Non-visual opsins were discovered in the early 1990s. These genes play roles in circadian rhythm in mammals, seasonal reproduction in birds, light avoidance in amphibian larvae, and neural development in fish. However, the interpretation of such studies and the success of future work are compromised by the fact that non-visual opsin repertoires have not been properly characterized in any of these lineages. Here, we show that non-visual opsins from tetrapods and ray-finned fish are distributed among 18 monophyletic subfamilies. An amphibian sequence occurs in every subfamily, whereas mammalian orthologs occur in only seven. Species in the major ray-finned fish lineages, Holostei, Osteoglossomorpha, Otomorpha, Protacanthopterygii, and Neoteleostei, have large numbers of non-visual opsins (22-32 genes) as a result of gene duplication events including, but not limited to, the teleost genome duplication (TGD). In contrast to visual opsins, where lineage-specific duplication is common, the ray-finned fish non-visual opsin repertoire appears to have stabilized shortly after the TGD event and consequently even distantly related species have repertoires of similar size and composition. Most non-visual opsins have been named without the benefit of a phylogenetic perspective and, accordingly, major revisions are proposed.

Variable expression of visual pigment proteins (opsins) in cone photoreceptors of the vertebrate retina is a primary determinant of vision plasticity. Switches in opsin expression or variable co-expression of opsins within differentiated cones have been documented for a few rodents and fishes, but the extent of photoreceptor types affected and potential functional significance are largely unknown. Here, we show that both single and double cones in the retina of a flatfish, the starry flounder (Platichthys stellatus), undergo visual pigment changes through opsin switches or variable opsin co-expression. As the post-metamorphic juvenile (i.e., the young asymmetric flatfish with both eyes on one side of the body) grows from ~5 g to ~196 g, some single cones and one member of unequal double cones switched from a visual pigment with maximum wavelength of absorbance, λmax, at shorter wavelengths (437 nm and 527 nm) to one with longer λmax (456 nm and 545 nm, respectively) whereas other cones had intermediate visual pigments (λmax at 445 nm or 536 nm) suggesting co-expression of two opsins. The shift toward longer wavelength absorbing visual pigments was in line with maximizing sensitivity to the restricted light spectrum at greater depths and achromatic detection of overhead targets.

Vertebrate color vision relies on the differential expression of visual pigment proteins (opsins) in cone photoreceptors of the retina. The diversity of opsins and their retinal expression patterns appear greatest for animals that experience variable light habitats, as is the case with flatfishes. Yet, opsin repertoires and expression patterns in this group of fishes are poorly described. Here, we unveil the visual opsin expression patterns of juvenile starry flounder (Platichthys stellatus) and describe the localization of cone types, their visual pigments and opsin expression. Juvenile starry flounder express eight opsins (Rh1, Sws1, Sws2A1, Sws2A2, Sws2B, Rh2A1, Rh2A2, Lws) and possess a corresponding number of photoreceptor visual pigments, with peak absorbance ranging from 369 to 557 nm. Retinal (vitamin A1) was the only chromophore detected in the retina. Intraretinal variation in opsin abundance consisted of greater expression of both RH2, and lesser expression of SWS1 and both SWS2A, opsin transcripts in the dorsal compared to the ventral retina. Overall cone density was greater in the dorsal retina which was also characterized by a larger proportion of unequal double cones compared with the ventral retina. Together, our results suggest that large opsin repertoires serve to optimize visual function under variable light environments by differential expression of opsin subsets with retinal location.

tom iwanicki 2 , Jamie Sickles 3 & tamara M. frank 3 extraocular photoreception, the ability to detect and respond to light outside of the eye, has not been previously described in deep-sea invertebrates. Here, we investigate photosensitivity in the bioluminescent light organs (photophores) of deep-sea shrimp, an autogenic system in which the organism possesses the substrates and enzymes to produce light. through the integration of transcriptomics, in situ hybridization and immunohistochemistry we find evidence for the expression of opsins and phototransduction genes known to play a role in light detection in most animals. Subsequent shipboard light exposure experiments showed ultrastructural changes in the photophore similar to those seen in crustacean eyes, providing further evidence that photophores are light sensitive. in many deep-sea species, it has long been documented that photophores emit light to aid in counterillumination-a dynamic form of camouflage that requires adjusting the organ's light intensity to "hide" their silhouettes from predators below. However, it remains a mystery how animals fine-tune their photophore luminescence to match the intensity of downwelling light. Photophore photosensitivity allows us to reconsider the organ's role in counterillumination-not only in light emission but also light detection and regulation. Photoreceptor cells inside the complex eyes of animals are responsible for light detection and subsequent signaling cascades linked to vision. Though light detection in animals is typically associated with ocular photoreceptors, the ability to detect and respond to light can also occur in extraocular tissues and structures 1. Extraocular photoreception has been documented across a range of structures and taxa, including the dermal chromatophores of cephalopods and fish, tube feet of echinoderms, pineal organs in fish and the central nervous systems of arthropods 2-6. Despite the occurrence across diverse metazoans, knowledge regarding the functionality of extraocular photoreceptors remains limited. Bioluminescent light organs, called photophores, provide a unique opportunity to study extraocular photosensitivity, as evidence suggests these structures not only emit light but can also detect it 7. Photophores are complex organs composed of bioluminescent cells (photocytes), and sometimes pigments, reflectors, and filtering structures 8. They can be divided into two types: bacterial or autogenic, where the light is produced by either symbiotic bacteria living within the structure or by the animal itself. In some species, photophores assist in a form of camouflage known as counterillumination. During this process, photophore emissions mimic the downwelling light blocked by the animal's body, thereby camouflaging the animal's profile that would otherwise be detectable to predators below 9,10. Deep-sea shrimp of the family Oplophoridae possess autogenic photophores in three (Systellaspis, Oplophorus and Janicella) of the ten genera 11. Within these three genera, species vertically...

Knowledge of crustacean vision is lacking compared to the more well-studied vertebrates and insects. While crustacean visual systems are typically conserved morphologically, the molecular components (i.e. opsins) remain understudied. This review aims to characterize opsin diversity across crustacean lineages for an integrated view of visual system evolution. Using publicly available data from 95 species, we identified opsin sequences and classified them by clade. Our analysis produced 485 putative visual opsins and 141 non-visual opsins. The visual opsins were separated into six clades: long wavelength sensitive (LWS), middle wavelength sensitive (MWS) 1 and 2, short wavelength or ultraviolet sensitive (SWS/UVS) and a clade of thecostracan opsins, with multiple LWS and MWS opsin copies observed. The SWS/UVS opsins were relatively conserved in most species. The crustacean classes Cephalocarida, Remipedia and Hexanauplia exhibited reduced visual opsin diversity compared to others, with the malacostracan decapods having the highest opsin diversity. Non-visual opsins were identified from all investigated classes except Cephalocarida. Additionally, a novel clade of non-visual crustacean-specific, R-type opsins (Rc) was discovered. This review aims to provide a framework for future research on crustacean vision, with an emphasis on the need for more work in spectral characterization and molecular analysis. This article is part of the theme issue ‘Understanding colour vision: molecular, physiological, neuronal and behavioural studies in arthropods’.

The hypothesis that size-selective mortality in the first marine year is a major regulator of recruitment in Pacific salmon Oncorhynchus spp. has led to interest in assessing the recent growth of field-caught fish. Understanding differences in relative growth across years, regions, habitats, and prey fields may provide insights into factors influencing survival. Plasma insulin-like growth factor 1 (IGF1), muscle RNA : DNA ratio (RD), and scale circulus spacing have all been used as indices of recent growth in juvenile salmonids. We concurrently assessed these growth rate indices in a laboratory study of postsmolt, young-of-the-year, ocean-type Chinook Salmon O. tshawytscha. We synthesized results with previous work to inform selection of appropriate growth rate indices for field studies on juvenile salmonids. Muscle samples suitable for RD analysis were obtained nonlethally and without subsequent growth impacts, even for very small juvenile salmon (75-99 mm FL). Plasma IGF1 concentration was strongly correlated with growth rate (R 2 = 0.79), while log e (RD) and mean spacing of the outer two circuli were moderately correlated with growth rate Subject editor: Milo Adkison, University of Alaska-Fairbanks, Juneau

Correlations between variation in opsin expression and variation in vision are often assumed but rarely tested. We exposed starry flounder ( Platichthys stellatus ) to either broad spectrum sunlight or green-filtered light in outdoor aquaria for seven weeks and then combined digital-PCR and camouflage experiments to test two hypotheses: i) short-wavelength sensitive opsin expression decreases in a green light environment, and ii) if observed, this change in opsin expression influences colour vision as estimated using a camouflage-based behavioural assay. Of the eight visual opsins measured, Sws1 (UV sensitive) and Sws2B (blue sensitive) expression was significantly lower in fish exposed to green light. However, opsin expression in fish transferred to an arena illuminated with white LED light for three hours after the green light treatment did not differ from broad spectrum controls. Changes in opsin expression in response to artificial light environments have been reported before, but rapid changes over three hours rather than days or weeks is unprecedented. We did not observe a significant difference in a flounder’s camouflage response based on light environment, although broad spectrum fish increased and green-filter fish decreased the pattern contrast when on the blue-green substrate, and this difference approached significance. This pattern is intriguing considering green-filter fish expressed fewer UV and blue opsins and we recommend increased statistical power for future experiments. Together, our results show that starry flounder opsin expression changes rapidly in response to changes in light environment, however, there is no apparent effect on their visually mediated camouflage.

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