Ontogenetic transitions, biomechanical trade-offs and macroevolution of scyphozoan medusae swimming patterns.

Ephyrae, the early stages of scyphozoan jellyfish, possess a conserved morphology among species. However, ontogenetic transitions lead to morphologically different shapes among scyphozoan lineages, with important consequences for swimming biomechanics, bioenergetics and ecology. We used high-speed imaging to analyse biomechanical and kinematic variables of swimming in 17 species of Scyphozoa (1 Coronatae, 8 "Semaeostomeae" and 8 Rhizostomeae) at different developmental stages. Swimming kinematics of early ephyrae were similar, in general, but differences related to major lineages emerged through development. Rhizostomeae medusae have more prolate bells, shorter pulse cycles and higher swimming performances. Medusae of "Semaeostomeae", in turn, have more variable bell shapes and most species had lower swimming performances. Despite these differences, both groups travelled the same distance per pulse suggesting that each pulse is hydrodynamically similar. Therefore, higher swimming velocities are achieved in species with higher pulsation frequencies. Our results suggest that medusae of Rhizostomeae and "Semaeostomeae" have evolved bell kinematics with different optimized traits, rhizostomes optimize rapid fluid processing, through faster pulsations, while "semaeostomes" optimize swimming efficiency, through longer interpulse intervals that enhance mechanisms of passive energy recapture.

Localization of Multiple Jellyfish Toxins Shows Specificity for Functionally Distinct Polyps and Nematocyst Types in a Colonial Hydrozoan.

Hydractinia symbiolongicarpus is a colonial hydrozoan that displays a division of labor through morphologically distinct and functionally specialized polyp types. As with all cnidarians, their venoms are housed in nematocysts, which are scattered across an individual. Here, we investigate the spatial distribution of a specific protein family, jellyfish toxins, in which multiple paralogs are differentially expressed across the functionally specialized polyps. Jellyfish toxins (JFTs) are known pore-forming toxins in the venoms of medically relevant species such as box jellyfish (class Cubozoa), but their role in other medusozoan venoms is less clear. Utilizing a publicly available single-cell dataset, we confirmed that four distinct H. symbiolongicarpus JFT paralogs are expressed in nematocyst-associated clusters, supporting these as true venom components in H. symbiolongicarpus. In situ hybridization and immunohistochemistry were used to localize the expression of these JFTs across the colony. These expression patterns, in conjunction with known nematocyst type distributions, suggest that two of these JFTs, HsymJFT1c-I and HsymJFT1c-II, are localized to specific types of nematocysts. We further interpret JFT expression patterns in the context of known regions of nematogenesis and differential rates of nematocyst turnover. Overall, we show that JFT expression patterns in H. symbiolongicarpus are consistent with the subfunctionalization of JFT paralogs across a partitioned venom system within the colony, such that each JFT is expressed within a specific set of functionally distinct polyp types and, in some cases, specific nematocyst types.

A new subfamily of ulmarid scyphomedusae, the Santjordiinae, with a description of Santjordia pagesi gen. et sp. nov. (Cnidaria: Scyphozoa: Discomedusae: Semaeostomeae: Ulmaridae) from the Sumisu Caldera, Ogasawara Islands, Japan.

An undescribed species of ulmarid medusa was observed in situ and captured at 812 m depth within the Sumisu Caldera, Ogasawara Islands, Japan. Morphological and molecular evidence points to it being distinct from other ulmarid medusae and a new species (pagesi), genus (Santjordia) and subfamily (Santjordiinae) are herein erected to contain it. This new subfamily of semaeostome ulmarid medusae has both marginal and subumbrellar rhopalia, making it unique within the order Semaeostomeae. Although the combination of subumbrellar tentacles and the lack of branched canals should warrant the erection of a new family within the Semaeostomeae, a lack of information on the gonad structure and poor bootstrap support in the molecular phylogenetic tree cause us to relegate it to the catch-all family Ulmaridae, until greater taxon sampling and phylogenetic analyses are carried out for the Semaeostomeae.

A morphological review of the jellyfish genus Nausithoe Klliker, 1853 (Nausithoideae, Coronatae, Scyphozoa, Cnidaria).

In this study we address the diversity of the scyphozoan jellyfish genus Nausithoe Klliker, 1853 (Nausithoidae, Coronatae), questioning the feasibility of using some characters of the medusa stage to identify species and filling in gaps concerning species of the genus and their distributions. Like most scyphozoans, the vast majority of the 21 Nausithoe species have a metagenetic life cycle, but similarity of most polyps within the genus highlights the need for studying morphology of the medusa stage. By analyzing morphological features on preserved and live specimens (polyps and medusae) and comparing these data with the original descriptions, we were able to validate twenty species of the group, providing new information for some of them.

siRNA-mediated gene knockdown via electroporation in hydrozoan jellyfish embryos.

As the sister group to bilaterians, cnidarians stand in a unique phylogenetic position that provides insight into evolutionary aspects of animal development, physiology, and behavior. While cnidarians are classified into two types, sessile polyps and free-swimming medusae, most studies at the cellular and molecular levels have been conducted on representative polyp-type cnidarians and have focused on establishing techniques of genetic manipulation. Recently, gene knockdown by delivery of short hairpin RNAs into eggs via electroporation has been introduced in two polyp-type cnidarians, Nematostella vectensis and Hydractinia symbiolongicarpus, enabling systematic loss-of-function experiments. By contrast, current methods of genetic manipulation for most medusa-type cnidarians, or jellyfish, are quite limited, except for Clytia hemisphaerica, and reliable techniques are required to interrogate function of specific genes in different jellyfish species. Here, we present a method to knock down target genes by delivering small interfering RNA (siRNA) into fertilized eggs via electroporation, using the hydrozoan jellyfish, Clytia hemisphaerica and Cladonema paciificum. We show that siRNAs targeting endogenous GFP1 and Wnt3 in Clytia efficiently knock down gene expression and result in known planula phenotypes: loss of green fluorescence and defects in axial patterning, respectively. We also successfully knock down endogenous Wnt3 in Cladonema by siRNA electroporation, which circumvents the technical difficulty of microinjecting small eggs. Wnt3 knockdown in Cladonema causes gene expression changes in axial markers, suggesting a conserved Wnt/ß-catenin-mediated pathway that controls axial polarity during embryogenesis. Our gene-targeting siRNA electroporation method is applicable to other animals, including and beyond jellyfish species, and will facilitate the investigation and understanding of myriad aspects of animal development.

Environmental drivers of the occurrence and abundance of the Irukandji jellyfish (Carukia barnesi).

Understanding the links between species and their environment is critical for species management. This is particularly true for organisms of medical and/or economic significance. The 'Irukandji' jellyfish (Carukia barnesi) is well known for its small size, cryptic nature, and highly venomous sting. Being the namesake of the Irukandji syndrome, contact with this marine stinger often leads to hospitalization and can be fatal. Consequently, the annual occurrence of this organism is believed to cost the Australian government an estimated $AUD3 billion annually in medical costs and losses for tourism. Despite its economic importance the logistical difficulties related to surveying C.barnesi in situ has led to a paucity of knowledge regarding its ecology and significantly impeded management strategies to date. In this study, we use six years of direct C. barnesi capture data to explore patterns pertaining to the annual occurrence and abundance of this species in the nearshore waters of the Cairns coast. We provide novel insights into trends in medusae aggregations and size distribution and primarily focus on the potential role of environmental drivers for annual C. barnesi occurrence patterns. Using a two-part hurdle model, eight environmental parameters were investigated over four time periods for associations with records of medusa presence and abundance. Final models showed a small amount of variation in medusa presence and abundance patterns could be accounted for by long-term trends pertaining to rainfall and wind direction. However, the assessed environmental parameters could not explain high annual variation or site location effects. Ultimately best-fit models had very low statistical inference power explaining between 16 and 20% of the variance in the data, leaving approximately 80% of all variation in medusa presence and abundance unexplained.

Phylogenetics of swimming behaviour in Medusozoa: the role of giant axons and their possible evolutionary origin.

Although neural tissues in cnidarian hydroids have a nerve net structure, some cnidarian medusae contain well-defined nerve tracts. As an example, the hydrozoan medusa Aglantha digitale has neural feeding circuits that show an alignment and condensation, which is absent in its relatives Aequorea victoria and Clytia hemisphaerica. In some cases, neural condensations take the form of fast propagating giant axons concerned with escape or evasion. Such giant axons appear to have developed from the fusion of many, much finer units. Ribosomal DNA analysis has identified the lineage leading to giant axon-based escape swimming in Aglantha and other members of the Aglaura clade of trachymedusan jellyfish. The Aglaura, along with sister subclades that include species such as Colobonema sericeum, have the distinctive ability to perform dual swimming, i.e. to swim at either high or low speeds. However, the form of dual swimming exhibited by Colobonema differs both biomechanically and physiologically from that in Aglantha and is not giant axon based. Comparisons between the genomes of such closely related species might provide a means to determine the molecular basis of giant axon formation and other neural condensations. The molecular mechanism responsible may involve 'fusogens', small molecules possibly derived from viruses, which draw membranes together prior to fusion. Identifying these fusogen-based mechanisms using genome analysis may be hindered by the many changes in anatomy and physiology that followed giant axon evolution, but the genomic signal-to-noise ratio may be improved by examining the convergent evolution of giant axons in other hydrozoa, such as the subclass Siphonophora.

Caltsacoryne setouchiensis (Hydrozoa, Anthoathecata) a new genus and species of hydrozoan jellyfish from Japan.

We report a new genus and species of hydrozoan jellyfish belonging to the order Anthoathecata collected from the Seto Inland Sea, western Japan. Caltsacoryne setouchiensis gen. et sp. n. can be distinguished from other species of Corynidae based on the following combination of morphological characters: number of tentacles, cnidocyst pads, manubrium length, and the shape of the gonad and tentacles. A table comparing the primary diagnostic characters of this new genus of Corynidae is presented.

Neurotoxicity of Olindias sambaquiensis and Chiropsalmus quadrumanus extracts in sympathetic nervous system.

Cnidarians are equipped with nematocysts, which are specialized organelles used to inoculate venom during prey capturing and defense. Their venoms are rich in toxins and a potential source of bioactive compounds, however, poorly explored so far. In this work, the activity of the methanolic extracts from the hydromedusa Olindias sambaquiensis and the cubozoan jellyfish Chiropsalmus quadrumanus were studied in sympathetic neurotransmission. For that, bisected rat vas deferens - a classic model of sympathetic neurotransmission - were incubated with the extracts for further myographic and histopathological analysis. The O. sambaquiensis extract, at 0.1 µg/mL, facilitated the neurogenic contractions of the noradrenergic-rich epididymal portion, while reducing the noradrenaline (NA) potency, which suggests an interaction with postsynaptic α1-adrenoceptors. On the other hand, a higher concentration (1 µg/mL) leads to time- and frequency-dependent blockade of nerve-evoked contractions without significantly changing the response to exogenous NA. In turn, the C. quadrumanus extract at 0.1 µg/mL induced blockade of nerve-evoked noradrenergic contractions while reducing the potency to exogenous NA. Both extracts did not affect the purinergic neurotransmission or induce muscle damages. Our results demonstrate that O. sambaquiensis and C. quadrumanus extracts significantly interfere with the noradrenergic neurotransmission without altering purinergic response or smooth muscle structure on rat vas deferens. Such results bring to light the pharmacological potential of O. sambaquiensis and C. quadrumanus molecules for therapeutics focusing on noradrenergic neurotransmission.

Exumbrellar Surface of Jellyfish: A Comparative Fine Structure Study with Remarks on Surface Reflectance.

The exumbrellar surfaces of six pelagic cnidarians from three classes were ultra-structurally compared to reveal their structural diversity in relation to their gelatinous, transparent bodies. We examined two hydrozoans (Diphyes chamissonis and Colobonema sericeum), a cubozoan (Chironex yamaguchii), and three scyphozoans (Atolla vanhöffeni, Aurelia coerulea, and Mastigias papua). The exumbrellar surfaces of the mesoglea in D. chamissonis, Ch. yamaguchii, Au. coerulea, and M. papua were covered with a simple epidermis; the shapes of the epidermal cells were remarkably different among the species. The epidermal cells of Ch. yamaguchii and M. papua possessed an array of microvilli on the apical side. The array possibly reduced light reflectance and provided some other surface properties, as seen for the cuticular nipple array in tunicates, considering the length, width, and pitch of the microvilli. The reduction of light reflectance on the array of microvilli was supported by the simulation with rigorous coupled wave analysis (RCWA). Microvilli were sparse and did not form an array in metephyrae of Au. coerulea. The mesoglea matrix beneath the basal side of the epidermis was loose in all of the species. The exumbrellar side of the mesoglea was exposed only in the mesopelagic species, At. vanhöffeni and Co. sericeum, and electron-dense layer(s) covered the surface of the mesoglea. It is uncertain whether the exumbrellar epidermis is absent in these species or the epidermal cells are completely exfoliated during the sampling and handling processes. In the latter case, the electron-dense layer(s) on the mesoglea surface might originally underlie the epidermis.

Jellyfish anaphylaxis: A wide spectrum of sensitization routes.

Background: Recent studies demonstrated that, in the past few years, the number of jellyfish species is increasing worldwide; this increase can be explained by environmental and climatic reasons. Contacts with jellyfish can cause acute and chronic effects, including allergic reactions. Although anaphylaxis caused by jellyfish is a rare event, repetitive stings during bathing as well as marine sports and job activities represent important risk factors that can increase the probability of sensitization. Recently, it was also pointed out the possibility of anaphylaxis caused by jellyfish ingestion. In these cases, the sensitization could also be related to previous stings. In cases in which there is no history of jellyfish contact or ingestion, it has been hypothesized that there is a sensitization to an unknown cross-reactive antigen. Objective: The purpose of this work was to collect and review published studies and cases of anaphylaxis associated with jellyfish. Methods: We performed a medical literature data base search, which included English language articles published until September 2019, by using the key words "jellyfish" associated with "anaphylaxis" or "anaphylactic shock." Results: The results of our research showed that dangerous reactions can be caused both by contact and ingestion. Moreover, the latest changes in food habits, life style, and globalization could lead to a more frequent exposure to jellyfish both by contact and ingestion, and, consequently, to a higher probability of sensitization. Conclusion: Prospective studies and well-structured research are needed to better understand all the potential immunologic elements of jellyfish, to clarify its role in sensitization, and to avoid possible dangerous allergic reactions caused by cross-reactivity.

Nematocyst types and venom effects of Aurelia aurita and Velella velella from the Mediterranean Sea.

Natural substances produced by venomous marine organisms are thought to be possible sources of useful compounds and new drugs having the potential to open new ways for pharmacology, nutrition and environmental applications. In this framework, cnidarians are very interesting being widely distributed and all are venomous organisms; so, a deep knowledge of their occurrence, morphology of venomous structures and of effects of venoms at cellular level is fundamental to evaluate the possible utilization of venomous compounds or extracts. In this research, the morphology and occurrence of nematocysts in two cnidarian species (Aurelia aurita, Velella velella), and the preliminary evaluation of the cytotoxicity of V. velella crude extract, of which cytotoxicity on cell cultures at present is unknown, were considered. The specimens were sampled in Güllük Bay, Southwestern coast of Turkey, and in the Gulf of Genova, Northwestern coast of Italy. Six nematocyst types (a-isorhiza, A-isorhiza, O-isorhiza, eurytele, polyspiras, birhopaloid) having different sizes, were observed in A. aurita, and two types (eurytele and stenotele) in V. velella. The crude extract from V. velella showed cytotoxic activity against cultured fibroblasts L929 at high doses, while inducing cell proliferation at low doses. The protein content in the extract increased remarkably after disruption of nematocysts.

Macroscale abundance patterns of hydromedusae in the temperate Southwestern Atlantic (27°-56° S).

Gelatinous organisms are crucial components of marine ecosystems and some species imply social and economic consequences. However, certain geographic areas, such as the temperate Southwestern Atlantic (SWA, 27° - 56° S), remain understudied in terms of jellyfish ecological data. We analyzed 3,727 plankton samples collected along ~6.7 million km2 over a 31-year period (1983-2014) to determine the occurrence, abundance, and diversity patterns of hydromedusae in the SWA. Analyses were made at both community and species levels. Two abundance hot spots of hydromedusae were identified, where values up to 2,480 ind. m-3 were recorded between 2003 and 2014. Liriope tetraphylla and Obelia spp. were the main responsible for recurrent peaks. Diversity indexes were in the range of those published for temperate areas worldwide, and some coastal zones showed values that can be considered moderate to high for a temperate neritic region. The community analysis yielded 10 groups following previously determined biogeographic schemes throughout the study area. This work enhances the knowledge of hydromedusae in the SWA and provides essential information about the current global warming context and the gelatinous zooplankton data necessity.

Widespread utilization of passive energy recapture in swimming medusae.

Recently, it has been shown that some medusae are capable of swimming very efficiently, i.e. with a low cost of transport, and that this is in part due to passive energy recapture (PER) which occurs during bell relaxation. We compared the swimming kinematics among a diverse array of medusae, varying in taxonomy, morphology and propulsive and foraging modes, in order to evaluate the prevalence of PER in medusae. We found that while PER was common among taxa, the magnitude of the contribution to overall swimming varied greatly. The ability of medusae to utilize PER was not related to morphology and swimming performance but was controlled by their swimming kinematics. Utilizing PER required the medusae to pause after bell expansion and individuals could modulate their PER by changing their pause duration. PER can greatly enhance swimming efficiency but there appear to be trade-offs associated with utilizing PER.

Assessing the Efficacy of First-Aid Measures in Physalia sp. Envenomation, Using Solution- and Blood Agarose-Based Models.

Stings from the hydrozoan species in the genus Physalia cause intense, immediate skin pain and elicit serious systemic effects. There has been much scientific debate about the most appropriate first aid for these stings, particularly with regard to whether vinegar use is appropriate (most current recommendations recommend against vinegar). We found that only a small percentage (≤1.0%) of tentacle cnidae discharge during a sting event using an ex vivo tissue model which elicits spontaneous stinging from live cnidarian tentacles. We then tested a variety of rinse solutions on both Atlantic and Pacific Physalia species to determine if they elicit cnidae discharge, further investigating any that did not cause immediate significant discharge to determine if they are able to inhibit cnidae discharge in response to chemical and physical stimuli. We found commercially available vinegars, as well as the recently developed Sting No More® Spray, were the most effective rinse solutions, as they irreversibly inhibited cnidae discharge. However, even slight dilution of vinegar reduced its protective effects. Alcohols and folk remedies, such as urine, baking soda and shaving cream, caused varying amounts of immediate cnidae discharge and failed to inhibit further discharge, and thus likely worsen stings.

Gramine-derived Bromo-alkaloids Activating NF--kB-dependent Transcription from the Marine Hydroid Abietinaria abietina.

Two new natural products, 6-bromogramine (1) and bis-6-bromogramine (2) were isolated from the marine hydroid A bietinaria abietina and their structures were established using NMR and MS analysis. Compounds 1 and 2 activate NF-cB-dependent transcriptional activity in JB6 Cl 41 NF-KB cells at 1.6 AM concentrations.

Phylogenomic Analyses Support Traditional Relationships within Cnidaria.

Cnidaria, the sister group to Bilateria, is a highly diverse group of animals in terms of morphology, lifecycles, ecology, and development. How this diversity originated and evolved is not well understood because phylogenetic relationships among major cnidarian lineages are unclear, and recent studies present contrasting phylogenetic hypotheses. Here, we use transcriptome data from 15 newly-sequenced species in combination with 26 publicly available genomes and transcriptomes to assess phylogenetic relationships among major cnidarian lineages. Phylogenetic analyses using different partition schemes and models of molecular evolution, as well as topology tests for alternative phylogenetic relationships, support the monophyly of Medusozoa, Anthozoa, Octocorallia, Hydrozoa, and a clade consisting of Staurozoa, Cubozoa, and Scyphozoa. Support for the monophyly of Hexacorallia is weak due to the equivocal position of Ceriantharia. Taken together, these results further resolve deep cnidarian relationships, largely support traditional phylogenetic views on relationships, and provide a historical framework for studying the evolutionary processes involved in one of the most ancient animal radiations.

Pain from bluebottle jellyfish stings.

An 11-year-old girl presented to the emergency department with severe pain after a jellyfish sting at a New South Wales beach. Bluebottle (Physalia) jellyfish was deemed the most likely cause considering her geographical location. The Australian Resuscitation Council Guideline (2010) suggests immersing in water as hot as can be tolerated for 20 min for treating pain from jellyfish stings. This guideline was written based on past case reports, books and randomised controlled trials (RCTs). We performed a search to assess the most current evidence for relief of pain from Bluebottle jellyfish stings, which yielded two systematic reviews and seven RCTs. Both systematic reviews had similar conclusions, with one of the RCTs used in both reviews showing the most relevance to our presenting patient in terms of demographics, location and jellyfish type. This journal club article is an appraisal of this RCT by Loten et al. and the validity of its conclusion that hot water immersion is most effective for the relief of pain from Bluebottle stings.

Radioactivity in three species of eastern Mediterranean jellyfish.

Activity concentrations of (137)Cs, (40)K, (210)Po, (210)Pb, (234)U and (238)U were determined in umbrella and oral arms of three widely distributed jellyfish species; namely Rhopilema nomadica Galil, 1990, Aurelia aurita Linne, 1758 and Aequorea forskalea Péron & Lesueur, 1810 collected from February 2011 to January 2012 in four sampling locations along the Syrian coast (Eastern Mediterranean Sea). The results have shown significant variations in radionuclides activity concentrations amongst the species. The average activity concentrations of (40)K, (210)Po, (210)Pb, (234)U and (238)U in the umbrella of R. nomadica species were higher than the average activity concentrations in the umbrella of A. aurita species by about 3.2, 1.4, 1.8, 3.2 and 3.2 folds, and A. forskalea species by about 45.5, 15.4, 19, 7.4 and 7.6 folds, respectively. The average activity concentrations of (40)K, (210)Po, (210)Pb, (234)U and (238)U in oral arms of R. nomadica species were higher than the average activity concentrations in oral arms of A. aurita species by about 3.8, 1.7, 1.9, 2.8 and 2.9 folds, respectively. (137)Cs activity concentrations were below the detection limit in all measured samples. In addition, activity concentrations of (137)Cs, (40)K, (210)Po, (210)Pb, (234)U and (238)U were also determined in 44 surface seawater samples and the activity concentrations ranged between 10.6 and 11.9 Bq l(-1) for (40)K, 1.1 and 1.4 mBq l(-1) for (210)Po, 0.5 and 0.7 mBq l(-1) for (210)Pb, 40.8 and 44.5 mBq l(-1) for (234)U, and 36.9 and 38.4 mBq l(-1) for (238)U, while (137)Cs activity concentrations were below the detection limit in all measured samples. Moreover, the umbrella and oral arms readily accumulated (40)K, (210)Po, (210)Pb, (234)U and (238)U above ambient seawater levels in the sequence of (210)Po > (210)Pb > (4) K > (234)U and (238)U. Concentration ratio (CR) values were relatively high for (210)Po and (210)Pb and reached 10(3) and 10(2), respectively for the jellyfish R. nomadica species compared to A. aurita and A. forskalea species. Therefore, R. nomadica can be used as biomonitor for these two radionuclides in the Eastern Mediterranean Sea. However, the obtained data can be considered the first reported baseline values for radioactivity in jellyfish.

Exploring the therapeutic potential of jellyfish venom.

The venom of certain jellyfish has long been known to be potentially fatal to humans, but it is only recently that details of the proteomes of these fascinating creatures are emerging. The molecular contents of the nematocysts from several jellyfish species have now been analyzed using proteomic MS approaches and include the analysis of Chironex fleckeri, one of the most venomous jellyfish known. These studies suggest that some species contain toxins related to peptides and proteins found in other venomous creatures. The detailed characterization of jellyfish venom is likely to provide insight into the diversification of toxins and might be a valuable resource in drug design.