within about 1 s and did not wait to adapt their visual system to the low light intensities. When moving from a bright facility into an environment with 0.5 log unit lower illuminance, the budgerigars detected targets with a luminance of 0.825 cd m−2 on a black background. When moving into an environment with 1.7 or 3.5 log units lower illuminance, they detected targets with luminances between 0.106 and 0.136 cd m−2. In tests with two simultaneously displayed targets, the birds discriminated similar luminance differences between the targets (Weber fraction of 0.41–0.54) in all light levels. Our results support the notion that partial adaptation of bird eyes to the lower illumination occurring within 1 s allows them to safely detect and feed their chicks. [ABSTRACT FROM AUTHOR]

We investigated the formation of an abstract concept of same/different in a harbor seal by means of a two-item same/different task. Stimuli were presented on a TFT monitor. The subject was trained to respond according to whether two horizontally aligned white shapes presented on a black background were the same, or different from each other, by giving a no-go or go response. Training comprised of four stages. First, the same/different task was trained with two shapes forming two same problems (A-A and B-B) and two different problems (A-B and B-A). After the learning criterion was reached, training proceeded with new pairs of shapes. In the second experimental stage, every problem was presented just five times before new problems were introduced. We showed that training to criterion with just two shapes resulted in item-specific learning, whereas reducing the number of presentations to five per problem led to the formation of a same/different learning set as well as some restricted relational learning. Training with trial-unique problems in the third stage of this study resulted in the formation of an abstract concept of same/different which was indicated by a highly significant performance in transfer tests with 120 novel problems. Finally, extra-dimensional transfer of the concept was tested. The harbor seal showed a significantly correct performance on transfer tests with 30 unfamiliar pattern and 60 unfamiliar brightness same/different problems, thus demonstrating that the concept is not restricted to the shape dimension originally learned, but can be generalized across stimulus dimensions. [ABSTRACT FROM AUTHOR]

Most visual systems are more sensitive to luminance than to colour signals. Animals resolve finer spatial detail and temporal changes through achromatic signals than through chromatic ones. Probably, this explains that detection of small, distant, or moving objects is typically mediated through achromatic signals. Macroglossum stellatarum are fast flying nectarivorous hawkmoths that inspect flowers with their long proboscis while hovering. They can visually control this behaviour using floral markings known as nectar guides. Here, we investigate whether this is mediated by chromatic or achromatic cues. We evaluated proboscis placement, foraging efficiency, and inspection learning of naïve moths foraging on flower models with coloured markings that offered either chromatic, achromatic or both contrasts. Hummingbird hawkmoths could use either achromatic or chromatic signals to inspect models while hovering. We identified three, apparently independent, components controlling proboscis placement: After initial contact, 1) moths directed their probing towards the yellow colour irrespectively of luminance signals, suggesting a dominant role of chromatic signals; and 2) moths tended to probe mainly on the brighter areas of models that offered only achromatic signals. 3) During the establishment of the first contact, naïve moths showed a tendency to direct their proboscis towards the small floral marks independent of their colour or luminance. Moths learned to find nectar faster, but their foraging efficiency depended on the flower model they foraged on. Our results imply that M. stellatarum can perceive small patterns through colour vision. We discuss how the different informational contents of chromatic and luminance signals can be significant for the control of flower inspection, and visually guided behaviours in general. [ABSTRACT FROM AUTHOR]

Recent research shows that a nocturnal hawkmoth, Manduca sexta, inspects flowers in search for nectar by means of a series of hovering and proboscis movements controlled by different sensory modalities, mainly vision and mechanoreception. The diurnal Macroglossum stellatarum is a closely related hawkmoth challenged with the same task but under illuminances 6–8 orders of magnitude higher. Here, we use flower models presenting color markings, 3D features, or both to study innate flower movements and sensory cues involved in the innate inspection behavior of M. stellatarum. On flat plain colored models, moths scanned the whole corolla with their proboscis, attaining intermediate success levels. When models presented color markings, moths biased proboscis placement, which affected inspection efficiency. Three-dimensional features affected inspection behavior and efficiency, by provoking strong “diving” responses. When both visual and tactile features were present, visual cues showed prevalence in the control of the inspection behavior. We evaluated learning abilities of moths, finding that M. stellatarum cannot only improve their efficiency by calibrating innate responses but also by changing their innate strategy. We discuss results on flower inspection behavior of M. stellatarum comparatively with known aspects of this behavior in M. sexta. Motor responses appear to be shared by these hawkmoths, but their sensory control shows important differences. Future studies will evaluate whether these differences are the result of an evolutionary divergence of their innate behavior associated with a switch to diurnal activity or the result of a flexible use of sensory information by the naïve animals. [ABSTRACT FROM PUBLISHER]

Hornets, the largest social wasps, have a reputation of being facultatively nocturnal. Here we confirm flight activity of hornet workers in dim twilight. We studied the eyes and ocelli of European hornets (Vespa crabro) and common wasps (Vespula vulgaris) with the goal to find the optical and anatomical adaptations that enable them to fly in dim light. Adaptations described for obligately nocturnal hymenoptera such as the bees Xylocopa tranquebarica and Megalopta genalis and the wasp Apoica pallens include large ocelli and compound eyes with wide rhabdoms and large facet lenses. Interestingly, we did not find any such adaptations in hornet eyes or ocelli. On the contrary, their eyes are even less sensitive than those of the obligately diurnal common wasps. Therefore we conclude that hornets, like several facultatively nocturnal bee species such as Apis mellifera adansonii, A. dorsata and X. tenuiscapa are capable of seeing in dim light simply due to the large body and thus eye size. We propose that neural pooling strategies and behavioural adaptations precede anatomical adaptations in the eyes and ocelli when insects with apposition compound eyes turn to dim light activity. [ABSTRACT FROM AUTHOR]

Humans and most vertebrates have duplex retinae with multiple cone types for colour vision in bright light, and one single rod type for achromatic vision in dim light. Instead of comparing signals from multiple spectral types of photoreceptors, such species use one highly sensitive receptor type thus improving the signal-to-noise ratio at night. However, the nocturnal hawkmoth Deilephila elpenor, the nocturnal bee Xylocopa tranquebarica and the nocturnal gecko Tarentola chazaliae can discriminate colours at extremely dim light intensities. To be able to do so, they sacrifice spatial and temporal resolution in favour of colour vision. We review what is known about colour vision in dim light, and compare colour vision thresholds with the optical sensitivity of the photoreceptors in selected animal species with lens and compound eyes. [ABSTRACT FROM AUTHOR]

Nectar-feeding insects find flowers by 2 means, innate preferences and learned associations. When insects that hibernate as imagos (i.e., adults) start foraging after a long winter break, what guides them to new nectar rewards? Are innate preferences kept over such a long period? And are learned associations useful after long breaks? In a series of experiments I show here that, depending on previous experience, the European hummingbird hawkmoth, Macroglossum stellatarum can use both types of information to choose a first flower after periods of 1 or 3 weeks. What is remembered seems to depend on innate preferences of the moths. Moths trained to feed from 1 of 2 colors that are equally attractive to naive moths keep the learned preference. Animals trained to prefer a less attractive color to an innately preferred color loose the learned preference and return to the innate choice behavior. I conclude that hummingbird hawkmoths can keep innate and learned preferences over a long period and are able to use both types of information when searching nectar rewards after long periods of hibernation. [ABSTRACT FROM AUTHOR]

We have used behavioural tests to determine the intensity thresholds of colour vision in Bourke's parrots (Neopsephotus bourkii) and budgerigars (Melopsittacus undulatus). We have also examined the relationship between these threstpolds and the optical sensitivities of single photoreceptors using morphological methods. Bourke's parrots lose colour vision in brighter light (0.4cdm2) than budgerigars (0.1 cdm2) and both birds lose colour vision in brighter light ('end of civil twilight') than humans (0.02cdm2, 'moonlight'). The optical sensitivities of single cones are similar in both birds (budgerigar 0.27 μm2sr, Bourke's parrot 0.25 μm2sr) but Bourke's parrots have more (cone to rod ratio, 1.2:1.0), thinner (2.8 μm) and longer rods (18.5 μm) than budgerigars (2.1:1.0, 3.4 μm, 13.3 μm). Bourke's parrots thus have an eye type that, with a flexible pooling mechanism, allows for high resolution or high absolute sensitivity depending on the light conditions. The results nicely agree with the activity patterns of the birds, Bourke's parrots being active during the day and in twilight while budgerigars are not normally active before sunrise and after sunset. However, Bourke's parrots have fewer cones than budgerigars, which implies that a smaller number of cones are pooled within each retinal integration area. That could explain why Bourke's parrots have a higher intensity threshold of colour vision than budgerigars. Furthermore, the study emphasises the need to expand the sensitivity measure so that photoreceptor integration units are used rather than single receptors. [ABSTRACT FROM AUTHOR]

Abstract: Colour vision models require measurement of receptor noise and the absorbance of visual pigments, oil droplets, and ocular media. We have studied how variation in these parameters influences colour matching, spectral sensitivity, and colour discrimination predictions in four bird species. While colour match predictions are sensitive to variation in visual pigment and oil droplet absorbance data, discrimination predictions are mostly sensitive to variation in receptor noise. Ocular media transmittance influences only modelled spectral sensitivities at short wavelengths. A comparison between predicted and measured spectral sensitivities in domestic fowl and duck revealed large discrepancies, likely because of influences from achromatic mechanisms. [Copyright &y& Elsevier]

Most bees are diurnal, with behaviour that is largely visually mediated, but several groups have made evolutionary shifts to nocturnality, despite having apposition compound eyes unsuited to vision in dim light. We compared the anatomy and optics of the apposition eyes and the ocelli of the nocturnal carpenter bee, Xylocopa tranquebarica, with two sympatric species, the strictly diurnal X. leucothorax and the occasionally crepuscular X. tenuiscapa. The ocelli of the nocturnal X. tranquebarica are unusually large (diameter ca. 1 mm) and poorly focussed. Moreover, their apposition eyes show specific visual adaptations for vision in dim light, including large size, large facets and very wide rhabdoms, which together make these eyes 9 times more sensitive than those of X. tenuiscapa and 27 times more sensitive than those of X. leucothorax. These differences in optical sensitivity are surprisingly small considering that X. tranquebarica can fly on moonless nights when background luminance is as low as 10−5 cd m−2, implying that this bee must employ additional visual strategies to forage and find its way back to the nest. These strategies may include photoreceptors with longer integration times and higher contrast gains as well as higher neural summation mechanisms for increasing visual reliability in dim light. [ABSTRACT FROM AUTHOR]

The moth Macroglossum stellatarum can learn the color and sometimes the odor of a rewarding food source. We present data from 20 different experiments with different combinations of blue and yellow artificial flowers and the two odors, honeysuckle and lavender. The experiments show that learning about the odors depends on the color used. By training on different color-odor combinations and testing on others, it becomes possible to investigate the exact relation between the two modalities during learning. Three computational models were tested in the same experimental situations as the real moths and their predictions were compared with the experimental data. The average error over all experiments as well as the largest deviation from the experimental data were calculated. Neither the Rescorla-Wagner model nor a learning model with independent learning for each stimulus component were able to explain the experimental data. We present the new hawkmoth learning model, which assumes that the moth learns a template for the sensory attributes of the rewarding stimulus. This model produces behavior that closely matches that of the real moth in all 20 experiments. [ABSTRACT FROM AUTHOR]

In animal eyes of the camera type longitudinal chromatic aberration causes defocus that is particularly severe in species with short depth of focus. In a variety of vertebrates, multifocal optical systems compensate for longitudinal chromatic aberration by concentric zones of different.refractive powers. Since a constricting circular pupil blocks peripheral zones, eyes with multifocal optical systems often have slit pupils that allow light to pass through all zones, irrespective of the state of pupil constriction. Birds have circular pupils and were therefore assumed to have monofocal optical systems. We examined the eyes of 45 species (12 orders) of bird using videorefractometry, and the results are surprising: 29 species (10 orders) have multifocal systems, and only five species (five orders) have monofocal systems. The results from 11 species (four orders) are inconclusive. We propose that pupils 'switching' between being fully opened (multifocal principle) to maximally closed (pinhole principle) can make multifocal optical systems useful for animals with circular pupils. Previous results indicate that mice have both multifocal optical systems and switching pupils. Our results suggest that parrots may use a similar mechanism. By contrast, owl pupils responded weakly to changes in illumination and stayed remarkably wide even in full daylight. Moreover, the parrots opened their pupils at higher light levels than owls, which correlates with the differences in sensitivity between diurnal and nocturnal eyes. [ABSTRACT FROM AUTHOR]

Abstract: The brightness discrimination ability of a male harbor seal was investigated at an ambient illumination of 0.9lx. The visual stimuli consisted of circular gray discs that were presented on a black background on a TFT monitor. Eight standard intensities were tested against sets of lower comparison intensities. In accordance with Weber’s law we observed a constant gain of the just noticeable intensity difference with increasing intensity of the standard stimulus. The calculated Weber fraction is 0.14. This result indicates that the brightness discrimination ability of the harbor seal is comparable to that of humans. [Copyright &y& Elsevier]

Abstract: New data on the phylogenetic relationships of various arthropod groups have spurred interesting attempts to reconstruct the evolution of arthropod nervous and visual systems. Some of the relevant new data are cell identities and developmental processes in the nervous and sensory systems, which is particularly useful for reconstructing the evolution of these systems. Here, we focus on the structure of compound eye ommatidia, and make an evolutionary analysis with functional arguments. We investigate possible routes of evolution that can be understood in terms of selection for improved visual function, and arrive at a number of conclusions that are discussed in the light of recent phylogenetic hypotheses. On the basis of ommatidial focusing structures and the arrangement of receptor cells we show that the evolution of compound eyes proceeded largely independently along at least two lineages from very primitive ancestors. A common ancestor of insects and crustaceans is likely to have had ommatidia with focusing crystalline cones, and colour and/or polarization vision. In contrast, the compound eyes in myriapods and chelicerates are likely to date back to ancestors with corneal lenses and probably without the ability to discriminate colour and polarization. [Copyright &y& Elsevier]

Abstract: Nocturnal bees, wasps and ants have considerably larger ocelli than their diurnal relatives, suggesting an active role in vision at night. In a first step to understanding what this role might be, the morphology and physiological optics of ocelli were investigated in three tropical rainforest species – the nocturnal sweat bee Megalopta genalis, the nocturnal paper wasp Apoica pallens and the diurnal paper wasp Polistes occidentalis – using hanging-drop techniques and standard histological methods. Ocellar image quality, in addition to lens focal length and back focal distance, was determined in all three species. During flight, the ocellar receptive fields of both nocturnal species are centred very dorsally, possibly in order to maximise sensitivity to the narrow dorsal field of light that enters through gaps in the rainforest canopy. Since all ocelli investigated had a slightly oval shape, images were found to be astigmatic: images formed by the major axis of the ocellus were located further from the proximal surface of the lens than images formed by the minor axis. Despite being astigmatic, images formed at either focal plane were reasonably sharp in all ocelli investigated. When compared to the position of the retina below the lens, measurements of back focal distance reveal that the ocelli of Megalopta are highly underfocused and unable to resolve spatial detail. This together with their very large and tightly packed rhabdoms suggests a role in making sensitive measurements of ambient light intensity. In contrast, the ocelli of the two wasps form images near the proximal boundary of the retina, suggesting the potential for modest resolving power. In light of these results, possible roles for ocelli in nocturnal bees and wasps are discussed, including the hypothesis that they might be involved in nocturnal homing and navigation, using two main cues: the spatial pattern of bright patches of daylight visible through the rainforest canopy, and compass information obtained from polarised skylight (from the setting sun or the moon) that penetrates these patches. [Copyright &y& Elsevier]

The basic precondition for color vision is the presence of at least two receptor types with different spectral sensitivities. The sensitivity of a receptor is mostly defined by the opsin-based visual pigment expressed in it. We show here, through behavioral experiments, that the nymphalid butterfly Heliconius erato, although it expresses short and medium wavelength opsins and only one long wavelength opsin, discriminates colors in the long-wavelength range (590 nm, 620 nm and 640 nm), whereas another nymphalid, Vanessa atalanta, despite having color vision, is unable to do so. In the eyes of H. erato we identified filtering pigments very close to the rhabdom which differ between ommatidia and produce the yellow and red ommatidial reflection seen under orthodromic illumination. The eyes of V. atalanta lack the filtering pigments, and reflect a homogeneous orange. We hypothesize that the filtering pigments found in the eyes of H. erato may shift the spectral sensitivity peak of the long wavelength receptors in some ommatidia towards longer wavelengths. The comparison of the signals between the two new receptor types makes color discrimination in the red range possible. To our knowledge, this is the first behavioral proof of color vision based on receptors expressing the same opsin. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The dual retina of humans and most vertebrates consists of multiple types of cone for colour vision in bright light and one single type of rod, leaving these animals colour-blind at night. Instead of comparing the signals from different spectral types of photoreceptors, they use one highly sensitive receptor, thus improving the signal-to-noise ratio. However, nocturnal moths and geckos can discriminate colours at extremely dim light intensities when humans are colour-blind, by sacrificing spatial and temporal rather than spectral resolution. The advantages of colour vision are just as obvious at night as they are during the day. Colour vision is much more reliable than achromatic contrast, not only under changing light intensities, but also under the colour changes occurring during dusk and dawn. It can be expected that nocturnal animals other than moths and geckos make use of the highly reliable colour signals in dim light. [ABSTRACT FROM AUTHOR]

Diurnal and nocturnal hawkmoths have been shown to use colour vision for flower discrimination. Here, we present evidence that the nocturnal hawkmoth Deilephila elpenor and the diurnal hawkmoth Macroglossum stellatarum also have colour constancy. Colour constancy was shown in D. elpenor in two multiple-choice experiments with five different bluish colour patches under white and blue illumination and with five yellowish colour patches under white, blue and yellow illumination. The mechanism underlying colour constancy in both species was investigated in two dual-choice experiments. The choice behaviour is consistent with the use of the von Kries coefficient law. Although the moths have colour constancy, they react to the colour of the illumination. They make fewer choices when tested under the changed illumination, where they never receive a reward, compared with the training illumination. Even if colour constancy can be explained by avon Kries adaptation mechanism, the fact that the animals discriminate between different illuminations indicates that some additional process must be involved. [ABSTRACT FROM AUTHOR]

Background: Some bees and wasps have evolved nocturnal behavior, presumably to exploit night-flowering plants or avoid predators. Like their day-active relatives, they have apposition compound eyes, a design usually found in diurnal insects. The insensitive optics of apposition eyes are not well suited for nocturnal vision. How well then do nocturnal bees and wasps see? What optical and neural adaptations have they evolved for nocturnal vision?Results: We studied female tropical nocturnal sweat bees (Megalopta genalis) and discovered that they are able to learn landmarks around their nest entrance prior to nocturnal foraging trips and to use them to locate the nest upon return. The morphology and optics of the eye, and the physiological properties of the photoreceptors, have evolved to give Megalopta''s eyes almost 30 times greater sensitivity to light than the eyes of diurnal worker honeybees, but this alone does not explain their nocturnal visual behavior. This implies that sensitivity is improved by a strategy of photon summation in time and in space, the latter of which requires the presence of specialized cells that laterally connect ommatidia into groups. First-order interneurons, with significantly wider lateral branching than those found in diurnal bees, have been identified in the first optic ganglion (the lamina ganglionaris) of Megalopta''s optic lobe. We believe that these cells have the potential to mediate spatial summation.Conclusions: Despite the scarcity of photons, Megalopta is able to visually orient to landmarks at night in a dark forest understory, an ability permitted by unusually sensitive apposition eyes and neural photon summation. [Copyright &y& Elsevier]

In behavioral experiments, the hawkmoth Deilephila elpenor can learn both the color and the position of artificial flowers. When very similar colors are used, moths select the correct color during the first test on a given day, thus using a stimulus strategy, but after repeated rewards, they switch to a place strategy and choose the flower in the position where they received the reward. When dissimilar colors are used, the moths continue to select flowers based on color and ignore position. We show how a computational model can reproduce the behavior in the experimental situation. The aim of the model is to investigate which learning and behavior selection strategies are necessary to reproduce the behavior observed in the experiment. The model is based on behavioral data and the sensitivities of the moth photoreceptors. The model consists of a number of interacting behavior systems that are triggered by specific stimuli and control specific behaviors. The ability of the moth to learn the colors of different flowers and the adaptive processes involved in the choice between stimulus-approach and place-approach strategies are reproduced very accurately by the model. The model has implications both for further studies of the ecology of the animal and for robotic systems. [ABSTRACT FROM AUTHOR]

Diurnal and nocturnal hawkmoths (Sphingidae, Lepidoptera) have three spectral types of receptor sensitive to ultraviolet, blue and green light. As avid flower visitors and pollinators, they use olfactory and visual cues to find and recognise flowers. Moths of the diurnal species Macroglossum stellatarum and the nocturnal species Deilephila elpenor, Hyles lineata and Hyles gallii use and learn the colour of flowers. Nocturnal species can discriminate flowers at starlight intensities when humans and honeybees are colour-blind. M. stellatarum can use achromatic, intensity-related cues if colour cues are absent, and this is probably also true for D. elpenor. Both species can recognise colours even under a changed illumination colour. [ABSTRACT FROM AUTHOR]

Provides an overview of the methods used to study animal color vision and discusses how quantitative modeling can suggest how photoreceptor signals are combined and compared to allow for the discrimination of biologically relevant stimuli. Color vision concepts; Uses of chromatic signals; Tests of visual mechanisms; Model thresholds for color discrimination.

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[sup -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. [ABSTRACT FROM AUTHOR]

Studies the color vision orientation of Papilio butteflies. Discrimination between vertically and horizontally polarized light of the same color in the context of oviposition and feeding; Ecological relevance of polarization-dependent color vision; Examples of other eyes that have similar structure.

Focuses on a study which developed simple models to describe the choice behaviour of ovipositing butterflies in terms of linear interactions between the different spectral types of photoreceptors. Colour-mediated behaviours in insects; Methodology; Discussion on the results.

Studies the innate preferences for flower features in hawkmoths. Choice between colors; Influence of hue and intensity; Role of flower size; Pattern preferences; Importance of parameters to moth choice; Choice curves plotting; Components of flower patterns.

Evaluates the learning performance of hawkmoth Macroglossum stellatarum, using the colours of artificial flowers as the cue to be learned. Testing methodology; How the learning behavior was influenced; How Macroglossum stellatarum L. (Lepidoptera, Sphingidae) was bred; What the study revealed; Colour learning and `forgetting'; Avoidance of unrewarded colour; Discussion and concluding remarks.

Examines the structure and function of learning flights in bees and wasps. Invariant elements and their consequences for information processing; Control of flight behavior; Relationship between learning flights and landmark guidance.

Colour vision allows animals to use the information contained in the spectrum of light to control important behavioural decisions such as selection of habitats, food or mates. Among arthropods, the largest animal phylum, we find completely colour-blind species as well as species with up to 40 different opsin genes or more than 10 spectral types of photoreceptors, we find a large diversity of optical methods shaping spectral sensitivity, we find eyes with different colour vision systems looking into the dorsal and ventral hemisphere, and species in which males and females see the world in different colours. The behavioural use of colour vision shows an equally astonishing diversity. Only the neural mechanisms underlying this sensory ability seems surprisingly conserved—not only within the phylum, but even between arthropods and the other well-studied phylum, chordates. The papers in this special issue allow a glimpse into the colourful world of arthropod colour vision, and besides giving an overview this introduction highlights how much more research is needed to fill in the many missing pieces of this large puzzle. This article is part of the theme issue 'Understanding colour vision: molecular, physiological, neuronal and behavioural studies in arthropods'. [ABSTRACT FROM AUTHOR]

The family Apidae, which is amongst the largest bee families, are important pollinators globally and have been well studied for their visual adaptations and visually guided behaviors. This review is a synthesis of what is known about their eyes and visual capabilities. There are many species-specific differences, however, the relationship between body size, eye size, resolution, and sensitivity shows common patterns. Salient differences between castes and sexes are evident in important visually guided behaviors such as nest defense and mate search. We highlight that Apis mellifera and Bombus terrestris are popular bee models employed in the majority of studies that have contributed immensely to our understanding vision in bees. However, other species, specifically the tropical and many non-social Apidae, merit further investigation for a better understanding of the influence of ecological conditions on the evolution of bee vision. [ABSTRACT FROM AUTHOR]

The article offers information on advances in physiological mechanisms of navigation in several organisms like tiny insects, snails, humans and robots. Topics discussed include navigational abilities of animals by using use visual landmarks, olfaction, magnetic maps, and celestial cues; brain centers involved in navigation by mammals and insects; and views on general principles of navigation that involves compass navigation, course control, odometer, spatial and olfactory memory.

• Raptors have large eyes allowing for high absolute sensitivity in nocturnal and high acuity in diurnal species. • Diurnal hunters have a deep central and a shallow temporal fovea, scavengers only a central and owls only a temporal fovea. • The spatial resolution of some large raptor species is the highest known among animals, but differs highly among species. • Visual fields of raptors reflect foraging strategies and depend on the divergence of optical axes and on head structures • More comparative studies on raptor retinae (preferably with non-invasive methods) and on visual pathways are desirable. Raptors have always fascinated mankind, owls for their highly sensitive vision, and eagles for their high visual acuity. We summarize what is presently known about the eyes as well as the visual abilities of these birds, and point out knowledge gaps. We discuss visual fields, eye movements, accommodation, ocular media transmittance, spectral sensitivity, retinal anatomy and what is known about visual pathways. The specific adaptations of owls to dim-light vision include large corneal diameters compared to axial (and focal) length, a rod-dominated retina and low spatial and temporal resolution of vision. Adaptations of diurnal raptors to high acuity vision in bright light include rod- and double cone-free foveae, high cone and retinal ganglion cell densities and high temporal resolution. We point out that more studies, preferably using behavioural and non-invasive methods, are desirable. [ABSTRACT FROM AUTHOR]

The amount of short wavelength (ultraviolet (UV), violet and blue) light that reaches the retina depends on the transmittance properties of the ocular media, especially the lens, and varies greatly across species in all vertebrate groups studied previously. We measured the lens transmittance in 32 anuran amphibians with different habits, geographical distributions and phylogenetic positions and used them together with eye size and pupil shape to evaluate the relationship with diel activity pattern, elevation and latitude. We found an unusually high lens UV transmittance in the most basal species, and a cut-off range that extends into the visible spectrum for the rest of the sample, with lenses even absorbing violet light in some diurnal species. However, other diurnal frogs had lenses that transmit UV light like the nocturnal species. This unclear pattern in the segregation of ocular media transmittance and diel activity is shared with other vertebrates and is consistent with the absence of significant correlations in our statistical analyses. Although we did not detect a significant phylogenetic effect, closely related species tend to have similar transmittances, irrespective of whether they share the same diel pattern or not, suggesting that anuran ocular media transmittance properties might be related to phylogeny. [ABSTRACT FROM AUTHOR]

Summary Butterfly eyes are random mosaics built of three ommatidia types, each with a different set of photoreceptors and pigments. What defines the combined features in each ommatidium? A new study has solved the puzzle. [ABSTRACT FROM AUTHOR]

The binocular field of vision differs widely in birds depending on ecological traits such as foraging. Owls (Strigiformes) have been considered to have a unique binocular field, but whether it is related to foraging has remained unknown. While taking into account allometry and phylogeny, we hypothesized that both daily activity cycle and diet determine the size and shape of the binocular field in owls. Here, we compared the binocular field configuration of 23 species of owls. While we found no effect of allometry and phylogeny, ecological traits strongly influence the binocular field shape and size. Binocular field shape of owls significantly differed from that of diurnal raptors. Among owls, binocular field shape was relatively conserved, but binocular field size differed among species depending on ecological traits, with larger binocular fields in species living in dense habitat and foraging on invertebrates. Our results suggest that (i) binocular field shape is associated with the time of foraging in the daily cycle (owls versus diurnal raptors) and (ii) that binocular field size differs between closely related owl species even though the general shape is conserved, possibly because the field of view is partially restricted by feathers, in a trade-off with auditory localization. [ABSTRACT FROM AUTHOR]

The binocular field of vision differs widely in birds depending on ecological traits such as foraging. Owls (Strigiformes) have been considered to have a unique binocular field, but whether it is related to foraging has remained unknown. While taking into account allometry and phylogeny, we hypothesized that both daily activity cycle and diet determine the size and shape of the binocular field in owls. Here, we compared the binocular field configuration of 23 species of owls. While we found no effect of allometry and phylogeny, ecological traits strongly influence the binocular field shape and size. Binocular field shape of owls significantly differed from that of diurnal raptors. Among owls, binocular field shape was relatively conserved, but binocular field size differed among species depending on ecological traits, with larger binocular fields in species living in dense habitat and foraging on invertebrates. Our results suggest that (i) binocular field shape is associated with the time of foraging in the daily cycle (owls versus diurnal raptors) and (ii) that binocular field size differs between closely related owl species even though the general shape is conserved, possibly because the field of view is partially restricted by feathers, in a trade-off with auditory localization. [ABSTRACT FROM AUTHOR]

We discovered nocturnal colour vision in the Asian giant honeybee Apis dorsata—a facultatively nocturnal species—at mesopic light intensities, down to half-moon light levels (approx. 10−2 cd m−2). The visual threshold of nocturnality aligns with their reported nocturnal activity down to the same light levels. Nocturnal colour vision in A. dorsata is interesting because, despite being primarily diurnal, its colour vision capabilities extend into dim light, while the 'model' European honeybee Apis mellifera is reported to be colour-blind at twilight. By employing behavioural experiments with naturally nesting A. dorsata colonies, we show discrimination of the trained colour from other stimuli during the day, and significantly, even at night. Nocturnal colour vision in bees has so far only been reported in the obligately nocturnal carpenter bee Xylocopa tranquebarica. The discovery of colour vision in these two bee species, despite differences in the extent of their nocturnality and the limitations of their apposition compound eye optics, opens avenues for future studies on visual adaptations for dim-light colour vision, their role in pollination of flowers at night, and the effect of light pollution on nocturnal activity in A. dorsata, a ubiquitous pollinator in natural, agricultural and urban habitats in the Asian tropics and sub-tropics. [ABSTRACT FROM AUTHOR]

Animals are thought to use achromatic signals to detect small (or distant) objects and chromatic signals for large (or nearby) objects. While the spatial resolution of the achromatic channel has been widely studied, the spatial resolution of the chromatic channel has rarely been estimated. Using an operant conditioning method, we determined (i) the achromatic contrast sensitivity function and (ii) the spatial resolution of the chromatic channel of a diurnal raptor, the Harris's hawk Parabuteo unicinctus. The maximal spatial resolution for achromatic gratings was 62.3 c deg-1, but the contrast sensitivity was relatively low (10.8-12.7). The spatial resolution for isoluminant redgreen gratings was 21.6 c deg-1--lower than that of the achromatic channel, but the highest found in the animal kingdom to date. Our study reveals that Harris's hawks have high spatial resolving power for both achromatic and chromatic vision, suggesting the importance of colour vision for foraging. By contrast, similar to other bird species, Harris's hawks have low contrast sensitivity possibly suggesting a trade-off with chromatic sensitivity. The result is interesting in the light of the recent finding that double cones--thought to mediate high-resolution vision in birds--are absent in the central fovea of raptors. [ABSTRACT FROM AUTHOR]

Little is known about the development of vision in wild birds. It is unknown, for example, whether the ability to see can be predicted by the level of prenatal growth or whether the eyes are open at hatching in a particular species. In this study, we investigated the growth of eyes, the formation of retinal ganglion cell topography, and the appearance of simple, visually guided behaviours in chicks of a small procellariiform seabird, Leach's storm petrel (Oceanodroma leucorhoa). This semi-precocial species, which has a well-developed sense of smell, nests in underground burrows where adults provision chicks for 6-8 weeks in the dark before fledging. Retinal ganglion cell topographic maps revealed that fine-tuning of cell distribution does not happen early in development, but rather that the ganglion cell layer continues to mature throughout provisioning and probably even after fledging. While the olfactory bulbs reached adult size around 7 weeks after hatching, the eyes and telencephalon continued to grow. Optokinetic head response and artificial burrow finding experiments indicated that chicks in the 2nd week after hatching lack even the most basic visually guided behaviours and are probably blind. Thus, vision in Leach's storm petrel chicks starts to function sometime around the 3rd week after hatching, well after the eyes have opened and the olfactory system is functional. [ABSTRACT FROM AUTHOR]