Opposite patterns of diurnal activity in the box jellyfish Tripedalia cystophora and Copula sivickisi.

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.

Visually guided obstacle avoidance in the box jellyfish Tripedalia cystophora and Chiropsella bronzie.

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.

The lens eyes of the box jellyfish Tripedalia cystophora and Chiropsalmus sp. are slow and color-blind.

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.

The ring nerve of the box jellyfish Tripedalia cystophora.

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.

Bilaterally symmetrical rhopalial nervous system of the box jellyfish Tripedalia cystophora.

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.

Rhopalia are integrated parts of the central nervous system in box jellyfish.

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.

Photoreceptor evolution: ancient siblings serve different tasks.

Photoreceptor cells of vertebrate eyes are fundamentally different from those of invertebrate eyes. New work on the brain of a ragworm now suggests that ancestral bilaterians possessed both types of photoreceptor cell.

Visual pigments: trading noise for fast recovery.

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.

Visual discrimination: Seeing the third quality of light.

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.

Absorption of white light in photoreceptors.

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.

Placement and restoration of implants by predoctoral students: the Creighton experience.

Teaching of implant dentistry in the predoctoral dental curriculum has evolved dramatically over the past 20 years. In 1974, only one third of American dental schools addressed the topic of implants. Today, 48 of the 54 American dental schools have predoctoral curricula. The Creighton University experience offers some unique and instructive insights into a 10-year process of developing and implementing a predoctoral implant dentistry curriculum. All interested students may perform both the surgical placement and restoration of implant prostheses. Clinical instruction involves all restorative and surgical faculty members. Favorable 3-year (91%) and 5-year (87%) surgical success rates have been maintained. This article presents one university's program for examination and discussion.

Eye ancestry: old genes for new eyes.

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.

A pessimistic estimate of the time required for an eye to evolve.

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.

Late effects following central nervous system radiation in a pediatric population.

Between 1970 and 1986, 120 children with central nervous system malignancy were treated with radiation therapy. These included 44 low-grade astrocytomas, 11 high grade astrocytomas, 32 medulloblastomas, 15 ependymomas/ependymoblastomas, 3 primitive neuroectodermal tumors and 8 pineal tumors. Seven children were treated without biopsy. Fifty-one treated children were evaluated for the effects of therapy on growth, endocrine function, IQ and hair regrowth. Mean height was 1.5 standard deviations below the mean height for the patient's age at study (range 0-5.7). Height was significantly less in patients receiving radiation to the pituitary and those with somatomedin-C deficiency. Height was also decreased with whole CNS radiation and spine dose > 20 Gy but not to a significant degree. Pituitary radiation in any dose increased the chance of endocrine deficiency (p = 0.004) and 21 of 51 patients had somatomedin-C deficiency. Mean IQ was 92.7 (+/- 18.8), with a slight trend toward decreased IQ with increasing whole brain dose of radiation. Hair regrowth was complete in 20 of 46 evaluated patients, diminished regrowth occurring with increasing volume and dose of radiation. No difference in the measured late effects could be detected with respect to age at treatment, sex, histology or location of tumor.

From cornea to retinal image in invertebrate eyes.

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.

The ontogenetic development of refracting superposition eyes in crustaceans: Transformation of optical design.

We have investigated, comparatively, the ontogenetic development of the compound eye in larvae of a mysid (Neomysis) and a euphausiid (Thysanoessa) species and found it to be close to identical in the two species. The larval eye is of apposition type with special adaptations for planktonic life. The elongated dioptric apparatus is devoid of screening pigment and instead has a proximal lens optically isolating the ommatidium. The pigmented retina is extremely compressed making the eye largely transparent and presumably suitable for a planktonic life. The presence of this specialized type of eye in the planktonic larvae of euphausiids was known before but it is intriguing to find exactly the same type in mysids, spending their entire larval life as embryos in the female marsupium. A possible explanation is offered if mysids earlier in evolution had planktonic larvae. Upon reduction of free-living larvae, the transparent type of eye may have been preserved because there is no selection pressure on the larva to change it. In late larval life, both species transform their eyes to a refracting superposition type typical for adult mysids and euphausiids. The process of transformation and the functional connection between transparent apposition and superposition is described.

The compound eye of Leptodora kindtii (Cladocera). An adaptation to planktonic life.

Each of the approximately 500 ommatidia in the compound eye of the cladoceran crustacean Leptodora kindtii has a crystalline cone consisting of five cells. Five retinula cells are also present, one of which contributes to the distal 1-2 micrometer of the rhabdom only; the other four retinula cells from a continuous rhabdom. Throughout the rhabdom its cross section displays two separate halves with the axis of the microvilli in one half perpendicular to that in the other (orthogonal pattern). Interferometric analysis of the refractive index of the crystalline cone revealed an inhomogeneous system with one distal and one proximal gradient. The gradient system was found to exclude rays entering from adjacent facets, thus maintaining the optical isolation. Consequently, these optics replace distal screening pigment, which is absent in the eye. The long and unscreened crystalline cones give rise to an almost transparent eye in conformity with the overall transparency of this plank-tonic animal. The morphological characteristics of the eye of this species deviate from other cladoceran eyes, but the optical design closely resembles that of some pelagic marine amphipod crustaceans.