Formation of Chaotic Patterns of the Gastrovascular System in the Ontogenesis of the Medusa Aurelia aurita

Only the first branching of the perradial and interradial channels of the gastrovascular system of Aurelia aurita (L.) (Scyphozoa: Ulmaridae) in the metaephyra stage is always uniform. Further branch formation is variable and random. New branches originate most often at the annular channel, but some of them grow from the radial channels. The early formation of irregular and chaotic patterns of the radial channel branching is caused by the asynchrony and topographic variability of the generation of new branches of different channels. During further morphogenesis, the spatial and temporal variability of branching increases, which leads to an increased state of chaos in the morphology of the branching channels of each individual. Lack of rigid determinancy of the gastrovascular system channel branching and plasticity of this system in the course of the entire ontogenesis provide possible evidence of its adaptive responses, for example, to the disruption of tetraradial symmetry or rearrangements after damage.     

Molecular characterization of aspartylglucosaminidase,a lysosomal hydrolase upregulated during strobilation in the moon jellyfish,Aurelia aurita

The life cycle of the moon jellyfish, Aurelia aurita, alternates between a benthic asexual polyp stage and a planktonic sexual medusa (jellyfish) stage. Transition from polyp to medusa is called strobilation. To investigate the molecular mechanisms of strobilation, we screened for genes that are upregulated during strobilation using the differential display method and we identified aspartylglucosaminidase (AGA), which encodes a lysosomal hydrolase. Similar to AGAs from other species, Aurelia AGA possessed an N-terminal signal peptide and potential N-glycosylation sites. The genomic region of Aurelia AGA was approximately 9.8 kb in length and contained 12 exons and 11 introns. Quantitative RT-PCR analysis revealed that AGA expression increased during strobilation, and was then decreased in medusae. To inhibit AGA function, we administered the lysosomal acidification inhibitors, chloroquine or bafilomycin A1, to animals during strobilation. Both inhibitors disturbed medusa morphogenesis at the oral end, suggesting involvement of lysosomal hydrolases in strobilation.     

Mesogleal cells of the jellyfish Aurelia aurita are involved in the formation of mesogleal fibres

The extracellular matrix of the jellyfish Aurelia aurita (Scyphozoa, Cnidaria), known as the mesoglea, is populated by numerous mesogleal cells (Mc). We determined the pattern of the Mc and the mesoglea, raised polyclonal antibodies (RA47) against the major mesogleal protein pA47 (47 kDa) and checked their specificity. In the mesoglea, RA47 stains pA47 itself. In immunoblots of Mc, RA47 stains bands of 120 kDa and 80 kDa; weaker staining is observed at pA47. The same staining pattern is seen on blots of jellyfish epidermal cells and of whole Hydra (Hydrozoa) or isolated mesoglea of Hydra. Our data indicate that pA47 is synthesized by Mc and epidermal cells as high molecular precursors. Using immunostaining techniques, we showed Mc to be involved in the formation of mesogleal non-collagenous (called "elastic" in classic morphological studies) fibres. The biochemical and morphological data suggest that Mc originate from the epidermis.     

On natural metamorphosis inducers of the cnidarians Hydractinia echinata (Hydrozoa) and Aurelia aurita (Scyphozoa)

 Hydractinia echinata and Aurelia aurita produce motile larvae which undergo metamorphosis to sessile polyps when induced by external cues. The polyps are found at restricted sites, A. aurita predominantly on rocks close to the shore, H. echinata on shells inhabited by hermit crabs. It has been argued that the differential distribution of the polyps in their natural environment largely reflects the distribution of the natural metamorphosis-inducing cues. In the case of H. echinata, bacteria of the genus Alteromonas were argued to meet these conditions. We found that almost all substrates collected in the littoral to induce metamorphosis in H. echinata, and several bacterial strains isolated from the sea, including the common E. coli, induce metamorphosis efficiently. In A. aurita metamorphosis may be induced by the water–air interface, whereby metamorphosis precedes (final) settlement. Received: 7 December 1998 / Accepted: 8 July 1999     

Size-based Prey Selectivity and Dietary Shifts in the Jellyfish, Aurelia aurita

Gut content analyses on field-caught Aurelia aurita showed bothquantitative and qualitative change in diet as a function ofmedusa size. Larger medusae tended towards greater numbersand diversity of prey (up to 1550 individual prey representingas many as 13 different prey groups). We also found that medusasize was a good predictor of prey diversity recovered from themedusa gut. While a shift toward greater prey diversity inlarger medusae might be explained by increased contact rateswith 'rare' prey taxa, we also found size-based prey selectivitychanges in A. aurita. We used in situ gut content data to describeselectivity by A. aurita for three prey types representing varyingdegrees of swimming or escape velocity. Fish eggs were usedas a non-swiming prey, and small (     

Aurelin, a novel antimicrobial peptide from jellyfish Aurelia aurita with structural features of defensins and channel-blocking toxins

A novel 40-residue antimicrobial peptide, aurelin, exhibiting activity against Gram-positive and Gram-negative bacteria, was purified from the mesoglea of a scyphoid jellyfish Aurelia aurita by preparative gel electrophoresis and RP-HPLC. Molecular mass (4296.95 Da) and complete amino acid sequence of aurelin (AACSDRAHGHICESFKSFCKDSGRNGVKLRANCKKTCGLC) were determined. Aurelin has six cysteines forming three disulfide bonds. The total RNA was isolated from the jellyfish mesoglea, RT-PCR and cloning were performed, and cDNA was sequenced. A 84-residue preproaurelin contains a putative signal peptide (22 amino acids) and a propiece of the same size (22 amino acids). Aurelin has no structural homology with any previously identified antimicrobial peptides but reveals partial similarity both with defensins and K+ channel-blocking toxins of sea anemones and belongs to ShKT domain family.     

Induction of metamorphosis from the larval to the polyp stage is similar in Hydrozoa and a subgroup of Scyphozoa (Cnidaria, Semaeostomeae)

Larvae of cnidarians need an external cue for metamorphosis to start. The larvae of various hydrozoa, in particular of Hydractinia echinata, respond to Cs⁺, Li⁺, NH₄ ⁺ and seawater in which the concentration of Mg²⁺ ions is reduced. They further respond to the phorbolester, tetradecanoyl-phorbol-13-acetate (TPA) and the diacylglycerol (DAG) diC8, which both are argued to stimulate a protein kinase C. The only well-studied scyphozoa, Cassiopea spp., respond differently, i.e. to TPA and diC8 only. We found that larvae of the scyphozoa Aurelia aurita, Chrysaora hysoscella and Cyanea lamarckii respond to all the compounds mentioned. Trigonelline (N-methylnicotinic acid), a metamorphosis inhibitor found in Hydractinia larvae, is assumed to act by delivering a methyl group for transmethylation processes antagonising metamorphosis induction in Chrysaora hysoscella and Cyanea lamarckii. The three species tested are scyphozoa belonging to the subgroup of semaeostomeae, while Cassiopea spp. belong to the rhizostomeae. The results obtained may contribute to the discussion concerning the evolution of cnidarians and may help to clarify whether the way metamorphosis can be induced in rhizostomeae as a whole is different from that in hydrozoa and those scyphozoa belonging to the subgroup semaeostomeae. Electronic Publication     

In situ estimation of ephyrae liberated from polyps of Aurelia aurita using settling plates in Tokyo Bay, Japan

The continuous changes in the number of newly established polyps of Aurelia aurita (L.) on settling plates under natural conditions were observed from August 1998 to September 1999 in Tokyo Bay, Japan. A sharp decline in survivorship of newly settled polyps was observed within the first few days, however, survivorship of polyps settled in October increased by budding up to 399% after two months. The number of discs in each strobila varied from 1 to 6, however, most of the strobilae formed single discs. The percentage ratios of the total number of ephyrae to the initial number of polyps on settling plates were generally lower than 10%, but the highest ratio of 594.4% was estimated for the polyps settled in October. It is considered that most of the liberated ephyrae originate from the polyps settled in October in Tokyo Bay. This study suggests that the occurrence of ripe medusae with planula larvae throughout the year contributes to the success of settlement and growth of the polyp stage in Tokyo Bay.     

Plate in the zone of oocyte and germinal epithelium contact in Scyphomedusa Aurelia aurita binds antibodies to ZP-domain protein mesoglein

The morphological features of oocyte and germinal epithelium (epithelial wall of germinal sinus) contact area in Scyphomedusa Aurelia aurita are described. Growing oocytes were divided into seven stages based on oocyte size. The structure revealed the area of contact between the oocyte and the germinal epithelium called the contact plate. During oocyte growth, single granules are fused into the homogeneous mass area of oocyte contact with epithelium. Plate components bound antibodies to mesoglein. It was assumed that the plate material contained ZP-domain proteins. Electrophoresis and immunoblotting results show that proteins immunologically similar to mesoglein have higher molecular masses, probably due to post-translation modifications, which are common for extracellular proteins. On the other hand, however, gonad proteins may be other representatives of jellyfish ZP-domain proteins. Further experiments should be conducted to clarify which alternative is true.     

Behavioural response of the scyphozoan jellyfish Aurelia aurita (L.) upon contact with the predatory jellyfish Cyanea capillata (L.)

Underwater manipulative experiments were carried out in situ to investigate the sensibility of the jellyfish Amelia aurita (L.) to contact with the tentacles of Cyanea capillata (L), commonly known as a predator on A. aurita. Movements of individual medusae touched by tentacles of C. capillata and other objects were video‐recorded during SCUBA dives. The behavioural variable studied was change in swim pulse frequency. The results showed that A. aurita was highly susceptible to the tentacles of C. capillata and responded with an increased swim pulse frequency when touched at the umbrellar margin but not at the central exumbrella. Contact with other objects also induced a behavioural response in A. aurita.     

Genetic characterization of the scyphozoan jellyfish Aurelia spp. in Chinese coastal waters using mitochondrial markers

Blooms of the moon jellyfish Aurelia spp. have occurred in the harbors and coastal waters around the world. The phylogenetic relationship and genetic characterization of Aurelia spp. was determined along the Chinese coastal waters based on sequences of the mitochondrial cytochrome c oxidase subunit I (COI) gene. The molecular analysis confirmed that all samples collected in Chinese coastal waters were Aurelia sp.1. We also analyzed the phylogenetic and population genetic structure of Aurelia sp.1 using the newly generated sequences supplemented with existing data from previous studies. The phylogenetic analyses of the COI regions did not support geographically restricted groups among the global samples of Aurelia sp.1. Analysis of molecular variance (AMOVA) indicated a complex genetic population structure and pattern of connectivity. Populations of Aurelia sp.1 were highly structured between most sampling sites over distances as small as 100 km (Rizhao and Qingdao) in certain cases. However, non-significant pairwise F_ST values were also observed between short geographic distances (Yantai, Rongcheng and Qingdao) and relatively distant sampling sites (Caofeidian, Rizhao and Japan). The life-cycle characteristics, together with the prevailing ocean currents in this region and possible anthropogenic introduction, were proposed and discussed as the main factors that determined the genetic patterns of Aurelia sp.1.     

Food and feeding of Aurelia aurita in Tokyo Bay with an analysis of stomach contents and a measurement of digestion times

In situ seasonal variations in stomach contents of Aurelia aurita (L.) in Tokyo Bay, Japan, were analyzed. Copepods, such as Oithona davisae Ferrari & Orsi were the predominant food items of A. aurita from June to November. The mean digestion time measured in incubation experiments was 0.95 h. Daily rations calculated using stomach content data and digestion times were 2.2–21.8 mg C ind^–1 corresponding to 0.58–5.56% of body carbon. The ingestion rate increased significantly with an increase in medusa size, although no significant relationship was found between medusa size and carbon specific daily ration. The zooplankton community in Tokyo Bay is characterized by the significant dominance of O. davisae and it is assumed that the prosperity of A. aurita is caused by the high abundance of the O. davisae population. It is suggested that a food chain comprised of microflagellates, cyclopoid copepods O. davisae, and A. aurita is the most significant one in Tokyo Bay and only a small portion of production is transferred to fish.     

Mass mortality of Sardinella aurita Valenciennes (Pisces, Clupeidae) in Thessaloniki Bay (Macedonia, Greece)

Mass mortality of Sardinella aurita was observed in Thessaloniki Bay during an abrupt temperature fall in February 1991. The mass of dead fish was estimated to be about 79.4 tonnes.     

Bioluminescence of Aequorea macrodactyla, a Common Jellyfish Species in the East China Sea

Studies of the bioluminescent mechanisms of jellyfish have been mainly confined to one species, Aequorea victoria. We describe the luminescent system of another species, Aequorea macrodactyla, which is commonly found in the warmer waters on the coastal region of East China Sea. The luminescent system of this species consists of a green fluorescent protein (GFP) and one or more aequorins. The GFP gene is 1042 bp. It encompasses a coding sequence of 717 bp organized as 3 exons, and it is predicted to specify a 27-kDa peptide, which shares 80% amino acid sequence identity with the GFP of A. victoria. The entire coding sequence was cloned into the pTO-T7 expression vector and expressed in Escherichia coli. Compared with GFP of A. victoria, the purified expressed protein exhibited an excitation peak at a higher wavelength of 476 nm and an emission peak at a lower wavelength of 496 nm, with a higher quantum yield of 1.0. The other photoprotein, aequorin, is encoded in a single open reading frame of 585 bp specifying a 23-kDa apoprotein. The gene was cloned in to the same expression vector and expressed in E. coli. The activity of the photoprotein was reconstituted by incubating the expressed apoprotein with coelenterazine f. In the presence of Ca²⁺ the reconstituted aequorin exhibits an emission peak at 470 nm. The kinetics of regeneration and the photoactivities of the reconstituted aequorins of the 2 species of jellyfish are similar. Nevertheless, Aequorea macrodactyla is expected to appear brighter and more ``blue' than Aequorea victorea because of the differences in the photoactivity of their GFPs.     

Massive leaping of Sardinella aurita valenciennes, 1847 under probable risk of predation in Northern Aegean sea (Greece)

On the morning of 15th December 2002, several hundred live round sardinella (Sardinella aurita) individuals leaped out of the water and became stranded on the breakwater of Nea Iraklitsa port (Kavala Gulf, northern Aegean Sea, Greece). The phenomenon was most likely caused by the attack of the cetacean predator Tursiops truncatus (bottlenose dolphin), which is abundant in the area and feeds upon pelagic fish. The ability of round sardinella to leap out of the water has never been reported for other pelagic fishes.     

A method for in situ estimation of prey selectivity and predation rate in large plankton, exemplified with the jellyfish Aurelia aurita (L.)

A method to estimate predation rates of large predatory zooplankton, such as jellyfish and ctenophores, is outlined. Large plankton size allows direct visual tracking of the predator during the process of foraging. The presented method is novel in the sense that it measures predation rate of a specific individual plankton predator in situ.After prey has been evacuated from the gut of an individual predator, the predator is incubated in situ, and observed by SCUBA-divers who recapture the individual after a defined time. Given that this incubation time is shorter than prey digestion time, predation rate can be calculated as increase in gut content over time. Clearance rates for different prey can be calculated from predation rates and prey concentrations in the water, allowing accurate estimates of prey selectivity. Thus, the problem of unknown feeding history and feeding environment, which can otherwise be a problem in prey selectivity studies of in situ-captured predators, is circumvented. Benefits and limitations of the method are discussed.The method was applied to adult medusae of the common jellyfish Aurelia aurita. A large variation in number of captured prey was detected both among individual jellyfish and among the various oral arms and gastric pouches within individuals. Clearance rates varied strongly with prey type. The medusae selected large crustacean prey (cladocerans and copepods/copepodites) over echinoderm larvae and copepod nauplii. Prey distribution within the medusae indicates that both tentacles and oral arms were used as prey capturing sites. Food passage time from prey capturing organs to gastric pouches was estimated.     

Sea warming and fish distribution: the case of the small pelagic fish, Sardinella aurita, in the western Mediterranean

This study analyses the temporal and spatial changes in abundance and distribution of the warm water species round sardinella (Sardinella aurita) in the western Mediterranean over the last decades in relation to sea water temperature. In the western Mediterranean basin (1950–2003), a significant positive relationship was found between round sardinella landings and temperature anomalies. Along a latitudinal gradient off the Mediterranean Iberian coast (1989–2004), a gradual increase in species abundance was observed from south to north, with a certain time lag going northwards, associated with the increase in sea water temperature. The abundance of round sardinella in the two warmest and southernmost areas was positively and significantly correlated with sea surface temperature registered during the start of gonad maturation the previous year. In addition, the positive relationship established between water temperature and abundance of round sardinella in the coldest and northernmost study area demonstrates that there is a temperature limit for the distribution of this species in the western Mediterranean. In addition, this study analyses round sardinella larvae distribution and abundance in the summers of 2003 and 2004, and conducts a comparison with the situation 20 years ago (summer 1983). Results show a marked increase in larval abundance during the last decades and the present appearance of larvae in the northernmost study areas, where they did not occur 20 years ago. This indicates the successful reproduction of round sardinella in the northern part of the Mediterranean, where the species has expanded, confirming its establishment in the area.     

Organization of the subumbrellar musculature in the ephyra,juvenile, and adult stages of Aurelia aurita Medusae

We used fluorescently labeled phalloidin to examine the subumbrellar musculature of the scyphozoan jellyfish Aurelia aurita in a developmental series from ephyra to adult medusa. In the ephyra, the swim musculature includes a disc‐like sheet of circular muscle, in addition to two radial bands of muscle in each of the eight ephyral arms. The radial muscle bands join with the circular muscle, and both circular and radial muscle act together during each swim contraction. As the ephyra grows into a juvenile medusa, arms tissue is resorbed as the bell tissue grows outward, so eventually, the ephyral arms disappear. During this process, the circular muscle disc also grows outward and the radial muscle bands of the arms also disappear. At this time, a marginal gap appears at the bell margin, which is devoid of circular muscle cells, but has a loose arrangement of radial muscle fibers. This marginal gap is preserved as the medusa grows, and contributes to the floppy nature of the bell margin. Radial distortions in the circular muscle layer involve muscle fibers that run in random directions, with a primarily radial orientation. These are believed to be remnants of the radial muscle of the ephyral arms, and the distortions decrease in number and extent as the medusa grows. Since the mechanics of swimming changes from drag‐based paddling in the ephyra to marginal rowing in the adult medusa, the development of the marginal gap and the presence of radial distortions should be considered in terms of this mechanical transition.     

The ontogeny of swimming behavior in the scyphozoan, Aurelia aurita. I. Electrophysiological analysis.

1. Electrical correlates of behavioral activity were observed in the lip and tentacles of the polyp, but none were detected during column contraction. The tentacles are the most electrically active tissue, and the potentials are conducted along the length of the tentacle, but conduction to other parts of the animal were not observed. 2. Although the tentacles of the polyp and the rhopalia of the medusa are probably homologous, the development of pacemaker activity during strobilation is not a smooth transition from tentacle contraction potentials (TCPs) to marginal ganglion potentials (MGPs). This result indicates that each pacemaker activity develops de novo. 3. Two types of behavior were observed in the polyp: local responses, and coordinated activity which involved integrated responses in several body parts. The coordinated responses indicate that neurological coordination can take place in the polyp. Furthermore, feeding and spasm in the ephyra are similar to feeding and the protective response in the polyp. This similarity suggests that both coordinated responses in the polyp are coordinated by interneural facilitation in the diffuse nerve net (DNN) as in the ephyra. 4. Swimming in the ephyra is a medusoid behavior but feeding and spasm are coordinated by the DNN and are polypoid responses. Therefore, the ephyra is a mixture of polypoid and medusoid behaviors. As the ephyra matures into an adult medusa both polypoid responses are lost, but the DNN remains to modulate pacemaker output and control marginal tentacle contractions. As development proceeds from polyp, to ephyra, to medusa, each subsequent stage acquires some new behavior while retaining some aspect from the previous stage.