Relay-ray way of hydroplasm movement in hydroid colonies

In this study, we continued the investigation of the distribution system of colonial hydroids in the course of its development, starting with its emergence during the planula metamorphosis and ending with the formed colony. The hydroplasmic stream system of two species of colonial hydroids—Perigonimus abyssi G.O. Sars, 1874, and Stauridia producta Wright, 1858—was studied. We found that the main principle by which hydroplasma moves in these colonies, which form no shoots, is the relay-race from one hydranth to another through a stolon fragment connecting them or it moves past the neighbouring hydranth to the next one, etc. We show that the efficiency of the conducting (distribution) system does not depend on the level of complexity of the colonial structure. The results of this study confirm the absence of general colonial processes of integration and self-regulation in hydroid polyps.     

Molecular Evidence Shows Low Species Diversity of Coral-Associated Hydroids in Acropora Corals

A novel symbiosis between scleractinians and hydroids (Zanclea spp.) was recently discovered using taxonomic approaches for hydroid species identification. In this study, we address the question whether this is a species-specific symbiosis or a cosmopolitan association between Zanclea and its coral hosts. Three molecular markers, including mitochondrial 16S and nuclear 28S ribosomal genes, and internal transcribed spacer (ITS), were utilized to examine the existence of Zanclea species from 14 Acropora species and 4 other Acroporidae genera including 142 coral samples collected from reefs in Kenting and the Penghu Islands, Taiwan, Togian Island, Indonesia, and Osprey Reef and Orpheus Island on the Great Barrier Reef, Australia. Molecular phylogenetic analyses of the 16S and 28S genes showed that Acropora-associated Zanclea was monophyletic, but the genus Zanclea was not. Analysis of the ITS, and 16S and 28S genes showed either identical or extremely low genetic diversity (with mean pairwise distances of 0.009 and 0.006 base substitutions per site for the 16S and 28S genes, respectively) among Zanclea spp. collected from diverse Acropora hosts in different geographic locations, suggesting that a cosmopolitan and probably genus-specific association occurs between Zanclea hydroids and their coral hosts.     

Reproductive costs of the symbiotic hydroid Hydractinia symbiolongicarpus (Buss and Yund) to its host hermit crab Pagurus longicarpus (Say)

Studies on the interaction between the hermit crab Pagurus longicarpus and its symbiotic hydroid Hydractinia symbiolongicarpus have focused on positive effects of hydroids on their host hermit crabs (e.g., protection from predators). Yet, these benefits may be balanced with reproductive costs, which are rarely studied. Results from field observations, laboratory trials, and a mesocosm experiment indicate that female hermit crabs in hydroid-colonized shells exhibit depressed ovigery, smaller clutch sizes, and increased clutch failure relative to females in bare shells. Frequent switching between bare and hydroid-colonized shells may alleviate negative effects when the density of hydroids in the environment is low, but at high densities Hydractinia may significantly impact hermit crab reproduction.     

Potential effect of floating dock communities on a South Carolina estuary

A 4-yr study on the motile epibionts colonizing hydroids on a floating dock in a South Carolina estuary indicated that the motile epibiotic community may impact benthic and nektonic community processes. Permanent members of the epibiotic community were preyed upon by fish, especially when the Atlantic silverside (Menidia menidia (L.)) became reproductively active and deposited eggs among the hydroids during the spring. When fish predation reduced the number of amphipods within the epibiotic community, crab megalopa colonized the hydroids. Crabs developed into juveniles and emigrated from the epibiotic community after 6–8 wk residence. The community on floating docks may be an important food source for some nekton and may recruit some benthic invertebrates.Although preferential larval settlement and fish predation are intuitive processes associated with floating dock communities, insufficient research has addressed the relationships in soft-bottom areas devoid of naturally occurring hard substrata.     

Distribution system formation in hydroid polyp development

The formation of the pulsatory distribution system, which is the basis of the colonial organization in hydroids, was studied in Gonothyrea loveni (Allm., 1859). The patterns of the distribution system functioning were established at different developmental stages.     

Feeding ecology and trophic impact of the hydroid Obelia dichotoma in the Kongsfjorden (Spitsbergen, Arctic)

Obelia dichotoma is a thecate hydroid with a worldwide distribution, occurring mainly on shallow water hard substrates. Since the trophic ecology of hydroids in polar waters is badly understood, the aim of the present work was to study qualitatively and quantitatively the diet of these organisms in an Arctic environment and to determine their trophic significance. For this purpose, the density of the hydroid population was documented, and simultaneously, zooplankton was sampled in two different years (1997 and 1998). Prey capture rates were estimated by analysing the gastrovascular content of the polyps in a diurnal cycle. Additionally, the digestion time of O. dichotoma was measured by laboratory feeding experiments using diatoms as food items. The analyses of the gastrovascular cavities of the polyps sampled during the diurnal cycles showed that O. dichotoma fed mainly on faecal pellets, organic matter and microalgae. Zooplankton prey was also observed, but gastrovascular contents and zooplankton abundance did not show any correlation in both years. The consumption rates of the hydroid populations differed between the 2 years. It was almost double (8.9 mg Carbon m^?2) in 1998 compared to 1997 (5.5 mg Carbon m^?2). The significance of the environmental variability in the feeding ecology and population dynamics of hydroids under Arctic conditions is discussed.     

Organizer regions in marine colonial hydrozoans

Organizers are specific tissue regions of developing organisms that provide accuracy and robustness to the body plan formation. Hydrozoan cnidarians (both solitary and colonial) require organizer regions for maintaining the regular body patterning during continuous tissue dynamics during asexual reproduction and growth. While the hypostomal organizer of the solitary Hydra has been studied relatively well, our knowledge of organizers in colonial hydrozoans remains fragmentary and incomplete. As colonial hydrozoans demonstrate an amazing diversity of morphological and life history traits, it is of special interest to investigate the organizers specific for particular ontogenetic stages and particular types of colonies. In the present study we aimed to assess the inductive capacities of several candidate organizer regions in hydroids with different colony organization. We carried out grafting experiments on colonial hydrozoans belonging to Leptothecata and Anthoathecata. We confirmed that the hypostome tip is an organizer in the colonial Anthoathecata as it is in the solitary polyp Hydra. We also found that the posterior tip of the larva is an organizer in hydroids regardless of the peculiarities of their metamorphosis mode and colony structure. We show for the first time that the shoot growing tip, which can be considered a key evolutionary novelty of Leptothecata, is an organizer region. Taken together, our data demonstrate that organizers function throughout the larval and polypoid stages in colonial hydroids.     

Hydroid defenses against predators: the importance of secondary metabolites versus nematocysts

Marine hydroids are commonly thought to be defended by stinging organelles called nematocysts that penetrate predator tissues and inject proteinaceous venoms, but not all hydroids possess these nematocysts. Although an increasing number of bioactive secondary metabolites have been isolated from marine hydroids, ecological roles of these compounds are poorly known. To test the hypothesis that nematocysts and noxious secondary metabolites represent alternative defenses against predation, we examined hydroids from North Carolina, United States for: (1) the palatability of whole polyps before and after nematocysts had been deactivated; (2) the palatability of their chemical extracts; and (3) their nutritional value in terms of organic content, protein content, and levels of refractory structural material (chitin). All hydroids were avoided by a generalist predator, the pinfish Lagodon rhomboides, compared with palatable control foods. Two of these (Halocordyle disticha and Tubularia crocea) became palatable after being treated with potassium chloride to discharge their nematocysts, suggesting that these species rely on nematocysts for defenses against predators. Chemical extracts from nematocyst-defended species had no effect on fish feeding. The four species that remained unpalatable after nematocysts had been discharged (Corydendrium parasiticum, Eudendrium carneum, Hydractinia symbiolongicarpus, Tridentata marginata) possessed chemical extracts that deterred feeding by pinfish. We have isolated and characterized the structures of the deterrent metabolites in two of these species. We found no differences in nutritional content or levels of chitin between nematocyst-defended and chemically defended species, and no evidence that either of these played a role in the rejection of hydroids as prey. Our results suggest that, among hydroids, chemical defenses may be at least as common as nematocyst-based defenses and that the two may represent largely alternative defensive strategies. The four hydroid species with deterrent extracts represent four families and both sub-orders of hydroids, suggesting that chemical defenses in this group may be widespread and have multiple origins. Received: 25 May 1999 / Accepted: 1 February 2000     

Potential antifouling strategies for marine finfish aquaculture: the effects of physical and chemical treatments on the settlement and survival of the hydroid Ectopleura larynx

The hydroid Ectopleura larynx is a common fouling organism on aquaculture nets. To contribute to the development of novel cleaning methods, laboratory and field studies determined the effects of heat (30, 40, 50 and 60°C for immersion times of 1 and 3 s) and acetic acid (0.2 and 2.0% for immersion times of 1, 3 and 10 s, 1 and 5 min) on the settlement of actinulae and the survival of juvenile and adult E. larynx. Laboratory studies showed that, regardless of immersion time, a temperature of 50°C was effective in preventing the settlement of actinulae and the survival of juveniles, while ≤12% of adult hydroids could survive. A temperature of 60°C killed all adult hydroids. For an acetic acid concentration of 0.2%, an immersion time of 1 min substantially reduced the settlement of actinulae and the survival of juvenile and adult hydroids, and none of the juvenile and adult hydroids survived after 5 min. For an acetic acid concentration of 2.0%, all immersion times were effective and reduced the mean settlement of actinulae and the survival of juvenile and adult hydroids to ≤10%. Field studies with fouled net panels exposed to selected heat or acetic acid treatments showed small reductions in mean wet weight and net aperture occlusion of the net panels 2 and 5 days after treatment. Visual inspections of the net panels showed that hydranths of the hydroids were shed, but the dead stolons of the hydroids remained on the treated net panels. Novel cleaning methods and devices may utilise these results to effectively kill E. larynx on aquaculture nets, while further studies are needed to determine the necessity of removing the dead hydroids before further biofouling accumulates on thenets.     

The composition and zonation of a Fucus serratus community in strangford lough,co. down

The flora and fauna associated with the serrated wrack, Fucus serratus L., in Strangford Lough have been studied. Fucus plants were larger where they were most abundant and their distribution was determined largely by the availability of suitable substratum; however, plant size was reduced under turbulent conditions. The fauna contained 79 taxa, of which eleven were common: these comprised four polyzoans, two tunicates, two sponges, two hydroids and a serpulid. All except three were more abundant on plants with low silt loads in turbulent, weedy areas. All species except Electra pilosa (L.) tended to occur on the concave surfaces of the plant. Evidence of zonation in each species' distribution along the plants was obtained. Analysis of association coefficients for the commonest species suggested that there are two distinct species groupings within the community.     

The hydroid fossil record and analytical techniques for assessing the affinities of putative hydrozoans and possible hemichordates

Hydrozoan cnidarians are widespread in modern environments, but their polyps or hydroids, when not biomineralized, are generally rare in the fossil record. To assess the affinities of four hydrozoan taxa previously described on the basis of supposed fossils of non‐biomineralized hydroids, we re‐analysed the type specimens of these taxa using a combination of light and electron microscopic tools, including backscattered electron (BSE) scanning electron microscopy (SEM) and energy‐dispersive X‐ray spectroscopy (EDS). New morphological, ultrastructural and taphonomic data were generated for Archaeoantennularia byersi from the Devonian of Michigan, Archaeocryptolaria compacta from the Ordovician of Virginia, and Mazohydra megabertha and Drevotella proteana from the Carboniferous Mazon Creek Lagerstätte of Illinois, which are preserved as carbonaceous fossils, aluminosilicate films and iron carbonate minerals in siderite concretions, respectively. In the context of these results, we provide a review of the fossil record of non‐biomineralized hydroids, describe possible biases and changes through time in their occurrence and preservation, and evaluate the criteria commonly used to identify and interpret their fossils. Although hydroids have been reported from Phanerozoic (particularly lower Palaeozoic) rocks around the world, many putative hydroids from the Palaeozoic are poorly substantiated and may actually be hemichordates. Indeed, none of the type specimens in this study represent unequivocal hydrozoans. As shown in BSE images, metatype specimens of A. byersi possess autothecae, fusellae, stolons, stolothecae and bithecae, which decisively indicate that they are dendroid graptolites rather than hydroids. The analyses yielded no evidence that A. compacta, Mazohydra and Drevotella are hydrozoans, as their holotypes lack the diagnostic morphological, taphonomic and ecological features characteristic of purported hydroid analogues. Consequently, our results suggest that many Palaeozoic hydroids may be hemichordates and that interpretations of hydroid fossils should be tested and refined using data collected via in situ analytical techniques like BSE‐SEM and EDS.     

Sediment “whips”: Amphipod artefacts from the rocky sublittoral in Britain

Sediment whips are reported attached to hydroids from circalittoral boulder grounds off St. Abbs, Scotland which are thought to be artefacts constructed by the amphipod Dyopedos porrectus Bate. Whips have been photographed in situ supporting primarily Dyopedos, but additionally caprellid amphipods and the arcturid isopod Astacilla longicomis (Sowerby). Under the scanning electron microscope the surface of the whip is seen to be criss-crossed with a meshwork of microfibrils thought to be derived from dactylar secretions of peraeopods 3 and 4 of Dyopedos. These impart significant strength and flexibility to whips underwater. The function of a whip is thought to be that of a vantage point for suspension-feeding.     

The growth patterns of some hydroids that are obligate epiphytes of seagrass leaves

The growth patterns of the hydroids Campanularia asymmetrica, Sertularia perpusilla and Algaophenia harpago on the Mediterranean seagrass Posidonia oceanica, and of Laomedea angulata on the temperate seagrass Zostera marina, were examined. The seagrass leaves grow from a basal meristem and have a short life which presents problems for these long-lived obligate epiphytes. Downward growth onto younger leaf tissue, shown by S. perpusilla and C. asymmetrica, reduces the risk of interspecific competition with other epiphytes, and places the hydroids on substrates that survive longer than older distal leaf tissue, close to short young leaves which they can colonize by stolon transfer. S. perpusilla was found only on the basal areas of leaves, possibly because of the progressive degeneration of their most apical tissue. C. asymmetrica showed no such degeneration and was found over most of the length of the leaves. The hydrorhizae of most small A. harpago and L. angulata grew both upward and downward along the leaves. Both species were most abundant on the apical halves of older leaves. This distal distribution is considered to be advantageous for these large branched hydroids which are suited for capturing planktonic prey, and to enhance their probability of colonizing neighbouring plants. Its disadvantages include the reduced probability of maintaining the hydroids on the same plant and an increased risk of being eaten by seagrass grazers.     

Hydrolysed walls in the water-conducting cells of Dendroligotrichum (bryophyta): Histochemistry and ultrastructure

Summary Histochemical techniques and electron microscopy have been used to investigate the nature of the oblique primary end-walls of the water-conducting cells (hydroids) of Dendroligotrichum dendroides. (Hedw.) Broth. The observed properties (weakly birefringent; IKI-H₂SO₄-positive; periodic acid/Schiff negative; toluidine blue O-negative) support the conclusion that these end-walls are the cellulose residue of a primary wall that has been hydrolysed during autolysis of the hydroids. The walls are now referred to as hydrolysed end-walls. The unhydrolysed lateral-walls appear to be protected from hydrolytic attack by lignin or a lignin-like compound within those walls. The similarities between the hydrolysed end-walls of the hydroids and the hydrolysed walls of vascular plant tracheary elements are discussed.Contribution No. 109 from the Department of Biology, The Pennsylvania State University.     

Automated in situ detection (AISD) of biomolecules

 In order to facilitate in situ detection of biomolecules in large sample series the processing of whole-mount specimens has been automated. A freely programmable liquid handling system is described by which embryos or similar biological materials are processed. Possible applications include in situ hybridization (ISH), immunocytochemistry (ICC) or reporter gene assays. Process times required for the preparation of whole-mount in situ hybridizations in Drosophila, Xenopus, Gallus and in hydroids were – in part – significantly reduced as compared with manual processing. Application of automated in situ detection (AISD) in random screening is demonstrated in hydroids. Potential further applications are discussed. Received: 13 May 1997 / Accepted: 27 June 1997     

Morphology versus molecules in moss phylogeny: new insights (or controversies) from placental and vascular anatomy in Oedipodium griffithianum

New data on the placenta and water-conducting cells of Oedipodium griffithianum challenge current ideas on moss phylogeny. The placental region in O.?griffithianum consists of cells with highly convolute wall labyrinths on both the sporophytic and gametophytic side. This type of placenta distinguishes an assemblage of mosses including Tetraphis, Buxbaumia and all arthrodontous mosses but is not found in basal lineages including polytrichopsid mosses. Other features that distinguish Oedipodium from polytrichopsid mosses are the foot entirely ensheathed by the parenchymatous tissue of the gametophyte vaginula, the lack of a necrotic foot tip, the lack of intercellular spaces in the foot parenchyma and the presence of typical bryopsid hydroids with uniformly thin cell walls in the leafy shoot. These results do not support molecular phylogenies resolving Oedipodium as the sister group to polytrichopsid mosses or to all peristomate mosses, but are compatible with a sister relationship to a clade encompassing Tetraphis, Buxbaumia and arthrodontous mosses.     

Allatotropin: An Ancestral Myotropic Neuropeptide Involved in Feeding

Background

Cell-cell interactions are a basic principle for the organization of tissues and organs allowing them to perform integrated functions and to organize themselves spatially and temporally. Peptidic molecules secreted by neurons and epithelial cells play fundamental roles in cell-cell interactions, acting as local neuromodulators, neurohormones, as well as endocrine and paracrine messengers. Allatotropin (AT) is a neuropeptide originally described as a regulator of Juvenile Hormone synthesis, which plays multiple neural, endocrine and myoactive roles in insects and other organisms.

Methods

A combination of immunohistochemistry using AT-antibodies and AT-Qdot nanocrystal conjugates was used to identify immunoreactive nerve cells containing the peptide and epithelial-muscular cells targeted by AT in Hydra plagiodesmica. Physiological assays using AT and AT- antibodies revealed that while AT stimulated the extrusion of the hypostome in a dose-response fashion in starved hydroids, the activity of hypostome in hydroids challenged with food was blocked by treatments with different doses of AT-antibodies.

Conclusions

AT antibodies immunolabeled nerve cells in the stalk, pedal disc, tentacles and hypostome. AT-Qdot conjugates recognized epithelial-muscular cell in the same tissues, suggesting the existence of anatomical and functional relationships between these two cell populations. Physiological assays indicated that the AT-like peptide is facilitating food ingestion.

Significance

Immunochemical, physiological and bioinformatics evidence advocates that AT is an ancestral neuropeptide involved in myoregulatory activities associated with meal ingestion and digestion.     

The distribution of hydroids (Cnidaria, Hydrozoa) from micro- to macro-scale: Spatial patterns on habitat-forming algae

Scaling up from local, short-term experiments to larger-area and longer-term ones is crucial to address the role of scale in ecology. Few studies, however, examined large-scale spatial variability in the distribution and abundance of marine organisms, with rare attempts to directly compare spatial variation at local (centimetres-metres) vs. regional (1000's of kilometres) scale. Here, we used a hierarchical design to describe the spatial distribution of the hydroids epiphitic of the brown alga Cystoseira amentacea, a habitat-forming species that provides a continuous, extensive settling substrate at regional scale along the rocky coasts in the Mediterranean Sea. This continuity provides the potential to deal with scale-related variability, increasing area of investigation without adding differences deriving from habitat heterogeneity or changes in topographic complexity. Hydroids were selected for their abundance and for their life cycle features (rapid growth, small body size, early sexual or asexual reproduction and short life span), allowing rapid responses to changes in environmental conditions. The aim of this study was to analyse whether the structure of hydroid assemblages living on C. amentacea had a consistent pattern of variation among three portions of the algal thallus (i.e., basal, middle, and distal) across a spectrum of scales and whether having or not a pelagic stage could exert a significant influence on the distribution patterns of the species. A total of 32 species were identified. Multivariate analyses showed that hydroid colonization of Cystoseira occurs differently along each thallus, with patterns of variation in the structure of assemblages differing at an even smaller spatial scale than that of single plants. However, such differences varied from patch to patch. Among the 14 species identified as “important” to define the hydroid assemblage inhabiting Cystoseira, only one (Clytia hemisphaerica) has free medusae, the other species reproducing by fixed gonophores or by short-lived medusoids. Univariate analysis showed significant differences among portions of thalli in terms of spatial variability at the various scales investigated, thus suggesting that patterns of multivariate variation along the three portions of thalli might vary across scale. Overall, our results suggest that patterns of distribution of hydroids along C. amentacea thalli significantly vary across spatial scales but that the observed differences can be hardly interpreted on the basis of life-cycle patterns.     

The growth and degeneration of the hydroid Sertularia perpusilla,an obligate epiphyte of leaves of the seagrass Posidonia oceanica

The hydrorhizae of Sertularia perpusilla colonies that originate from stolon attachments initially grow up and down Posidonia leaves. After a short time the distal hydroid tissue degenerates concurrently with the downward growth of the proximal hydrorhiza onto younger sections of the leaves. Consequently the hydroids move down the leaves which grow upward from a basal meristem. In situ inversion of leaves had no effect on either the orientation or the rates of growth and degeneration. This indicates that orientation may be in response to age related features of leaf tissue rather than to light, gravity or water movement, the directions of which were reversed by inverting the leaves. However, orientation of growth of new hydroids from attached stolons is immediate, probably before the hydroid colony is long enough to be able to detect an age gradient. This paradox could be explained if initial orientation were based on the direction of gravity or light but subsequent growth were directed along the age gradient. For hydroids in the size range studied there was a relation between growth rate and number of hydranths. Thus growth rate may be limited by food rather than by there being only one growing tip of the hydrorhizae.     

Association of a luminous Vibrio sp., taxonomically related to Vibrio harveyi, with Clytia linearis (Thornely, 1900) (Hydrozoa, Cnidaria)

A previously unknown association between a luminous Vibrio sp., taxonomically related to the species Vibrio harveyi and a common member of the shallow/mid water communities of the Mediterranean Sea, the hydrozoan Clytia linearis is described. All the specimens of C. linearis observed under blue light excitation showed both a natural luminescence appearing as a series of fine dots due to clytin, and a clear fluorescence on the external side of the perisarc around the colonies due to the presence of luminous bacteria. Luminous bacteria were isolated from the surface of C. linearis, their phenotypic characterization as isolates was performed by several morphological, biochemical, and cultural tests, completed with 16S rDNA sequence analysis. All the isolates were referred to a Vibrio sp. taxonomically related to V. harveyi. The association of the V. harveyi-related species with C. linearis, as already suggested for another hydroid, Aglaophenia octodonta, could be explained with the activity of these bacteria of feeding on the chitinous structures present in these hydroids. Moreover, the adhesion of the luminous bacterium (here referred to as Vibrio sp. CL1) on C. linearis may contribute to the survival of this Vibrio species in the marine environment providing a suitable growth habitat.