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.

Cubozoans differ from other cnidarians by their body architecture and nervous system structure. In the medusa stage they possess the most advanced visual system within the phylum, located in sophisticated sensory structures, rhopalia. The rhopalium is a club-shaped structure with paired pit-shaped pigment cup eyes, paired slit-shaped pigment cup eyes, and two complex camera-type eyes: one small upper lens eye and one large lower lens eye. The medusa carries four rhopalia and visual processing and locomotor rhythm generation takes place in the rhopalia. We show here a bilaterally symmetric organization of neurons, with commissures connecting the two sides, in the rhopalium of the cubozoan Tripedalia cystophora. The fortuitous observation that a subset of neurons is strongly immunoreactive for a PCNA (proliferating cell nuclear antigen)-like epitope allowed us to analyze the organization of these neurons in detail. Distinct PCNA-immunoreactive (PCNA-ir) nuclei form six bilateral pairs that are associated with the slit eyes, pit eyes, upper lens eye, and the posterior wall of the rhopalium. Three commissures connect the clusters of the two sides and all clusters in the rhopalium have connections to the area around the base of the stalk. This neuronal system provides an anatomical substrate for integration of visual signals from the different eyes.

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.

The four visual sensory structures of a cubomedusa, the rhopalia, display a surprisingly elaborate organization by containing two lens eyes and four bilaterally paired pigment cup eyes. Peptides containing the peptide sequence Arg-Phe-NH2 (RFamide) occur in close association with visual structures of cnidarians, including the rhopalia and rhopalial stalk of cubomedusae, suggesting that RFamide functions as a neuronal marker for certain parts of the visual system of medusae. Using immunofluorescence we give a detailed description of the organization of the RFamide-immunoreactive (ir) nervous system in the rhopalia and rhopalial stalk of the cubomedusae Tripedalia cystophora and Carybdea marsupialis. The bilaterally symmetric RFamide-ir nervous system contains four cell groups and three morphologically different cell types. Neurites spread throughout the rhopalia and occur in close vicinity of the pigment cup eyes and the lower lens eye. Two commissures connect the two sides of the system and neurites of one rhopalial cell group extend into the rhopalial stalk. The RFamide-ir nervous system in the rhopalia of cubomedusae is more widespread and comprises more cells than earlier discerned. We suggest that the system might not only integrate visual input but also signals from other senses. One of the RFamide-ir cell groups is favorably situated to represent pacemaker neurons that set the swimming rhythm of the medusa.

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