Regression and replacement of hydranths in thecate hydroids,and the structure of hydrothecae

In 1953 the author described a regression and replacement cycle of hydranths in Laomedea flexuosa and proposed that this is a common feature of thecate hydroids. Here examples are presented of replacement of hydranths in species from 5 families of thecates, but it has not been demonstrated that this is as common as had been thought. More continuous studies are needed. A brief consideration is given to the origin of hydrothecae and their development.     

Basis of ontogenetic and evolutionary transformations of thecate hydroids

The high diversity of spatial organization of shoots in colonies of thecate hydroids (Cnidaria, Hydroidomedusa, Leptomedusae) is determined by their modular organization, which is characterized by the cyclic morphogenesis in the colony. It is attempted to show that evolutionary and ontogenetic changes in the spatial organization of hydroids of this group are based on the allometric growth of modules of colony shoots. An increase in size of a developing module provides prerequisites for earlier initiation of the growing tips of succeeding moduls (heterochrony). In some cases, heterochronies determined transition from cyclic to acyclic morphogenesis. The earlier emergence of new growing tips allowed integration of several primary modules into secondary modules, resulting among other things in changes in relative positions of primary modules (heterotopy). In complex colonies, these changes are traced in the ontogeny of a single colony.     

Changes in the patterning of a hydroid colony

It is a widely held view that colonial hydroids (Cnidaria, Hydroidomedusae) are formed on the basis of a repetition of uniform elements. The dominant opinion is that the equal spatial organisation of the colony exists during all stages of its development except the primary polyp, which develops from the settled larva. However, the complex structure and large dimensions of shoots in certain thecate species (subcl. Leptomedusae) suggest that the organisation of the primary shoot differs strongly from that of established colonies. The present study based on a thorough collection and examination of the collected material allowed to describe the entire sequence of the colony ontogeny in Hydrallmania falcata (Sertulariidae). The established shoots of this species are characterised by relatively large size, spiral arrangement of pinnate branches over the shoot stem, and hydranths arranged in one row along the upper side of branches. We showed that the primary shoot developing from the larva has much smaller dimensions and an alternate arrangement of hydranths. During further colony development the shoot organisation undergoes a gradual transformation ending with the emergence of large shoots with 'characteristic' species-specific features. The discovered sequence of changes in shoot patterning shows certain correlations with alterations of the growing tip dimensions. The dimensions of the growing tip seem to determine the patterning in accordance with the particular spatial location of the tip. This finding implies the necessity of a detailed reinvestigation of the entire colony development in thecate hydroids, which would make a significant contribution to the understanding of the morphogenetic evolution and patterning mechanisms within this group of colonial organisms.     

Basic morphogenetic processes in Hydrozoa and their evolutionary implications: an exercise in rational taxonomy

Species-specific morphology in thecate hydroids is considered as a function of 2 fundamental morphogenetic characteristics: parameters of growth pulsations and the relation between the migratory activities of the endo- and ectodermal cells of the growing tips. Comparative, experimental and modelling data are presented suggesting that increases in the values of these parameters lead to gradual transformation of the narrow tubular rudiments of primitive thecates to the more transversely extended and later bilaterally symmetrical morphologies of advanced forms. There is a corresponding change in the mode of branching, from stolonal through alternate to opposite, with densely packed hydranths and hydrothecae. The relations between the traditional systematic approach to this group and the present ontogenetically based interpretation are discussed.     

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.     

Probable advanced hydroid from the Early Ordovician of China

A fossil of a colonial organism with pyritized soft tissues of elongated fusiform zooids, found in the middle part of the early Floian Fenxiang Formation in Hubei Province of China, probably represents the oldest reliable record of a hydroid cnidarian. The preservation of the fossil is sufficiently different from that of associated carbonized skeletons of graptolites to exclude affinities with this group. The fossil is unlikely to be a bryozoan because of the mode of budding from proximal, not distal, parts of parent zooids, which is typical rather of hydroids. Although no thecae are preserved, the fossil, named Sinobryon elongatum gen. et sp. nov., is suggested to be a thecate hydroid, possibly related to the Haleciidae. The apparent presence of an advanced member of the thecaphoran Macrocolonia clade in strata 470 Ma old means that much of the hydroid (and cnidarian) diversification preceded the Middle Ordovician.     

Cell Movements Associated with Terminal Growth in Colonial Hydroids

A series of studies into the nature or leimiual growth movementsin hydroids (chiefly the thecate Campanularia) is reviewed.Both stolon and pedicel tips of all hydroids observed grow byan endless repetition of "growth cycles." In thecate stolons,the geometry of the tip's excursions is relatively simple andsuccessive cycles are predictable in their duration and percycle growth. Conversely, athecate stolons are much more complexor variable in each of these respects. In Campanularia bothcycle time and per cycle growth differ among genetic stocks. Such environmental or environment-related factors as temperature,organic contaminants in the sea water, and the nutritive conditionof the colony each affect cycle time and/or per cycle growth.Also, ingestion of a large meal temporarily inhibits the retractionsthat characterize most growth cycles. Growth movements are a property of the stolon tip, since isolatedtips less than 0.5 mm long move in basically normal fashion.However, two aspects of the cycle are, modified by more distantregions: the cycle is accelerated by a pacemaker lying 0.5 to2.0 mm from the tip, and the extent of each cycle's retractionphase is determined by intrastolonic pressure generated by hydroplasmicflow. Certain neuroinhibitor drugs duplicate in intact stolonsthe cycle changes seen in surgical isolates. Synchronized with each growth cycle is a pattern of epidermalthickening and thinning at the stolon tip, this layer beingthinnest just after the crest and thickest about two-thirdsof the way through the cycle. Cells behind the tip also movein relation to the growth cycle, generally approaching the tipduring epidermal thickening and retreating during epidermalthinning. Several types of metabolic inhibitors markedly affect the stolonicgrowth movements, indicating the need of oxidative metabolismand of protein synthesis using a short-lived mRNA for the tip'sactivities. Campanularia pedicel growth cycles differ from stolon growthcycles in having a shorter and more variable duration, a lesserper cycle growth, and a shallower retraction phase, with changesin these features along the length of the pedicel correlatedwith changes in the type of growth occurring. These studies and those of other investigators are related,their contributions to our understanding of elongation are discussed,and persisting problems are highlighted.     

Shaping of colony elements in Laomedea flexuosa Hinks (Hydrozoa, Thecaphora) includes a temporal and spatial control of skeleton hardening.

The colonies of thecate hydroids are covered with a chitinous tubelike outer skeleton, the perisarc. The perisarc shows a species-specific pattern of annuli, curvatures, and smooth parts. This pattern is exclusively formed at the growing tips at which the soft perisarc material is expelled by the underlying epithelium. Just behind the apex of the tip, this material hardens. We treated growing cultures of Laomedea flexuosa with substances we suspected would interfere with the hardening of the perisarc (L-cysteine, phenylthiourea) and those we expected would stimulate it (dopamine, N-acetyldopamine). We found that the former caused a widening of and the latter a reduction in the diameter of the perisarc tube. At the same time, the length of the structure elements changed so that the volume remained almost constant. We propose that normal development involves a spatial and temporal regulation of the hardening process. When the hardening occurs close to the apex, the diameter of the tube decreases. When it takes place farther from the apex, the innate tendency of the tip tissue to expand causes a widening of the skeleton tube. An oscillation of the position at which hardening takes place causes the formation of annuli.     

Swimming medusoid gonophores in two species of Sertularella (Cnidaria,Hydrozoa, Sertulariidae) from Réunion Island,Indian Ocean

Summary

Two life cycle patterns predominate in thecate hydroids, those with alternating polyp and medusa stages, and those with fixed gonophores. Relatively few species release short-lived medusoids, also called swimming gonophores, that carry the gametes. In the family Sertulariidae, with its large number of taxa, this mode of metagenesis has been documented previously for only two species. It is described here for two others, Sertularella diaphana (Allman, 1885) and Sertularella n. sp., from the coastal waters of Reunion Island. Freshly collected fertile male and female colonies released medusoids near dawn, one per gonotheca. Strong contractions of the bell in the swimming movements induced the shedding of gametes. Once gametes were shed, medusoids quickly degenerated and died. The medusoids were similar morphologically in these two related species, and no obvious sexual dimorphism was noted. They were ovoid in shape, the spadix was eccentric, and a velum was present; but they lacked a mouth, tentacles, sense organs, and radial and circular canals as described earlier in Amphisbetia operculata (Linneus, 1758) and Sertularia marginata Kirchenpauer, 1864. However, the medusoids described here lacked exumbrellar nematocysts. Moreover, the spadix with its dense mass of gametes was different from all other described in being stuck from base to top on one side of the subumbrella in spite of hanging freely in the subumbrellar cavity. The hypothesis that this unusual reproductive mode in sertulariids has been underestimated is confirmed by these findings.     

Structure,composition, and biogenesis of prasinophyte cell coverings

Summary The cell body and flagellar surfaces of certain green flagellates are covered by non-mineralized scales. Scale structure has been widely used in the systematics of this group of algae commonly known as the Prasinophyceae. The special importance of the flagellar hairs as a taxonomic marker is discussed. We summarize current knowledge about the structure and chemical composition of these scales with emphasis on thecate flagellates. Scales consist mainly of acidic polysaccharides involving unusual 2-keto sugar acids. Glycoproteins as minor components are mainly involved in mediating scale subunit and scale-membrane interactions and species specific glycosylation patterns exist. In thecate prasinophytes the elaboration of 3-deoxy-manno-2-octulosonic acid and galacturonic acid side chains presumably favours a complex of thecal scales with calcium ions and thus extracellular coalescence of the scales to a rigid cell wall. Scales are formed within the Golgi apparatus (GA) and especially in thecate prasinophytes scale formation (i.e., during flagellar regeneration) represents an excellent model system for GA function. Movement of developing scales through the GA requires cisternal progression. Biogenesis of scales involves mainly polysaccharide synthesis, whereas about 50% of the scale-associated glycoproteins are added from a pre-existing pool. Possible functions of prasinophyte scales are briefly discussed.Abbreviations Dha 3-deoxy-lyxo-2-heptulosaric acid - DSA Datura stramonium agglutinin - ER endoplasmic reticulum - GA Golgi apparatus - GNA Galanthus nivalis agglutinin - Kdo 3-deoxy-manno-2-octulosonic acid - MeKdo 3-deoxy-5-O-methyl-manno-2-octulosonic acid - SDS-PAGE sodium dodecyl sulphate polyacrylamide gel electrophoresis     

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 the nets.     

Mixoscyphus antarcticus gen. nov., sp. nov. (Cnidaria, Hydrozoa, Sertulariidae), the first truly endemic genus of Antarctic benthic hydroids

A genus and species of Antarctic benthic hydroids new to science, Mixoscyphus antarcticus gen. nov., sp. nov., is described and figured. Its systematic position amongst allied genera of the family Sertulariidae is discussed. The studied material originates from the South Shetland Islands area (West Antarctica), which was collected by several Spanish and US Antarctic expeditions. Mixoscyphus gen. nov. currently represents the only complete endemic genus of Antarctic benthic hydrozoans. A discussion of other genera of benthic hydroids that are largely endemic to the Antarctic is also provided.     

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.     

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     

How rich is the deep-sea Antarctic benthic hydroid fauna?

The bathymetric distribution of the 155 known species of Antarctic benthic hydroids has been studied with the aim of determining bathymetric groups and estimating the richness of the deep-sea Antarctic benthic hydroid fauna. Six bathymetric groups could be recognized on the basis of vertical zonation patterns. Most species have wide bathymetric ranges, but must nevertheless be considered shelf species. Although 62 species of benthic hydroids have been found in the Antarctic deep-sea, only 10 are strict inhabitants of that area. An inverse relative dominance between anthoathecate and leptothecate hydroids has been observed, with the former being more frequent in shallow waters.     

Patterns of hydroid (Cnidaria, Hydrozoa) species richness and distribution in an Arctic glaciated fjord

The consequences of global warming are particularly evident in high polar areas. Deglaciation phenomenon—negative mass balance of Svalbard glaciers and recession of tidal glaciers—results in landscape and shoreline change. These areas of very dynamic conditions are now open for primary colonists, among them hydroids, typical early colonists of the vacant substratum. This study aims to explore the patterns of Hydrozoan diversity and distribution in Hornsund (west Spitsbergen). Hydroids associated with shallow water kelp beds as well as those occurring on deeper subtidal soft bottom were collected at sites located along gradients of glacial disturbance (i.e., high mineral sedimentation, ice-berg scouring). Samples were collected by scuba diving (three sites of different distance to active tidal glaciers), van Veen grabs (two sites located in the inner and outer fjord basin), and dredges taken from along a fjord transect. Hydroid diversity differed significantly between sites located in the vicinity of glaciers fronts in glaciated bays and sites comparatively free from glacier disturbance. Glacial disturbance results in low frequencies of occurrence and high levels of rarity of hydroids at sites located close to glacier fronts. The species richness of hydroids colonizing the hard substrate elements present in deeper subtidal decreases along the fjord axis (i.e., along the glacial sedimentation gradient).     

Phagotrophic Mechanisms and Prey Selection in Free-living Dinoflagellates1

Three types of feeding mechanisms are known in dinoflagellates: pallium feeding, tube feeding, and direct engulfment. Pallium feeding has only been described for heterotrophic thecate species (Protoperidinium, Diplopsalis group). Tube feeding is commonly found among both naked and thecate species of mixotrophic and heterotrophic dinoflagellates (e.g. Amphidinium, Dinophysis, Gyrodinium, Peridiniopsis). Direct engulfment is mainly found among naked species (e.g. Gymnodinium, Gyrodinium, Noctiluca): recently, however, some thecate species have been shown to use this feeding mechanism as well. Feeding behavior in dinoflagellates involves several steps prior to actual ingestion, including precapture, capture, and prey manipulation. As feeding mechanisms allow the ingestion of relatively large prey or parts thereof, dinoflagellates are regarded as raptorial feeders. While prey size plays an important role in the ability of dinoflagellates to ingest food, this alone cannot explain observed prey preferences. Some dinoflagellate species can be very selective in their choice of prey, while others show a remarkable versatility.