Soft skeletons transmit force with variable gearing.

A hydrostatic skeleton allows a soft body to transmit muscular force via internal pressure. A human's tongue, an octopus' arm and a nematode's body illustrate the pervasive presence of hydrostatic skeletons among animals, which has inspired the design of soft engineered actuators. However, there is a need for a theoretical basis for understanding how hydrostatic skeletons apply mechanical work. We therefore modeled the shape change and mechanics of natural and engineered hydrostatic skeletons to determine their mechanical advantage (MA) and displacement advantage (DA). These models apply to a variety of biological structures, but we explicitly consider the tube feet of a sea star and the body segments of an earthworm, and contrast them with a hydraulic press and a McKibben actuator. A helical winding of stiff, elastic fibers around these soft actuators plays a critical role in their mechanics by maintaining a cylindrical shape, distributing forces throughout the structure and storing elastic energy. In contrast to a single-joint lever system, soft hydrostats exhibit variable gearing with changes in MA generated by deformation in the skeleton. We found that this gearing is affected by the transmission efficiency of mechanical work (MA×DA) or, equivalently, the ratio of output to input work. The transmission efficiency changes with the capacity to store elastic energy within helically wrapped fibers or associated musculature. This modeling offers a conceptual basis for understanding the relationship between the morphology of hydrostatic skeletons and their mechanical performance.

"Indirect development" increases reproductive plasticity and contributes to the success of scyphozoan jellyfish in the oceans.

Ecologists and evolutionary biologists have been looking for the key(s) to the success of scyphomedusae through their long evolutionary history in multiple habitats. Their ability to generate young medusae (ephyrae) via two distinct reproductive strategies, strobilation or direct development from planula into ephyra without a polyp stage, has been a potential explanation. In addition to these reproductive modes, here we provide evidence of a third ephyral production which has been rarely observed and often confused with direct development from planula into ephyra. Planulae of Aurelia relicta Scorrano et al. 2017 and Cotylorhiza tuberculata (Macri 1778) settled and formed fully-grown polyps which transformed into ephyrae within several days. In distinction to monodisk strobilation, the basal polyp of indirect development was merely a non-tentaculate stalk that dissolved shortly after detachment of the ephyra. We provide a fully detailed description of this variant that increases reproductive plasticity within scyphozoan life cycles and is different than either true direct development or the monodisk strobilation. Our observations of this pattern in co-occurrence with mono- and polydisk strobilation in Aurelia spp. suggest that this reproductive mode may be crucial for the survival of some scyphozoan populations within the frame of a bet-hedging strategy and contribute to their long evolutionary success throughout the varied conditions of past and future oceans.

Cassiosomes are stinging-cell structures in the mucus of the upside-down jellyfish Cassiopea xamachana.

Snorkelers in mangrove forest waters inhabited by the upside-down jellyfish Cassiopea xamachana report discomfort due to a sensation known as stinging water, the cause of which is unknown. Using a combination of histology, microscopy, microfluidics, videography, molecular biology, and mass spectrometry-based proteomics, we describe C. xamachana stinging-cell structures that we term cassiosomes. These structures are released within C. xamachana mucus and are capable of killing prey. Cassiosomes consist of an outer epithelial layer mainly composed of nematocytes surrounding a core filled by endosymbiotic dinoflagellates hosted within amoebocytes and presumptive mesoglea. Furthermore, we report cassiosome structures in four additional jellyfish species in the same taxonomic group as C. xamachana (Class Scyphozoa; Order Rhizostomeae), categorized as either motile (ciliated) or nonmotile types. This inaugural study provides a qualitative assessment of the stinging contents of C. xamachana mucus and implicates mucus containing cassiosomes and free intact nematocytes as the cause of stinging water.

From single neurons to behavior in the jellyfish Aurelia aurita.

Jellyfish nerve nets provide insight into the origins of nervous systems, as both their taxonomic position and their evolutionary age imply that jellyfish resemble some of the earliest neuron-bearing, actively-swimming animals. Here, we develop the first neuronal network model for the nerve nets of jellyfish. Specifically, we focus on the moon jelly Aurelia aurita and the control of its energy-efficient swimming motion. The proposed single neuron model disentangles the contributions of different currents to a spike. The network model identifies factors ensuring non-pathological activity and suggests an optimization for the transmission of signals. After modeling the jellyfish's muscle system and its bell in a hydrodynamic environment, we explore the swimming elicited by neural activity. We find that different delays between nerve net activations lead to well-controlled, differently directed movements. Our model bridges the scales from single neurons to behavior, allowing for a comprehensive understanding of jellyfish neural control of locomotion.

Medusa: A Review of an Ancient Cnidarian Body Form.

Medusae (aka jellyfish) have multiphasic life cycles and a propensity to adapt to, and proliferate in, a plethora of aquatic habitats, connecting them to a number of ecological and societal issues. Now, in the midst of the genomics era, affordable next-generation sequencing (NGS) platforms coupled with publically available bioinformatics tools present the much-anticipated opportunity to explore medusa taxa as potential model systems. Genome-wide studies of medusae would provide a remarkable opportunity to address long-standing questions related to the biology, physiology, and nervous system of some of the earliest pelagic animals. Furthermore, medusae have become key targets in the exploration of marine natural products, in the development of marine biomarkers, and for their application to the biomedical and robotics fields. Presented here is a synopsis of the current state of medusa research, highlighting insights provided by multi-omics studies, as well as existing knowledge gaps, calling upon the scientific community to adopt a number of medusa taxa as model systems in forthcoming research endeavors.

First description of the life cycle of the jellyfish Rhizostoma luteum (Scyphozoa: Rhizostomeae).

Jellyfish blooms are a significant environmental problem that is increasing and may be influenced by anthropocentric practices such as overfishing, pollution, eutrophication, translocation, climate change, and ocean acidification. Many jellyfish have unknown life cycles leading to these blooms. We describe for the first time, the life cycle of scyphozoan jellyfish Rhizostoma luteum from the planula to the young medusa stages, based on laboratory observations. We also provide a preliminary assessment of temperature related to life stages. Comparisons were made with early life history stages of its sibling species Rhizostoma pulmo and Rhizostoma octopus. The life cycle of R. luteum follows the general pattern of metagenesis of scyphozoans. Scyphistoma culture was maintained in filtered seawater at 17-17.5 °C, salinity 37 and light photoperiod (12:12 h light:dark). Scyphistomae were exposed to an experimental temperature descent for two days to test their survival capacity under severe winter conditions. Only one asexual reproduction mode was observed, which is employed for propagation, consisting of podocyst formation with excystment, subsequent development of scyphistoma, strobilation and liberation of viable ephyra. The development of the ephyra to metaephyra was photodocumented, reaching the metaephyra stage in approximately 21-25 days. Young medusae grow rapidly and maturity was reached after a 3-month post-liberation period with a mean bell diameter of 13.27 ± 2.26 cm and wet weight of 181.53 ± 53 g. The life cycle of R. luteum resembles that of its congeners, with the distinction that it has the unique features of being a brooding species (internal fertilisation with subsequent release of planulae) and under the conditions tested, the predominantly strobilation type observed was monodisc, and not polydisc as with the other two species in the genus Rhizostoma. As R. luteum shows sufficient requisite to form blooms if environmental circumstances change, it is important to understand its life cycle.

Tentacle extract from the jellyfish Cyanea capillata increases proliferation and migration of human umbilical vein endothelial cells through the ERK1/2 signaling pathway.

Wound healing is a complex biological process, and current research finds that jellyfish have a great capacity for promoting growth and healing. However, the underlying mechanisms remain unclear. Thus, this study was conducted to investigate the molecular mechanisms and effects of a tentacle extract (TE) from the jellyfish Cyanea capillata (C. capillata) on cell proliferation and migration in human umbilical vein endothelial cells (HUVECs). First, our results showed that TE at the concentration of 1 µg/ml could promote cell proliferation over various durations, induce a transition of the cells from the G1-phase to the S/G2-phase of the cell cycle, and increase the expression of cell cycle proteins (CyclinB1 and CyclinD1). Second, we found that TE could activate the PI3K/Akt, ERK1/2 and JNK MAPK signaling pathways but not the NF-κB signaling pathway or the apoptosis signaling cascade. Finally, we demonstrated that the TE-induced expression of cell cycle proteins was decreased by ERK1/2 inhibitor PD98059 but not by PI3K inhibitor LY294002 or JNK inhibitor SP600125. Similarly, the TE-enhanced migration ability of HUVECs was also markedly attenuated by PD98059. Taken together, our findings indicate that TE-induced proliferation and migration in HUVECs mainly occurred through the ERK1/2 MAPK signaling pathway. These results are instructively important for further research on the isolation and purification of growth-promoting factors from C. capillata and are hopeful as a means to improve human wound repair in unfavorable conditions.

Local versus Generalized Phenotypes in Two Sympatric Aurelia Species: Understanding Jellyfish Ecology Using Genetics and Morphometrics.

For individuals living in environmentally heterogeneous environments, a key component for adaptation and persistence is the extent of phenotypic differentiation in response to local environmental conditions. In order to determine the extent of environmentally induced morphological variation in a natural population distributed along environmental gradients, it is necessary to account for potential genetic differences contributing to morphological differentiation. In this study, we set out to quantify geographic morphological variation in the moon jellyfish Aurelia exposed at the extremes of a latitudinal environmental gradient in the Gulf of Mexico (GoM). We used morphological data based on 28 characters, and genetic data taken from mitochondrial cytochrome oxidase I (COI) and nuclear internal transcribed spacer 1 (ITS-1). Molecular analyses revealed the presence of two genetically distinct species of Aurelia co-occurring in the GoM: Aurelia sp. 9 and Aurelia c.f. sp. 2, named for its divergence from (for COI) and similarity to (for ITS-1) Aurelia sp. 2 (Brazil). Neither species exhibited significant population genetic structure between the Northern and the Southeastern Gulf of Mexico; however, they differed greatly in the degree of geographic morphological variation. The morphology of Aurelia sp. 9 exhibited ecophenotypic plasticity and varied significantly between locations, while morphology of Aurelia c.f. sp. 2 was geographically invariant (i.e., canalized). The plastic, generalist medusae of Aurelia sp. 9 are likely able to produce environmentally-induced, "optimal" phenotypes that confer high relative fitness in different environments. In contrast, the non-plastic generalist individuals of Aurelia c.f. sp. 2 likely produce environmentally-independent phenotypes that provide the highest fitness across environments. These findings suggest the two Aurelia lineages co-occurring in the GoM were likely exposed to different past environmental conditions (i.e., different selective pressures) and evolved different strategies to cope with environmental variation. This study highlights the importance of using genetics and morphometric data to understand jellyfish ecology, evolution and systematics.

Three routes to crypsis: Stasis, convergence, and parallelism in the Mastigias species complex (Scyphozoa, Rhizostomeae).

Evolutionary inference can be complicated by morphological crypsis, particularly in open marine systems that may rapidly dissipate signals of evolutionary processes. These complications may be alleviated by studying systems with simpler histories and clearer boundaries, such as marine lakes-small bodies of seawater entirely surrounded by land. As an example, we consider the jellyfish Mastigias spp. which occurs in two ecotypes, one in marine lakes and one in coastal oceanic habitats, throughout the Indo-West Pacific (IWP). We tested three evolutionary hypotheses to explain the current distribution of the ecotypes: (H1) the ecotypes originated from an ancient divergence; (H2) the lake ecotype was derived recently from the ocean ecotype during a single divergence event; and (H3) the lake ecotype was derived from multiple, recent, independent, divergences. We collected specimens from 21 locations throughout the IWP, reconstructed multilocus phylogenetic and intraspecific relationships, and measured variation in up to 40 morphological characters. The species tree reveals three reciprocally monophyletic regional clades, two of which contain ocean and lake ecotypes, suggesting repeated, independent evolution of coastal ancestors into marine lake ecotypes, consistent with H3; hypothesis testing and an intraspecific haplotype network analysis of samples from Palau reaffirms this result. Phylogenetic character mapping strongly correlates morphology to environment rather than lineage (r=0.7512, p<0.00001). Considering also the deeper relationships among regional clades, morphological similarity in Mastigias spp. clearly results from three separate patterns of evolution: morphological stasis in ocean medusae, convergence of lake morphology across distinct species and parallelism between lake morphologies within species. That three evolutionary routes each result in crypsis illustrates the challenges of interpreting evolutionary processes from patterns of biogeography and diversity in the seas. Identifying cryptic species is only the first step in understanding these processes; an equally important second step is exploring and understanding the processes and patterns that create crypsis.

Symmetrization in jellyfish: reorganization to regain function, and not lost parts.

We recently reported a previously unidentified strategy of self-repair in the moon jellyfish Aurelia aurita. Rather than regenerating lost parts, juvenile Aurelia reorganize remaining parts to regain essential body symmetry. This process that we called symmetrization is rapid and frequent, and is not driven by cell proliferation or cell death. Instead, the swimming machinery generates mechanical forces that drive symmetrization. We found evidence for symmetrization across three other species of jellyfish (Chrysaora pacifica, Mastigias sp., and Cotylorhiza tuberculata). We propose reorganization to regain function without recovery of initial morphology as a potentially broad class of self-repair strategy beyond radially symmetrical animals, and discuss the implications of this finding on the evolution of self-repair strategies in animals.

Length Is Associated with Pain: Jellyfish with Painful Sting Have Longer Nematocyst Tubules than Harmless Jellyfish.

A large number of humans are stung by jellyfish all over the world. The stings cause acute pain followed by persistent pain and local inflammation. Harmful jellyfish species typically cause strong pain, whereas harmless jellyfish cause subtle or no pain. Jellyfish sting humans by injecting a tubule, contained in the nematocyst, the stinging organ of jellyfish. The tubule penetrates into the skin leading to venom injection. The detailed morphology of the nematocyst tubule and molecular structure of the venom in the nematocyst has been reported; however, the mechanism responsible for the difference in pain that is caused by harmful and harmless jellyfish sting has not yet been explored or explained. Therefore, we hypothesized that differences in the length of the nematocyst tubule leads to different degrees of epithelial damage. The initial acute pain might be generated by penetration of the tubule, which stimulates pain receptor neurons, whilst persistent pain might be caused by injection of venom into the epithelium. To test this hypothesis we compared the lengths of discharged nematocyst tubules from harmful and harmless jellyfish species and evaluated their ability to penetrate human skin. The results showed that the harmful jellyfish species, Chrysaora pacifica, Carybdea brevipedalia, and Chironex yamaguchii, causing moderate to severe pain, have nematocyst tubules longer than 200 µm, compared with a jellyfish species that cause little or no pain, Aurelia aurita. The majority of the tubules of harmful jellyfishes, C. yamaguchii and C. brevipedalia, were sufficiently long to penetrate the human epidermis and physically stimulate the free nerve endings of Aδ pain receptor fibers around plexuses to cause acute pain and inject the venom into the human skin epithelium to cause persistent pain and inflammation.

Structural and Developmental Disparity in the Tentacles of the Moon Jellyfish Aurelia sp.1.

Tentacles armed with stinging cells (cnidocytes) are a defining trait of the cnidarians, a phylum that includes sea anemones, corals, jellyfish, and hydras. While cnidarian tentacles are generally characterized as structures evolved for feeding and defense, significant variation exists between the tentacles of different species, and within the same species across different life stages and/or body regions. Such diversity suggests cryptic distinctions exist in tentacle function. In this paper, we use confocal and transmission electron microscopy to contrast the structure and development of tentacles in the moon jellyfish, Aurelia species 1. We show that polyp oral tentacles and medusa marginal tentacles display markedly different cellular and muscular architecture, as well as distinct patterns of cellular proliferation during growth. Many structural differences between these tentacle types may reflect biomechanical solutions to different feeding strategies, although further work would be required for a precise mechanistic understanding. However, differences in cell proliferation dynamics suggests that the two tentacle forms lack a conserved mechanism of development, challenging the textbook-notion that cnidarian tentacles can be homologized into a conserved bauplan.

Cyanea capillata bell kinematics analysis through corrected in situ imaging and modeling using strategic discretization techniques.

Obtaining accurate kinematic data of animals is essential for many biological studies and bio-inspired engineering. Many animals, however, are either too large or too delicate to transport to controlled environments where accurate kinematic data can be easily obtained. Often, in situ recordings are the only means available but are often subject to multi-axis motion and relative magnification changes with time leading to large discrepancies in the animal kinematics. Techniques to compensate for these artifacts were applied to a large jellyfish, Cyanea capillata, freely swimming in ocean waters. The bell kinematics were captured by digitizing exumbrella profiles for two full swimming cycles. Magnification was accounted for by tracking a reference point on the ocean floor and by observing the C. capillata exumbrella arclength in order to have a constant scale through the swimming cycles. A linear fit of the top bell section was used to find the body angle with respect to the camera coordinate system. Bell margin trajectories over two swimming cycles confirmed the accuracy of the correction techniques. The corrected profiles were filtered and interpolated to provide a set of time-dependent points along the bell. Discrete models of the exumbrella were used to analyze the bell kinematics. Exumbrella discretization was conducted using three different methods. Fourier series were fitted to the discretized models and subsequently used to analyze the bell kinematics of the C. capillata. The analysis showed that the bell did not deform uniformly over time with different segments lagging behind each other. Looping of the bell trajectory between contraction and relaxation was also present through most of the exumbrella. The bell margin had the largest looping with an outer path during contraction and inner path during relaxation. The subumbrella volume was approximated based on the exumbrella kinematics and was found to increase during contraction.

Pelagia benovici sp. nov. (Cnidaria, Scyphozoa): a new jellyfish in the Mediterranean Sea.

A bloom of an unknown semaestome jellyfish species was recorded in the North Adriatic Sea from September 2013 to early 2014. Morphological analysis of several specimens showed distinct differences from other known semaestome species in the Mediterranean Sea and unquestionably identified them as belonging to a new pelagiid species within genus Pelagia. The new species is morphologically distinct from P. noctiluca, currently the only recognized valid species in the genus, and from other doubtful Pelagia species recorded from other areas of the world. Molecular analyses of mitochondrial cytochrome c oxidase subunit I (COI) and nuclear 28S ribosomal DNA genes corroborate its specific distinction from P. noctiluca and other pelagiid taxa, supporting the monophyly of Pelagiidae. Thus, we describe Pelagia benovici sp. nov. Piraino, Aglieri, Scorrano & Boero.

Organization of the ectodermal nervous structures in jellyfish: scyphomedusae.

Antibodies to α- or ß-tubulin and to the bioactive peptide FMRFamide were used to investigate the organization of the ectodermal nervous structures in five species of scyphomedusae. Within the swim system, morphological evidence, including a developmental sequence, suggests that the tubulin-immunoreactive nerve net in the subumbrella is the Giant Fiber Nerve Net (Motor Nerve Net) that directly activates the swim musculature, and the FMRFamide-immunoreactive nerve net is the Diffuse Nerve Net that serves a sensory function and also enhances swim muscle activity. Similar dual labeling was found in other structures, including those involved in feeding and protective reactions (pedalia and tentacles, radial strips of smooth muscle), and in the exumbrella, where the networks were associated with batteries of nematocysts. In addition, FMRFamide immuno-staining in the rhopalia and rhopalial niches suggests that sensory components of these networks may aid in the gravitational sense of scyphomedusae.

Stable hovering of a jellyfish-like flying machine.

Ornithopters, or flapping-wing aircraft, offer an alternative to helicopters in achieving manoeuvrability at small scales, although stabilizing such aerial vehicles remains a key challenge. Here, we present a hovering machine that achieves self-righting flight using flapping wings alone, without relying on additional aerodynamic surfaces and without feedback control. We design, construct and test-fly a prototype that opens and closes four wings, resembling the motions of swimming jellyfish more so than any insect or bird. Measurements of lift show the benefits of wing flexing and the importance of selecting a wing size appropriate to the motor. Furthermore, we use high-speed video and motion tracking to show that the body orientation is stable during ascending, forward and hovering flight modes. Our experimental measurements are used to inform an aerodynamic model of stability that reveals the importance of centre-of-mass location and the coupling of body translation and rotation. These results show the promise of flapping-flight strategies beyond those that directly mimic the wing motions of flying animals.

Internal anatomy of Haliclystus antarcticus (Cnidaria, Staurozoa) with a discussion on histological features used in Staurozoan taxonomy.

Stauromedusae have relatively few macromorphological characters, making both their taxonomy and identification difficult. For this reason, histological characters are also employed in the taxonomy of the group. This study presents a detailed description of the histomorphology of Haliclystus antarcticus Pfeffer, 1889 (Cnidaria, Staurozoa). We make new observations for the species and for the class, and address functional, taxonomical, and evolutionary aspects of staurozoan histo-anatomy. A complete reconstruction of H. antarcticus body plan is used to guide a more detailed observation, based on light microscopy, of structures rarely cited in the literature, such as the intertentacular lobules, the ostia between adjacent perradial pockets, and the male and female gonadal vesicles. Two possible regions of nematocyst formation are hypothesized and discussed. We also provide a review of the current use of histological characters in the taxonomy of the group. Understanding the body plan of stauromedusae is a challenge, because each single individual presents characters found in medusae and in polyps of other medusozoans. Comprehensive histological descriptions are important to establish relations of homology within Staurozoa and Cnidaria, providing crucial data on their evolution.

Jellyfish body plans provide allometric advantages beyond low carbon content.

Jellyfish form spectacular blooms throughout the world's oceans. Jellyfish body plans are characterised by high water and low carbon contents which enables them to grow much larger than non-gelatinous animals of equivalent carbon content and to deviate from non-gelatinous pelagic animals when incorporated into allometric relationships. Jellyfish have, however, been argued to conform to allometric relationships when carbon content is used as the metric for comparison. Here we test the hypothesis that differences in allometric relationships for several key functional parameters remain for jellyfish even after their body sizes are scaled to their carbon content. Data on carbon and nitrogen contents, rates of respiration, excretion, growth, longevity and swimming velocity of jellyfish and other pelagic animals were assembled. Allometric relationships between each variable and the equivalent spherical diameters of jellyfish and other pelagic animals were compared before and after sizes of jellyfish were standardised for their carbon content. Before standardisation, the slopes of the allometric relationships for respiration, excretion and growth were the same for jellyfish and other pelagic taxa but the intercepts differed. After standardisation, slopes and intercepts for respiration were similar but excretion rates of jellyfish were 10× slower, and growth rates 2× faster than those of other pelagic animals. Longevity of jellyfish was independent of size. The slope of the allometric relationship of swimming velocity of jellyfish differed from that of other pelagic animals but because they are larger jellyfish operate at Reynolds numbers approximately 10× greater than those of other pelagic animals of comparable carbon content. We conclude that low carbon and high water contents alone do not explain the differences in the intercepts or slopes of the allometric relationships of jellyfish and other pelagic animals and that the evolutionary longevity of jellyfish and their propensity to form blooms is facilitated by their unique body plans.

Embryos, polyps and medusae of the Early Cambrian scyphozoan Olivooides.

The Early Cambrian organism Olivooides is known from both embryonic and post-embryonic stages and, consequently, it has the potential to yield vital insights into developmental evolution at the time that animal body plans were established. However, this potential can only be realized if the phylogenetic relationships of Olivooides can be constrained. The affinities of Olivooides have proved controversial because of the lack of knowledge of the internal anatomy and the limited range of developmental stages known. Here, we describe rare embryonic specimens in which internal anatomical features are preserved. We also present a fuller sequence of fossilized developmental stages of Olivooides, including associated specimens that we interpret as budding ephyrae (juvenile medusae), all of which display a clear pentaradial symmetry. Within the framework of a cnidarian interpretation, the new data serve to pinpoint the phylogenetic position of Olivooides to the scyphozoan stem group. Hypotheses about scalidophoran or echinoderm affinities of Olivooides can be rejected.