Localization and quantification of carbonic anhydrase activity in the symbiotic Scyphozoan Cassiopea xamachana

The relationship between density and location of zooxanthellae and levels of carbonic anhydrase (CA) activity was examined in Cassiopea xamachana. In freshly collected symbiotic animals, high densities of zooxanthellae corresponded with high levels of CA activity in host bell and oral arm tissues. Bleaching resulted in a significant loss of zooxanthellae and CA activity. Recolonization resulted in full restoration of zooxanthellar densities but only partial restoration of CA activity. High levels of CA activity were also seen in structures with inherently higher zooxanthellar densities, such as oral arm tissues. Similarly, the oral epidermal layer of bell tissue had significantly higher zooxanthellar densities and levels of CA activity than did aboral bell tissues. Fluorescent labeling, using 5-dimethylaminonapthalene-1-sulfonamide (DNSA) also reflected this tight-knit relationship between the presence and density of zooxanthellae, as DNSA-CA fluorescence intensity was greatest in host oral epithelial cells directly overlying zooxanthellae. However, the presence and density of zooxanthellae did not always correspond with enzyme activity levels. A transect of bell tissue from the margin to the manubrium revealed a gradient of CA activity, with the highest values at the bell margin and the lowest at the manubrium, despite an even distribution of zooxanthellae. Thus, abiotic factors may also influence the distribution of CA and the levels of CA activity.     

Predation by the phyllosoma larva of Ibacus novemdentatus on various kinds of venomous jellyfish

The phyllosoma, a larva of spiny and slipper lobsters, has an exceptionally flat body and long appendages. It is known to associate with several species of cnidarian jellyfish, a behavior that is not rare in crustaceans. Indeed, phyllosomas clinging onto jellyfish have been observed both in the laboratory and in the natural environment. Wild phyllosomas have been found to contain jellyfish tissues in their hepatopancreas and feces, suggesting that the larvae utilize jellyfish as a food source; however, how they capture jellyfish and what species of jellyfish they prefer have rarely been investigated. The few previous studies conducted have suggested that phyllosomas have a high specificity for jellyfish (preying on only a few species); in contrast, the results of our study indicate that specificity is low. We show that phyllosomas prey on a variety of jellyfish species including deadly stinging types, on a variety of jellyfish developmental stages, and on various parts of the jellyfish body. When making contact with a jellyfish, phyllosomas first cling onto its exumbrella, feed on its tentacles or oral arms, and then consume the exumbrella. Phyllosomas may be capable of defending themselves against any types of nematocyst sting, and it is likely that they have evolved to utilize venomous jellyfish as a food in the open sea, where food may be scarce.     

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.     

Large eddy simulation of the unsteady flow-field in an idealized human mouth-throat configuration

The present study concerns the simulation and analysis of the flow field in the upper human respiratory system in order to gain an improved understanding of the complex flow field with respect to the process affecting drug delivery for medical treatment of the human air system. For this purpose, large eddy simulation (LES) is chosen because of its powerful performance in the transitional range of laminar and turbulent flow fields. The average gas velocity in a constricted tube is compared with experimental data (Ahmed and Giddens, 1983) and numerical data from Reynolds-averaged Navier-Stokes (RANS) equations coupled with low Reynolds number (LRN) κ-ω model (Zhang and Kleinstreuer, 2003) and LRN shear-stress transport κ-ω model (Jayaraju et al., 2007), for model validation. The present study emphasizes on the instantaneous flow field, where the simulations capture different scales of secondary vortices in different flow zones including recirculation zones, the laryngeal jet zone, the mixing zone, and the wall shear layer. It is observed that the laryngeal jet tail breaks up, and the unsteady motion of laryngeal jet is coupled with the unsteady distribution of secondary vortices in the jet boundary. The present results show that it is essential to study the unsteady flow field since it strongly affects the particle flow in the human upper respiratory system associated with drug delivery for medical treatment.     

Computational fluid dynamics in the oral cavity of ram suspension-feeding fishes

We have modeled steady, three-dimensional flow with a no-slip boundary condition in cylindrical and conical oral cavities possessing vertical or slanted branchial slits. These numerical simulations illustrate the transport of food particles toward the esophagus, as well as the velocity profiles of water exiting the oral cavity via the branchial slits. The maximum and average velocities are highest for flow exiting the most posterior branchial slit. The highest volume flow rates also occur in the most posterior slit for the cylindrical simulations, but occur in the most anterior slit for the conical simulations. Along the midline, there is a pronounced bilaterally symmetrical vortex in the posterodorsal region of the cylindrical and conical oral cavities and a second bilaterally symmetrical vortex in the posteroventral region of the cylinder. Particles entrained in the vortices will recirculate in the posterior oral cavity, increasing the probability of encounter with sticky, mucus-covered surfaces such as the oral roof, gill arches, or gill rakers. The posterodorsal vortex could serve to concentrate particles near the entrances of the epibranchial organs. The ventral vortex could be involved in sequestering dense inorganic particles that sink toward the floor of the oral cavity. All vortices are absent in the conical simulation with vertical branchial slits, indicating that the slanted branchial slits between the gill arches are responsible for the formation of the vortex in the conical oral cavity. Experiments using in vivo flow visualization techniques are needed to determine whether ram suspension feeders, pump suspension feeders, and non-suspension-feeding fishes possess vortices in the posterior oral cavity that contribute to particle transport, food particle encounter with sticky surfaces, and inorganic particle rejection.     

Frequency tuning in animal locomotion

In locomotion that involves repetitive motion of propulsive structures (arms, legs, fins, wings) there are resonant frequencies f(*) at which the energy consumption is a minimum. As animals need to change their speed, they can maintain this energy minimum by tuning their body resonances. We discuss the physical principles of frequency tuning, and how it relates to forces, damping, and oscillation amplitude. The resonant frequency of pendulum-type oscillators (e.g. swinging arms and legs) may be changed by varying the mass moment of inertia, or the vertical acceleration of the pendulum pivot. The frequency of elastic vibrations (e.g. the bell of a jellyfish) can be tuned with a non-linear modulus of elasticity: soft for low deflection amplitudes (low resonant frequency), and stiff for large displacements (high resonant frequency). Tuning of elastic oscillations can also be achieved by changing the effective length or cross-sectional area of the elastic members, or by allowing springs in parallel or in series to become active. We propose that swimming and flying animals generate oscillating propulsive forces from precisely placed shed vortices and that these tuned motions can only occur when vortex shedding and the simple harmonic motion of the elastic elements of the propulsive structures are in resonance.     

Stranding events provide indirect insights into the seasonality and persistence of jellyfish medusae (Cnidaria: Scyphozoa)

It is becoming increasingly evident that jellyfish (Cnidaria: Scyphozoa) play an important role within marine ecosystems, yet our knowledge of their seasonality and reproductive strategies is far from complete. Here, we explore a number of life history hypotheses for three common, yet poorly understood scyphozoan jellyfish (Rhizostoma octopus; Chrysaora hysoscella; Cyanea capillata) found throughout the Irish and Celtic Seas. Specifically, we tested whether (1) the bell diameter/wet weight of stranded medusae increased over time in a manner that suggested a single synchronised reproductive cohort; or (2) whether the range of sizes/weights remained broad throughout the stranding period suggesting the protracted release of ephyrae over many months. Stranding data were collected at five sites between 2003 and 2006 (n = 431 surveys; n = 2401 jellyfish). The relationship between bell diameter and wet weight was determined for each species (using fresh specimens collected at sea) so that estimates of wet weight could also be made for stranded individuals. For each species, the broad size and weight ranges of stranded jellyfish implied that the release of ephyrae may be protracted (albeit to different extents) in each species, with individuals of all sizes present in the water column during the summer months. For R. octopus, there was a general increase in both mean bell diameter and wet weight from January through to June which was driven by an increase in the variance and overall range of both variables during the summer. Lastly, we provide further evidence that rhizostome jellyfish may over-wintering as pelagic medusa which we hypothesise may enable them to capitalise on prey available earlier in the year. Handling editor: K. Martens     

Reynolds number limits for jet propulsion: a numerical study of simplified jellyfish

The Scallop theorem states that reciprocal methods of locomotion, such as jet propulsion or paddling, will not work in Stokes flow (Reynolds number=0). In nature the effective limit of jet propulsion is still in the range where inertial forces are significant. It appears that almost all animals that use jet propulsion swim at Reynolds numbers (Re) of about 5 or more. Juvenile squid and octopods hatch from the egg already swimming in this inertial regime. Juvenile jellyfish, or ephyrae, break off from polyps swimming at Re greater than 5. Many other organisms, such as scallops, rarely swim at Re less than 100. The limitations of jet propulsion at intermediate Re is explored here using the immersed boundary method to solve the 2D Navier-Stokes equations coupled to the motion of a simplified jellyfish. The contraction and expansion kinematics are prescribed, but the forward and backward swimming motions of the idealized jellyfish are emergent properties determined by the resulting fluid dynamics. Simulations are performed for both an oblate bell shape using a paddling mode of swimming and a prolate bell shape using jet propulsion. Average forward velocities and work put into the system are calculated for Re between 1 and 320. The results show that forward velocities rapidly decay with decreasing Re for all bell shapes when Re<10. Similarly, the work required to generate the pulsing motion increases significantly for Re<10. When compared to actual organisms, the swimming velocities and vortex separation patterns for the model prolate agree with those observed in Nemopsis bachei. The forward swimming velocities of the model oblate jellyfish after two pulse cycles are comparable to those reported for Aurelia aurita, but discrepancies are observed in the vortex dynamics between when the 2D model oblate jellyfish and the organism. This discrepancy is likely due to a combination of the differences between the 3D reality of the jellyfish and the 2D simplification, as well as the rigidity of the time varying geometry imposed by the idealized model.     

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.     

A biomimetic robotic jellyfish (Robojelly) actuated by shape memory alloy composite actuators

An analysis is conducted on the design, fabrication and performance of an underwater vehicle mimicking the propulsion mechanism and physical appearance of a medusa (jellyfish). The robotic jellyfish called Robojelly mimics the morphology and kinematics of the Aurelia aurita species. Robojelly actuates using bio-inspired shape memory alloy composite actuators. A systematic fabrication technique was developed to replicate the essential structural features of A. aurita. Robojelly's body was fabricated from RTV silicone having a total mass of 242 g and bell diameter of 164 mm. Robojelly was able to generate enough thrust in static water conditions to propel itself and achieve a proficiency of 0.19 s(-1) while the A. aurita achieves a proficiency of around 0.25 s(-1). A thrust analysis based on empirical measurements for a natural jellyfish was used to compare the performance of the different robotic configurations. The configuration with best performance was a Robojelly with segmented bell and a passive flap structure. Robojelly was found to consume an average power on the order of 17 W with the actuators not having fully reached a thermal steady state.     

Contrast and rate of light intensity decrease control directional swimming in the box jellyfish Tripedalia cystophora (Cnidaria, Cubomedusae)

Box jellyfish respond to visual stimuli by changing the dynamics and frequency of bell contractions. In this study, we determined how the contrast and the dimming time of a simple visual stimulus affected bell contraction dynamics in the box jellyfish Tripedalia cystophora. Animals were tethered in an experimental chamber where the vertical walls formed the light stimuli. Two neighbouring walls were darkened and the contraction of the bell was monitored by high-speed video. We found that (1) bell contraction frequency increased with increasing contrast and decreasing dimming time. Furthermore, (2) when increasing the contrast and decreasing the dimming time pulses with an off-centred opening had a better defined direction and (3) the number of centred pulses decreased. Only weak effects were found on the relative diameter of the contracted bell and no correlation was found for the duration of bell contraction. Our observations show that visual stimuli modulate swim speed in T. cystophora by changing the swim pulse frequency. Furthermore, the direction of swimming is better defined when the animal perceives a high-contrast, or fast dimming, stimulus.     

大型水母声学观测与评估技术研究进展

20世纪末以来,世界多个海域频繁出现大型水母暴发现象,对海洋生态系统、海洋渔业、沿海工业和滨海旅游业带来了巨大的灾难。为了研究大型水母生态习性,进而揭示其暴发机理并进行灾害的预警防治,近些年来国内外学者开展了大量的采用网具、目视、水下摄像、声学技术、航空影像等多种手段的大型水母监测调查工作,其中使用声学技术对大型水母进行资源评估和行为跟踪目前在欧美、日本、韩国等渔业发达国家已经开展了相关应用,在资源评估、运动学规律等研究中展现出较好的观测效果和应用潜力。目前我国在大型水母声学观测研究应用领域鲜有文献报道,通过介绍国际上利用声学技术对大型水母进行资源调查与评估、空间分布监测、运动规律等研究成果,为今后我国开展大型水母声学调查研究提供理论基础和科学依据。通过本文的分析,建议可以借鉴国际上采用科学鱼探仪、高分辨率成像声呐、声学信标等方法对大型水母进行监测调查和资源评估的研究成果,结合实际情况将声学技术逐步研究并应用到我国大型水母资源调查与评估、自然生态习性研究、重点水域大型水母动态监测预警中去,完善我国大型水母监测调查体系。     

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.     

A quasi three-dimensional visualization of unsteady wake flow in human undulatory swimming

Human undulatory underwater swimming (UUS) is an underwater propelling technique in competitive swimming and its propulsive mechanism is poorly understood. The purpose of this study was to visualize the three-dimensional (3D) flow field in the wake region during human UUS in a water flume. A national level male swimmer performed 41 UUS trials in a water flume. A motion capture system and stereo particle image velocimetry (PIV) equipment were used to investigate the 3D coordinates of the swimmer and 3D flow fields in the wake region. After one kick cycle was divided into eight phases, we conducted coordinate transformations and phase averaging method to construct quasi 3D flow fields. At the end of the downward kick, the lower limbs external rotations of the lower limbs were observed, and the feet approached towards each other. A strong downstream flow, i.e. a jet was observed in the wake region during the downward kick, and the paired vortex structure was accompanied by a jet. In the vortex structure, a cluster of vortices and a jet were generated in the wake during the downward kick, and the vortices were subsequently shed from the feet by the rotated leg motion. This suggested that the swimmer gained a thrust by creating vortices around the foot during the downward kick, which collided to form a jet. This paper describes, illustrates, and explains the propulsive mechanism of human UUS.     

Morphology of the nematocysts of the medusae of two scyphozoans, Catostylus mosaicus and Phyllorhiza punctata (Rhizostomeae): implications for capture of prey

Abstract. We examined the cnidomes (total complement of nematocysts) of medusae of the zooxanthellate and azooxanthellate jellyfishes Phyllorhiza punctata and Catostylus mosaicus (Rhizostomeae, Scyphozoa), and compared the assemblage of zooplankton captured on the oral arms of each species to determine whether differences in the types or amount of zooplankton captured were consistent with possible differences in the cnidomes. Cnidomes were described using light and scanning electron microscopy. Each species had a distinct cnidome and, in general, specimens of P. punctata appeared to have far fewer nematocysts than those of C. mosaicus. Four types of nematocysts were identified in medusae of C. mosaicus; 2 types of holotrichous isorhizae, rhopaloids, and birhopaloids. In C. mosaicus, the oral arms and bell margins possessed all of these types, but the cnidomes of the 2 regions differed in relative abundances and sizes of isorhizae and rhopaloids. Five types of nematocysts were identified in medusae of P. punctata, although not all types were found in all specimens. Round holotrichous isorhizae were found only in the bell, while oval holotrichous isorhizae, rhopaloids of 2 distinct size ranges, and birhopaloids were found in the bell and oral arms. Cnidomes of the bell and oral arms in specimens of P. punctata also differed in the relative abundance and sizes of oval isorhizae and rhopaloids. Although there were clear differences in the overall cnidomes and absolute abundances of nematocysts in each species, the oral arms (feeding appendages) of specimens of both C. mosaicus and P. punctata had similar types and relative abundances of nematocysts. Zooplankton sampled from the oral arms of each species showed that both species preyed predominantly on copepod nauplii and larvae of gastropods and bivalves. Medusae of C. mosaicus captured ～10 × more gastropod larvae and 5 × more bivalve larvae than those of P. punctata. Specimens of P. punctata captured approximately twice as many copepod nauplii as those of C. mosaicus. Differences in the relative abundance of types of zooplankton captured by each species could not be adequately explained by differences in the cnidomes of the oral arms.     

Influence of glottic aperture on the tracheal flow

The extra-thoracic mouth-throat area has a major influence on the aerosol delivery to the proximal or peripheral intra-thoracic airways. To characterize the particle deposition in this area, it is important to investigate first the flow structures in this crucial--in relation to the aerosol deposition--region. The glottis, which is delimited by the vocal cords and therefore has the narrowest passage, generates a laryngeal jet and a reverse flow downstream the glottis. It is generally assumed that the glottis has different shapes and cross-sectional areas at different moments during the respiratory cycle and also depends on the average inspiratory flow rate. Therefore, the influence of a circular glottal aperture, with a cross-sectional area of 90 mm2 and an elliptical and triangular shape, both with an area of 45 mm2, on the flow is investigated. However, the area of the circular aperture is twice as big as the area of the elliptical one, it has almost no influence on the flow structures. On the other hand, the triangular glottal aperture shifts the laryngeal jet in the direction of the posterior wall, and generates two pairs of counter rotating secondary vortices downstream the glottis, where the circular and elliptical only aperture generates one pair of vortices. The difference in pressure drop is more dominated by the cross-sectional area than by the shape of the glottis. This suggests the need for rendering geometry of future upper airway models even more realistic as the appropriate three-dimensional (3D) medical imaging techniques are becoming available.     

Unsteady flow field around a human hand and propulsive force in swimming

Much effort has been undertaken for the estimation of propulsive force of swimmers in the front crawl. Estimation is typically based on steady flow theory: the so-called quasi-steady analysis. Flow fields around a swimmer, however, are extremely unsteady because the change direction of hand produces unsteady vortex motions. To evaluate the force correctly, it is necessary to know the unsteady properties determined from the vortex dynamics because that unsteadiness is known to make the force greater. Unsteady flow measurements were made for this study using a sophisticated technique called particle image velocimetry (PIV) in several horizontal planes for subjects swimming in a flume. Using that method, a 100 time-sequential flow fields are obtainable simultaneously. Each flow field was calculated from two particle images using the cross-correlation method. The intensity of vortices and their locations were identified. A strong vortex was generated near the hand and then shed by directional change of the hand in the transition phase from in-sweep to out-sweep. When the vortex was shed, a new vortex rotating in the opposite direction around the hand was created. The pair of vortices induced the velocity component in the direction opposite to the swimming. Results of this study show that the momentum change attributable to the increase in this velocity component is the origin of thrust force by the hand.     

Evolution of striated muscle: jellyfish and the origin of triploblasty

The larval and polyp stages of extant Cnidaria are bi-layered with an absence of mesoderm and its differentiation products. This anatomy originally prompted the diploblast classification of the cnidarian phylum. The medusa stage, or jellyfish, however, has a more complex anatomy characterized by a swimming bell with a well-developed striated muscle layer. Based on developmental histology of the hydrozoan medusa this muscle derives from the entocodon, a mesoderm-like third cell layer established at the onset of medusa formation. According to recent molecular studies cnidarian homologs to bilaterian mesoderm and myogenic regulators are expressed in the larval and polyp stages as well as in the entocodon and derived striated muscle. Moreover striated and smooth muscle cells may have evolved directly and independently from non-muscle cells as indicated by phylogenetic analysis of myosin heavy chain genes (MHC class II). To accommodate all evidences we propose that striated muscle-based locomotion coevolved with the nervous and digestive systems in a basic metazoan Bauplan from which the ancestors of the Ctenophora (comb jellyfish), Cnidaria (jellyfish and polyps), as well as the Bilateria are derived. We argue for a motile tri-layered cnidarian ancestor and a monophyletic descent of striated muscle in Cnidaria and Bilateria. As a consequence, diploblasty evolved secondarily in cnidarian larvae and polyps.     

Synthetic peptides inducing metamorphosis in a tropical jellyfish: a quantitative structure-activity relationship study

Synthetic peptides are known to trigger metamorphosis to the polyp stage in larvae and larva-like buds of the jellyfish Cassiopea andromeda. Coupling of a hydrophobic moiety to the amino terminus of such peptides was found to increase their efficiency. Hydrophobicity can thus be considered an important factor in the biological activity of peptidic inducers. By establishing the n-octanol/water partition coefficient in order to characterize the hydrophobicity of bioactive peptides, we demonstrate that this parameter and efficiency are positively correlated in peptides of the same amino acid sequence. Hydrophobicity proved to be of much lesser importance than amino acid sequence when data of structurally disparate inducer peptides were considered.