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Most animals are visually oriented, and their eyes provide their 'window to the world'. Eye size correlates positively with visual performance, because larger eyes can house larger pupils that increase photon catch and contrast discrimination, particularly under dim light, which have positive effects on behaviours that enhance fitness, including predator avoidance and foraging. Recent studies have linked predation risk to selection for larger eyes and pupils, and such changes should be of importance for the majority of teleost fishes as they have a pupil that is fixed in size (eyes lack a pupillary sphincter muscle) and, hence, do not respond to changes in light conditions. Here, we quantify eye and pupil size of individual crucian carp, a common freshwater fish, following controlled manipulations of perceived predation risk (presence/absence). We also tested if crucian carp responded to increased predation risk by shifts in diel activity patterns. We found that crucian carp show phenotypic plasticity with regards to pupil size, but not eye size, as pupil size increased when exposed to predators (pike). Predator-exposed crucian carp also shifted from diurnal to nocturnal activity. Using a modelling exercise, we moreover show that the plastically enlarged pupils significantly increase visual range, especially for small objects under dim light conditions. Overall, our results provide compelling evidence for predator-induced pupil enlargement resulting in enhanced visual capabilities in a teleost fish. Pupil size plasticity in combination with the observed shift towards nocturnal activity may allow for efficient foraging also under dark conditions when predation risk from diurnal and visually oriented predators is reduced. The data highlight the powerful role of predation risk for eye development and evolution.

Millipedes are a species-rich and ancient arthropod clade which typically bear a pair of lateral compound eyes with a small number of large facets. To understand the visual tasks that underlie the evolution of millipede eyes, their spatial resolving performance is of key importance. We here investigate the spatial resolution of the millipede Cylindroiulus punctatus using behavioural assays. Individual animals were placed in the centre of a cylindrical arena under bright downwelling light, with dark stimuli of varying angular dimensions placed on the arena wall. We used continuous isoluminant stimuli based on a difference of Gaussians signal to test for orientation to the dark target via object taxis. Headings of individual animals were tracked in relation to the stimuli to determine whether the animals oriented towards the stimulus. We implemented a multilevel logistic regression model to identify the arc width of the stimulus that animals could resolve. We then modelled the angular sensitivity needed to identify this. We also related the visual performance to the 3D anatomy of the eye. We found that C. punctatus can resolve a stimulus of 56° period (sufficient to detect a 20° dark target). Assuming a contrast threshold of 10%, this requires a receptor acceptance angle of 72° or narrower. Spatial resolving power this low would only suffice for the simplest visual tasks, such as shelter-seeking.

Every aspect of vision, from the opsin proteins to the eyes and the ways that they serve animal behavior, is incredibly diverse. It is only with an evolutionary perspective that this diversity can be understood and fully appreciated. In this review, I describe and explain the diversity at each level and try to convey an understanding of how the origin of the first opsin some 800 million years ago could initiate the avalanche that produced the astonishing diversity of eyes and vision that we see today. Despite the diversity, many types of photoreceptors, eyes, and visual roles have evolved multiple times independently in different animals, revealing a pattern of eye evolution strictly guided by functional constraints and driven by the evolution of gradually more demanding behaviors. I conclude the review by introducing a novel distinction between active and passive vision that points to uncharted territories in vision research.

[Graphic][1] Mike Land contemplating discoveries after a long day on board the British Research vessel RRS Discovery, 1987, in the Atlantic outside West Africa. Michael Francis Land – known to his colleagues and friends as Mike – was born in Dartmouth, UK, on 12 April 1942 and died

Calibration procedures, method tests and useful tips

Simple roles for photoreception are likely to have preceded more demanding ones such as vision. The driving force behind this evolution is the improvement and elaboration of animal behaviors using photoreceptor input. Because the basic role for all senses aimed at the external world is to guide behavior, we argue here that understanding this "behavioral drive" is essential for unraveling the evolutionary past of the senses. Photoreception serves many different types of behavior, from simple shadow responses to visual communication. Based on minimum performance requirements for different types of tasks, photoreceptors have been argued to have evolved from non-directional receptors, via directional receptors, to low-resolution vision, and finally to high-resolution vision. Through this sequence, the performance requirements on the photoreceptors have gradually changed from broad to narrow angular sensitivity, from slow to fast response, and from low to high contrast sensitivity during the evolution from simple to more advanced and demanding behaviors. New behaviors would only evolve if their sensory performance requirements to some degree overlap with the requirements of already existing behaviors. This need for sensory "performance continuity" must have determined the order by which behaviors have evolved and thus been an important factor guiding animal evolution. Naturally, new behaviors are most likely to evolve from already existing behaviors with similar neural processing needs and similar motor responses, pointing to "neural continuity" as another guiding factor in sensory evolution. Here we use these principles to derive an evolutionary tree for behaviors driven by photoreceptor input.

Eyes are not unique to animals. As described by Nilsson and Marshall, prominent eyes, complete with retina and lens, have unexpectedly evolved in single cell dinoflagellates.

Cubozoan medusae have a stereotypic set of 24 eyes, some of which are structurally similar to vertebrate and cephalopod eyes. Across the approximately 25 described species, this set of eyes varies surprisingly little, suggesting that they are involved in an equally stereotypic set of visual tasks. During the day Tripedalia cystophora is found at the edge of mangrove lagoons where it accumulates close to the surface in sun-lit patches between the prop roots. Copula sivickisi (formerly named Carybdea sivickisi) is associated with coral reefs and has been observed to be active at night. At least superficially, the eyes of the two species are close to identical. We studied the diurnal activity pattern of these two species both in the wild and under controlled conditions in laboratory experiments. Despite the very similar visual systems, we found that they display opposite patterns of diurnal activity. T. cystophora is active exclusively during the day, whereas C. sivickisi is actively swimming at night, when it forages and mates. At night T. cystophora is found on the muddy bottom of the mangrove lagoon. C. sivickisi spends the day attached to structures such as the underside of stones and coral skeletons. This species difference seems to have evolved to optimize foraging, since the patterns of activity follow those of the available prey items in their respective habitats.

Box jellyfish (Cubomedusae) are visually orientating animals which possess a total of 24 eyes of 4 morphological types; 2 pigment cup eyes (pit eye and slit eye) and 2 lens eyes [upper lens-eye (ule) and lower lens-eye (lle)]. In this study, we use electroretinograms (ERGs) to explore temporal properties of the two lens eyes. We find that the ERG of both lens eyes are complex and using sinusoidal flicker stimuli we find that both lens eyes have slow temporal resolution. The average flicker fusion frequency (FFF) was found to be approximately 10 Hz for the ule and 8 Hz for the lle. Differences in the FFF and response patterns between the two lens eyes suggest that the ule and lle filter information differently in the temporal domain and thus are tuned to perform different visual tasks. The data collected in this study support the idea that the visual system of box jellyfish is a collection of special purpose eyes.

Cubomedusae have a total of 24 eyes of four morphologically different types. Two of these eye types are camera-type eyes (upper and lower lens-eye), while the other two eye types are simpler pigment pit eyes (pit and slit eye). Here, we give a description of the visual system of the box jellyfish species Chiropsella bronzie and the optics of the lens eyes in this species. One aim of this study is to distinguish between general cubozoan features and species-specific features in the layout and optics of the eyes. We find that both types of lens eyes are more severely under-focused in C. bronzie than those in the previously investigated species Tripedalia cystophora. In the lower lens-eye of C. bronzie, blur circles subtend 20 and 52 degrees for closed and open pupil, respectively, effectively removing all but the coarsest structures of the image. Histology reveals that the retina of the lower lens-eye, in addition to pigmented photoreceptors, also contains long pigment-cells, with both dark and white pigment, where the dark pigment migrates on light/dark adaptation. Unlike the upper lens-eye lens of T.cystophora, the same eye in C.bronzie did not display any significant optical power.

The visual system of box jellyfish comprises a total of 24 eyes. These are of four types and each probably has a special function. To investigate this hypothesis the morphology and optics of the lesser eyes, the pit and slit eyes, were examined. The pit eyes hold one cell type only and are probably mere light meters. The slit eyes, comprising four cell types, are complex and highly asymmetric. They also hold a lens-like structure, but its optical power is minute. Optical modeling suggests spatial resolution, but only in one plane. These unique and intriguing traits support strong peripheral filtering.

Box jellyfish have the most elaborate sensory system and behavioural repertoire of all cnidarians. Sensory input largely comes from 24 eyes situated on four club-shaped sensory structures, the rhopalia, and behaviour includes obstacle avoidance, light shaft attractance and mating. To process the sensory input and convert it into the appropriate behaviour, the box jellyfish have a central nervous system (CNS) but this is still poorly understood. The CNS has two major components: the rhopalial nervous system and the ring nerve. The rhopalial nervous system is situated within the rhopalia in close connection with the eyes, whereas the ring nerve encircles the bell. We describe the morphology of the ring nerve of the box jellyfish Tripedalia cystophora as ascertained by normal histological techniques, immunohistochemistry and transmission electron microscopy. By light microscopy, we have estimated the number of cells in the ring nerve by counting their nuclei. In cross sections at the ultrastructural level, the ring nerve appears to have three types of neurites: (1) small "normal"-looking neurites, (2) medium-sized neurites almost completely filled by electron-lucent vacuoles and (3) giant neurites. In general, only one giant neurite is seen on each section; this type displays the most synapses. Epithelial cells divide the ring nerve into compartments, each having a tendency to contain neurites of similar morphology. The number and arrangement of the compartments vary along the length of the ring nerve.

Cubozoans, or box jellyfish, each have twenty-four eyes of four types, but no central brain for information processing. An investigation of these eyes reveals optics as sophisticated as in vertebrates. Despite this, the retina is out of focus and the sharp image is not used to provide acute vision, but as a way of processing visual information. ‘Blurred’ vision may be perfect for avoiding large stationary objects without focusing on small floating objects and plankton. This may also be a pointer to a missing link in the early evolution of animal visual systems, likely to have involved eyes performing a single visual task only. The cover shows the two lens eyes and two pairs of pigment pit eyes in the bizarre sensory club of Chiropsalmus sp., larger but similar to those of Tripedalia cystophora used in the optical study. Cubozoans, or box jellyfish, differ from all other cnidarians by an active fish-like behaviour and an elaborate sensory apparatus1,2. Each of the four sides of the animal carries a conspicuous sensory club (the rhopalium), which has evolved into a bizarre cluster of different eyes3. Two of the eyes on each rhopalium have long been known to resemble eyes of higher animals, but the function and performance of these eyes have remained unknown4. Here we show that box-jellyfish lenses contain a finely tuned refractive index gradient producing nearly aberration-free imaging. This demonstrates that even simple animals have been able to evolve the sophisticated visual optics previously known only from a few advanced bilaterian phyla. However, the position of the retina does not coincide with the sharp image, leading to very wide and complex receptive fields in individual photoreceptors. We argue that this may be useful in eyes serving a single visual task. The findings indicate that tailoring of complex receptive fields might have been one of the original driving forces in the evolution of animal lenses.

Earlier detailed studies of cnidarian planula larvae have revealed a simple nervous system but no eyes or identifiable light sensing structures. Here, we describe the planula of a box jellyfish, Tripedalia cystophora, and report that these larvae have an extremely simple organization with no nervous system at all. Their only advanced feature is the presence of 10-15 pigment-cup ocelli, evenly spaced across the posterior half of the larval ectoderm. The ocelli are single cell structures containing a cup of screening pigment filled with presumably photosensory microvilli. These rhabdomeric photoreceptors have no neural connections to any other cells, but each has a well-developed motor-cilium, appearing to be the only means by which light can control the behaviour of the larva. The ocelli are thus self-contained sensory-motor entities, making a nervous system superfluous.

Some insects and vertebrates use the pattern of polarized light in the sky as an optical compass1,2,3,4,5. Only a small section of clear sky needs to be visible for bees and ants to obtain a compass bearing for accurate navigation5,6. The receptors involved in the polarization compass are confined to a small part of the retina, and the eyes are built predominantly for other visual tasks7. Here we report the discovery of a unique compass organ in the spider Drassodes cupreus, where a pair of specialized secondary eyes cooperate to analyse skylight polarization. These eyes do not form images, but use a built-in polarization filter to determine precisely the direction of polarization. Measurements using a model eye indicate that the compass organ is best suited for navigation at dusk and dawn. Behavioural experiments show that the spiders are primarily active after sunset and that they use polarization cues to find their way back to the nest after foraging trips. A similar organization of the secondary eyes in several spider families indicates that such compass organs may not be an isolated phenomenon.

The fraction F of incident light absorbed by a photoreceptor of length l has traditionally been given by F =1 -e −kl , where k is the absorption coefficient of the photoreceptor. Unfortunately, this widely-used expression is incorrect for absorption of the type of light most common in natural scenes—broad spectrum “white” light—and significantly over-estimates absorption. This is because the measured values of k are only valid at the absorbance peak wavelength of rhodopsin, whereas at other wavelengths (which the eye may also see) k is lower. We have accounted for the wavelength dependence of k and calculated the absorption of white light from four different natural radiant sources: the quantal irradiances of natural daylight and a patch of very blue sky, and the quantal reflections of soil and green foliage irradiated by natural daylight. Based on these results, a simple averaged correction for white light stimulation is derived, F = kl /(2.3 + kl ), which is valid for a wide range of k and l , and therefore applicable to both vertebrate and invertebrate photoreceptors.

The luminance range over which the pupil mechanism operates was measured with pupil reflectometry in 11 species of butterflies and 13 species of dipteran flies. The different species were selected to be as different as possible regarding the range of ambient luminances in which they are active. Habitat luminance ranges were also measured and correlated to luminances in the experimental situation. The pupil mechanism in butterflies operates in the centre of the luminance range in which the different species are active. Three distinct groups of butterflies with pupil sensitivities matched to their specific types of activity pattern were identified: species active only in direct sunlight, species active also in shaded places and species extending their activity into dawn and dusk. Quite differently, the pupil mechanisms of dipteran flies operate in the upper end of the ambient luminances, and in some species well above the luminances normally encountered by the animal. All fly pupils start to close roughly at the same luminance, irrespective of the luminances in which the species are active. The results suggest that the most important role for the pupil mechanism in many of the butterfly species is to maximize acuity over a wide range of luminances, whereas in flies it is to avoid saturation of transduction units and thereby maximize the photoreceptor's signal-to-noise ratio at high light intensities.

Eye evolution is driven by the evolution of visually guided behavior. Accumulation of gradually more demanding behaviors have continuously increased the performance requirements on the photoreceptor organs. Starting with nondirectional photoreception, I argue for an evolutionary sequence continuing with directional photoreception, low-resolution vision, and finally, high-resolution vision. Calculations of the physical requirements for these four sensory tasks show that they correlate with major innovations in eye evolution and thus work as a relevant classification for a functional analysis of eye evolution. Together with existing molecular and morphological data, the functional analysis suggests that urbilateria had a simple set of rhabdomeric and ciliary receptors used for directional photoreception, and that organ duplications, positional shifts and functional shifts account for the diverse patterns of eyes and photoreceptors seen in extant animals. The analysis also suggests that directional photoreception evolved independently at least twice before the last common ancestor of bilateria and proceeded several times independently to true vision in different bilaterian and cnidarian groups. This scenario is compatible withPax-gene expression in eye development in the different animal groups. The whole process from the first opsin to high-resolution vision took about 170 million years and was largely completed by the onset of the Cambrian, about 530 million years ago. Evolution from shadow detectors to multiple directional photoreceptors has further led to secondary cases of eye evolution in bivalves, fan worms, and chitons.

The vast differences between vertebrate and arthropod eyes suggest that the recently discovered homologous master control genes for eye development had another function before eyes evolved in the early Cambrian.

The pupil mechanism in the acone apposition eye of the semi-aquatic common backswimmer Notonecta glauca (Hemiptera) was investigated with infrared reflectometry of the pseudopupil. This method allows non-invasive continuous measurements of pupil responses in the living animal. The dynamic range of the pupil sensitivity is about 7 log units during daytime and 6 log units at night. During the day, the sensitivity range of the pupil covers the normal daylight intensities in the animal’s habitat, just under the water surface (I50=1019.2 photons m−2 sr−1 s−1). At night, the sensitivity is 1 log unit lower (I50=1020.2 photons m−2 sr−1 s−1), ensuring that the pupil is maximally open when light intensities are low. During daytime, light adaptation is completed in slightly less than 40 min, and dark adaptation takes approximately 50 min. The pupil response is only slightly slower at night. The speed of the response as well as the pupil sensitivity are dependent on the preceding adaptation history. An endogenous circadian rhythm determines the control range of the pupil aperture. However, the rhythm is easily disturbed, especially within a 3 h period before dusk and dawn. The results are compared with corresponding results from other insects with the same type of pupil mechanism.

Infrared reflectometry of the deep pseudopupil was used for measuring the absolute sensitivity and the dynamics of the pupil mechanism in the acone apposition eye of tipulid flies. The following species were studied: Tipula luteipennis, T. fusca, T. pagana, T. paludosa and T. staegeri. Pupil responses plotted against light intensity produced S-shaped R log I curves that were fitted by Lipetz equations with I50-values (intensity corresponding to 50% of the maximal response) ranging between 1017.4 and 1020.2 photons m−2 str−1 s−1. The sensitivity of the pupil is considerably higher (1–2 log units) during daytime than at night in all examined species. Differences in the sensitivity between different tipulid species are explained by the different light intensities in their natural environment. The dynamic range of the pupil mechanism is exceptionally large. In T. luteipennis, the pupil operates over a total range of about 8.5 log units of light intensity. This implies that the pupil mechanism of T. luteipennis is functional at light intensities ranging from bright sunlight down to moonlight. The speed of the pupillary movements are slower at night than during the day. During daytime, light adaptation is completed in about 30 min and dark adaptation in about 50 min. At night, the corresponding values are 45 and 60 min. The differences in speed and sensitivity of the pupil mechanism between day and night are due to a circadian rhythm, setting the control range of the pupil aperture.

Fossil feathers, hairs and eyes are regularly preserved as carbonized traces comprised of masses of micrometre-sized bodies that are spherical, oblate or elongate in shape. For a long time, these minute structures were regarded as the remains of biofilms of keratinophilic bacteria, but recently they have been reinterpreted as melanosomes; that is, colour-bearing organelles. Resolving this fundamental difference in interpretation is crucial: if endogenous then the fossil microbodies would represent a significant advancement in the fields of palaeontology and evolutionary biology given, for example, the possibility to reconstruct integumentary colours and plumage colour patterns. It has previously been shown that certain trace elements occur in fossils as organometallic compounds, and hence may be used as biomarkers for melanin pigments. Here we expand this knowledge by demonstrating the presence of molecularly preserved melanin in intimate association with melanosome-like microbodies isolated from an argentinoid fish eye from the early Eocene of Denmark.

Den här skriften handlar om ljus och ljusets betydelse för oss människor. Tolv forskare med vitt skilda bakgrunder samlas och diskuterar frågor som skrider över gränserna. Vad är ljus egentligen? Hur kan vi förstå ljus? Hur påverkar det oss? På vilket sätt kan vi använda oss av det? Vilka tekniska möjligheter har vi att skapa och styra olika sorters ljus? Vilka möjligheter ger det oss inför framtiden? I centrum för dialogen står samarbete och nytänkande. Här ges en inblick i perspektiv på ljus från områden som biologi, teknik, material, fysik, estetik, filosofi, medicin och psykologi, vilka sammantaget, på ett åskådligt och lättbegripligt sätt, presenterar den flervetenskapliga ljusrelaterade kompetensen i Lund. Gruppen av experter hoppas att de frågor som ställs och de svar som ges i den här skriften ska inspirera och leda vidare inom fältet ljusforskning. Tillsammans med Pufendorfinstitutet, genom vilket expertgruppens initiativ getts möjlighet att växa och utvecklas, vill de med andra ord att sätta ljuset på...just ”ljuset”!

Objects can differ in brightness and colour. At least that is what our own visual system tells us. It now seems that stomatopod shrimps, and possibly also cephalopod molluscs, can see the direction of the electric vector of light, in much the same way we see colour.

SummaryRecent work on the expression of retinal transcription factors and other molecular cues delivers interesting but partly contradictory information on the early phases of eye evolution.

The optical information processing that takes place in an eye involves a large variety of very different optical components that are put together to solve a task which, in its basic nature, does not differ much from one species to another. The principal task of an eye is to sort the incoming photons so that they excite specific sensory neurons depending on angle of incidence, wavelength and plane of polarization. Despite the apparent simplicity of the task, the solutions to it are often complex and the variation between species is enormous. The pinhole camera, the Keplerian and Galilean telescopes, the corner reflector, optical fibres, and interference filters, are all names of optical devices invented by man. It now appears that all of these devices, and many more, exist in various combinations in the optics of invertebrate eyes. The similarity between man's and nature's optical engineering has been useful in many ways. For the study of eyes, it has helped to understand biological design principles in unparalleled detail.

SUMMARY Box jellyfish, cubomedusae, possess an impressive total of 24 eyes of four morphologically different types. Two of these eye types, called the upper and lower lens eyes, are camera-type eyes with spherical fish-like lenses. Compared with other cnidarians, cubomedusae also have an elaborate behavioral repertoire, which seems to be predominantly visually guided. Still, positive phototaxis is the only behavior described so far that is likely to be correlated with the eyes. We have explored the obstacle avoidance response of the Caribbean species Tripedalia cystophora and the Australian species Chiropsella bronzie in a flow chamber. Our results show that obstacle avoidance is visually guided. Avoidance behavior is triggered when the obstacle takes up a certain angle in the visual field. The results do not allow conclusions on whether color vision is involved but the strength of the response had a tendency to follow the intensity contrast between the obstacle and the surroundings (chamber walls). In the flow chamber Tripedalia cystophora displayed a stronger obstacle avoidance response than Chiropsella bronzie since they had less contact with the obstacles. This seems to follow differences in their habitats.

Cubomedusae, or box jellyfish, have the most elaborate visual system of all cnidarians. They have 24 eyes of four morphological types, distributed on four sensory structures called rhopalia. Box jellyfish also display complex, probably visually guided behaviors such as obstacle avoidance and fast directional swimming. Here we describe the strikingly complex and partially bilaterally symmetrical nervous system found in each rhopalium of the box jellyfish, Tripedalia cystophora, and present the rhopalial neuroanatomy in an atlas-like series of drawings. Discrete populations of neurons and commissures connecting the left and the right side along with two populations of nonneuronal cells were visualized using several different histochemical staining techniques and electron microscopy. The number of rhopalial nerve cells and their overall arrangement indicates that visual processing and integration at least partly happen within the rhopalia. The larger of the two nonneuronal cell populations comprises approximately 2,000 likely undifferentiated cells and may support a rapid cell turnover in the rhopalial nervous system.

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.

Box jellyfish, or cubomedusae, possess an impressive total of 24 eyes of four morphologically different types. Compared to other cnidarians they also have an elaborate behavioral repertoire, which for a large part seems to be visually guided. Two of the four types of cubomedusean eyes, called the upper and the lower lens eye, are camera type eyes with spherical fish-like lenses. Here we explore the electroretinograms of the lens eyes of the Caribbean species, Tripedalia cystophora, and the Australian species, Chiropsalmus sp. using suction electrodes. We show that the photoreceptors of the lens eyes of both species have dynamic ranges of about 3 log units and slow responses. The spectral sensitivity curves for all eyes peak in the blue-green region, but the lower lens eye of T. cystophora has a small additional peak in the near UV range. All spectral sensitivity curves agree well with the theoretical absorbance curve of a single opsin, strongly suggesting color-blind vision in box jellyfish with a single receptor type. A single opsin is supported by selective adaptation experiments.

In cubomedusae, the central nervous system (CNS) is found both in the bell (the ring nerve) and in the four eye-bearing sensory structures (the rhopalia). The ring nerve and the rhopalia are connected via the rhopalial stalks and examination of the structure of the rhopalial stalks therefore becomes important when trying to comprehend visual processing. In the present study, the rhopalial stalk of the cubomedusae Tripedalia cystophora has been examined by light microscopy, transmission electron microscopy, and electrophysiology. A major part of the ring nerve is shown to continue into the stalk and to contact the rhopalial neuropil directly. Ultrastructural analysis of synapse distribution in the rhopalial stalk has failed to show any clustering, which indicates that integration of the visual input is probably spread throughout the CNS. Together, the results indicate that cubomedusae have one coherent CNS including the rhopalia. Additionally, a novel gastrodermal nerve has been found in the stalk; this nerve is not involved in visual processing but is likely to be mechanosensory and part of a proprioceptory system.

Spontaneous activation of rhodopsin without light absorption occurs at a much lower rate in rod photoreceptors and insect rhabdoms than in cones. The difference lies in the pigment molecules themselves, and has implications for the design of visual photoreceptors.

Animal eyes have long served as a classical example of independent origin followed by convergence of structures onto a few different solutions. During the past decade this view has been challenged by the discovery of shared developmental regulatory genes. The Pax6 gene in particular is almost universally employed for eye formation in bilaterian animals, despite widely different embryological origins. The resulting controversy on the multiple or single origins of animal eyes has gradually been sharpened by continuing discoveries of further general similarities in the genetic regulatory circuits of eye development. Recent work on gene expression in specified cell types, together with comparative studies of developmental genes in cnidarians, now show some promise to a solution of the controversy.

Visual performance and visual interactions in pelagic animals are notoriously hard to investigate because of our restricted access to the habitat. The pelagic visual world is also dramatically different from benthic or terrestrial habitats, and our intuition is less helpful in understanding vision in unfamiliar environments. Here, we develop a computational approach to investigate visual ecology in the pelagic realm. Using information on eye size, key retinal properties, optical properties of the water and radiance, we develop expressions for calculating the visual range for detection of important types of pelagic targets. We also briefly apply the computations to a number of central questions in pelagic visual ecology, such as the relationship between eye size and visual performance, the maximum depth at which daylight is useful for vision, visual range relations between prey and predators, counter-illumination and the importance of various aspects of retinal physiology. We also argue that our present addition to computational visual ecology can be developed further, and that a computational approach offers plenty of unused potential for investigations of visual ecology in both aquatic and terrestrial habitats.

Background We recently reported (Curr Biol 22:683–688, 2012) that the eyes of giant and colossal squid can grow to three times the diameter of the eyes of any other animal, including large fishes and whales. As an explanation to this extreme absolute eye size, we developed a theory for visual performance in aquatic habitats, leading to the conclusion that the huge eyes of giant and colossal squid are uniquely suited for detection of sperm whales, which are important squid-predators in the depths where these squid live. A paper in this journal by Schmitz et al. (BMC Evol Biol 13:45, 2013) refutes our conclusions on the basis of two claims: (1) using allometric data they argue that the eyes of giant and colossal squid are not unexpectedly large for the size of the squid, and (2) a revision of the values used for modelling indicates that large eyes are not better for detection of approaching sperm whales than they are for any other task. Results and conclusions We agree with Schmitz et al. that their revised values for intensity and abundance of planktonic bioluminescence may be more realistic, or at least more appropriately conservative, but argue that their conclusions are incorrect because they have not considered some of the main arguments put forward in our paper. We also present new modelling to demonstrate that our conclusions remain robust, even with the revised input values suggested by Schmitz et al.

Dragonflies of the genus Sympetrum have compound eyes conspicuously divided into dorsal and ventral regions. Using anatomical, optical, electrophysiological, in-vivo photochemical and microspectrophotometrical methods, we have investigated the design and physiology of the dorsal part which is characterized by a pale yellow-orange screening pigment and extremely large facets. The upper part of the yellow dorsal region is a pronounced fovea with interommatidial angles approaching 0.3°, contrasting to the much larger values of 1.5°–2° in the rest of the eye. The dorsal eye part is exclusively sensitive to short wavelengths (below 520 nm). It contains predominantly blue-receptors with a sensitivity maximum at 420 nm, and a smaller amount of UV-receptors. The metarhodopsin of the blue-receptors absorbs maximally at 535 nm. The yellow screening pigment transmits longwavelength light (cut-on 580 nm), which increases the conversion rate from metarhodopsin to rhodopsin (see Fig. 11a). We demonstrate that because of the yellow pigment screen nearly all of the photopigment is in the rhodopsin state under natural conditions, thus maximizing sensitivity. Theoretical considerations show that the extremely long rhabdoms (1.1 mm) in the dorsal fovea are motivated for absorption reasons alone. A surprising consequence of the long rhabdoms is that the sensitivity gain, caused by pumping photopigment into the rhodopsin state, is small. To explain this puzzling fact we present arguments for a mechanism producing a gradient of rhodopsin concentration along the rhabdom, which would minimize saturation of transduction units, and hence improve the signal-to-noise ratio at high intensities. The latter is of special importance for the short integration time and high contrast sensitivity these animals need for spotting small prey at long distances.

Observations of the infrared deep pseudopupil, optical determinations of the corneal nodal point, and histological methods were used to relate the visual fields of individual rhabdomeres to the array of ommatidial optical axes in four insects with open rhabdoms: the tenebrionid beetle Zophobas morio, the earwig Forficula auricularia, the crane fly Tipula pruinosa, and the backswimmer Notonecta glauca.

The energy dependence of the pupil pigment-migrations in the fly Musca domestica was studied in live animals, using optical techniques and nitrogen-gas induced anoxia. The results obtained can be summarized in 3 points: 1. Energy deficiency can make the pupil mechanism stop in any state, extreme or intermediate. 2. Anoxia induced during intermittent stimulation makes the pupil stop in the closed state (aggregated pigment granules). 3. During long-term anoxia the pupil very slowly opens (dispersal of pigment granules), irrespective of ambient intensity.

Theoretical considerations of eye design allow us to find routes along which the optical structures of eyes may have evolved. If selection constantly favours an increase in the amount of detectable spatial information, a light-sensitive patch will gradually turn into a focused lens eye through continuous small improvements of design. An upper limit for the number of generations required for the complete transformation can be calculated with a minimum of assumptions. Even with a consistently pessimistic approach the time required becomes amazingly short: only a few hundred thousand years.

Using infrared reflectometry of the deep pseudopupil, we have measured the absolute sensitivity, the dynamic range and the speed of the pupil mechanism in the acone apposition eye of two tenebrionid beetles: Zophobas morio F. and Tenebrio molitor L. The following conclusions are made from the results: 1. There is a substantial difference in sensitivity of the pupil mechanism between the two beetle species. The pupil is about 5.3 log units more sensitive in Zophobas than in Tenebrio. 2. There is also a difference in sensitivity between day and night. Surprisingly, the sensitivity is higher at day-time, and the difference is about 0.5 log units in both Zophobas and Tenebrio. 3. Light adaptation is completed faster during daytime than at night in both Zophobas and Tenebrio, whereas dark adaptation is completed about equally fast both day and night in both species. The speed of the pupil response, however, is dependent on the preceding adaptation history. 4. The pupil mechanism in both species is under the influence of a circadian rhythm, which determines the size of the pupil aperture in such a way that the pupil is maximally open when dark-adapted at night, but only partially open when dark-adapted at daytime.

In the superposition eyes of the sphingid moth Deilephila and the neuropteran Ascalaphus, adjustment to different intensities is subserved by longitudinal migrations of screening pigment in specialized pigment cells. Using ophthalmoscopic techniques we have localized the light-sensitive trigger that controls pigment position. In both species, local illumination of a small spot anywhere within the eye glow of a dark-adapted eye evokes local light adaptation in the ommatidia whose facets receive the light. Details of the response pattern demonstrate that a distal light-sensitive trigger is located axially in the ommatidium, just beneath the crystalline cone, and extends with less sensitivity deep into the clear zone. The distal trigger in Deilephila was shown to be predominantly UV sensitive, and a UV-absorbing structure, presumably the distal trigger, was observed near the proximal tip of the crystalline cone. In Ascalaphus we also found another trigger located more proximally, which causes local pigment reaction in the ommatidia whose rhabdoms are illuminated (the centre of the eye glow). The light-sensitive trigger for this response appears to be the rhabdom itself.

Compound eyes of the decapod shrimp Gennadas, the hermit crab Dardanus, and the syncarid crustacean Anaspides are studied histologically and with optical experiments. The results demonstrate that these three crustaceans all have refracting superposition eyes. The conclusion is based on the following observations: Euphausiids and mysids were previously thought to be the only crustaceans with refracting superposition eyes, whereas the species investigated here were assumed to have reflecting superposition eyes (decapod shrimps) or apposition eyes (hermit crabs and syncarids). The present findings increase more than twofold the number of crustacean groups that are known to have developed refracting superposition optics. It also provides insight into the evolutionary mechanisms that may have led to the development of this type of imaging optics.