The retinal topography of three species of coleoid cephalopod (one cuttlefish, one squid and one octopus) was investigated to examine and compare the structure, density and organization of the photoreceptors. The aim was to determine if there were areas of increased cell density and/or cell specialization that might be related to lifestyle or phylogeny. The orientation of photoreceptors around the curved surface of the retina was also mapped to reveal how the overall arrangement of cell microvilli might enable the perception of polarized light stimuli. It was found that all species possessed an increase in photoreceptor density in a horizontal streak approximately placed at the position of a potential horizon in the habitat. The overall arrangement of photoreceptor microvillar arrangements followed lines of latitude and longitude in a global projection that has been rotated by 908. This arrangement seems to map polarization sensitivities on the outside world in a vertical and horizontal grid. The potential significance of this and other retinal specializations is discussed in the context of phylogenetic and habitat differences between species.

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SUMMARYThe existence of polarization sensitivity (PS), most likely resulting from the orthogonal arrangement of microvilli in photoreceptors, has been proposed in cephalopods for some time, although it has rarely been examined behaviourally. Here, we tested the mourning cuttlefish, Sepia plangon, and the reaper cuttlefish, Sepia mestus, for polarization sensitivity using a largefield optomotor stimulus containing polarization contrast. Polaroid filter drums with stripes producing alternating e-vectors were rotated around free-moving animals. Polarized optomotor responses were displayed, and these responses were similar to those performed in response to a black-and-white, vertically-striped drum, whereas no responses were displayed to a plain polarizing control drum producing just a vertical e-vector. This indicates that the animals are able to see the contrast between adjacent stripes in the polarizing drum. To our knowledge, this is the first demonstration of functional polarization sensitivity in cuttlefish.

SUMMARY Coleoid cephalopods (octopus, cuttlefish and squid) potentially possess polarization sensitivity (PS) based on photoreceptor structure, but this idea has rarely been tested behaviourally. Here, we use a polarized, striped optokinetic stimulus to demonstrate PS in the striped pyjama squid, Sepioloidea lineolata. This species displayed strong, consistent optokinetic nystagmic eye movements in response to a drum with stripes producing e-vectors set to 0 deg, 45 deg, 90 deg and 135 deg that would only be visible to an animal with PS. This is the first behavioural demonstration of a polarized optokinetic response in any species of cephalopod. This species, which typically sits beneath the substrate surface looking upwards for potential predators and prey, possesses a dorsally shifted horizontal pupil slit. Accordingly, it was found to possess a horizontal strip of high-density photoreceptors shifted ventrally in the retina, suggesting modifications such as a change in sensitivity or resolution to the dorsal visual field.

The cornea is the first optical element in the path of light entering the eye, playing a role in image formation and protection. Corneas of vertebrate simple camera-type eyes possess microprojections on the outer surface in the form of microridges, microvilli, and microplicae. Corneas of invertebrates, which have simple or compound eyes, or both, may be featureless or may possess microprojections in the form of nipples. It was previously unknown whether cephalopods (invertebrates with camera-type eyes like vertebrates) possess corneal microprojections and, if so, of what form. Using scanning electron microscopy, we examined corneas of a range of cephalopods and discovered nipple-like microprojections in all species. In some species, nipples were like those described on arthropod compound eyes, with a regular hexagonal arrangement and sizes ranging from 75 to 103 nm in diameter. In others, nipples were nodule shaped and irregularly distributed. Although terrestrial invertebrate nipples create an antireflective surface that may play a role in camouflage, no such optical function can be assigned to cephalopod nipples due to refractive index similarities of corneas and water. Their function may be to increase surface-area-to-volume ratio of corneal epithelial cells to increase nutrient, gas, and metabolite exchange, and/or stabilize the corneal mucous layer, as proposed for corneal microprojections of vertebrates.