Telomerase activity is not related to life history stage in the jellyfish Cassiopea sp.

The polyp (scyphistoma) of the jellyfish Cassiopea sp. can be maintained in culture for a long time, as polyps repeatedly reproduce asexually via formation of vegetative buds or propagules. The medusa, which is the sexually reproducing stage, typically has a relatively short life span. As a first step to understand the difference in life spans of the polyp and medusa stages of Cassiopea sp., we measured telomerase activity in different life cycle stages. We found telomerase activity in tissues of aposymbiotic polyps and propagules and symbiotic ephyrae (newly budded medusae) and adult medusae. No significant difference in telomerase activity was found between polyps and the bell region of the medusae. The cloned elongation products of the stretch PCR contained the TTAGGG repeats suggesting that the jellyfish has the ‘vertebrate’ telomere motif (TTAGGG)_n. This is the first study to show that somatic tissues of both polyp and medusa stages of a cnidarian had telomerase activity. Telomerase activity in somatic tissues may be related to the presence of multipotent interstitial cells and high regenerative capacity of cnidarians.     

Field observations of four Aurelia labiata jellyfish behaviours: swimming down in response to low salinity pre-empted swimming up in response to touch,but animal and plant materials were captured equally

Jellyfish live in complex environments and must continually make behavioural choices. In field observations, adult Aurelia labiata were confronted with a conflict between swimming up elicited by touch of the manubrium and swimming down elicited by low salinity. Following a touch, downward-swimming medusae (1.5–2.0 m deep) turned and swam to within 0.5 m of the surface when the salinity in the top 1.5 m of the water column was greater than 20 ppt but medusae uniformly refused to swim up into the top 1.25 m when the salinity was less than 20 ppt even after being touched three times. The central nervous system of A. labiata appears to have neural circuitry that specifies their response when medusae encounter stimuli that elicit incompatible behaviours. Upward-swimming adult medusae had animal, vegetable or cellulose (paper) material dispersed ahead of them. Medusae captured each material on the bell margin and transported it to a gastric pouch. Medusae displayed only minor behavioural differences in the process. Having sensory, neural and muscular systems organized to capture and pass to the stomach, a huge variety of materials allows medusae to survive in different seasons and environments.     

Induction of segmentation in polyps of Aurelia aurita (Scyphozoa, Cnidaria) into medusae and formation of mirror-image medusa anlagen

Polyps of Aurelia aurita can transform into several medusae (jellyfish) in a process of sequential subdivision. During this transformation, two processes take place which are well known to play a key role in the formation of various higher metazoa: segmentation and metamorphosis. In order to compare these processes in bilaterians and cnidarians we studied the control and the kinetics of these processes in Aurelia aurita. Segmentation and metamorphosis visibly start at the polyp's head and proceed down the body column but do not reach the basal disc. The small piece of polyp which remains will develop into a new polyp. The commitment to the medusa stage moves down the body column and precedes the visible onset of segmentation by about one day. Segmentation and metamorphosis can start at the cut surface of transversely cut body columns, leading to a mirror-image pattern of sequentially developing medusae.     

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.     

Synchronized release of medusae from three species of hydrozoan fire corals

 The release of medusae from three hydrozoan fire corals, Millepora dichotoma, M. murrayi and M. platyphylla, was investigated at three sites in southern Taiwan from February 1994 to July 1995. All three species were gonochoristic, and developed and released several batches of medusae between April and May. The duration of open ampulla appearing on the surface of coralla was short, about three months, and could be used to infer the reproductive season of the fire corals between April and May. No obvious lunar cycles of medusa release were found in these species. Medusa release started before dark at approximately 17:00 h and continued for several hours. Males began releasing medusae earlier than females. Synchronization of medusa release between colonies, i.e., the probability of occurring on the same nights, was significantly higher within a species than between different species. Hybridization in nature among the three species is, therefore, unlikely due to segregation in the spawning dates. Moreover, the synchronization within each species was often significantly higher within versus between sites. The free-swimming medusae released gametes within approximately one hour, and the spent medusae lived for a few more hours. Medusae may facilitate fertilization rates as a result of an apparently negatively geotactic swimming response that results in medusa aggregation at the surface. No differences in the sizes of medusae, eggs and sperm were detected among the three species; however, some characteristic differences of medusa nematocysts were found. Accepted: 25 September 1997     

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.     

Protein composition of mesoglea and mesogloeal cells of medusa Aurelia aurita

Protein composition of mesoglea of the scyphomedusa Aurelia aurita was revealed in SDS-PAGE. Some major bands are visible in mesoglea of a mature medusa: 30, 45-47, 85 kDa, three bands between 100-200 kDa, and several bands with molecular weights > 300 kDa. Polyclonal antisera RA45/47 against protein 45 kDa were raised. RA45/47 react with 45-47 kDa protein in mesogleal sample and protein 120 kDa in mesogleal cells on immunoblot. Immunohistochemical analysis of A. aurita histological sections of young and mature medusae showed antigen localization in mesogleal cell granules and in the apical part of ectodermal cells. In mature medusae, the antigen was localized also in elastic fibers. We can conclude that in A. aurita mesogleal cells, along with ectodermal cells, take part in the formation of extracellular matrix of mesoglea.     

Neurobiology of Stomotoca. II. Pacemakers and conduction pathways.

Evidence is presented for separate conduction pathways for swimming and for tentacle coordination in the marginal nerves of the jellyfish Stomotoca. The effector muscles are fired through junctions sensitive to excess Mg++, probably represented by the neuromuscular synapses observed by electron microscopy. The swimming effector (striated muscle) fires one-to-one with nerve input signals and myoid conduction occurs. Tentacle responses (smooth muscle contractions) involve facilitation, presumably at the neuro-effector junction; responses are graded and nonpropagating. Electrical correlates of two further conducting systems using the marginal nerves have been recorded. Their functions are unknown. One, the bridge system, extends up the four radii and encircles the peduncle; the other (ring system) is confined to the margin. A fifth conducting system is inferred in the case of the pointing response and its distribution is plotted. Signals have not been obtained from it. Pointing is accompanied by a burst of muscle potentials in the radial smooth muscles and is exhibited after a lengthy latency, indicating a local pacemaker. A sixth conducting pathway is the epithelial system, which mediates crumpling, a response involving the radial muscles without pacemaker intervention. Characteristic conduction velocities and wave forms are noted for the first four systems and for epithelial pulses. All systems, except perhaps the pointing conduction system, through-conduct under excess Mg++. Spontaneous activity patterns are described for the swimming, tentacle pulse, and ring systems. Abrupt increases in light intensity inhibit spontaneous activity, sudden decreases augmenting it. In the absence of specialized photoreceptors, light is presumed to act directly on central neurons. Epithelial pulses inhibit swimming, apparently by blocking the generation or conduction of the primary nervous events. This observation, taken in conjunction with evidence of feedback inhibition of the primary swimming system by the cells it fires, is discussed in relation to possible mechanisms whereby the output of nerve cells might be altered by activity in the excitable epithelial cells which envelop them.     

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 life cycle of Hydrichthys tnirus (Gnidaria: Hydrozoa: Anthomedusae: Pandeidae)

The previously unknown life cycle of the parasitic hydroid Hydrichthys mirus is described. The adult medusa has 5–6 tentacles and could be referred to Leuckartiara. Another species of Hydrichthys has previously been shown to have a Stomotoca -like medusa, characterized by the possession of two tentacles. It is proposed that Hydrichthys originated from the Leuckartiara lineage and that, through paedomorphosis in at least one species, the medusa retained the two-tentacle state of the newly-released medusa of all pandeids, thus becoming referable to Stomotoca. It is suggested that Stomotoca has hitherto constituted a polyphyletic taxon, embracing parallel lineages of pandeids, each retaining juvenile features. An additional genus, Larsonia , is introduced to accommodate species with Stomotoca-like medusae and Hydrichthys -like hydroids.     

Composition of gelatin from developmental stages of the sea nettle,chrysaora quinquecirrha

• 1.1. Gelatin, extracted by autoclaving animals after removing free amino acids with 80% ethanol, ranges from 43% of total protein in the scyphistoma to 80% in the adult medusa.
• 2.2. Glutamic acid is predominant in gelatin from the scyphistoma, whereas adult gelatin is more typical of collagens from other cnidarians.
• 3.3. Glycine, hydroxylysine and hydroxyproline are all lower than in other cnidarian collagens, suggesting a low degree of cross-linking in this species.

     

Neurobiology of Stomotoca. II. Pacemakers and conduction pathways

Evidence is presented for separate conduction pathways for swimming and for tentacle coordination in the marginal nerves of the jellyfish Stomotoca. The effector muscles are fired through junctions sensitive to excess Mg⁺⁺, probably represented by the neuromuscular synapses observed by electron microscopy. The swimming effector (striated muscle) fires one-to-one with nerve input signals and myoid conduction occurs. Tentacle responses (smooth muscle contractions) involve facilitation, presumably at the neuro-effector junction; responses are graded and nonpropagating. Electrical correlates of two further conducting systems using the marginal nerves have been recorded. Their functions are unknown. One, the bridge system, extends up the four radii and encircles the peduncle; the other (ring system) is confined to the margin. A fifth conducting system is inferred in the case of the pointing response and its distribution is plotted. Signals have not been obtained from it. Pointing is accompanied by a burst of muscle potentials in the radial smooth muscles and is exhibited after a lengthy latency, indicating a local pacemaker. A sixth conducting pathway is the epithelial system, which mediates crumpling, a response involving the radial muscles without pacemaker intervention. Characteristic conduction velocities and wave forms are noted for the first four systems and for epithelial pulses. All systems, except perhaps the pointing conduction system, through-conduct under excess Mg⁺⁺. Spontaneous activity patterns are described for the swimming, tentacle pulse, and ring systems. Abrupt increases in light intensity inhibit spontaneous activity, sudden decreases augmenting it. In the absence of specialized photoreceptors, light is presumed to act directly on central neurons. Epithelial pulses inhibit swimming, apparently by blocking the generation or conduction of the primary nervous events. This observation, taken in conjunction with evidence of feedback inhibition of the primary swimming system by the cells it fires, is discussed in relation to possible mechanisms whereby the output of nerve cells might be altered by activity in the excitable epithelial cells which envelop them.     

Controlled activation of species typical behaviour to low salinity,seawater movement and seawater depth in <Emphasis Type="Italic">Aurelia labiata</Emphasis> (Scyphozoa) Jellyfish in Roscoe Bay,Canada

Controlled observations in Roscoe Bay examined whether Aurelia labiata medusae would respond to three environmentally significant stimuli: low salinity, seawater movement and seawater depth. Significantly more upward-swimming medusae turned and swam down or sideways when they encountered an experimentally created low-salinity plume than did upward-swimming medusae that encountered a seawater plume or a seawater plume containing milk powder. Downward-swimming medusae that encountered a plume containing freshwater continued swimming down. Significantly more upward-swimming medusae exposed to an experimentally created stream of seawater that gently pushed them horizontally turned and swam down than did upward-swimming medusae not exposed to the seawater stream. Downward-swimming medusae continued to swim down when pushed horizontally. A stronger seawater stream that tumbled as well as pushed medusae horizontally was less effective. In a shallow area near the time of a low tide, most medusae were in the top metre of a 2 m water column whilst in an adjacent area where the water was 4–6 m deep, almost all medusae were below 1.5 m. These observations add to a growing body of evidence that the ecological distribution of A. labiata is influenced by their ability to respond adaptively to significant environmental stimuli.     

In situ time budgets of the scyphomedusae Aurelia aurita, Cyanea sp., and Chrysaora quinquecirrha

The in situ behavior of three scyphomedusan species was videorecorded by scuba divers in natural daytime lighting with minimalinterference to the medusae. The mean percentage of time thatindividual medusae spent swimming ranged from 93 to 100%. Therewere no significant differences in the percent time spent swimmingbetween life stages of a species (ephyra, adult) or betweenspecies. The predominance of swimming activity by medusae indicatesthat swimming, and hence the creation of fluid motions responsiblefor prey entrainment and capture, plays a widespread functionalrole in feeding by scyphomedusae.     

Muscle and nerve net organization in stalked jellyfish (Medusozoa: Staurozoa)

Staurozoan cnidarians display an unusual combination of polyp and medusa characteristics and their morphology may be informative about the evolutionary origin of medusae. We studied neuromuscular morphology of two staurozoans, Haliclystus ‘sanjuanensis ’ and Manania handi , using whole mount immunohistochemistry with antibodies against FMRFamide and α‐tubulin to label neurons and phalloidin to label muscles. All muscles appeared to lack striations. Longitudinal interradial muscles are probable homologues of stalk muscles in scyphopolyps, but in adult staurozoans they are elaborated to inwardly flex marginal lobes of the calyx during prey capture; these muscles are pennate in M. handi . Manubrial perradial muscles, like the manubrium itself, are an innovation shared with pelagic medusae and manubrial interradial muscles are shared with scyphozoan ephyra. Marginal muscles of M. handi displayed occasional synchronous contraction reminiscent of a medusa swim pulse, but contractions were not repetitive. The nerve net in both species showed regional variation in density and orientation of neurons. Some areas labeled predominantly by α‐tubulin antibodies (exumbrellar epidermis), other areas labeled exclusively by FMRFamide antibodies (dense plexus of neurites surrounding the base of secondary tentacles, neuronal concentration at the base of transformed primary tentacles; gastrodermal nerve net), but most areas showed a mix of neurons labeled by these two antibodies and frequent co‐labeling of neurons. Transformed primary tentacles had a concentration of FMRFamide‐immunoreactive neurons at their base that was associated with a pigment spot in M. handi; this is consistent with their homology with rhopalia of medusae, which are also derived from primary tentacles. The muscular system of these staurozoans embodies characteristics of both scyphopolyps and pelagic medusae. However, their nerve net is more polyp‐like, although marginal concentrations of the net associated with primary and secondary tentacles may facilitate the richer behavioral repertoire of staurozoans relative to polyps of other medusozoans. J. Morphol. 278:29–49, 2017. ©© 2016 Wiley Periodicals,Inc.     

EVOLUTION OF LIFE CYCLE,COLONY MORPHOLOGY,AND HOST SPECIFICITY IN THE FAMILY HYDRACTINIIDAE (HYDROZOA,CNIDARIA)

Biased transitions are common throughout the tree of life. The class hydrozoa is no exception, having lost the feeding medusa stage at least 70 times. The family hydractiniidae includes one lineage with pelagic medusae (Podocoryna) and several without (e.g., Hydractinia). The benthic colony stage also varies widely in host specificity and in colony form. The five‐gene phylogeny presented here requires multiple transitions between character states for medusae, host specificity, and colony phenotype. Significant phylogenetic correlations exist between medusoid form, colony morphology, and host specificity. Species with nonfeeding medusae are usually specialized on a single host type, and reticulate colonies are correlated with nonmotile hosts. The history of feeding medusae is less certain. Podocoryna is nested within five lineages lacking medusae. This requires either repeated losses of medusae, or the remarkable re‐evolution of a feeding medusa after at least 150 million years. Traditional ancestral reconstruction favors medusa regain, but a likelihood framework testing biased transitions cannot distinguish between multiple losses versus regain. A hypothesis of multiple losses of feeding medusae requires transient selection pressure favoring such a loss. Populations of species with feeding medusae are always locally rare and lack of feeding medusae does not result in restricted species distribution around the world.     

A newly discovered oxidant defence system and its involvement in the development of Aurelia aurita (Scyphozoa, Cnidaria): reactive oxygen species and elemental iodine control medusa formation

In Aurelia aurita, applied iodine induces medusa formation (strobilation). This process also occurs when the temperature is lowered. This was found to increase oxidative stress resulting in an increased production of iodine from iodide. One polyp produces several medusae (initially termed ephyrae) starting at the polyp's oral end. The spreading of strobilation down the body column is controlled by a feedback loop: ephyra anlagen decrease the tyrosine content in adjacent polyp tissue by producing melanin from tyrosine. Endogenous tyrosine is able to remove iodine by forming iodiferous tyrosine compounds. The reduced level of tyrosine causes the ephyra-polyp-border to move towards the basal end of the former polyp. We argue that an oxidant defence system may exist which makes use of iodide and tyrosine. Like other marine invertebrates, polyps of Aurelia contain iodide ions. Inevitably produced peroxides oxidise iodide into iodine. The danger to be harmed by iodine is strongly decreased by endogenous tyrosine which reacts with iodine to form iodiferous tyrosine compounds including thyroxin. Both substances together, iodide and tyrosine, form an efficient oxidant defence system which shields the tissue against damage by reactive oxygen species. In the course of evolution (from a species at the basis of the animal kingdom like Aurelia to a highly evolved species like man) the waste product thyroxin (indicating a high metabolic rate) has developed into a hormone which controls the metabolic rate.     

Notes on Stauridiosarsia producta (Hydrozoa, Corynidae), a New Addition to the New England Marine Fauna1

The hydroid and medusa of Stauridiosarsia producta (Wright, 1858) are reported for the first time from the New England coast of North America. The laboratory reared hydroid exhibits variation encompassing the previously reported forms from England and Helgoland. Nematocysts of the hydroid are stenoteles of two sizes. Young medusae have nematocysts irregularly scattered over the exumbrella. These are lost after 5–7 days. The characteristic umbilical canal of the medusa becomes distinct about 24 hours after liberation. Development to sexual maturity requires about 30 days. In addition to the two sizes of stenoteles, the medusa also possesses desmonemes.     

Central neural circuitry in the jellyfish Aglantha: a model 'simple nervous system'

Like other hydrozoan medusae, Aglantha lacks a brain, but the two marginal nerve rings function together as a central nervous system. Twelve neuronal and two excitable epithelial conduction systems are described and their interactions summarized. Aglantha differs from most medusae in having giant axons. It can swim and contract its tentacles in two distinct ways (escape and slow). Escape responses are mediated primarily by giant axons but conventional interneurons are also involved in transmission of information within the nerve rings during one form of escape behavior. Surprisingly, giant axons provide the motor pathway to the swim muscles in both escape and slow swimming. This is possible because these axons can conduct calcium spikes as well as sodium spikes and do so on an either/or basis without overlap. The synaptic and ionic bases for these responses are reviewed. During feeding, the manubrium performs highly accurate flexions to points at the margin. At the same time, the oral lips flare open. The directional flexions are conducted by FMRFamide immunoreactive nerves, the lip flaring by an excitable epithelium lining the radial canals. Inhibition of swimming during feeding is due to impulses propagated centrifugally in the same epithelium. Aglantha probably evolved from an ancestor possessing a relatively simple wiring plan, as seen in other hydromedusae. Acquisition of giant axons resulted in considerable modification of this basic plan, and required novel solutions to the problems of integrating escape with non-escape circuitry.     

Systematics and biogeography of the jellyfish Aurelia labiata (Cnidaria: Scyphozoa)

The hypothesis that the common eastern North Pacific Aurelia is A. aurita is falsified with morphological analysis. The name Aurelia labiata is resurrected, and the species is redescribed, to refer to medusae differing from A. aurita by a suite of characters related to a broad and elongated manubrium. Specifically, the oral arms are short, separated by and arising from the base of the fleshy manubrium, and the planulae are brooded upon the manubrium itself, rather than on the oral arms. Aurelia aurita possesses no corresponding enlarged structure. Furthermore, the number of radial canals is typically much greater in A. labiata, and thus the canals often appear more anastomosed than in A. aurita. Finally, most A. labiata medusae possess a 16-scalloped bell margin, whereas the margin is 8-scalloped in most A. aurita. Separation of the two forms has previously been noted on the basis of allozyme and isozyme analyses and on the histology of the neuromuscular system. Partial 18S rDNA sequencing corroborates these findings. Three distinct morphotypes of A. labiata, corresponding to separate marine bioprovinces, have been identified among 17 populations from San Diego, California, to Prince William Sound, Alaska. The long-undisputed species A. limbata may be simply a color morph of A. labiata, or a species within a yet-unelaborated A. labiata species complex. The first known introduction of Aurelia cf. aurita into southern California waters is documented. Although traditional jellyfish taxonomy tends to recognize many species as cosmopolitan or nearly so, these results indicate that coastal species, such as A. labiata, may experience rapid divergence among isolated populations, and that the taxonomy of such species should therefore be scrutinized with special care.